New England Wild Flower Society's Flora Novae Angliae: A Manual for the Identification of Native and Naturalized Higher Vascular Plants of New England 9780300184754

Table of contents :
Contents
Preface
Acknowledgments
Introduction
Glossary of Terminology
Key to the Families
Lycophytes
Monilophytes
Gymnosperms
Magnoliids
Monocots
Tricolpates
Literature Cited
Index

Citation preview

Flora Novae Angliae

New England Wild Flower Society’s

Flora Novae Angliae A Manual for the Identification of Native and Naturalized Higher Vascular Plants of New England

Arthur Haines Illustrated by Elizabeth Farnsworth and Gordon Morrison



 

N e w H a v e n a n d Lo n d o n

This book is dedicated to the Standing People. hic est res secretissima quam nemo novit, (radix radicis et gemma gemmae hic est et caelum caeli arboris vitae dictae; quae crescit altior quam anima sperare potest aut animus abdere potest) et hoc est miraculum quod stellas separat tuum cor gero (id gero in corde meo)

The mission of New England Wild Flower Society is to conserve and promote the region’s native plants to ensure healthy, biologically diverse landscapes. Founded in 1900, the Society is the nation’s oldest plant conservation organization and a recognized leader in native plant conservation, horticulture, and education.

Copyright © 2011 by the New England Wild Flower Society. All rights reserved. This book may not be reproduced, in whole or in part, including illustrations, in any form (beyond that copying permitted by Sections 107 and 108 of the U.S. Copyright Law and except by reviewers for the public press), without written permission from the publishers.

Yale University Press books may be purchased in quantity for educational, business, or promotional use. For information, please e-mail [email protected] (U.S. office) or [email protected] (U.K. office).

Library of Congress Control Number: 2011922019 ISBN 978-0-300-17154-9 (hardcover : alk. paper) A catalogue record for this book is available from the British Library.

This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Printed in the United States of America.

10 9 8 7 6 5 4 3 2 1

Contents

Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi Physical Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xii Taxonomy and Philosophy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii How to Use This Manual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xv Glossary of Terminology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xvii Key to the Families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Group 1 (lycophytes, monilophytes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Group 2 (gymnosperms) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Group 3 (monocots) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Group 4 (woody angiosperms with 2 or more leaves per node) . . . . . . . . . . . . . . . . . . . . . . . . 11 Group 5 (woody angiosperms with 1 leaf per node) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Group 6 (herbaceous angiosperms with inferior ovaries) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Group 7 (herbaceous angiosperms with superior ovaries and zygomorphic flowers) . . . . 23 Group 8 (herbaceous angiosperms with superior ovaries, actinomorphic flowers, and 2 or more distinct carpels) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Group 9 (herbaceous angiosperms with superior ovaries, actinomorphic flowers, connate petals, and a solitary carpel or 2 or more connate carpels) . . . . . . . . . . . . . . . . . . 27 Group 10 (herbaceous angiosperms with superior ovaries, actinomorphic flowers, distinct petals or the petals lacking, and 2 or more connate carpels) . . . . . . . . . . . . . . . . . . 30 Lycophytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Monilophytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Gymnosperms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Magnoliids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 Monocots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 Tricolpates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307 Literature Cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 891 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 917

Aroostook

Piscataquis

Somerset

Maine

Penobscot

Washington Franklin

Grand Isle

Franklin

Orleans

Essex Hancock

Coos

Lamoille

Waldo

Caledonia

Chittenden

Androscoggin

Oxford Washington

Grafton

Orange

Addison

New Carroll Hampshire

Vermont Rutland

Windsor

Kennebec

Cumberland Sagadahoc

Belknap

York

Sullivan

Hillsborough

Rockingham

ra

Merrimack

St rd

Bennington

f fo

Windham Cheshire

Essex Franklin

Middlesex

Massachusetts Hampshire

Tolland

Hartford

Windham Providence

Mid

Plymouth

Kent

Barnstable

New London

d le

Newport

sex

New Haven

Bristol Bristol

Connecticut

Fairfield

Norfolk

Norfolk

Hampden

Litchfield

Suffolk

Suffolk Norfolk

Worcester

Berkshire

Washington

Knox Lincoln

Rhode Island

Dukes

Nantucket

vii

Preface

New England Wild Flower Society is proud to be publishing the first comprehensive flora of New England in fifty years. The result of ten years of research by Arthur Haines, the volume reflects the changing face of the region, the dynamic nature of the flora, and advances in taxonomy and nomenclature. Like so many long-term projects, it began as a more modest, although important, research initiative. In 2001 the Society hired Arthur to visit herbaria throughout New England to seek specimens of rare plant species and to determine if they were correctly identified and named. Over a two-year period, he found that one of every twelve specimens required some annotation, either to correct a misidentification or to identify a new species or infraspecific taxon that had been named since the plant was collected. That discovery convinced Arthur and the Society’s Director of Conservation, Bill Brumback, of the need for a new flora. Although there are a number of floras that include New England in their coverage, the last comprehensive flora to focus exclusively on the region was written by Frank Conklin Seymour in 1969 (with later updates), which was based largely on Merritt Lyndon Fernald’s 1950 (8th) edition of Gray’s Manual of Botany. One of the most significant contributions of this new flora is the updated information about plant distribution within the New England states. Arthur’s dedicated research and his collaboration with the best botanists in the region has led to the discovery of a number of taxa not previously attributed to the region and to the elimination of a number of taxa erroneously believed to be

a part of our flora. As so often happens, a fact—in this case a mistake about a plant’s range—tends to be repeated in all subsequent publications; this manual removes attributions based primarily on anecdotes rather than specimens. While Flora Novae Angliae incorporates a half-century of changes on the landscape and in the profession, it is also a milestone from which to measure what lies ahead. Human disturbance on the land and of our climate may accelerate the dynamism on the ground. As new species move in, and others perish or migrate to more suitable areas, this manual will stand as the definitive resource by which these changes can be tracked. Of course, the conservation staff at New England Wild Flower Society will lead those trackers. The team is developing an interactive, online tool that builds on the information captured between these covers and offers a platform for disseminating updates, announcing discoveries, and teaching botany to a new generation. For their dedication and contribution to botanical knowledge, I wish to thank Arthur Haines, his collaborators, and the book’s illustrators, Elizabeth Farnsworth and Gordon Morrison. The research and publication would not have been possible without the generous support over several years of the Stratford Foundation, the Bromley Charitable Trust, and an anonymous friend. We are fortunate as well that Yale University Press embraced us and offered a well-designed path to publication. —Debbi Edelstein   Executive Director   New England Wild Flower Society

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ix

Acknowledgments

I am grateful to many people, institutions, and organizations for their assistance in collecting and organizing all of the information necessary to make this manual possible. The help came in so many forms, from outright support to encouragement or review of the manuscript. Major thanks are given to the New England Wild Flower Society for believing in the need of this work and seeking the funding needed for writing and research expenses. I am especially grateful to the Stratford Foundation, Bromley Charitable Trust, and an anonymous friend of the Society for their generous support.

Taxonomic Assistance This manual has drawn on the research and experience of many experts in the field and would not have been possible without their efforts. Their comments, herbarium specimen determinations, and discussions have been critical to forming the taxonomic scheme presented here. As with any work of this kind, the responsibility of errors and misinterpretation of data rest solely with the author. Ihsan Al-Shehbaz (Brassicaceae) George Argus (Salix) Mary Barkworth (Poaceae) David Boufford (Circaea) Dick Brummitt (Calystegia, nomenclature) Julian Campbell (Elymus) Jacques Cayouette (Carex, Eriophorum, Panicum) Yousheng Chen (Viola) William Eddie (Campanula) Robert Faden (Tradescantia) Robert Freckmann (Dichanthelium) Kanchi Gandhi (nomenclature) Llorenç Gonyalons (Linaria) Raul Gutierrez (Proboscidea) Robert Haynes (Sagittaria) Barre Hellquist (aquatic tracheophytes, Potamogetonaceae) Stewart Hinsley (Malvaceae)

Peter Hoch (Onagraceae) Mingjuan Huang (Trichostema) Zdenek Kaplan (Potamogetonaceae) Thomas Lammers (Campanula) Walter Lewis (Rosa) Joel McNeal (Cuscuta) John McNeill (Minuartia, Spergularia) James Montgomery (Dryopteris) Michael Moody (Myriophyllum) Robbin Moran (Monilophytes) John Morton (Silene) Guy Nesom (Erigeron, Oligoneuron) Jörg Ochsmann (Centaurea) James Phipps (Crataegus) Mark Porter (Gilia) James Pringle (Bartonia, Gypsophila, Syringa) Richard Rabeler (Spergularia) Anton Reznicek (Carex) Norman Robson (Hypericum) Roger Sanders (Agastache) John Semple (Solidago, Symphyotrichum) Robert Soreng (Poa) Richard Spjut (Taxus) Tyler Smith (Carex) John Strother (Hieracium, Vernonia) Carl Taylor (Isoetes) Arthur Tucker (Mentha) Gordon Tucker (Cyperus) Sam Vander Kloet (Vaccinium) David Werier (Carex) Alan Whittemore (Berberis) Dieter Wilken (Gilia)

Regional Floristic Assistance I am extremely grateful for the collaboration of the following persons, most of whom contributed many hours toward this work. Their use of the draft manual was crucial for field testing the accuracy of the identification keys. As importantly, they supplied many new records, herbarium specimens for review, and literature sources that were central to bringing together current information for New England tracheophytes.

x ackn ow le dgme nts

Ray Angelo (New England) Robert Bertin (Massachusetts) David Boufford (New England) Don Cameron (Maine) Bryan Conolly (Massachusetts, Connecticut) Melissa Cullina (Massachusetts) Rick Enser (Rhode Island) Arthur Gilman (Vermont) John Kartesz (New England) Peter Lockwood (Rhode Island) David Lovejoy (Massachusetts) Leslie Mehrhoff (Connecticut) William Moorhead (Connecticut) Bill Nichols (New Hampshire) Eleanor Saulys (Connecticut) Bruce Sorrie (Massachusetts) Dan Sperduto (New Hampshire) Francis Underwood (Rhode Island)

Collections Assistance The following curators and work staff provided herbarium loans, images of specimens, and/or made collections available for study. They are thanked profusely for their assistance. David Barrington (herbarium loans, vt) Deborah Bell (herbarium research, US) Christopher Campbell (herbarium loans, Maine) Robert Capers (herbarium loans, conn) Chad Jones (herbarium loans, CCNL) Kieth Killingbeck (herbarium research, KIRI) Walter Kittredge (herbarium research, GH, NEBC) Maryanne Laukaitis (herbarium research, mass) Roberta Lombardi (herbarium research, Mass) Bronwyn Murre (herbarium research, Mass) Karen Searcy (herbarium loans, Mass) Mark Strong (herbarium research, US) Janet Sullivan (herbarium research, NHA) Patrick Sweeney (herbarium research, YU) Barbara Thiers (herbarium loans, NY) Emily Wood (herbarium loans, GH, NEBC)

Literature Research Pulling together the vast amount of contemporary research was a critical part of the study for this work. Without the assistance of the following people, this manual would be decades out of date. Anthony Brach (Flora of China) Lisa DeCesare (library materials, Botany Libraries of the Gray Herbarium) Gretchen Wade (library materials, Botany Libraries of the Gray Herbarium) Mary Walker (journal materials)

Editorial, Administrative, and Language Assistance Though many people pointed out potential errata and contributed helpful suggestions, the following persons are recognized for their major contributions to the manual. Their willingness to invest time in this project is greatly appreciated. Barbara Grunden (editing) Tom Hayward (Latin translation) Glen Mittelhauser (editing) Alice Schori (editing) Thomas Vining (manuscript comments, Latin translation) Gray Wexelblat (indexing)

xi

Introduction

New England is composed of the six northeastern-most states—Connecticut, Maine, Massachusetts, New Hampshire, Rhode Island, and Vermont (map, p. vi). Though it is somewhat of an arbitrarily defined area, it does have some degree of natural boundaries, including the Atlantic Ocean on much of its eastern border, the St. John River on portions of its northern and northeastern border, Lake Champlain on its northwestern border, the Berkshires and included watersheds (e.g., Housatonic River) on or near its western border, and Long Island sound on its southern border. It is considered to have a well-studied flora, and several manuals have been written that treat the tracheophytes (i.e., vascular plants excluding mosses) occurring in the region. However, it is important to note that no comprehensive flora has been written with a strong focus on updating the taxonomy (i.e., names and circumscription of taxa) and distribution of New England’s tracheophytes since Fernald (1950b). Though Seymour (1982) and Magee and Ahles (1999) are more recent works that made important contributions to the knowledge of the region’s plants, the former relied heavily on the taxonomy set forth by Fernald (1950b) and the latter used plant distributions generated primarily from the herbarium surveys performed by the late Harry Ahles in the 1970s. Because of these facts, significant aspects of each manual were out-dated when printed. Major focus has been placed on updating (and in many cases verifying) the accumulated knowledge of the regional flora. However, as with any project, limitations in time and funding have meant that not every question could be explored and many issues remain to be answered. Flora Novae Angliae is primarily to serve as a checklist of the approximately 3520 native and naturalized tracheophytes (or higher vascular plants) that occur in New England and a manual for their identification. Additionally, this work also provides common names, synonymy, local distribution, ecology, and regional conservation status for the

included plants and treats approximately 320 hybrids. Further references and discussions are provided for some taxa, especially those groups or species that are difficult and/or have confusing taxonomic histories. Illustrations are found for approximately one in every three species. The illustrations usually focus on diagnostic characters, especially those that may be difficult to explain by words alone. For this first edition of Flora Novae Angliae, the precision of plant distributions is to state level. All distribution statements are, insofar as possible, vouchered by herbarium specimens (in a few cases, correspondence and journal articles have been used where experienced botanists discuss species that are unlikely to be misidentified). This level of precision was chosen due to the constraints (e.g., time, funding) to properly preparing plant records to the county level. Even for accurate reporting of plant distribution to state level, numerous problems have arisen. A major issue centers on regional floras that were not prepared by persons who understood the critical need for museum specimen vouchering—Fernald (1922a) discussed one such flora and the problems it created (and still creates today with the current species lists for Rhode Island). Another major problem is the continued listing of species that have been shown not to occur in the region (often as a result of specimen annotation or taxonomic realignment). This issue is especially prevalent with authors who do not frequent herbaria or are not intimately familiar with the pertinent literature of their region. This situation is exacerbated when authors do not perform any research to identify if the records are legitimate. As Fernald stated in volume 44 of Rhodora: “Errors once born never die but, on the contrary, by others not situated to know the facts are continually mistaken for the truth and consequently perpetuated.” Many erroneous reports are noted in this manual with the purpose of

xii IN TROD UCTIO N

ending their continued mention through the generations of botanical literature. No work is perfect and the mention of an author’s erroneous report is not intended as insult or suggestion the work is without merit. Many potentially erroneous reports have also been noted. These are distribution statements by various authors that may be properly vouchered. These reports are noted with the comment that specimens are unknown (i.e., I was unable to find a voucher specimen for a given state in the time allotted for this project). In some cases, significant effort was put forth to locate specimens and many regional herbaria were searched. In other cases, only one or two major herbaria were searched and additional effort may locate a voucher specimen. References are provided throughout this manual to enable users to continue their study beyond the information provided here. Selected references have been chosen because they highlight changes in taxonomy (i.e., they were the sources for names used in the flora), provide detailed descriptions of taxa (sometimes with illustrations), and/ or give excellent distribution information through maps or specimen citations. Therefore, some references cited in this manual are out-of-date as to names and circumscription of taxa, but they provide detailed (and valuable) information nonetheless.

Physical Region Flora Novae Angliae is a manual for the identification of the tracheophytes growing outside of cultivation in New England and is not intended to provide a detailed treatise on the geology, climate, and ecoregions of the six member states. However, some discussion of the physical region is important to setting the stage for the diversity of the flora. The following characterization is largely adapted from Seymour (1982). New England is characterized by hilly and mountainous terrain, topographic features that are part of the Appalachian Mountains. The northern tier of states (Maine, New Hampshire, and Vermont) are by far the most mountainous and possess the highest peaks—Katahdin at 1606 m (ME), Mount Washington at 1917 m (NH), and Mount Mansfield at 1339 m (VT). The geology of New England’s peaks is highly variable and ranges from primarily acidic rock (e.g.,

granite, schist) to basic rock (e.g., limestone, marble). The mountains and hills have been highly altered by glaciation and all but a tiny portion of New England (southern Marthas Vineyard) was covered during the last glacier. Alpine habitat is most abundant in Maine and New Hampshire. Lakes and ponds are frequent over most of New England, especially the northern states and the Cape Cod region of Massachusetts. Lake Champlain in Vermont is the largest in the region (approximately 1130 km2) and was, at one time, an arm of the Atlantic Ocean. Other large lakes include Moosehead Lake in Maine (311 km2), Lake Winnipesauke in New Hampshire (184 km2), and the Quabbin Resevoir in Massachusetts (100 km2). Rivers are also numerous in New England. The largest is the Connecticut River which originates north of New England and flows north to south the entire length of New England into Long Island sound. Maine has additional large rivers (Penobscot, Kennebec, Androscoggin). Several rivers in New England harbor identifiable assemblages of plants due to their underlying bedrock and/or associated climate. These include the St. John River and Aroostook River in northern Maine with their ice-scoured, Laurentian shorelines, the Housatonic and Hoosic Rivers in western Massachusetts and Connecticut that flow through limestone and marble dominated bedrock regions, and the lower reaches of the Connecticut River (CT), Merrimack River (MA), and Kennebec River (ME) with large stretches of fresh and brackish tidal habitat. The coastal plain of New England harbors some species that are typical of the midAtlantic and southeastern United States. Coastal plain communities are most prevalent in Rhode Island and eastern Massachusetts. This plain extends up the Connecticut River into Massachusetts where some species occur that are otherwise unknown from inland locations. Of note is that the region of Maine with hydrologic features most resembling coastal plain pondshores is found inland in the southwestern portion of the state. All but one state in New England (Vermont) border the Atlantic Ocean. This greatly affects the hydrology, salinity, geology, and climate of the near-coastal regions. In descending order, the total length of tidal shoreline for the five states that border the Atlantic Ocean is: Maine

i n t ro d uc t i o n  xiii

with 5565 km, Massachusetts with 2430 km, Connecticut with 989 km, Rhode Island with 614 km, and New Hampshire with 210 km. Saline, brackish, and fresh tidal habitats are found in these five states. Mean annual temperature varies considerably from southern to northern New England. Southern Connecticut has a mean temperature of approximately 10.5 C. This varies to approximately 3.8 C in northern Maine and New Hampshire. Total annual precipitation for the New England states is mostly between 101 cm and 127 cm (exceptions occur). This is comparable to much of the northeastern and centraleastern United States. Humidity is highest along the southeastern coastal areas of New England and northern New Hampshire and northwestern Maine at approximately 80%. The humidity decreases in large areas of interior New England to 70%. Again, this is comparable to much of the central and eastern United States. Given the high degree of variation in the physical region of New England, it is easy to perceive why a great diversity exists in the regional flora. Of note is that several taxa are endemic to New England, including (but not limited to) Carex oronensis, Crataegus bicknellii, Crataegus schizophylla, Digitaria filiformis var. laeviglume, Eupatorium novaeangliae, and Potentilla robbinsiana. Several near-endemics are also known, including Eleocharis diandra, Geum peckii, Pedicularis furbishiae, and Persicaria puritanorum. Additional species that occur in the region are still being described new to science. Examples from the past ten years include Eleocharis aestuum and Carex reznicekii. This is noteworthy given the extensive study this flora has received. Undoubtedly more species await discovery.

Taxonomy and Philosophy As previously stated, well-supported advances in plant systematics have been incorporated into this manual. These advancements are important for furthering our understanding of the species growing outside of cultivation on the New England landscape. They sometimes affect the definition of species and can have implications for conservation (e.g., a species considered to be rare in one or more states may prove to be an environmental form of a more common and wider ranging species).

These advancements do not always simplify the taxonomy and identification of local plants. In fact, some changes made in this flora will not be seen as favorable because they create additional challenges for amateur and professional botanists (e.g., intellectual challenges of understanding new definitions of species, memorization challenge of large families that have separated into numerous smaller families). However, the complexity created by these changes is not considered to be a sufficient cause to ignore them—advancement of our understanding of what has occurred and is occurring on the landscape is the priority. Taxonomy is not intended to make simplified lists of plants, it is intended to reflect our current understanding of the flora. As with other works that I have authored (e.g., Haines 2003a), the nomenclature used in this manual has attempted to adhere to three policies, in addition to following obvious practices (e.g., conforming to rules of nomenclature). The policies are: monophyly; non-arbitrariness; and consistency (see Haines 2003a for expanded discussion of these policies). This flora is not a final work, but instead, a contribution to the taxonomy of tracheophytes in New England. Ideas and principles printed herein may require alteration as new data are uncovered by later workers. Nomenclature is not meant to be a static system, contrary to popular desire. It needs to change to reflect our best estimate of evolutionary relationships. If nomenclature remained fixed, we would still be recognizing mountain holly (Ilex mucronatus) as belonging to the cranberry genus (Vaccinium) and sweetfern (Comptonia peregrina) as belonging to the sweet-gum genus (Liquidambar). Many authors follow a general trend of recognizing fewer or more taxonomic entities (i.e., lumping and splitting, respectively). Though many would argue that this manual follows a splitting philosophy, I would argue that the taxonomic concepts presented here are the result of the biosystematics data that have been collected. This type of information (and this information alone) should drive what we recognize. How easy or difficult it will be to remember, what the traditional circumscription of the taxa has been, or how an authority in a group prefers to present the material have nothing to do with the evolutionary history of a group of plants or the evolutionary processes that are occurring

xiv INTRODUCTION

today. Such information has been ignored in this manual. It is important to incorporate the results of research papers and monographs into the working vocabulary of regional botanists. If this is not accomplished, two taxonomies result—one used by the experts who study specific groups and one used by the people who work in the field within a state or region. This approach always leads to the failure to protect some rare taxa because those working in the field are, on average, recognizing fewer entities than those who are most familiar with a given group. The typical long delay in accepting new information (“taxonomic inertia”) only serves to hamper our ability to understand the flora and find and protect species of conservation concern. For taxa above the rank of species, only monophyletic entities are used except where research is ongoing (i.e., no clear answer has been generated from the existing data). Use of the monophyly criterion helps eliminate some of the arbitrariness of naming systems and recognizes real groups of organisms. To this end, effort has been made to find family and genus names that are demonstrably monophyletic as well as valid and effectively published and possessing a morphological definition (in addition to the genetic one). Family treatment generally follows Judd et al. (2008) except for the notable exceptions below: Lycophytes: Lycopodiaceae sensu lato (Haines 2003a). Monilophytes: Polypodiaceae sensu lato Smith et al. (2006). Monocots: Potamogetonaceae (Lindquist et al. 2006). Tricolpates: Linderniaceae (Albach et al. 2005), Phyrmaceae and related families (Beardsley and Olmstead 2002), Primulaceae sensu lato (Källersjö et al. 2000), Santalaceae and related families (Nickrent et al. 2010). Species concepts are changing and while morphological distinction is still very important, other types of data are being provided more weight than historically given. In other words, not all species recognized in this manual have the same level of morphogical separation from their congenerics. Some taxa may possess only one or two obvious differences from the most

similar species but have marked differences in phenology (e.g., Solidago aestivalis from S. rugosa), ecology (e.g., Ranunculus caricetorum from R. hispidus), modern or historical geography (e.g., Stachys pilosa from S. palustris), and/or ploidy level (e.g., Juncus ambiguus from J. bufonius). Botanists who prefer simplified treatements, such as Gleason and Cronquist (1991), where large amounts of meaningful variation in morphology and ecology were lost to broadly defined taxa, will find Flora Novae Angliae to be substantially more complicated in some genera. However, I encourage them (and other botanists) to persevere in their study. There is no group of plants in New England that is beyond anyone’s capabilities (though some groups will take longer to gain an appreciation for). Learning the tracheophyte species of New England requires field, museum, and literature study. Simply put, if you are willing to dedicate time to a group, you will be rewarded with increased understanding of that group. Some authors debate how to treat species that may have weak support for their recognition. It is important to remember that the null hypothesis is one of no difference. This means that if a taxon lacks support it should not be recognized. Lack of support can be suggested by numerous intermediates that are the result of confluent morphologies (i.e., not the result of hybridization). However, it is important to take into account the observations of species range-wide. Intermediate morphologies in a small portion of a species’ range is not necessarily justification to avoid recognition of the species. For example, Minuartia glabra appears to be a well-defined species over most of its range, particularly in the southeastern United States. However, in Maine it is difficult to identify apart from M. groenlandica at some sites because the typical morphological and ecological distinctions between them break down. These two species are still treated as separate in this manual because they are distinct over most of their range. Examination of species range-wide helps to avoid regional bias. Infraspecific taxa are (i.e., subspecies, variety) are frequently recognized for species that display recognizable and sortable variation that can be attributed to underlying genetic differences. For many of these species the variation has been named as

in t ro d uc t i o n  xv

subspecies (ssp.) or variety (var.). Therefore, a choice needs to be made with regard to what rank the variation will be recognized at. Even a brief survey of floras and monographs by interested persons will reveal tremendous inconsistency in the use of infraspecific ranks, even within the work of a single author. Even when an author has chosen to use only one of these taxonomic ranks, they are using the rank for all manners of differences— morphological, geographical, chromosomal, ecological, phenological, etc. This creates confusion and leads to naming systems that are more arbitrary. For this manual, infraspecific taxa that display morphological differences that are accompanied by complete or near complete geographical separation and/or differences in ploidy level are recognized at the subspecies level. Those infraspecific taxa that display morphological differences but are largely or wholly sympatric and possess similar ploidy level are recognized as varieties. Many non-native species in New England have their origins in Europe and/or Asia. These species often display variation that can be treated at the infraspecific level (i.e., the taxa possess valid subspecies or varieties). Unfortunately, in numerous cases, the nonnative plants have either not been studied carefully to identify the infraspecific taxa present in North America or the variation displayed in North America does not match known subspecies or varieties. For these reasons, most of the non-native species are not further categorized into infraspecific taxa. Determining the origin of a species (i.e., whether or not the taxon is native or nonnative) is extremely complicated and has not been given proper credit in many works. Many authors rely largely on the date of collection to determine the nativeness of a species. This simplistic approach fails to take into account that our floras are dynamic and constantly changing. All species, including rare taxa, are capable of moving around on the landscape and taking advantage of human-disturbed habitats. I prefer an approach that examines more information, including habitat use, substrate, dispersal biology, and level of disjunction, as well as the date of collection. Species that move into new areas in habitats that are similar to or mimic their natural habitat in nearby states are here considered native (e.g., Lycopodiella alopecuroides, a species of peatlands and coastal plain pond shores, is considered

native in ME in a hydric powerline right-ofway with peaty soil). Given that time has not permitted a thorough assessment of each taxon’s origin in New England, there are undoubtedly some inconsistencies in how species are treated in this manual. Non-native species included in this flora are those that have been collected outside of cultivation. This means that the species in question was not intentionally planted or, if planted, as spread beyond its original site. Proof of naturalization for planted species is provided by disjunction of populations (i.e., new colonies have emerged at distance from the original colonies). This ensures that seedlings or saplings of a non-native species observed at a site are not merely new shoots arising from a rhizome of a parent plant. Street trees, garden plants, produce from cultivated fields, and similar items are not included here. Naturalization is sometimes difficult to determine, especially given the paucity of information that often accompanies herbarium labels of older collections. Further, long-persistent populations, especially those that are tolerant of shade of the encroaching forest, are easily confused with truly naturalized. The problem of determining naturalization is made worse by species that were collected once or a few times and are no longer found in New England (e.g., wool waste plants). It is sometimes impossible to ascertain how long the plants existed at a location and if they produced new generations of plants. The absence of plants at a site is not evidence that the plants were not part of the flora—consider that some native species are no longer found in New England (again highlighting the dynamic nature of the flora). Because of these difficulties, I have been liberal with the species included in this flora. This manual can be seen partly as a historical record of what has been collected and became or may have become naturalized.

How to Use This Manual There are two ways to reach species entries in this manual—identification key and index. For plants where the identity is unknown, the identification key is necessary and can be accessed at multiple points depending on how familiar the user is with the plant in hand (e.g., the user must begin with the key to the families for a complete unknown, the user can begin with the key to the species

xvi IN TROD U CTIO N

for a plant known to the genus). The index can be used when the identity of the plant is known to access information such as distribution and ecology or back key in the identification key to find morphological information. Each species treated in the flora is listed alphabetically in numbered sequence and has the following information provided: accepted scientific name, origin, common name, synonyms (where applicable), distribution, ecology, and miscellaneous comments. Additional references may be found for species with miscellaneous comments. Specimen citation information may also be given for reports of species that were based on misidentified herbarium vouchers. Three items may appear at the upper right of each species entry—origin, regional conservation status, and figure number. All species will be identified as native (N) or non-native (E) in the flora (or a combination of both for some species). Some species in the flora have two or more infraspecific taxa that may have different origins. These will be discussed in the comments that follow the key to subspecies or varieties. Those species that are of regional conservation concern (i.e., rare throughout the region) will have the letter C follow the origin designation. Species are considered to be regionally rare when they meet criteria presented in Brumback and Mehrhoff (1996). Briefly, these are primarily species that are globally rare, have 20 or fewer occurrences in New England, or historically occurred in New England. For those species that are illustrated, a figure number will follow the origin or the designation of regional conservation concern notation. The number that precedes each species is presented to help determine parentage of nothospecies (i.e., hybrids). Therefore, all nothospecies included in the manual are provided with a numeric formula (e.g., 1 × 3) in addition to the scientific name (when present). The numbering of species also extends to infraspecific taxa and then appears as a letter following the number of the species. The lettering assists with tracking synonyms to a particular subspecies or variety. For example, if the third species is a genus is subdivided into three varieties, it will have a number and letter precede its varietal name in the key to the infraspecific taxa. In this case, there will be a 3a, 3b,

and 3c. These numbers and letters will appear in the synonymy of the species to show users which scientific names apply to which variety. Not all letters appear in the synonymy each time because some infraspecific taxa do not have synonyms. This manual is not a comprehensive source of common names applied to the tracheophyte species in New England. Further, common name choice has attempted to supply a uniform name to each genus (where possible) so that the common names have a measure of phylogenetic utility (i.e., the names do not suggest an incorrect relationship). For this reason, users of this manual may find common names difficult to use for indexing. The phylogenetic utility extends to both providing a unique name and a consistent root name for each genus. For example (unique name), Symphyotrichum cordifolium has long been known as heartleaved aster. However, it does not belong to the genus Aster, where the common name aster properly applies. Symphyotrichum has become known as American-aster. Therefore, this species is referred to as heart-leaved American-aster (the hyphen in the final word of the common name indicating false relationship; i.e., this species is not a true aster). For another example (consistent root name), Lobelia cardinalis is usually referred to as cardinal flower in New England. However, given that most members of the genus are referred to as lobelia, the common name red lobelia was chosen for this plant. Unfortunately these guidelines cannot be applied to all groups as some genera defy consistent use of root names (short of fabricating new names; e.g., Viburnum). Some species complexes will be very difficult to learn by the information presented in this manual alone. This is due to a number of factors, such as relative differences (e.g., darker leaves, denser arrangement of stems), difficult to articulate characters, and species that do not always display the total suite of characters that define them (this often the result of hybrid origin). Side-by-side comparisons are a must in these cases. Though this is possible in the field, it is often easiest to accomplish in the herbarium setting. Without this approach, there may be some species complexes found in this manual that will be difficult to understand.

xvii

Glossary of Terminology

a- : Latin prefix meaning not, without. abaxial : on the side away from the axis, usually refers to the underside of a leaf (compare with adaxial). acaulescent : without an upright, leafy stem. accrescent : increasing in size with age. accumbent cotyledons : cotyledons oriented such that one of their edges is set against the radicle (compare with incumbent cotyledons). achene : a dry, indehiscent, usually oneseeded, fruit. acroscopic : toward the distal end (compare with basiscopic). actinomorphic : radially symmetrical; a shape that can be divided by two or more planes into roughly equal halves, usually refers to the perianth of a flower (compare with zygomorphic). acuminate : tapering to a narrow point, more tapering than acute, less than attenuate. acute : condition of an apex with more or less straight sides that meet to form an angle of less than 90°, more pointed than obtuse, but less pointed than acuminate. adaxial : on the side toward the axis, usually refers to the top side of a leaf (compare with abaxial). adhere : to stick together, but not be actually fused, applies to unlike organs (compare with cohere, connivent). adnate : fused with a structure different from itself, as when stamens are adnate to petals (compare with connate, distinct, free). adpressed : same as appressed. adventive : not intentionally introduced and usually occurring sporadically here and there. aerenchyma : a type of plant tissue that is filled with large air spaces. aerial : above ground or water.

aggregate fruit : a class of compound fruits in which two or more ovaries from an apocarpous gynoecium are united to form a single fruit (compare with multiple fruit, simple fruit). alternate : referring to the arrangement of a node that bears only one similar organ, such as a leaf or flower (compare with opposite, whorled). alveolate : with alveoli. alveolus (plural: alveoli) : one of many pronounced, angular cavities that collectively provide a honeycomb appearance. ament : a type of inflorescence, usually with an elongate, non-fleshy axis, composed of flowers with a reduced or no perianth, subtended by bracts, and usually anemophilous. amphibious : capable of living in water and on land; often referring to plants capable of surviving when the water source has disappeared. amphistomic : with stomata on both surfaces of the leaves. amplexicaul : clasping, with two basal lobes that surround the supporting axis. ampliate : enlarged or expanded. androecium : the stamens taken collectively. androgynous : a spike with both staminate and carpellate flowers, the carpellate located at the base, below the staminate (compare with gynaecandrous). anemophilous : wind pollinated. angiosperm : a plant belong to a large group of tracheophytes characterized by possession of true flowers and fruits, includes the magnoliids, monocots, and tricolpates. annular : with the form of a ring. annulus : a ring of specialized cells encircling a sporangium that aids in sporangia dehiscence, found in leptosporangiate ferns.

xviii gloss a ry

ante : in front of, as with antepetalous stamens where the stamens are positioned directly in front of or opposite the petals, as opposed to alternating with them. anther : pollen-bearing structure usually at the end of a stalk called a filament. antherode : a sterile, modified anther. anthocarp : a compound unit found in the Nyctaginaceae consisting of the fruit, an achene or utricle, and the enclosing, indurate calyx. anthocyanic : pigmented with anthocyanins, this usually manifesting as a tinging or complete suffusion of pink, red, or purple anthocyanin : a pigment ranging in color from blue to purple to red. antrorse : upward or forward oriented (compare with retrorse). aphyllopodic : having basal sheaths without blades; with new shoots arising laterally from parent shoot (compare with phyllopodic). apical placentation : a type of placentation in which the ovule is borne at the apex of the locule. apiculate : having an apiculus. apiculus : an abrupt, short, very small, projected tip. apo- : prefix meaning the parts are distinct. apophysis (plural: apophyses) : portion of a scale of a megasporangiate strobilus, or seed cone, that is visible when the strobilus is closed. appressed : laying close to and pressed against a surface pointing in the distal direction. aquatic : living in fresh water. areole : a bounded space, such as the area surrounded by veinlets of leaves with reticulate venation. aril : a specialized appendage on a seed, often brightly colored, derived from the funiculus or seed coat. aristate : tipped with a slender bristle. armed : bearing a sharp projection such as a prickle, spine, or thorn (compare with unarmed). article : an individual segment (or mericarp) of a loment (or schizocarp), found in the Fabaceae. articulated : joined to something, often at a place of eventual separation.

ascending : diverging from an axis at an angle of 15°–45°, less upward than erect, less outward than spreading. attenuate : tapering very gradually to a prolonged tip. auricle : lobe-like appendage at the base of an organ. awn : a narrow bristle, commonly used to describe the terminal or sometimes abaxial bristle on the floral scales of some Poaceae, or the apical bristle on the scales of some Cyperaceae. axil : the position between an axis and a lateral organ from that axis, such as a leaf axil. axile placentation : a kind of placentation in which the ovules are borne on the central axis formed by the meeting of the partitions, found only in ovaries with two or more locules. axillary : borne in an axil. axis : the main stem of a structure, such as a plant or inflorescence. basal placentation : a type of placentation in which the ovule is borne at the base of the locule. basifixed : attached at the base (compare with mesifixed). basiscopic : toward the proximal end (compare with acroscopic). beak : a slender, terminal appendage on a three-dimensional organ. berry : an indehiscent, fleshy fruit with two or more seeds. bi- : prefix meaning two. bifacial : having both the ab- and adaxial surface visible, usually used to describe leaf blades (compare with unifacial). bifid : cleft into two more or less equal parts. bilabiate : condition of a gamosepalous calyx or a gamopetalous corolla with two lips, usually an upper and a lower one. bisexual : containing both pollenand ovule-producing organs (compare with unisexual). blade : the expanded, distal portion of an organ, such as a leaf or petal, as opposed to the narrow, basal portion (compare with claw). bloom : a white or white-blue powdery or waxy coating that can be rubbed away.

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bract : a modified leaf that subtends an inflorescence or flower that is not part of the flower proper, commonly of reduced size compared with the foliage leaves. bracteal : with the form and position of bracts. bracteate : having bracts. bracteolate : having bracteoles. bracteole : diminutive of bract; a bract of the second order. bractlet : same as bracteole. branchlet : an ultimate segment of a branch; a twig. bristle : a stiff hair. bud : an undeveloped shoot, inflorescence, or flower, in woody plants often covered by scales and serving as the overwintering stage. bulb : a group of modified leaves serving as a food-storage organ, borne on a short, vertical, underground stem (compare with corm). bulbiferous : bearing or having bulbs. bulblet : diminutive of bulb; a new bulb arising from a parent bulb. bulbil : a bulb-like structure borne in the leaf axils or in place of flowers. bulbous : swollen. bundle scar : one of the marks found within a leaf scar that indicate the number and position of the vascular bundles that passed from stem to leaf. bursicle : a pouch-like or flap-like portion of the stigma that envelopes part of the pollinium, found in the Orchidaceae ca. : about, approximately (Latin = circa). caducous : falling off early, as stipules that leave behind a scar. callosity : a hardened thickening. callous : with a thick, firm texture. callus (plural: calli) : a firm, thickened portion of an organ; the firm base of the lemma in the Poaceae. calyculate : possessing a set of small bracts at the base of an involucre. calyptra : a lid or cap that encloses or sits atop a structure and is eventually pushed off by the structure as it develops. calyptrate : possessing a calyptra. calyx : sepals of a given flower taken collectively.

campanulate : the shape of a corolla (or other structure) that is circular or nearly circular in outline and has a slender basal tube that expands above; bell-shaped. caniculate : channeled longitudinally. capillary : very fine, hair-like, not-flattened. capitate : abruptly expanded at the apex, thereby forming a knob-like tip. capitulescence : the arrangement of capitula on a plant. capitulum : a type of inflorescence, found mostly in the Asteraceae, composed of many flowers borne on an expanded receptacle and subtended by a common involucre. capsule : a dry, dehiscent, fruit derived from more than one carpel (therefore dehiscing by two or more valves, teeth, or pores) and lacking a replum. carpel : one type of hightly modified leaves of a flower, bearing the ovules and usually composed of an ovary, style, and stigma. carpellate : bearing carpels. carpophore : a stalk-like prolongation of the receptacle forming a central axis of the flower to which the base of the petals and filaments may be adnate. cartilaginous : nearly as firm as bone, but flexible. caruncle : an appendage at or near the hilum of some seeds. caryopsis : a specialized type of achene in which the seed coat is adnate to the pericarp, found in the Poaceae. cataphyll : a leaf sheath that does not bear a blade caudate : having a tail-like appendage; a leaf apex drawn out into a very long tapering point, more so than attenuate. caudex : firm, hardened, summit of a root mass that functions as a perennating organ. caudicle : the slender stalk of a pollinium in the Orchidaceae, also called a translator arm. caulescent : possessing a stem, usually an upright (or partially upright) and leafy one. cauline : of or pertaining to the above ground portion of the stem. cespitose : growing in a compact cluster with closely spaced stems; tufted. chaff : thin, dry scales borne on the receptacle in the Asteraceae. channeled : with a deep, rounded groove.

xx glossary

chartaceous : with the texture of paper. chasmogamous : flowers that are open and cross-pollinate (compare with cleistogamous). chlorophyll : the light-sensitive green pigment found in some plant organs. cilia : hairs found at the margin of an organ. ciliate : provided with cilia. ciliolate : diminutive of ciliate. circinate : a type of vernation with the tip curled up, such that the apex is the center of the coil. circumboreal : an encircling region of boreal habitats north of the temperate region and south of the arctic region. circumscissile : dehiscing by an encircling, transverse line so that the top comes off like a lid or cap. clathrate : with a pattern of parallel areoles; lattice-like. clavate : widened in the distal portion, like a baseball bat. claw : the narrow, basal portion of perianth parts (compare with blade). cleistogamous : flowers that remain closed and self-pollinate (compare with chasmogamous). coetaneous : the condition of the flowers and leaves expanding and developing at approximately the same time (compare with precocious, serotinous). cohere : to stick together, but not actually fused, applies to like organs (compare with adhere, connivent). coiled raceme : an indeterminate inflorescence where all the pedicels arise on the same side of the axis and the entire inflorescence is arched or coiled near the apex. coma : tuft of hairs that aids in wind dispersal of seeds. commissure : the surface where two carpels cohere, normally applied to schizocarpic fruits. comose : with a coma. compound : composed of multiple parts. compound leaf : a leaf that is divided to the midrib, with distinct, expanded portions called leaflets. concolored : with the parts or sides of similar or nearly similar color. concrescent : grown together. conduplicate : folded lengthwise into nearly equal parts.

conic : cone-shaped, used to describe a three-dimensional object. connate : describing two like parts that are fused (compare with adnate, distinct). connivent : converging and touching but not actually fused, applies to like organs (compare with adhere, cohere). convolute : arranged such that one edge is covered and the other is exposed, usually referring to petals in bud (compare with imbricate, valvate). cordate : with a rounded lobe on each side of a central sinus; heart-shaped. coriaceous : with a firm, leathery texture. coralloid : coral-like. corm : a short, vertical, enlarged, underground stem that serves as a food storage organ (compare with bulb). corpusculum : a sticky gland to which a pollinium is attached by way of a translator arm that aids in the pollinia’s dispersal, found in the Apocynaceae. corymb : an indeterminate inflorescence, somewhat similar to a raceme, that has elongate lower branches that create a more or less flat-topped inflorescence. costa (plural: costae) : a prominent midvein or midrib of a leaflet. costule : diminutive of costa; a second order costa, found on a leafule. crenate : with rounded teeth. crenulate : finely crenate. crest : an elevated, often complex, appendage or rib on an organ. crisped : a leaf margin that is undulate or crimped perpendicular to the plane of the leaf surface. crown : a form that pappus can take, in which the ovary is surrounded by a lobed or toothed rim. cucullate : arched and enfolding; hood-shaped. culm : a stem, often used for graminoid species. cuneate : tapering to the base with relatively straight, non-parallel margins; wedge-shaped. cupuliform : cup-shaped (also referred to as cupulate). cupule : a diminutive, cupulate structure. cyathiescence : the arrangement of cyathia on a plant, used in the Euphorbiaceae.

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cyathium (plural: cyathia) : a type of pseudanthial inflorescence in which carpellate and staminate flowers lacking perianth are subtended by a set of bracts that are connate into a cup-like structure and sometimes bearing petaloid appendages, the entire unit resembling a single flower, found in the Euphorbiaceae. cylindric : with the three-dimensional shape of a cylinder. cyme : a class of determinate inflorescences, in which the terminal flower of the inflorescence develops first and then each flower or pair of flowers enters anthesis in succession to the base. cymule : same as dichasium. cypsela : a special type of achene that has an adnate pappus, found in the Asteraceae and Caprifoliaceae. cystolith : a concretion of calcium carbonate, appearing as minute dots or segments on the leaf surfaces in some Urticaceae. deciduous : not persistent, falling off after the normal function. declined : curved downward. decompound : repeatedly compound into numerous segments, often in an irregular fashion. decorticated : with the bark removed. decumbent : a stem that is prostrate at the base and curves upward to have an erect or ascending, apical portion. decurrent : possessing an adnate line or wing that extends down the axis below the node, usually referring to leaves on a stem (compare with surcurrent). decurved : curved downward. decussate : arrangement of organs with two organs produced at each node, and each successive node with organs set at right angles to the orientation of those of the previous node, usually referring to branches or leaves. deflexed : abruptly bent downward. dehiscent : condition of a structure that splits open, thereby releasing contents such as seeds or pollen (compare with indehiscent). deltate : triangle-shaped. dentate : provided with outward oriented teeth (compare with crenate, serrate, undulate). denticulate : diminutive of dentate; with small, outward oriented teeth.

depauperate : poorly developed due to unfavorable conditions. determinate : growing to a fixed size, with the apex or apical portion developing first, then successively developing to the base (compare with indeterminate). diadelphous : having filaments connate in two groups, often one distinct and the others fused. dichasial cyme : a cyme that is composed of two or more dichasia or branches in a dichasial fashion. dichasium (plural: dichasia) : a cyme that bears three flowers—a central, earlier blooming flower and two lateral, later developing flowers positioned opposite one another (compare with monochasium). dichlamydeous : condition of a perianth composed of two series of parts. dichotomously branched : splitting into two essentially equivalent branches at each node. dimorphic : having two forms. dioecious : having only unisexual flowers of the same type (i.e., only carpellate or only staminate) on a given plant (compare with monoecious, polygamous, synoecious). distal : toward, or at the end away from, the point of attachment (compare with proximal). distichous : arranged along a stem in two ranks. distinct : not connate, separate from other similar organs (compare with adnate, connate). dithecal : the condition of an anther with two locules. divaricate : horizontally spreading. divergent : spreading, not parallel. divided : condition of an organ with sinuses extending all the way to the axis, the organ thus considered to be compound (compare with lobed). drupe : a fleshy fruit with a firm, often bony, endocarp that encloses the usually solitary seed. The endocarp may be composed of one unit or two or more closely associated units called pyrenes. e-, ex- : Latin prefix meaning without. echinate : with spines. eglandular : without glands.

xxii glossa ry

elaiosome : an oily or fatty body or appendage, usually on seeds, offering food to ants. ellipsoid : three-dimensional equivalent of an ellipse. elliptic : more or less shaped like an ellipse, widest in the middle and tapering to each end. emarginate : provided with a small notch at the apex. emergent : growing with roots and possibly lower stem in the water but with upper portions of the plant in the air. endocarp : the inner portion of the pericarp. ensiform : more or less linear with a sharply pointed apex; sword-shaped . entire : lacking teeth of on the margin, often applied to leaf blades and petals. entomophilous : insect pollinated. epicalyx : a set of bracts that are closely spaced and resemble the true calyx, creating the illusion of more sepals than are actually present. epigeal : upon or near the surface of the ground. epigynous : with an inferior ovary, with or without a hypanthium (compare with hypogynous, perigynous). epipetalous : attached to the petals. epistomic : with stomata on only the adaxial surface. equilateral : having each side of the basal portion of a leaf blade is similar size and shape (compare with inequilateral). equitant : with the edge, rather than the face, of conduplicate, unifacial leaves set toward the stem (as with the leaves of Iris). erect : diverging from an axis at an angle of up to 15°. erose : with a ragged edge. eusporangiate : having a sporangium two or more cells thick (i.e., opaque) that lacks an annulus and usually also lacks a stalk, found in several free-sporing families (e.g., Lycopodiaceae, Selaginellaceae, Equisetaceae, Ophioglossaceae; compare with leptosporangiate). evanescent : fading away and not reaching the margin or apex. excurrent : with the central rib or axis continuing or projecting beyond the organ. exfoliating – peeling or flaking, as in the bark of Betula papyrifera or Platanus occidentalis.

exserted : projecting beyond an enclosing structure (compare with included). extrorse : curved or directed outward (compare with introrse). falcate : having a form that is elongate, compressed, and curving; sickle-shaped false indusium : a modified tooth or reflexed margin of a fern leaf that covers the sorus (compare with indusium). farina : inflated hairs, commonly white or ochroleucous, that collectively create a mealy coating on a surface. farinose : with a white to grey or yellowwhite mealy coating, often created by inflated hairs (e.g., Chenopodium) but sometimes created by a crystalline exudate (e.g., Primula); provided with farina. fasciated : the condition of being abnormally proliferated in size and/or number, often with fusion of parts. fascicle : a bundle or compact cluster. fenestrate : with openings or windowlike slits. few : two to four (compare with several, many). fibrillose : the condition of leaf sheaths that are breaking down into fibers; appearing fibrous. fibrovascular bundle : a group of specialized conducting cells in the petioles of ferns. -fid : suffix meaning deeply divided, the number of parts given by the first portion of the word (e.g., bifid, trifid). filament : stalk supporting an anther. filiform : thread-like. flabellate : prominently dilated apically with a truncate to rounded apex; fan-shaped. flaccid : limp, lax, not retaining its form when removed from the water or supporting terrain. flexuous : condition of an elongate axis that arches or bends in alternating directions in a zig-zag fashion. floral leaves : leaves associated with flowers or found on the same branch. floricane : the second-year flowering stem of Rubus (compare with primocane). floriferous : bearing flowers. foliaceous : more or less resembling the texture and color and often shape of a leaf.

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folist : a soil dominated by organic material overlying bedrock and stones, formed as a result of reduced decomposition that is not the result of saturation by water (e.g., low mean annual temperature due to elevation). follicle : a dehiscent, unicarpellate fruit that splits along a single suture on one side of the mature carpel. fornix : a scale or appendage in the corolla tube of some Boraginaceae. free : the condition of a plant organ not attached or fused to another type of plant organ (compare with adnate, distinct). free-central placentation : arrangement of ovules attached to a column or axis extending from the base of a compound, unilocular ovary. fruit : the mature or ripened ovary and any associated structures that ripen and unite with it. funnelform : a constantly flaring tube, often used to describe a gamopetalous corolla; funnel-shaped. fusiform : narrowly tapering at both ends and thickest in the middle; spindle-shaped. galea : the concave, hood-like, upper lip of some corollas, as in the Lamiaceae and Orobanchaceae. galeate : possessing or having the form of a galea. gamo- : Greek prefix meaning the parts are connate (compare with apo-). gamophyllous : having connate leaf bases, such that the stem appears to pierce through the leaves. gemma (plural: gemmae) : a small vegetative bud that separates from the parent plant and forms a new plant. gemmiferous : bearing or producing gemmae. gibbous : conspicuously swollen on one side. girdle : an encircling mark or ridge; a zone on the immediate, distal side of an Isoetes megaspore. glabrate : nearly glabrous or becoming so. glabrous : lacking hairs. gland : a protuberance or depression on an organ or at the summit of a hair that produces a sticky or greasy substance. glandular : bearing glands (compare with eglandular). glaucous : covered with a bloom. globose : spherical; shaped like a globe.

glochid : a bristle or spine that bears one or more barbs, sometimes also referred to as a glochidium. glomerule : a compact cluster of two or more flowers. glume : scale that is empty and does not subtend flowers, found at the base of Poaceae spikelets, where it is usually paired, or at the base of Eleocharis spikes. glutinous : covered with a sticky or tacky substance. graminoid : with the form of grasses, specifically with narrow, linear leaves, often used to refer to species of the Cyperaceae, Juncaceae, and Poaceae. gymnosperms : a group of primitive plants characterized by the possession of seeds not enclosed in the integuments of an ovary (i.e., naked seeds), includes the families of conifers (e.g., Cupressaceae, Pinaceae, Taxaceae). gynecandrous : having both staminate and carpellate flowers on the same spike, the staminate located at the base, below the carpellate (compare with androgynous). gynobase : a prolonged portion of the receptacle present in many members of the Boraginaceae and Lamiaceae. gynobasic : appearing attached to the base of the carpels (as opposed to the apex), as in the style of many members of the Boraginaceae and Lamiaceae. gynoecium : the carpels of a flower taken collectively. gynophore : a stalk that elevates the gynoecium above the base of the flower. gynostegium : a short-cylindric structure formed from adnate filament, anther, and style material, found in some Apocynaceae. halophyte : a plant adapted to growing in a salty substrate. hastate : more or less triangular in outline with outward-oriented basal lobes; halberd-shaped. haustorium : a specialized, root-like connection to a host plant that a parasite uses to extract nourishment. hemi- : Greek prefix meaning half. hemiparasite : a partially parasitic plant that possesses chlorophyll (compare with holoparasite). herb : a plant that dies back to the ground at the end of each growing season.

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herbaceous : herb-like, not woody; having the texture of leaves. herbage : the above ground vegetative parts of a plant, such as stems and leaves. heterophyllous : having foliage leaves that are larger than bracteal leaves; having early (and sometimes late) season leaves that are of a different shape or division from the middle season leaves (compare with homophyllous). hilum : the scar at the point of attachment of a seed. hirsute : pubescent with coarse, somewhat stiff, usually curving hairs, coarser than villous but softer than hispid. hispid : pubescent with coarse, stiff hairs that may be uncomfortable to the touch, coarser than hirsute but softer than bristly. hispidulous : diminutive of hispid. holoparasite : a completely parasitic plant that lacks chlorophyll (compare with hemiparasite). homophyllous : having foliage leaves and bracteal leaves of similar size; having leaves throughout the season that are of a similar shape or division (compare with heterophyllous). hood : a tubular-shaped portion of the corona in some Apocynaceae. horn : a slender process borne at the base of the hoods of the corona in some Apocynaceae. hydathode : a multicellular structure within a leaf near its margin through which water is actively discharged, the structure appears in monilophytes as a termination and an expansion of the vein prior to the vein reaching the margin of the blade. hydric : wet (compare with mesic, xeric) hydrophilous : water-pollinated. hydrophyte : a plant adapted to growing in the water. hypanthium : a ring or cup-like section of tissue in a flower formed from the fusion of the basal portion of the sepals, petals, and stamens. hypogeal : within the ground or below the surface. hypogynous : with a superior ovary that lacks a hypanthium (compare with, epigynous, perigynous). hypostomic : with stomata on only the abaxial surface.

imbricate : overlapping, as shingles on a roof; arrangement of bud scales with the outer/lower ones overlying and partially concealing the inner/upper ones. included : shorter than and contained within an enclosing structure (compare with exserted). incumbent cotyledons : cotyledons oriented such that the flat surface of one is set against the radicle (compare with accumbent cotyledons). indehiscent : not separating or splitting open at maturity (compare with dehiscent). indeterminate : developing from the base and continuing to elongate or expand throughout the growing season (compare with determinate). indument : the epidermal outgrowths of a plant that are only one cell wide (i.e., hairs) taken collectively, this term does not include ramentum (of monilophytes, for example); same as indumentums. indurate : hard, firm. indusium : an epidermal outgrowth of tissue that covers a sorus, found in the ferns (compare with false indusium). inequilateral : describing the basal portion of leaf blades when one side is a size and/or shape that is different from that of the other side of the midvein (compare with equilateral). inferior ovary : an ovary attached to the receptacle below the insertion of all other flower parts (e.g., sepals, petals; compare with superior ovary). inflorescence : arrangement of flowers on a stem or branch. infructescence : arrangement of fruits on a stem or branch. internode : portion of a stem between two successive nodes. introrse : curved or directed inward (compare with extrorse). invest : to enclose or envelop. involucel : diminutive of involucre; an involucre of the second order. involucral bract : one bract of an involucre (sometimes also called a phyllary). involucre : a structure that basally surrounds another structure, used to describe a series of bracts at the base of an inflorescence. involute : rolled inward/upward, toward the adaxial surface (compare with revolute).

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isodiametric : approximately the same size in all dimensions. keel : a sharp, longitudinal ridge where two surfaces join; the two lower, united petals in the corolla of some Fabaceae. labellum : the modified, usually lowest, petal of the Orchidaceae. lacerate : appearing torn, with an irregular and jagged margin. lacuna (plural: lacunae) : a space or gap located within tissues; vein patterns that create the appearance of spaces or gaps within tissues. laminar : a description of a structure that is relatively thin with broad faces (e.g., a leaf blade), used also to describe the flattened filaments of some primitive dicot plants (e.g., Magnoliaceae) lanceolate : widest below the middle and tapering at both ends, narrower than ovate but wider than linear; lance-shaped. lateral : on the side(s), neither adaxial nor abaxial. latex : a clear or colored liquid produced in some plants, containing colloidal particles of terpenes in water. latitudinal : perpendicular to the main axis (compare with longitudinal). leaf : one of the lateral appendages of the axis of the plant, often flattened and serving the function of photosynthesis, composed of the blade, petiole, and stipule(s), any of which may be lacking in a given species. leaflet : one of the expanded, first-orderdivision portions of a compound leaf; an ultimate segment of a once-divided leaf. leafule : one of the expanded, second-orderdivision portions of a compound leaflet; an ultimate segment of a twice-divided leaf. legume : a dehiscent fruit that splits along one suture and is derived from a unicarpellate ovary, found in the Fabaceae. lemma : one of the pair of bracts that subtends each floret in the Poaceae, oriented so its adaxial surface faces the axis of the spikelet (compare with glume, palea). lenticel : a slightly raised portion of the bark of woody plants involved in gas exchange, often circular or oval and also often of different color from the remaining bark. lenticular : lens-shaped. lepidote : covered with small scales.

leptosporangiate : having a sporangium that is one cell layer thick (i.e., translucent) that has an annulus and a stalk, found in many fern families (e.g., Aspleniaceae, Blechnaceae, Dryopteridaceae, Marsileaceae, Woodsiaceae; compare with eusporangiate) liana : a woody plant with vine-like habit (compare with vine). ligule : an appendage at the junction of the sheath and blade of the adaxial surface of a leaf in many Poaceae and Cyperaceae; the flattened portion of the ray flower in the Asteraceae. limb : the expanded portion of a gamopetalous corolla, distal to the throat. linear : very narrow, with more or less parallel sides, narrower than lanceolate. lip : a projecting portion of a gamosepalous calyx or gamopetalous corolla. lobe : a projecting portion of an organ too large to be referred to as a tooth. lobed : possessing lobes, with the sinuses between the lobes not extending all the way to the axis (compare with divided). locule : a seed-containing chamber in an ovary; a chamber in any structure that contains one or more items. loculicidal : a type of dehiscence in which a capsule splits along the midrib or outer median line of each carpel such that the capsule dehisces through each carpel (compare with septicidal). lodicule : a tiny scale found in the flowers of the Poaceae that is the vestigial perianth. loment : a schizocarpic fruit of some Fabaceae that is composed of one-seeded segments that may disarticulate from one another. long shoot : a typical branch or shoot with remote leaves (compare with short shoot). longitudinal : parallel to the main axis (compare with latitudinal). lycophyte : a group of primitive tracheophytes marked by free-sporing habit, microphylls, and biflagellate sperm, marked also by eusporangiate condition; clubmosses, firmosses, spikemosses, and quillworts (compare with monilophyte). lunate : crescent-shaped. lustrous : shiny, possessing a luster. maculate : spotted or splotched.

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magnoliid : a group of primitive, dicotyledenous angiosperms with nontricolpate pollen, taken here to collectively refer to all such taxa, including the magnoliids proper (e.g., Magnoliaceae, Lauraceae, Aristolochiaceae) and members of earlier diverging lineages (e.g., Nymphaeaceae) malpighiaceous : condition of hairs when they are attached near their center and not at their base; mesifixed. mamilla (plural: mamillae) : a nippleshaped protrusion. many : 11 or more (compare with few, several). marcescent : withering but persistent and not falling. marginal : pertaining to or borne on or near the margin. marginal placentation : a type of placentation in which the ovules are borne at the connate margins of a simple carpel. megaspore : a spore that gives rise to a female gametophyte, usually much larger than a microspore. megasporophyll : a leaf, often modified (reduced and/or without chlorophyll), that bears megaspores in its axil. membranaceous : thin, flexible, almost translucent; membrane-like. mericarp : each half- or single-carpellate segment of a schizocarpic fruit. -merous : Greek suffix meaning members of a set or cycle, usually used to indicate the numbers comprising each set of the perianth. mesic : moist, neither dry nor wet (compare with hydric, xeric). mesifixed : attached in the middle (compare with basifixed). mesocarp : the middle layer of the pericarp of a fruit when three layers can be distinguished. microspore : a spore that gives rise to a male gametophyte, usually much smaller than a megaspore. mineral soil : soil composed mostly of degraded parent rock material such as clay, silt, and sand. mixed inflorescence : an inflorescence with an indeterminate, central axis and determinate (i.e., cymose) branches. monadelphous : having all of the filaments of stamens connate into a single group.

moniliform : arrangement of a set of spherical or ovoid bodies that are separate and non-overlapping on an elongate axis; bead-like. monilophyte : a group of primitive tracheophytes marked by free-sporing habit, megaphylls, and multiflagellate sperm, in New England usually also marked by leptosporangiate condition; ferns (compare with monilophyte) monocarpic : flowering and fruiting only once and then dying. monochasium (plural: monochasia) : a two-flowered cymule, with a terminal flower and a single, lateral flower (compare with dichasium). monochlamydeous : perianth comprised of one series of parts. monocots : a group of early arising angiosperms characterized by one seed leaf, perianth in cyles of three, parallelveined leaves, and monosulcate pollen (with modifications of these character states in several genera). monoecious : having unisexual flowers, with both the carpellate and staminate flowers borne on the same plant (compare with dioecious, polygamous, synoecious). monomorphic : having one form. monopodial : with a simple axis from which branches or appendages originate (compare with sympodial). monothecal : the condition of an anther with a single locule. mucro : a short, abrupt, projecting tip of an organ. mucronate : having a mucro. mucronulate : diminutive of mucronate; having a tiny mucro. multiple fruit : a fruit composed of two or more ovaries from separate flowers (compare with, aggregate fruit, simple fruit). muricate : with small, pointed projections. nectary : a structure that produces nectar. nectariferous : bearing nectar. nerve : a conspicuous vein of an organ. node : point of attachment of a leaves, peduncles, pedicels, and branches to a stem or articulation point. nut : a type of fruit with a single locule and single seed surrounded by a thick, hard pericarp, often subtended by an involucre. nutlet : diminutive of nut.

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ob- : Latin prefix indicating a reverse in direction or orientation (e.g., obovate : egg-shaped in cross-section with the wider end near the apex). oblique : emerging or joining at an angle other than parallel or perpendicular. obloid : three-dimensional equivalent of oblong. oblong : shaped similar to a rectangle that is longer than wide and with rounded ends. obsolete : reduced to the point of being undetectable or virtually absent, implies presence in ancestral forms (compare with rudimentary, vestigial). obtuse : bluntly pointed, with the margins forming an angle of greater than 90°, more pointed than rounded, but less pointed than acute. ocrea (plural: ocreae) : the tubular, sheathing stipule located apically to the petiole, found in the Polygonaceae. ocreola (plural: ocreolae) : the tubular bracteole subtending the flowers in the Polygonaceae. ochroleucous : a pale yellow-white color. oligotrophic : condition of lakes that lack considerable amounts of aquatic vegetation, usually deep, clear, and with sandy or rocky bottoms. opposite : positioned directly across from one another; referring to the arrangement of a node that bears two similar organs (compare with alternate, whorled). orbicular : circular in outline. organic soil : soil composed mostly of partially degraded plant material, such as peat and muck. orifice : an opening into a cavity. ornamented : with elaborate processes or patterns. oval : similar to a broadly elliptic shape with rounded ends. ovary : the expanded basal portion of a carpel that bears the ovules. ovate : two-dimensional equivalent of egg-shaped, widest below the middle and broadly tapering to each end, wider than lanceolate. ovoid : three-dimensional equivalent of ovate; egg-shaped. ovule : an immature seed. palate : a raised area on the lower lip of a bilabiate corolla that can obscure or close off the opening.

palea : the innermost of a pair of bracts subtending a single grass floret, oriented so its adaxial surface faces away from the axis of the spikelet (compare with glume, lemma). palmate : with structures radiating from a common point, like fingers or parts of a fan. pandurate : with a constriction near the middle of the organ. panicle : a branched, indeterminate inflorescence. papilionaceous : having a corolla with the architecture of most members of the Fabaceae; i.e., with an upright banner petal that sits outside all the other petals, two lateral wing petals, and two lower petals fused together to form a keel-like structure. papilla (plural: papillae) : a short, rounded or cylindrical projections. papillate : covered with papillae. papillose : same as papillate. pappus : the modified calyx in the Asteraceae crowning the ovary and appearing as bristles, scales, awns, or a rim of tissue, or sometimes a mixture of these. parasite : a plant that acquires its food and water through specialized attachments to a host plant. parietal placentation : a type of placentation in which the partitions extend only part way toward the center of the ovary and the ovules are borne on the partitions, found in ovaries composed of two or more carpels. pectinate : the condition of an unbranched axis with a single row of narrow appendages emitting from one or both sides. pedicel : stalk of a single flower or fruit in an inflorescence or infructescence that has two or more flowers or fruits, respectively (compare with peduncle). pedicellate : borne on a pedicel. peduncle : a stalk that supports an inflorescence composed of one or more flowers (compare with pedicel). pedunculate : borne on a peduncle. pellucid : transparent or translucent. peltate : attachment of an organ at its center rather than at its base; umbrella-like.

xxviii gloss ary

pendulous : limply hanging from near the base. penicillate : provided with a tuft of hairs at the apex. pentadelphous : having filaments connate into five groups. penultimate : next to last. pepo : a specialized type of berry with an inseparable, often leathery, rind derived from an inferior ovary, found in the Cucurbitaceae. perennate : to survive the winter season, often in a form different from that of the summer phase of the plant, such as winter buds, seeds, or corms. perennial : a plant that lives for more than two years. perfoliate leaf : a leaf that has the basal margins connate, creating the illusion that the stem pierces the middle of a leaf. perianth : the calyx, corolla, and corona taken collectively, one or more of which may be absent. pericarp : the mature ovary wall of a fruit. perigynium (plural: perigynia) : a specialized bract that subtends the carpellate flowers in Carex and whose margins are united thereby forming a saclike structure with an orifice at the apex through which the stigmas protrude. perigynous : with a superior ovary and a hypanthium (compare with epigynous, hypogynous). persistent : remaining attached through the year or after the normal function. perulate : covered with one or more scales, as in the winter buds of most woody plants. petal : one of the hightly modified leaves of a flower, usually pigmented and thereby attracting pollinators. petaliferous : producing or possessing petals. petaloid : resembling a petal (usually in shape and color). petiole : non-expanded, basal portion of a leaf; stalk of a leaf. petiolate : having a petiole. petiolulate : having a petiolule. petiolule : non-expanded, basal portion of a leaflet; stalk of a leaflet. photosynthetic : capable of photosynthesizing due to the presence of chlorophyll.

-phyll : Greek suffix meaning leaf (e.g., sporophyll, trophophyll). phyllary : one bract of the involucre that subtends a capitulum; an involucral bract associated with a capitulum. phyllopodic : having the basal sheaths blade-bearing; with new shoots arising from the center of parent shoot (compare with aphyllopodic). phyllotaxis : arrangement of nodes on the branchlet in alternate-leaved species. Phyllotaxis is given as two numbers separated by a slash, e.g., ⅖. The first number (2 in the example) refers to the number of revolutions around the twig that must be made to reach a position on exactly the same side of the twig that one started from. The second number refers to the number of nodes (5 in the example) that must be counted in a given direction to reach the same position that one started from. Example: Given a plant with a phyllotaxis of ⅖, an observer, starting at a node or winter bud, would have to make two revolutions around the twig and, in the process, count five nodes or winter buds to reach a node or winter bud oriented in the same direction as the one started from. phytomelan : a black crust found on seeds. pilose : with sparse, straight, spreading hairs. pin : one type of heterostylic flower in which the style is longer than the stamens (compare with thrum). pinnate : arranged along both sides of an elongate axis, such as the barbs on a feather. pinnatifid : pinnately lobed. pistil : the outward appearance of the carpel(s) of a flower. pith : the internal cylinder of parenchymous tissue in a stem or root. placentation : arrangement of ovules in an ovary, specifically refers to how they are attached. planoconvex : cross-sectional shape, for objects with two surfaces, that is flat on one surface and convexly rounded on the other. plumose : like a tuft of hairs or feathers; specifically applied to hairs or bristles that have lateral branches arising in a pinnate fashion. pollen : the microgametophyte of seed plants, contained in anthers.

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pollinarium (plural: pollinaria) : a pollinium with its associated structures (e.g., caudicle, stipe, viscidium) pollinium : a mass of pollen grains cohering as a single unit, as in the Orchidaceae and some Apocynaceae. polygamous : with both unisexual and bisexual flowers on the same plant (compare with dioecious, monoecious, synoecious). pome : a special type of berry found in some Rosaceae; specifically, a fleshy fruit derived from a compound, inferior ovary, with a papery, cartilaginous, or bony endocarp and usually two or more seeds. praemorse : abruptly ending in a truncated fashion. precocious : the condition of the flowers’ expanding and developing before the leaves (compare with coaetaneous, serotinous). prickle : a small, more or less sharp, epidermal outgrowth (compare with spine, thorn). primocane : the first-year, vegetative stem in Rubus (compare with floricane). process : a slender, protruding feature. procumbent : prostrate or trailing most of the length. propagule : a body with the function of reproduction, such as a seed, turion, or bulb. prophyll : a second- or higher-order bract subtending some portion of a branching inflorescence or structure; one of a pair of bracteoles that subtends the perianth in Juncus. proximal : toward or at the point of attachment (compare with distal). pseudanthium (plural: pseudanthia) : a cluster of small flowers that appears to be a single flower (e.g., a capitulum, a cyathium, an inflorescence of Benthamia). pseudoterminal bud : the distalmost bud on an indeterminate branchlet, identifiable by the presence of a twig scar adjacent to the bud (compare with terminal bud). pseudowhorl : falsely appearing to be whorled. ptyxis : the manner in which an individual leaf or petal is oriented in bud (e.g., conduplicate, involute, wrinkled), here considered part of definition of vernation (see also vernation).

puberulent : diminutive of pubescent; with minute hairs. pubescence : the collection of epidermal outgrowths only one cell wide, not as stiff to be prickles, nor flattened as scales; hairs. pubescent : with pubescence (i.e., with hairs). pulvinate : densely matted together in clumped colonies; cushion-like. punctate : dotted with small glands or pits. pustule : a blister. pustulose : having or with the form of pustules. pyrene : one of the units comprising the endocarp of a drupe, used when the endocarp is composed of two or more units. pyriform : pear-shaped. pyxis : a dehiscent fruit that is circumscissile around the middle, the top falling off and exposing the seed(s). raceme : an indeterminate inflorescence with a central axis, often elongate, and with pedicellate flowers. rachilla : diminutive of rachis; the axis of a sedge flower or the axis of the flowers in a grass spikelet. rachis : the central or main axis of a structure. rame : an elongate inflorescence that bears both sessile and pedicellate flowers, used in the Poaceae. ramentum : the collection of scales of a given plant taken collectively, usually used in the discussion of monilophytes (compare with indument). rank : a longitudinal position on a stem. ray : the flat, usually elongate, portion of a ray flower corolla. ray flower : the zygomorphic, ray-bearing flower in the Asteraceae. receptacle : an expanded portion at the apex of the pedicel or peduncle to which the parts of a flower are attached. reflexed : abruptly bent backward or downward. remote : separated to some extent from others of the same kind (e.g., remote flowers), antonym of approximate. reniform : wider than long, with a central, basal sinus; kidney-shaped. replum : a frame-like placenta positioned between the valves of a silique or a silicle.

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resupinate : possessing a 180-degree twist in the pedicel or ovary so that the flower is upside down. reticulate : with a visible network due to a pattern of splitting and rejoining. retrorse : downward or backward oriented (compare with antrorse). retuse : with a terminal notch in an otherwise blunt or rounded apex. revolute : rolled under, toward the abaxial surface (compare with involute). rhizomatous : bearing rhizomes. rhizome : a horizontal, underground stem. rhizophore : a modified leafless shoot that produce shoots. rhombic : diamond-shaped. rostellum : a cap-like or beak-like, sterile projection of the stigma. rotate : the shape of a corolla that is circular or nearly circular in outline and lacks a basal tube and has fully or nearly horizontally spreading lobes; platter-shaped. rotund : circular in outline. rudimentary : poorly developed and reflecting an earlier evolutionary stage, not a reduction of the feature (compare with obsolete, vestigial). rugose : wrinkled. rugulose : diminutive of rugose; minutely wrinkled. runner : an ultimate branch of a stolon or rhizome. saccate : bearing, or in the shape of, a pouch or bag. sagittate : arrow-shaped, specifically applied to structures with backward oriented basal lobes. salverform : shape of a corolla with a slender, basal tube and a more or less horizontally spreading limb. samara : a specialized type of achene that bears a wing or wings that aid in wind dispersal. scabrous : rough textured due to short, stiff hairs or minute prickles. scabrule : a single hair or prickle that collectively provides a scabrous texture. scale : a small, thin, flat epidermal outgrowth two or more cells wide. scape : a peduncle that arises from near the ground and lacks leaves (although it may possess small bracts), found in plants that lack aerial stems.

scapose : with the form of a scape. scarious : thin and dry, often chartaceous and translucent. schizocarp : a simple fruit composed of two or more carpels that separates at maturity into half- or unicarpellate segments called mericarps. scurfy : beset with small scales, providing a rough, irregular texture. scutelliform : the shape of a corolla that is circular or nearly circular in outline and lacks a basal tube and has somewhat ascending lobes (i.e., the corolla is slightly concave), more concave than rotate but not as concave as cupuliform; saucer-shaped. secund : with flowers or branches oriented or directed to one side of the axis; one-sided. seed : a mature ovule and perennating organ of many plants. senesce : to die back, as in the autumn. sepal : one of the hightly modified leaves of a flower, commonly a member of the outermost series and usually green to brown, lacking bright pigment. sepaloid : appearing like a sepal, usually in color, texture, and shape. septate : possessing septa. septum (plural: septa) : a partition; a partition formed by the walls of adjacent carpels in an ovary. septicidal : splitting through the septa, so that the capsule dehisces by separating the individual carpels (compare with loculicidal). sericeous : silky, due to the presence of long, slender, neatly appressed hairs. serotinous : the condition of the flowers’ expanding and developing after the formation of the leaves (compare with coaetaneous, precocious). serrate : a margin of a leaf provided with forward pointing, sharp teeth (compare with undulate, crenate, dentate). serrulate : diminutive of serrate; with small, forward pointing, marginal teeth. sessile : attached at the base, without a stalk of some form. seta (plural: setae) : a short, slender, relatively straight, firm bristle. setaceous : bristle-like. setose : covered with bristles. several : five to ten (compare with few, many).

g lossa ry  xxxi

sheath : the basal portion of leaves of bracts that surrounds and/or encloses the stem or branches. short shoot : a peg- or knob-like shoot with closely crowded leaves (compare with long shoot). shrub : a woody plant lacking a tree-like form, usually shorter than 6 m and with many stems at the base. silicle : similar to a silique, but up to three times as long as wide (compare with silique). silique : a fruit that dehisces by two valves, leaving the persistent replum to which the seeds are attached, found only in the Brassicaceae, this name is reserved for fruits greater than three times as long as wide (compare with silicle). simple : undivided. simple fruit : a fruit derived from a single ovary of a flower (compare with aggregate fruit, multiple fruit). sinus : the indented area between two lobes of an organ. sobol : a stem arising from axillary buds, near the ground, that is horizontal for a very short distance and then turns to become an upright, aerial stem. soboliferous : bearing sobols. sordid : with a dingy, gray coloration; appearing dirty. sorus (plural: sori) : aggregation of sporebearing structures of ferns. spadix : a spike-like inflorescence with flowers crowded on a fleshy axis, often associated with a spathe. spathe : a relatively large, solitary bract that subtends and commonly encloses the inflorescence, often associated with a spadix. spatulate : abruptly widened from a narrow base into a somewhat rounded apex; spoon-shaped. spicule : a minute, stiff, spine-like process on the margins of an organ. spike : an indeterminate inflorescence consisting of an elongate axis with sessile flowers. spikelet : diminutive of spike; the second order designation of a compound spike; one or more florets of a grass flower subtended by a pair of glumes. spine : a slender, firm, sharp structure derived from a leaf or a portion of a leaf (compare with prickle, thorn).

spinescent : with the form of or bearing a spine. spinose : bearing spines. spinule : diminutive of spine; a tiny spine. spinulose : diminutive of spinose; bearing tiny spines. sporangiaster : modified sporangium, forming an indument that partly covers the sorus. sporangiophore : a stalk that bears sporangia. sporangium : a structure that produces spores. spore : a one-celled reproductive structure in free-sporing plants (e.g., lycophytes, monilophytes) that gives rise to the gamete-bearing plant. sporocarp : a hardened, capsule-like structure derived from a modified leaflet that encloses the sorus in the Marsileaceae. sporophore : the fertile, sporebearing portion of the leaf blade in the Ophioglossaceae (compare with trophophore). sporophyll : a specialized leaf that bears a sporangium (compare with trophophyll). spreading : diverging from an axis by an angle of 45–90°. spur : a hollow, often elongate, process borne on the calyx or corolla; a slender process, often resembling a bristle, borne on the stamens of some species. squarrose : abruptly curving outward near the apex. stamen : one of the highly modified leaves in a flower that bear pollen; anther and filament collectively. staminate : describing a structure or plant that bears stamens. staminode : a sterile, modified stamen. stellate : condition of compound hairs that branch three or more times from a common point. sterigma (plural: sterigmata) : small, woody projection of a twig of some conifers to which the leaf is attached. stigma : pollen-receptor surface and associated tissue at the distal end of a style (when the latter is present). stipe : a stalk of a structure, used when more precise terms (such as petiole, peduncle, style, etc.) are inappropriate.

xxxii  gloss ary

stipel : an appendage, often paired, found at the base of petiolules in some species. stipitate : borne on a stipe. stipule : an appendage, often paired, found at the base of the petiole in some species of plants, a part of the leaf. stolon : a horizontal stem at or just above the surface of the ground. stoloniferous : producing and/or possessing stolons. stomate : an opening in the epidermal tissue of a leaf through which gas exchange occurs. stramineous : light yellow-brown; straw-colored. striate : ornamented with parallel lines. striga (plural: strigae) : a type of hair that is straight, appressed, and rigid. strigose : pubescent with straight, neatly appressed hairs. strobilus (plural: strobili) : the reproductive organ of the ferns, fern allies, and conifers, consisting of a cluster of sporophylls on an axis; a cone. strophiole : an appendage at the hilum of some seeds. style : the stalk that connects the stigma(s) to the ovary. stylopodium (plural: stylopodia) : a disklike or cylindrical enlargement of the basal portion of the style. sub- : Latin prefix meaning under or nearly so. submersed : below the surface of the water. subtend : positioned at the base of, as with bracts of a flower. subulate : narrowly tapering from the base to the apex; awl-shaped. succulent : fleshy and/or juicy. suffrutescent : somewhat woody or woody only near the base, firmer than herbaceous. suffruticose : somewhat woody. sulcate : with one or more narrow, elongate, and shallow depressions; grooved. superior ovary : an ovary attached to the receptacle above the insertion of all other flower parts (e.g., sepals, petals; compare with inferior ovary). surcurrent : possessing an adnate line or wing that extends up the axis above the node (compare with decurrent). suture : a line of fusion that will eventually split in a dehiscent structure such as a fruit.

sympodial : with a compound axis composed of one or more short, axillary branches (or leaf portions, as in Acorus) that supersede or overgrow the original, terminal bud of each axis segment (compare with monopodial). syncarpous : describing a gynoecium composed of connate carpels. synoecious : having only bisexual flowers (compare with dioecious, monoecious, polygamous). taproot : a large, central root that is noticeably larger than the lateral roots. tendril : a specialized leaf, branch, or inflorescence that aids a vine or liana in climbing. tepals : petals and sepals that are indistinguishable from each other, except by position, due to overall similarity. terete : circular in cross-section. terminal : the distal most position. terminal bud : the distalmost bud on a determinate branchlet, identified by the absence of a twig scar adjacent to the bud (compare with pseudoterminal bud). ternate : divided into three parts. terrestrial : pertaining to non-aquatic, land environments. testa : mature seed coat derived from mature integuments of the ovule. tetradynamous : description of a sixstamened androecium in which four stamens are longer than the other two, found commonly in Brassicaceae. tetragonous : four-angled, more or less square in cross-section. thalloid : appearing as a thallus. thallus : a body that is not differentiated into a stem and leaf. theca : a locule of an anther. thorn : a short, stiff, pointed process that is a modified branch (compare with prickle, spine). throat : the narrow, basal portion of a sympetalous corolla, proximal to the limb. thrum : one type of heterostylic flower in which the stamens nearly equal to exceed the length of the style(s) (compare with pin). thyrse : a mixed inflorescence with racemosely arranged cymes. tomentose : pubescent with curled, tangled, often matted, hairs; woolly.

g lossa ry  xxxiii

tomentulose : diminutive of tomentose; with short, fine, woolly hairs. tortuous : irregular due to bends and curves. torulose : the outline of an elongate structure with alternating constrictions and enlargements. torus : the receptacle of a flower or capitulum. trifid : cleft into three more or less equal parts. trigonous : triangular in cross-section. triquetrous : three-angled with each angle projecting in a thin, wing-like rib. tricolpates : a group of derived, dicotyledenous plants characterized by tricolpate pollen, includes most plants with two seed leaves trophophore : the sterile, photosynthetic portion of the leaf in the Ophioglossaceae (compare with sporophore). trophophyll : the sterile, photosynthetic leaf of lycophytes, usually scale-like and with a single nerve (compare with sporophyll). trullate : describing an organ that tapers with ± straight sides to each end but with the widest portion below the middle; trowel-shaped in outline truncate : with the base or apex of an organ latitudinally straight, appearing to have been cut off. tubercle : a swelling or projection borne on an organ, usually of different color and/ or texture. tuberculate : bearing tubercles. tuber : a thickened food-storage portion of a rhizome. tuberiferous : bearing tubers. tuberous : tuber-like. turbinoid : broad-obovoid-obonic; top shaped. turion : a small, bulbil-like, perennating organ. twig : same as branchlet. twining : growing in a curling, encircling fashion, thereby gaining support from other vegetation or structures. umbel : an indeterminate inflorescence, somewhat similar to a raceme, which has a greatly reduced central axis, such that the flowers appear to originate from the same point, usually resulting in a more or less flat-topped inflorescence, with a sequence of flowering from outside to inside. umbellet : diminutive of umbel; the ultimate unit of a compound umbel. umbilicate : having a small, circular depression in a broader, ± round surface.

umbo : a raised or thickened portion of the apophysis of a seed cone. umbraculiform : the shape of a corolla that is circular or nearly circular in outline and lacks a basal tube and has curvedascending segments so that a concave area exists inside the corolla segments; umbrella-shaped. unarmed : without any prickles, spines, or thorns. uncinate : hooked at the apex, similar to Velcro fasteners. undulate : wavy-margined, without prominent teeth. unifacial : leaf blades that have margins united so that only the abaxial surface is visible, as in the leaf of a species of Iris (compare with bifacial). unisexual : a flower that bears only stamens or only carpels; a flower that bears both stamens and carpels but one of which is non-functional or abortive and the flower is functionally unisexual (compare with bisexual). urceolate : constricted at a point just before an opening; urn-shaped. utricle : a specialized type of achene in which the pericarp is loosely attached and often readily removed. vallecular cavity : one of a set of longitudinal cavities in the stems of Equisetum external to the central cavity and internal to the carinal cavities. valvate : the condition of a structure that opens by valves; a position of petals in bud or winter bud scales that have the margins of each touching but not overlapping (compare with imbricate, convolute). valve : one portion of an ovary wall that separates when a capsule dehisces; one member of the inner set of accrescent tepals in Rumex. vascular bundle : a strand of xylem, phloem, and associated tissue, usually used in reference to fern petiole cross-sections or leaf scars in woody plants. vein : a visible vascular bundle passing through an organ; same as nerve. veinlet : the ultimate branch of a vein. velum : the thin, membranous tissue that covers the sporangium in Isoetes. velutinous : pubescent with fine, short, spreading hairs; velvety feeling. venation : the pattern of veins on an organ.

xxxiv gloss a ry

vernation : the orientation of leaves in bud or petals in bud, here used inclusively for the individual organs (ptyxis in the strict sense) and their relationship to one another (e.g., convolute, imbricate, valvate). verrucose : covered with small, wartlike projections versatile : an arrangement of anthers in which the filament is connected to the center of an anther; condition of a mesifixed anther (compare with basifixed). verticil : one whorled cycle of organs. verticillaster : a series of cymosely arranged flowers appearing to form a whorl that are distributed along an indeterminate axis, used to describe portions of the inflorescence in the Lamiaceae. verticillate : arranged in whorls. vestigial : poorly developed due to evolutionary reduction (compare with obsolete, rudimentary). villous : pubescent with long, soft, bent hairs, the hairs not crimped or tangled. vine : an herbaceous plant that does not fully support its own weight but rather has a stem that trails along the ground or climbs over other vegetation often by means of special climbing appendages or growth form.

virgate : a tall, slender, erect form. viscid : sticky, glutinous. viscidulous : diminutive of viscid. viscidulum (plural: viscidula) : a sticky gland to which the pollinia are attached that aids in the pollinia’s dispersal, found in the Orchidaceae. wanting : absent or nearly so. weeds : rapidly spreading plants of disturbed habitats. whorled : the arrangement of three or more similar organs at a node, such as leaves or flowers (compare with alternate, opposite). wing : a thin, flat projection from the side, top, or back of a structure; one of the two lateral petals in flowers of the Fabaceae; one of the two large, petaloid sepals in flowers of the Polygalaceae xeric : dry (compare with hydric, mesic). zygomorphic : bilaterally symmetrical; a shape that can be divided into equal halves by only one plane (compare with actinomorphic).

Key to the Families 1a. Plants typically reproducing by spores, seeds and fruits not produced; gametophyte independent of sporophyte; ferns and fern-like plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 1 1b. Plants typically reproducing by seeds, the seeds borne within a fruit or not; gametophyte dependent on sporophyte; seed plants 2a. Plants not producing true flowers; seeds commonly borne in strobili on the surface of a scale (embedded in a fleshy aril in Taxus), never enclosed in an ovary; styles and stigmas absent; trees and shrubs with narrow, scale- or needle-like, usually persistent, leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 2 2b. Plants usually producing true flowers; seeds enclosed in an ovary; stigma(s) and usually style(s) present, elevated above the ovary; woody or herbaceous plants with various types of leaves 3a. Leaf blades usually parallel-veined (or the plants thalloid in some Araceae); seeds with 1 cotyledon; perianth typically 3- or 6-merous; vascular bundles scattered throughout the stem; secondary growth absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 3 3b. Leaf blades usually pinnately veined; seeds with 2 cotyledons; perianth typically 4-, 5-, or more, -merous; vascular bundles arranged in a ring around a central pith; secondary growth absent or present 4a. Plants definitely woody; secondary growth present (though difficult to detect in some genera) 5a. Each node with 2 or more leaves or leaf scars (i.e., leaves opposite or whorled) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 4 5b. Each node with 1 leaf or leaf scar (i.e., leaves alternate; subopposite at some nodes in some Rhamnus and in Salix purpurea) . . . . . . . . . . . . . . . . . . . . . . . . Group 5 4b. Plants herbaceous or suffrutescent; secondary growth absent 6a. Flowers epigynous or partly so (i.e., ovary inferior) . . . . . . . . . . . . . . . . . . Group 6 6b. Flowers hypogynous or perigynous (i.e., ovary superior) or the flowers lacking a perianth 7a. Flowers zygomorphic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 7 7b. Flowers actinomorphic or without a perianth 8a. Gynoecium apocarpous, appearing as 2 or more distinct ovaries (i.e., the flower with 2 or more pistils) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 8 8b. Gynoecium composed of either a single carpel or 2 or more connate carpels, the ovary thereby superficially appearing as 1 (i.e., the flower with 1 pistil) 9a. Corolla gamopetalous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 9 9b. Corolla apopetalous, absent, or very inconspicuous . . . . . . . Group 10

2  ke y to th e familie s

Group 1 1a. Stems conspicuously jointed, bearing at each joint a small whorl of black or red-brown leaves (sometimes white-margined) that are united at the base; sporangia aggregated in a terminal strobilus [Fig. 25], borne on the underside of peltate sporangiophores . . . . . . . . . . . Equisetaceae 1b. Stems not conspicuously jointed, the leaves not arranged in distinct and remote whorls; sporangia aggregated in a terminal strobilus or not, but never borne on peltate sporangiophores 2a. Leaves or entire plant floating on the surface of the water (or stranded on mud as water levels decline); sporangia borne in sporocarps 3a. Plants rooted in the substrate, with 4-parted, clover-like leaves . . . . . . . . Marsileaceae 3b. Plants free-floating, with simple leaves 4a. Floating leaf blades mostly 5–15 mm long, with abundant and conspicuous branched hairs on the upper surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Salviniaceae 4b. Floating leaf blades 0.5–0.6 (–2) mm long, papillose, but lacking hairs on the upper surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Azollaceae 2b. Leaves not floating on surface of water (though often borne below the surface in Isoetaceae); sporangia borne on the surface of leaves, sometimes ± concealed 5a. Leaves narrow, with unexpanded, grass-like or quill-like blades; sporangia embedded in the leaf bases near the surface of the substrate [Fig. 7]; plants aquatic . . . . Isoetaceae 5b. Leaves various, but not quill-like; sporangia variously positioned, but usually elevated above the substrate; plants of upland or wetland communities 6a. Plants entirely filamentous, forming felt-like colonies on rock, in caves, and under sheltered overhangs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hymenophyllaceae 6b. Plants not filamentous, with moss-like or fern-like aspect, on various substrates 7a. Leaves small, up to 1.1 cm long, scale-like, with a single midvein; sporangia usually aggregated in a terminal strobili (arranged in alternating zones on the shoot in the Huperziaceae) 8a. Strobili quadrangular in cross-section (terete in Selaginella selaginoides); leaves with a small, adaxial ligule; spores of 2 sizes, the larger more than 0.3 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Selaginellaceae 8b. Strobili terete in cross-section or the sporophylls not aggregated into a strobilus; leaves without a ligule; spores of 1 size, less than 0.05 mm wide 9a. Sporophylls aggregated in a terminal strobilus [Figs. 12, 13], stalked, with lateral membranes, senescing after spore dispersal; gemmiphores not produced; plants anisotomously branched . . . . . . . . . . . . . . . . Lycopodiaceae 9b. Sporophylls arranged in alternating zones on the shoot [Fig. 2], unstalked, without lateral membranes, remaining green for the life of the plant; gemmiphores produced on mature plants; plants isotomously branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Huperziaceae 7b. Leaves 1–200 cm long, foliaceous, prominently veined; sporangia variously organized, but not borne in terminal strobili 10a. Plants vines, with palmately lobed leaf blades . . . . . . . . . . . . . Lygodiaceae 10b. Plants not vines, the leaf blades unlobed, pinnately lobed, or pinnately divided 11a. Sporangia thick-walled, lacking an annulus, commonly bearing more than 1000 spores; leaves not circinate in bud . . . . . . . . . . . Ophioglossaceae

g ro u p 1   3

11b. Sporangia thin-walled, with an annulus, bearing up to 512 spores; leaves circinate in bud 12a. Sporangium with an inconspicuous annulus composed of a group of apical, thick-walled cells on either side of the sporangial opening, bearing 256–512 spores; spore-bearing leaves or leaflets brown at maturity, conspicuously different from the vegetative leaves or leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmundaceae 12b. Sporangium with a well-developed annulus that forms a complete or nearly complete ring around the sporangium, usually bearing only 16–64 spores; spore-bearing leaves or leaflets normally green and similar to, or slightly different from, the vegetative leaves or leaflets (very different in Matteuccia and Onoclea) 13a. Sori located at the leaf margin; indusium formed, at least in part, by a revolute leaf margin or portion of the margin (i.e., false indusium) [Fig. 33]; spores tetrahedral or tetrahedral-globose 14a. Petiole apparently forking into 2 main divisions; sori borne on surface of the false indusium, this best viewed through dissection; ultimate segments without a distinct midrib . . . . (in part) Pteridaceae 14b. Petiole not appearing to fork into 2 divisions; sori borne on surface of leaf blade near margin and covered by false indusium; ultimate segments with a midrib 15a. Leaf blade broad-deltate to ovate-deltate, (25–) 40–70 cm wide; nectaries present at base of leaf divisions, appearing as dark, oval areas in early season; base of petiole with 10 or more vascular bundles . . . . . . . . . . . . . . . . . . (in part) Dennstaedtiaceae 15b. Leaf blade narrow-oblong to lanceolate or broad-lanceolate, 1.5–18 cm wide; nectaries absent; base of petiole with 1 or 2 vascular bundles 16a. Sori discrete, spherical, borne in circular or slightly bivalvate cups formed by fusion of the true indusium with the false indusium; false indusium merely a reflexed tooth or lobe . . . . . . . . . . . . . . . . . . . . . . . . (in part) Dennstaedtiaceae 16b. Sori becoming confluent as marginal bands, not borne in cup-like structures; false indusium formed by a ± elongate, revolute margin . . . . . . . . . . . . . . . . . . . . . . (in part) Pteridaceae 13b. Sori not located at the margin (except in Dryopteris marginalis) [Fig. 34]; indusium absent or present, but not formed by a revolute leaf margin (formed by a highly modified, revolute leaf margin in Matteuccia and Onoclea); spores oblong, reniform, bilateral, or winged 17a. Some of the sporangia modified to form sporangiasters (i.e., tiny, stalked, and stipitate-glandular bodies mixed within the sorus); indusium absent; leaves evergreen, coriaceous, once-pinnate [Figs. 30, 31] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polypodiaceae 17b. Sporangiasters absent; indusium usually present, sometimes inconspicuous (absent in Gymnocarpium and Phegopteris); leaves evergreen or deciduous, herbaceous to coriaceous, variously divided 18a. Sori closely aligned in a single, chain-like row immediately adjacent to and on either side of the costa or costule (depending on the leaf division); indusium elongate and flap-like, opening toward the costa or costule (depending on leaf division); veins of leaf blade rejoining to form 1 or 2 (or more) rows of

4  key to the famili es

aereoles along the costa, then forking until meeting the margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Blechnaceae 18b. Sori variously arranged, when in rows parallel to the costa or costule then not immediately adjacent to them, rather on or adjacent to veinlets; indusium round to elongate, when elongate then opening away from the associated veinlet; veins of leaf blade unforked or freely forking, but not rejoining to form 1 or more rows of aereoles (except in Asplenium rhizophyllum) 19a. Scales of the stem prominently clathrate-reticulate; indusia elongate and flap-like; petiole with 2 vascular bundles, these united distally into 1 X-shaped bundle . . . . . . . . . . Aspleniaceae 19b. Scales of the stem not prominently clathrate-reticulate; indusia elongate and flap-like, circular, reniform, lacerate, or absent; petiole with 2–7 vascular bundles, when 2, united into 1 U-shaped bundle distally 20a. Plants pubescent with needle-like, unicellular, transparent hairs (scales may be present as well) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelypteridaceae 20b. Plants glabrous or pubescent, when pubescent the hairs pigmented, glandular, multicellular and/or flattened (these flattened “hairs” are actually microscales) 21a. Leaves conspicuously dimorphic—fertile leaves bearing sporangia hidden by highly modified, indurate, revolute leaf segments and vegetative leaves either pinnately lobed (pinnate only at base) and with reticulate venation or vegetative leaves with lower leaflets greatly reduced in size compared with medial leaflets; spores green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Onocleaceae 21b. Leaves monomorphic to weakly dimorphic; sterile leaves with simple or forking veins that do not rejoin, at least once-divided, and with basal leaflets that do not very gradually reduce in size; spores yellow or light brown to brown 22a. Indusium inferior, composed of lacerate or filamentous segments at maturity; veins not reaching margin of leaf segments, ending in hydathodes near the margin; petioles articulated near base, this recognizable as a small, swollen node (not articulated in W. obtusa) . . . . . . . . . . . . . . . . (in part) Woodsiaceae 22b. Indusium absent or present, sometimes inconspicuous, superior when present, attached centrally or laterally, not divided; veins reaching margin of leaf segments (except in Dryopteris); petioles not articulated near base 23a. Indusium round to reniform; petiole base with 3–7 vascular bundles; teeth of leaf segments often bristle-tipped . . . . . . . . . . . . . . . . Dryopteridaceae 23b. Indusium broad-linear, hooked, or horseshoe-shaped, or absent [Figs. 34, 38]; petiole base with 2 vascular bundles; teeth of leaf segments not bristle-tipped . . . . . . . . . (in part) Woodsiaceae

g ro u p 2   5

Group 2 1a. Seeds borne singly, partly concealed by a red, fleshy aril; abaxial surface of the flat leaves bearing pale yellow, longitudinal, stomatal lines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taxaceae 1b. Seeds 1–400, borne in woody or fleshy strobili, lacking a fleshy, red aril [Figs. 43, 45, 48]; abaxial surface of the flat to tetragonous leaves without yellow lines (though sometimes with white or gray stomatal lines) 2a. Leaves opposite or whorled, imbricate and concealing the branchlet (the branchlet visible in Juniperus communis), persistent on dried specimens; strobili with peltate scales, or with opposite, basifixed scales, or the scales fleshy and coalesced into a berry-like form [Fig. 43]; each scale of a strobilus bearing 1–20 ovules, without free subtending bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cupressaceae 2b. Leaves alternate, fascicled, or in dense tufts on short shoots, not concealing the branchlet, deciduous from dried specimens; strobili with flat, alternately arranged, basifixed scales [Figs. 45, 48]; each scale of a strobilus bearing 2 ovules, subtended by a free bract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pinaceae

Group 3 1a. Plants thalloid, not differentiated into stems and leaves, 0.5–15 mm long, free floating on or near the surface of water [Fig. 61] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Araceae 1b. Plants not thalloid, differentiated into stems and leaves, usually much longer or taller; habit various, but when floating usually rooted in the substrate 2a. Flowers replaced by, or modified into, vegetative propagules 3a. Plants from a conspicuous bulb [Fig. 60], with a strong odor of onion or garlic; stems ± scapose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Alliaceae 3b. Plants without a conspicuous, bulbous base, lacking the odor of onion; stems with leaves 4a. Leaves terete or elliptic in cross-section and with complete, transverse septa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Juncaceae 4b. Leaves flat, without complete transverse septa (though partial septa may be present) 5a. Plants submersed aquatics, with flaccid underwater leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Potamogetonaceae 5b. Plants terrestrial, with firmer leaves 6a. Inflorescence a panicle [Fig. 259]; leaves arranged in 2 ranks, with collar-like ligules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Poaceae 6b. Inflorescence a spike, with flowers in an umbel-like cluster; leaves arranged in 3 ranks, with ligules, but these inconspicuous and not collar-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cyperaceae 2b. Flowers not replaced by vegetative bulbils 7a. Gynoecium composed of 2 or more distinct carpels and each carpel with a single stigma [Fig. 301]; plants aquatic or of hydric habitats

6  key to th e famili e s

8a. Leaf blades triquetrous in cross-section; perianth petaloid, composed of green-pink sepals and pink petals; flowers with 9 stamens and 6 carpels . . . . . . . . . . . Butomaceae 8b. Leaf blades either flat, ± terete, or capillary; perianth either absent, entirely sepaloid, or composed of green sepals and white petals; flowers with 1–many stamens and 3–many carpels, but not consistently 9 and 6, respectively 9a. Perianth present, showy, differentiated into sepals and petals; petals white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alismataceae 9b. Perianth, if present, reduced, not differentiated into sepals and petals; petals (or inner tepals), if present, usually green, yellow-green, or brown 10a. Plants emersed; leaves ± terete in cross-section 11a. Each carpel with 2 ovules; inflorescence bracteate; each leaf with a terminal pore [Fig. 304] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scheuchzeriaceae 11b. Each carpel with 1 ovule; inflorescence without bracts; leaves without terminal pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Juncaginaceae 10b. Plants submersed (except often the inflorescence, which is held above the water surface); leaves flat or capillary 12a. Each flower with 4 tepal-like appendages and 4 stamens; fruits sessile; stipules distinct from the leaves or basally connate with a distinct tip; plants primary of fresh or fresh-tidal waters . . . . . . . . . (in part) Potamogetonaceae 12b. Each flower with 0 tepals and 2 stamens; fruits long-stipitate; stipules completely connate to the leaves; plants of brackish waters . . . . . Ruppiaceae 7b. Gynoecium composed of 1 carpel or 2 or more connate carpels and then the ovary surmounted by 2 or more stigmas; plants aquatic or terrestrial 13a. Plants aquatic or marine, with thin, flaccid, often translucent, submersed or floating leaves; flowers unisexual 14a. Flowers in dense, spherical clusters [Fig. 306], each flower with 1 stigma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Typhaceae 14b. Flowers not in spherical clusters, each flower with 2–6 or more stigmas 15a. Flowers with a perianth composed of 3 sepals and 3 petals; androecium usually with 2 or 9 stamens; gynoecium composed of 3–6 weakly united carpels; leaves whorled (at least in part) or all basal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Hydrocharitaceae 15b. Flowers without a perianth; androecium with 1 stamen; gynoecium composed of a single, unilocular, uni- or bicarpellate ovary; leaves alternate or opposite 16a. Pollen filiform; gynoecium with 2 stigmas; leaves mostly alternate; plants perennial, with rhizomes, marine . . . . . . . . . . . . . . . . . . . . . Zosteraceae 16b. Pollen globose; gynoecium with 2–4 stigmas; leaves opposite; plants annual, without rhizomes, of fresh water habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Hydrocharitaceae 13b. Plants of upland or wetland habitats, with firmer leaves that are usually, at least partly, emersed (or at least the leaves holding their form when removed from the water); flowers bisexual, or, less commonly, unisexual 17a. Inflorescence a spadix 18a. Leaves ensiform, unifacial, parallel-veined; ovary 2- or 3-locular; anthers introrse; spathe-like bract (actually the distal portion of a sympodial leaf) narrow, grass-like, not concealing the spadix . . . . . . . . . . . . . . . . . . . . . Acoraceae

g ro u p 3   7

18b. Leaves with broad and flat, bifacial, pinnately veined blades; ovary unilocular; anthers extrorse; spathe broad, partially or entirely concealing the spadix (least so in Calla) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Araceae 17b. Inflorescence various, but not a spadix 19a. Flowers anemophilous or self-pollinated, without a perianth or with an inconspicuous perianth of scales, bristles, or sepaloid tepals 20a. Flowers arranged in a solitary, terminal, involucrate, white to gray, pseudanthial cluster; roots conspicuously septate . . . . . . . . . . Eriocaulaceae 20b. Flowers arranged otherwise; roots lacking conspicuous septa 21a. Inflorescence consisting of a dense, elongate, cylindrical spike, the carpellate portion below, the staminate portion above; fruit a wind-dispersed, 1-seeded follicle . . . . . . . . . . . . . . . . . . . (in part) Typhaceae 21b. Inflorescence otherwise; fruit a capsule, achene, utricle, or caryopsis 22a. Fruit a 3- or many-seeded capsule; perianth of 6 sepaloid tepals [Figs. 140, 147] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Juncaceae 22b. Fruit indehiscent; perianth of 2–6 inconspicuous tepals, scales, bristles, or absent altogether 23a. Ovary with 1 stigma (2 stigmas in Sparganium eurycarpum); perianth of 2–6 tepals; inflorescence a dense, spherical cluster [Fig. 306] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Typhaceae 23b. Ovary with 2 or 3 stigmas; perianth of scales, bristles, or absent; inflorescence variously open to compact 24a. Leaves arranged in 3 ranks or absent and represented by bladeless sheaths; stems usually solid, very often triangular in cross-section; flowers spirally arranged (distichous in Cyperus and Dulichium) [Figs. 73, 89, 125], ; fruit an achene [Figs. 97, 116, 126] . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cyperaceae 24b. Leaves arranged in 2 ranks; stems usually hollow, almost never triangular in cross-section; flowers and floral scales distichously arranged [Figs. 181, 239, 232]; fruit a caryopsis or rarely a utricle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Poaceae 19b. Flowers clearly entomophilous, with an evident, showy perianth 25a. Ovary inferior 26a. Flowers zygomorphic [Figs. 164, 174, 176], with 1 or 2 stamens; pollen cohering in pollinia; seeds minute, dust-like . . . . . . . . . . . . . . . Orchidaceae 26b. Flowers actinomorphic [Figs. 136, 138], with 3 or 6 stamens; pollen separate, not cohering in masses; seeds larger 27a. Plants vines, dioecious . . . . . . . . . . . . . . . . . . . . . . . . . Dioscoreaceae 27b. Plants upright, not twining, synoecious 28a. Inflorescence conspicuously white-tomentose; perianth tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haemodoraceae 28b. Inflorescence not tomentose; perianth glabrous or minutely pilose 29a. Flowers with 3 stamens; leaves equitant; seeds without phytomelan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Iridaceae 29b. Flowers with 6 stamens; leaves not equitant; seeds usually with phytomelan

8   ke y to th e familie s

30a. Perianth pubescent on the abaxial (i.e., outside) surface; leaves pubescent; plants from corms . . . . . . . . . Hypoxidaceae 30b. Perianth glabrous; leaves glabrous; plants from bulbs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amaryllidaceae 25b. Ovary superior 31a. Flowers borne in the axils of firm, imbricate bracts, aggregated in a dense, ellipsoid to ovoid, terminal spike; leaf blades grass-like; petals yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xyridaceae 31b. Flowers not from the axils of imbricate bracts, commonly in more open inflorescences; leaf blades various; petal color various 32a. Sepals sepaloid, green 33a. Leaf blades grass-like, with parallel venation; inflorescence composed of more than 1 flower; fruit a capsule . . . Commelinaceae 33b. Leaf blades broad, with pinnately branched secondary veins [Fig. 159]; inflorescence composed of a solitary, terminal flower; fruit a berry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Melanthiaceae 32b. Sepals petaloid, of a color similar to that of the petals 34a. Perianth zygomorphic [Fig. 286], the upper lip bearing 1 or 2 closely spaced yellow spots; stamens adnate to the corolla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Pontederiaceae 34b. Perianth actinomorphic, lacking yellow spots; stamens free or adnate to the corolla 35a. Leaves palmately veined (with pinnately branched secondary veins), with a well defined petiole and blade, with stipular tendrils [Fig. 305] . . . . . . . . . . . . . . . . . . . . . . . . Smilacaceae 35b. Leaves parallel-veined, generally lacking a well defined petiole and blade, without tendrils 36a. Actual leaves scale-like; upper branches filiform, green, simulating leaves, clustered in the leaf axils . . . . . Asparagaceae 36b. Leaves foliaceous; upper branches not clustered in the leaf axils 37a. Flowers with 3 stamens; plants aquatic, with flaccid leaves . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Pontederiaceae 37b. Flowers with 6 stamens (rarely only 4); plants of upland or seasonally saturated soils, with firmer leaves 38a. Tepals evidently connate; filaments adnate to the tepals or free in Hostaceae 39a. Tepals 4–18 cm long including the basal, connate portion; anthers versatile 40a. Leaf blades lanceolate to broad-ovate, 5–200 cm wide; inflorescence a raceme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hostaceae 40b. Leaf blades linear to lanceolate, 0.8–5 cm wide; inflorescence a solitary flower or an umbellike helicoid cyme 41a. Tepals yellow or orange, the basal, connate portion 1.5–3 cm long; plants mostly 5–15 dm tall, from tuberous roots, with flowers borne on an evident scape . . . . . . . . . Hemerocallidaceae

g ro u p 3   9

41b. Tepals pink to purple or white (rarely yellow), the basal, connate portion longer than 8 cm; plants shorter than 3 dm, from corms, with short, obscure, subterranean scapes . . . . . . . . . . . . . . . . . . . . . . . (in part) Colchicaceae 39b. Tepals 0.6–2.2 cm long; anthers basifixed or versatile 42a. Inflorescence axillary or a secund raceme; fruit a berry . . . . . . . . . . . . . . . . . (in part) Ruscaceae 42b. Inflorescence a raceme with spirally arranged flowers; fruit a capsule 43a. Perianth minutely roughened on the abaxial (i.e., outside) surface [Fig. 160]; pedicels ascending; leaf blades 5–26 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nartheciaceae 43b. Perianth not roughened on the abaxial surface; pedicels spreading-ascending to drooping, especially the lower ones [Fig. 128]; leaf blades 2–8 mm wide (10–20 mm wide in Chionodoxa) . . . . . . . . . . (in part) Hyacinthaceae 38b. Tepals distinct or nearly so; filaments free or adnate in Allium, Convallaria, and Streptopus 44a. Plants with a large rosette of firm leaves that have fibrous margins and a spinose apex; tepals thick and fleshy . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agavaceae 44b. Plants with or without basal rosettes, but the basal leaves (if present) lacking marginal fibers and apical spines; tepals relatively thinner 45a. Flowers in a many-flowered umbel at the summit of a scape; seeds with phytomelan; plants with the odor of onion or garlic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Alliaceae 45b. Flowers arranged otherwise (forming an umbel with 3–6 flowers in Clintonia) at the summit of a scape or leafy stem; seeds without phytomelan (except in Ornithogalum) plants without the odor of onion or garlic 46a. Scape viscid-pubescent; leaves both equitant and distichous . . . . . . . . . Tofieldiaceae 46b. Stem or scape glabrous or pubescent, but not viscid; leaves not set edge to stem, spirally arranged or, less commonly, distichous 47a. Leaves in 1 or 2 whorls; styles elongate, capillary, exceeding the tepals in length; fruit a purple to black berry . . . . . (in part) Liliaceae 47b. Leaves arranged otherwise; styles shorter; fruit a capsule or berry, the berry blue, red, or becoming red 48a. Styles 3, separate to the base; fruit a septicidal capsule (loculicidal in Stenanthium)

10   key to th e fa mili es

49a. Styles simple, unspotted, tepals 2-12 mm long; inflorescence a terminal raceme or panicle . . . . . . . . . . . . . . (in part) Melanthiaceae 49b. Styles distally bifid, maculate; tepals 20–25 mm long; inflorescence of few to several flowers in the axils of leaves . . . . . . . . . . . (in part) Liliaceae 48b. Styles 1, divided or lobed only in the apical portion; fruit a berry or loculicidal capsule 50a. Stems from a bulb or corm; fruit a capsule, circular or bluntly angled in cross-section 51a. Aerial stems leafy; tepals 30–100 mm long . . . . . . . . . . . . . . . . . (in part) Liliaceae 51b. Aerial stems scapose; tepals 12–33 mm long 52a. Tepals white with an abaxial green stripe or entirely blue; inflorescence a raceme (sometimes a solitary flower in Othocallis); leaf blades not mottled . . . . . . . . . (in part) Hyacinthaceae 52b. Tepals yellow; inflorescence a solitary flower; leaf blades adaxially mottled . . . . . . . . . . . . . . . (in part) Liliaceae 50b. Stems from rhizomes; fruit a berry, or a capsule with triquetrous cross-section in Uvularia 53a. Tepals white; inflorescence terminal, a raceme or panicle [Fig. 302] . . . . . . (in part) Ruscaceae 53b. Tepals yellow, green-white, yellow-green, or pink; inflorescence axillary, an umbel, or a solitary, terminal flower 54a. Tepals yellow; fruit a capsule; leaves borne on an aerial stem . . . . . . . . . . . (in part) Colchicaceae 54b. Tepals green-white, pink, or green-yellow; fruit a berry; leaves all basal or borne on an aerial stem . . . . . . . . . . (in part) Liliaceae

g ro u p 4   1 1

Group 4 1a. Plants 5–30 mm tall, parasitic on Picea, Larix, and Pinus [Fig. 939]; anthers sessile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Viscaceae 1b. Plants much taller or longer at maturity, not parasitic; anthers with filaments 2a. Leaves compound 3a. Fruit an inflated, thin-walled capsule with 3 lobes [Fig. 917]; gynoecium with 3 styles that are basally distinct and apically coherent . . . . . . . . . . . . . . . . . . . . . . . Staphyleaceae 3b. Fruit otherwise; gynoecium with 1 style that is often terminated by a lobed or branched stigmatic surface 4a. Perianth zygomorphic; fruit a capsule 5a. Plants trees with palmately compound leaves; capsule 3–5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Sapindaceae 5b. Plants lianas with pinnately compound leaves; capsule 10–15 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Bignoniaceae 4b. Perianth actinomorphic or absent; fruit an achene, schizocarp, drupe, or samara 6a. Plants lianas (though woody only near base); flowers with numerous stamens; fruit an achene terminated by an elongate, plumose style [Fig. 812] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Ranunculaceae 6b. Plants upright shrubs or trees; flowers with 2–12 stamens; fruit otherwise 7a. Leaflets punctate with aromatic glands; winter buds naked, enclosed by petiole base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Rutaceae 7b. Leaflets not punctate; winter buds covered by scales, not enclosed in petiole base 8a. Fruit a samaroid schizocarp; ovary bilobed, compressed; androecium composed of 4–12 stamens, commonly 8 . . . . . . . . . . . . (in part) Sapindaceae 8b. Fruit a samara or drupe; ovary not bilobed, not compressed; androecium composed of 2 or 5 stamens 9a. Perianth absent; androecium composed of 2 stamens; fruit a samara; flowers usually unisexual; trees at maturity . . . . . . . . . . . (in part) Oleaceae 9b. Perianth present and gamopetalous; androecium composed of 5 stamens; fruit a drupe; flowers bisexual; shrubs at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Adoxaceae 2b. Leaves simple 10a. Apical portions of the stem succulent; leaves minute and scale-like, 1–3 mm long [Fig. 339]; stems appearing jointed; plants of Atlantic coast shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Amaranthaceae 10b. Apical portions of the stem herbaceous to woody; leaves with foliaceous blades, mostly longer; stems not appearing jointed; plants of inland habitats 11a. Leaves and leaf scars whorled 12a. Perianth zygomorphic; fruit a slender, elongate capsule 20–45 cm long; seeds winged, the wings terminating in hairs . . . . . . . . . . . . . . . . . . . (in part) Bignoniaceae 12b. Perianth actinomorphic; fruit an achene-like schizocarp or a subglobose capsule 0.4–0.7 cm long; seeds without prominent wings

12   k ey to the fa milies

13a. Inflorescence a cyme [Fig. 714]; flowers each with 8 or 10 stamens; petals pink-purple, 10–15 mm long; stems woody in the basal portion, surrounded by loose, spongy tissue, the buds barely emerging through the tissue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lythraceae 13b. Inflorescence a dense, spherical cluster [Fig. 874]; flowers each with 4 stamens; petals white, 5–8 mm long; stems woody throughout, not surrounded by spongy tissue, the buds clearly visible . . . . . . . . . . . . . . . . . (in part) Rubiaceae 11b. Leaves and leaf scars opposite 14a. Leaves and branchlets conspicuously white- and/or brown-lepidote [Fig. 605]; plants dioecious . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Elaeagnaceae 14b. Leaves and branchlets without abundant scales; plants synoecious, polygamous, or dioecious 15a. Leaf blades 6–15 mm long, crowded and imbricate [Fig. 615]; pulvinate plants of exposed, alpines areas above 1000 m elevation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Diapensiaceae 15b. Leaf blades longer than 20 mm, not crowded; plants of lower elevation habitats (except sometimes Viburnum edule) 16a. Inflorescence a capitulum; fruit a resin-dotted cypsela . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Asteraceae 16b. Inflorescence not pseudanthial; fruit a schizocarp, capsule, follicle, berry, or drupe, or 10–35 achenes enclosed in a fleshy hypanthium 17a. Perianth of 15–30 brown-purple tepals; gynoecium with 10–35 carpels; androecium with 10–20 stamens and 10–25 staminodes seated inside the cycle of stamens . . . . . . . . . . . . . . . . . . . . . . . . . . Calycanthaceae 17b. Perianth with fewer petaloid segments; gynoecium with 1–5 carpels; androecium otherwise, mostly with fewer stamens 18a. Inflorescence mixed (i.e., thyrsoid), the indeterminate axis bearing cymose lateral branches, lateral branches manifesting as dense verticillasters of 6–10 flowers; fruit a schizocarp, separating into 4 achene-like mericarps; foliage pleasantly fragrant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lamiaceae 18b. Inflorescence not thyrsoid (appearing thyrsoid in Decodon, but then each cyme subtended by foliage leaves); fruit a capsule, follicle, or drupe, or a schizocarp in Acer that splits into 2 samara-like mericarps; foliage not especially fragrant 19a. Stems woody only in the basal portion and surrounded by loose, spongy tissue, the buds barely emerging through the tissue; stamens dimorphic, of 2 different lengths . . . . . (in part) Lythraceae 19b. Stems woody ± throughout, not surrounded by spongy tissue, the buds clearly visible; stamens monomorphic 20a. Fruit a samaroid schizocarp; ovary bilobed and compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Sapindaceae 20b. Fruit a capsule, follicle, or drupe; ovary not, or inconspicuously, lobed, not compressed 21a. Flowers without perianth; fruit a follicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cercidiphyllaceae 21b. Flowers with perianth; fruit a capsule, berry, achene, or drupe 22a. Perianth 5-merous

g ro u p 4   13

23a. Corolla 50–70 mm long; ovary superior; fruit a capsule 30–40 mm long; seeds winged . Paulowniaceae 23b. Corolla 3–50 mm long; ovary inferior; fruit a drupe, berry, or achene, or a capsule 8–15 mm long; seeds without evident wings 24a. Flowers with 8 or 10 stamens; corolla apopetalous . . . . . . . . . . . . . . (in part) Hydrangeaceae 24b. Flowers with 4 or 5 stamens; corolla gamopetalous 25a. Style absent or very short; corolla of fertile flowers actinomorphic, white, 4–8 mm long (some species with sterile, enlarged, marginal flowers) . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Adoxaceae 25b. Style elongate; corolla actinomorphic or zygomorphic, white, pink, yellow, red, or purple, 3–50 mm . . . . . . . . . . . . . . . (in part) Caprifoliaceae 22b. Perianth 4-merous 26a. Leaf blades sessile, punctate with pellucid dots; petals yellow . . . . . . . . . . . . . . . . . . (in part) Hypericaceae 26b. Leaf blades petiolate (though the petioles sometimes short), lacking pellucid dots; petals white, yellow, green-white, blue, or brown-purple 27a. Flowers with 2 or 4 stamens 28a. Corolla gamopetalous 29a. Flowers with 4 stamens; leaf blades white-tomentose on the abaxial surface . . . . . . . . . . . . . . . . . . . (in part) Scrophulariaceae 29b. Flowers with 2 stamens; leaf blades glabrous or pubescent on the abaxial surface, but not white-tomentose . . . (in part) Oleaceae 28a. Corolla apopetalous 30a. Fruit a drupe; leaf blades with arcuate venation; styles present; branches terete . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cornaceae 30b. Fruit a capsule; leaf blades not with arcuate venation; styles absent; branches often ± quadrangular . . . . . . . . (in part) Celastraceae 27b. Flowers with 5–many stamens; corolla apopetalous 31a. Petals yellow; leaf blades sessile and pellucidpunctate . . . . . . . . . . . . . . . . . (in part) Hypericaceae 31b. Petals white, brown-purple, green-white, or ± yellow in Euonymus alatus; leaf blades petiolate (though the petioles sometimes short), without pellucid dots 32a. Fruit a drupe; leaf blades entire, with arcuate venation [Fig. 594] . (in part) Cornaceae 32b. Fruit a capsule; leaf blades toothed, with pinnate venation . . . . . (in part) Hydrangeaceae

14   ke y to t he fami lie s

Group 5 1a. Inflorescence a capitulum; fruit a cypsela . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Asteraceae 1b. Inflorescence not pseudanthial; fruit various, but not a cypsela 2a. Leaves compound 3a. Perianth zygomorphic 4a. Corolla usually papillionaceous; filaments connate, conspicuously so in most genera (distinct in Cladrastis), not seated on a nectar disk; fruit a legume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Fabaceae 4b. Corolla not papillionaceous; filaments distinct, seated on a nectar disk; fruit a bladdery-inflated, 3-valved capsule . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Sapindaceae 3b. Perianth actinomorphic 5a. Leaves palmately compound 6a. Leaflets retuse at the apex; sepals numbering 3 or 6, petaloid, purple brown; gynoecium of 3 distinct carpels (i.e., the flowers with 3 pistils) . . . . Lardizabalaceae 6b. Leaflets tapering to the apex; sepals numbering 5, sepaloid, green; gynoecium of 2–5 connate carpels (i.e., the flowers with 1 pistil) 7a. Plants climbing by means of tendrils produced opposite the leaves, unarmed; inflorescence a panicle; ovary superior . . . . . . . . . . . . (in part) Vitaceae 7b. Plants not climbing, armed with prickles; ultimate unit of inflorescence an umbel; ovary inferior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Apiaceae 5b. Leaves pinnately compound 8a. Leaves twice pinnately compound (only once pinnately compound on spurs in Gleditsia); fruit a legume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Fabaceae 8b. Leaves once pinnately compound; fruit not a legume 9a. Inflorescence a unisexual ament, the staminate drooping; flowers small, with an inconspicuous perianth; fruit a drupe with a fibrous exocarp and nutlike endocarp [Fig. 689] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Juglandaceae 9b. Inflorescence not an ament, uni- or bisexual, none drooping; flowers with an evident perianth; fruit a fleshy drupe, samara, follicle, pome, or aggregate 10a. Leaf blades punctate with pellucid dots; fruit a fleshy follicle, an orbicular samara, or a drupe . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Rutaceae 10b. Leaf blades not punctate; fruit mainly otherwise, though a few genera with drupe fruits 11a. Leaflets entire except for 1 or more coarse teeth near the base; fruit a samara . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Simaroubaceae 11b. Leaflets toothed to lobed; fruit not a samara 12a. Ultimate unit of the inflorescence an umbel; styles swollen at the base forming a stylopodium . . . . . . . . . . . . . . . . . . . . . . (in part) Apiaceae 12b. Ultimate unit of the inflorescence not an umbel; styles without a stylopodium 13a. Perianth monochlamydeous, only the sepals present; wood yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Ranunculaceae

g ro u p 5   15

13b. Perianth dichlamydeous, both sepals and petals present; wood not yellow 14a. Flowers with a hypanthium, bisexual; plants with a watery sap; stems armed or unarmed . . . . . . . . . . . . . . . (in part) Rosaceae 14b. Flowers without a hypanthium, commonly unisexual; plants with a milky or watery sap (note: when watery, the sap poisonous); stems unarmed . . . . . . . . . . . . . . . . . . . Anacardiaceae 2b. Leaves simple 15a. Plants lianas 16a. Leaves with sheathing stipules; perianth segments with the midvein abaxially winged, especially in fruit; fruit a trigonous achene . . . . . . . . . (in part) Polygonaceae 16b. Leaves exstipulate, or minute and caducous (e.g., Actinidiaceae), or evident in the Vitaceae, but these not sheathing the entire circumference of the stem; perianth segments lacking an evident, abaxial, longitudinal wing; fruit a berry, drupe, or capsule 17a. Calyx gamosepalous, 3-lobed, with a strongly curved basal tube, petaloid [Fig. 50]; flowers with 6 stamens . . . . . . . . . . . . . . . . . . . . (in part) Aristolochiaceae 17b. Calyx gamosepalous, aposepalous, or obsolete, without a strongly curved tube, sepaloid; flowers with 5 or 12–100 stamens (when stamens are present) 18a. Corolla gamopetalous; anthers dehiscing by terminal pores or clefts; some of the leaf blades with a distinct pair of small lobes or leaflets [Fig. 914] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Solanaceae 18b. Corolla apopetalous (cohering at the summit in Vitis); anthers dehiscing by longitudinal slits; leaf blades simple or lobed, the lobes, when present, not confined to the base of the blade 19a. Leaf blades with peltate attachment of petioles; flowers with 6–9 petals and 12–24 stamens; fruit a drupe . . . . . . . . . . . . . . . . . . . . . . Menispermaceae 19b. Leaf blades with basal attachment of petioles; flowers with 5 petals and 5 or 18–100 stamens (when stamens are present); fruit a berry or capsule 20a. Leaves palmately veined or palmately lobed [Fig. 942]; tendrils and inflorescences produced opposite the leaves; fruit a berry, white to blue or dark blue to black (rarely red or green) . . . . . . . . . . . . . . . (in part) Vitaceae 20b. Leaves pinnately veined; tendrils absent; inflorescences terminal or axillary; fruit a capsule or green-yellow berry 21a. Stamens 5, the anthers basifixed; fruit an orange-yellow to orange capsule 6–13 mm long; pith continuous . . . . . . . (in part) Celastraceae 21b. Stamens 18–100, the anthers versatile; fruit a green-yellow berry 20–30 mm long; pith diaphragmed . . . . . . . . . . . . . . . . . . . Actinidiaceae 15b. Plants erect to mat-forming, but not climbing 22a. Plants with a milky latex; fruit a multiple, usually fleshy (dry in Broussonetia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Moraceae 22b. Plants with a watery sap; fruit simple or aggregate, dry or fleshy 23a. Stamens monadelphous; epicalyx of (6–) 8–15 bractlets present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Malvaceae 23b. Stamens distinct (connate for a short distance at the base in Halesia and cohered or connate in 2 species of Salix); epicalyx absent

16  k ey to the fa mili es

24a. Fruit a dark red to nearly black drupe with thin flesh on a peduncle (7–) 10–20 (–25) mm long; leaf blades asymmetrical at the base, usually strongly so, with 1 or both sides cordate . . . . . . . . . . . . . . . . . . . . . . . (in part) Cannabaceae 24b. Fruit otherwise; leaf blades ± symmetrical or, less commonly, asymmetrical at the base, narrow-cuneate to cordate 25a. Flowers unisexual 26a. Leaves palmately 3- to 5-lobed; carpellate flowers in dense, spherical clusters; fruit a spherical aggregate of achenes or capsules 27a. Bark exfoliating, creating a mottled pattern of colors; fruit an aggregate of narrow achenes; leaf blades with broad-triangular lobes that are coarsely and sparsely serrate . . . . . . . . . . . . . . . . . . Platanaceae 27b. Bark not exfoliating; fruit an aggregate of capsules; leaf blades with narrow-triangular to triangular lobes that are finely and closely serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Altingiaceae 26b. Leaves not palmately lobed (except in some 3-lobed leaves of Sassafras); flowers not in dense, spherical clusters; fruit otherwise 28a. Plants up to 50 cm tall, with numerous, crowded leaves with linear to narrow-elliptic blades 3–8 mm long [Fig. 610]; fruit a dry or fleshy drupe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Ericaceae 28b. Plants usually taller, with larger, broader leaf blades; fruit various 29a. Inflorescence an ament (the carpellate flowers solitary in in small clusters in the Fagaceae); perianth absent or tiny and inconspicuous 30a. Winter buds either covered by a single, cap-like scale or with the lowest scale centered over the leaf scar; fruit a capsule, containing comose seeds . . . . . . . . . . . . . . . . . . . . . . . . . . Salicaceae 30b. Winter buds with 2 or more scales, the lowest scale not centered over the leaf scar; fruit an achene, samara, or nut 31a. Foliage with abundant, aromatic, yellow or brown resin glands; plants dioecious . . . . . . . . . . . . . . . . . . . . . . . Myricaceae 31b. Foliage eglandular or with relatively nonaromatic glands; plants monoecious 32a. Ovary 3- or 6-locular; carpellate flowers solitary or in small clusters; fruit a nut [Figs. 658, 659] . . . . . . . Fagaceae 32b. Ovary 2-locular; carpellate flowers commonly in aments; fruit an achene, samara, or nut . . . . . . . Betulaceae 29b. Inflorescence not an ament; perianth present, evident 33a. Corolla of carpellate flowers (these usually with 8 sterile anthers) 15–20 mm long; perianth 4-merous; fruit a yellow-brown berry 20–40 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ebenaceae 33b. Corolla 1–15 mm long; perianth 4- to 10-merous; fruit a red, yellow, silver, or blue to black drupe or berry 4–15 mm long 34a. Pith diaphragmed; carpellate flowers epigynous; staminate flowers with 8–15 stamens . . . . (in part) Cornaceae 34b. Pith continuous; carpellate flowers hypogynous or perigynous; staminate flowers with 4–9 stamens

g ro u p 5   17

35a. Staminate flowers with 9 stamens; anthers dehiscing by uplifting valves; plants with aromatic foliage and wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lauraceae 35b. Staminate flowers with 4–8 stamens; anthers dehiscing by longitudinal slits; plants without aromatic foliage and wood 36a. Ovary with 2–4 stigmas, the style 2- to 4-lobed; stamens antepetalous . . . . . . . . . . . (in part) Rhamnaceae 36b. Ovary with 1 stigma, the style unbranched; stamens antesepalous 37a. Leaf blades and branchlets with abundant silver scales; plants sometimes armed with thorns; petals absent [Fig. 605]; styles elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Elaeagnaceae 37b. Leaves and branchlets not lepidote; plants unarmed; petals present; styles very short, nearly absent . . . . . . . . . . . . . . . . . . . . . (in part) Aquifoliaceae 25b. Flowers bisexual 38a. Leaf blades entire, with abundant silver scales; plants sometimes armed with thorns . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Elaeagnaceae 38b. Leaf blades entire, toothed, or lobed, not lepidote; plants unarmed or armed 39a. Corolla zygomorphic; fruit a legume . . . . . . . . . (in part) Fabaceae 39b. Corolla (or petaloid portions of flower) actinomorphic; fruit not a legume 40a. Corolla of 4 yellow, linear petals; plants flowering in the fall during and after the falling of the leaves . . . . . . . . Hamamelidaceae 40b. Corolla of 4–10 variously colored petals or petal-like members (petaloid hypanthium in Dirca), the petals not strap-shaped with parallel margins; plants flowering prior to leaf fall 41a. Fruit an elliptic to suborbicular samara (fruit unwinged in Ulmus parvifolia); flowers tiny, anemophilous, precocious (serotinous in Ulmus parvifolia); leaf blades double-serrate in most species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulmaceae 41b. Fruit not a samara; flowers variously pollinated, generally entomophilous, precocious to serotinous; leaf blades entire, serrate, or lobed 42a. Peduncle of the inflorescence adnate to a conspicuous, elongate bract; the numerous stamens commonly connate into 5 antepetalous groups . . . . . . . . . . . . . . (in part) Malvaceae 42b. Peduncle of the inflorescence without bracts or, when bracteate, free from the bract; stamens 4–many, distinct (shortly connate at the base in Halesia, but that species with 4-merous, gamopetalous flowers) 43a. Pith diaphragmed; gynoecium composed of a single carpel (very rarely 2 carpels) . . . . . . . . . . (in part) Cornaceae 43b. Pith continuous; gynoecium composed of 1–66 (–73) carpels

18  key to the fa mili es

44a. Gynoecium with (9–) 19–66 (–73) carpels [Fig. 52]; androecium with laminar filaments; fruit an aggregate of samaras or an agreggate of follicles that each bear a large seed covered by a pink to red or red-orange aril . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Magnoliaceae 44b. Gynoecium with 1–10 carpels; androecium usually with slender filaments; fruit otherwise, usually a drupe, berry, pome, achene, capsule, or aggregate (when an aggregate, as in some Rosaceae, usually with fleshy carpels) 45a. Perianth monochlamydeous 46a. Flowers with a conspicuous hypanthial tube, the petaloid members yellow or pink to purple (rarely white); stamens 8 . . . . . . . . Thymelaeaceae 46b. Flowers with a short, inconspicuous hypanthial tube, or lacking one altogether, petaloid members absent or white; stamens 4 or 5 47a. Stamens antepetalous; leaves commonly subopposite; branches often ending in a short, spine-like process . . . . . . . (in part) Rhamnaceae 47b. Stamens antesepalous; leaves alternate; branches without spines . . . . . . . . . . . . . . . . . . . . . . . (in part) Aquifoliaceae 45b. Perianth composed of 2 or more whorls, both the sepals and petals present 48a. Flowers perigynous or epigynous 49a. Stamens numbering more than the petals [Fig. 852] 50a. Stamens numbering more than 10, more than twice as many as the petals 51a. Stamens adnate to the base of the corolla; calyx light yellow, the lobes 1–1.5 mm long, ciliate on the margins; ovary epigynous; fruit a blue (rarely white) drupe 3–8 mm long . . . . . . Symplocaceae 51b. Stamens not inserted at the base of the corolla; calyx otherwise; ovary perigynous in flower (but becoming epigynous in fruit in some species); fruit not as above . . . . . . . . . (in part) Rosaceae 50b. Stamens numbering 10 or fewer, up to twice as many as the petals 52a. Stamens numbering 8, the filaments shortly connate at the base and distinct above; fruit 4-winged, dry, 1- to 3-seeded; flowers with 1 style . . . . . . . . Styracaceae 52b. Stamens 5–10, the filaments not monadelphous at the base; fruit otherwise; flowers with 2–5 styles . . . . . . . . . . . . . . . . . . . . . . (in part) Rosaceae

g ro u p 5   19

49b. Stamens as many as the petals [Fig. 676] 53a. Ovary inferior [Fig. 675] 54a. Plants with broad-based prickles along the stems and branches; inflorescence a corymb-like panicle of umbels . . . . . . . . . . . . . . (in part) Apiaceae 54b. Plants unarmed or with bristles or thin prickles; inflorescence a raceme or corymb (rarely a solitary flower) . . . . . . . . . . . . . . . . . . . . . . . Grossulariaceae 53b. Ovary superior (at least in flower) 55a. Plants usually armed with stout thorns; stamens antesepalous [Fig. 837] . . . . . . . . . . . . . . . . . . . . . . (in part) Rosaceae 55b. Plants unarmed or with a slender, spine-like process at the apex of some branches; stamens antepetalous . . . . . . . . . . . . . . . . . . . (in part) Rhamnaceae 48b. Flowers hypogynous 56a. Plants armed with spines [Fig. 479]; perianth composed of 6 sepals in 2 cycles and 6 petals in 2 cycles; anthers dehiscing by uplifting valves . . . . . . . . . . . . . . . . (in part) Berberidaceae 56b. Plants unarmed or with thorns in Lycium; perianth composed of fewer parts; anthers dehiscing by longitudinal slits or apical pores 57a. Sepals dimorphic, the 2 outer smaller than the 3 inner [Figs. 585, 586]; petals yellow or red in Lechea . . . . . . . . . . Cistaceae 57b. Sepals monomorphic; petals white, pink, red, purple, green, or green-white 58a. Stamens numbering the same as the petals and antepetalous, often enclosed by the petals (especially in bud), or the petals absent and the stamens alternating with the sepals . . . . (in part) Rhamnaceae 58b. Stamens fewer than, as many as, or more than the petals, alternating with the petals, not enclosed by the petals (recessed in pouches in the corolla in Kalmia) 59a. Stamens fewer than or as many as the petals or inner tepals 60a. Ovary 2-locular; fruit a red berry 10–20 mm in diameter; plants usually with thorns . . . . . . . . . . . . . . (in part) Solanaceae 60b. Ovary 3- to 8-locular; fruit otherwise; plants unarmed

20   ke y to the fami lie s

61a. Androecium of 4 stamens seated on a lobed, nectary disk; leaf blades 3–5 mm long, scalelike, subampexicaul; perianth 4-merous, with a corolla shorter than 2 mm . . . . . . . Tamaricaceae 61b. Androecium of 5–10 stamens (only 4 in some Ilex), not seated on a conspicuous nectar disk; leaf blades various, though usually longer and/or with a petiole; perianth mostly 5-merous, infrequently only 4-merous, the corolla longer than 2 mm 62a. Styles short, the stigma nearly sessile on the summit of the ovary; sepals present, though small and inconspicuous . . . . . . . (in part) Aquifoliaceae 62b. Styles elongate, the stigma elevated above the summit of the ovary; sepals usually evident . . . . . . . . . . . (in part) Ericaceae 59b. Stamens numbering more than the petals 63a. Plants evidently pubescent with compound hairs; ovary 3-locular; leaf blades with plane margins, deciduous . . . Clethraceae 63b. Plants glabrous or with simple hairs; ovary 2- to 9-locular; leaf blades sometimes with revolute margins and/or evergreen . . . . . . . . . . . . . . . . (in part) Ericaceae

Group 6 1a. Inflorescence a capitulum (the capitula with only 1 flower in Echinops); anthers of disk flowers connate in a ring around the style; fruit a cypsela . . . . . . . . . . . . . . (in part) Asteraceae 1b. Inflorescence not a capitulum (a capitulum-like cyme in some Caprifoliaceae); anthers not connate in a ring (except in Lobelia); fruit otherwise (except in some Caprifoliaceae) 2a. Flowers unisexual 3a. Plants vines, with tendrils; corolla gamopetalous [Fig. 600] . . . . . . . . . Cucurbitaceae 3b. Plants not vines, without tendrils; corolla apopetalous (very inconspicuous and reduced to a rim of sepals in Hippuris) 4a. Plants aquatic, with thin, flaccid, submersed leaves; flowers anemophilous, with an inconspicuous perianth

g ro u p 6   2 1

5a. Leaves pectinately divided into narrow segments or the leaves reduced to minute scales or bumps on the stem, mostly alternate or whorled [Figs. 678, 679, 680]; flowers with 4 or 8 stamens . . . . . . . . . . . . . . . . . . . . . . . (in part) Haloragaceae 5b. Leaves simple, whorled; flowers with a single stamen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Plantaginaceae 4b. Plants terrestrial, with thicker, firm leaves; flowers entomophilous, with a well-developed perianth 6a. Leaves simple; fruit a drupe; inflorescence an axillary cymule of 3 flowers— the 2 lateral flowers usually staminate, the central bisexual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Comandraceae 6b. Leaves compound or evidently lobed; fruit not a drupe; inflorescence otherwise 7a. Leaf blades 1- to 3-times pinnately compound; fruit an achene enclosed in an indurate hypanthium or a follicle . . . . . . . . . . . . . . . . . . . . . . (in part) Rosaceae 7b. Leaf blades palmately divided into 3–7 broad segments; fruit a schizocarp [Figs. 335, 336] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Apiaceae 2b. Flowers bisexual 8a. Stems very succulent, jointed, with flattened, oblong to suborbicular segments [Fig. 532], the areoles (i.e., modified nodes) with glochids and sometimes 1 or 2 spines; perianth consisting of numerous, spirally arranged tepals [Fig. 532] . . . . . . . . . Cactaceae 8b. Stems not consisting of succulent, pad-like segments, unarmed (mostly) or armed; perianth 2- to 6-merous 9a. Leaves usually 4–6 per node, entire, with arcuate venation; flowers pseudanthial, small, collectively subtended by 4 large, white, petaloid bracts . . . . (in part) Cornaceae 9b. Leaves 1–12 per node, entire, toothed, or lobed, without arcuate venation; flowers not pseudanthial, lacking petaloid bracts 10a. Calyx 3-merous, gamosepalous, petaloid . . . . . . . . (in part) Aristolochiaceae 10b. Calyx not 3-merous (except in some Galium), connate or not, sepaloid (petaloid sepals in Aizoaceae and petaloid tepals in Comandraceae), not red-brown 11a. Plants aquatic 12a. Plants with a rosette of floating leaves that have petioles swollen near midlength forming a float; hypanthium with 4 stout horns, these persistent on the fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lythraceae 12b. Plants without a conspicuous rosette of floating leaves, lacking specialized petioles with floats; hypanthium lacking horns 13a. Leaves with a prominent, sheathing base; inflorescence an umbel [Figs. 347, 348] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Apiaceae 13b. Leaves lacking a prominent, sheathing base; inflorescence of axillary flowers or a raceme 14a. Leaves compound, all basal, with 3 thick, fleshy leaflets [Fig. 724]; fruit a 2-valved capsule . . . . . . . . . . . . . . . . . . . . (in part) Menyanthaceae 14b. Leaves simple or compound, not confined to the base of a plant, not with 3 thick leaflets; fruit an achene-like drupe or a 4- or 5-valved capsule 15a. Leaves whorled, simple; flowers with a single stamen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Plantaginaceae 15b. Leaves alternate or opposite, simple, lobed, or compound; flowers with 3, 5, or 8 stamens

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16a. Perianth 4-merous; flowers with 4 stamens; fruit a 4-valved capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Onagraceae 16b. Perianth 3- or 5-merous; flowers with 3 or 5 stamens; fruit indehiscent or a 5-valved capsule 17a. Leaves pinnately lobed or pinnately compound (merely toothed in the emersed leaves of Proserpinaca palustris) [Figs. 677, 680, 681]; perianth composed of only 3 sepals; flowers with 3 antesepalous stamens . . . . . . . . . (in part) Haloragaceae 17b. Leaves simple and unlobed; perianth composed of 5 sepals and 5 petals; flowers with 5 antepetalous stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Theophrastaceae 11b. Plants of upland or wetland habitats, but typically not growing in standing water 18a. Inflorescence a capitulum-like cyme [Figs. 543, 544, 549]; ovary and fruit partially enclosed in a gamophyllous, tubular involucel that may be expanded apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Caprifoliaceae 18b. Inflorescence not resembling a capitulum; ovary and fruit not enclosed in a tubular involucel 19a. Gynoecium with 5–10 carpels; leaf blades conspicuously papillose, the papillae not requiring high magnification to see; fruit woody, subglobose, indehiscent, with 4–6 wings, each wing projecting into an apical horn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Aizoaceae 19b. Gynoecium with 2–6 carpels; leaf blades without easily viewed papillae; fruit otherwise 20a. Perianth 2- to 4-merous 21a. Leaf blades cordate-clasping the stem [Fig. 541]; ovary mostly trilocular; fruit usually a 3-valved capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Campanulaceae 21b. Leaf blades sessile or petiolate, but not clasping the stem; ovary bilocular or quadrilocular; fruit otherwise 22a. Corolla gamopetalous [Figs. 882, 883]; stamens 3 or 4; leaves opposite or whorled, stipulate (though the stipule sometimes reduced to an interpetiolar line) . . . . . . . . . . . . . (in part) Rubiaceae 22b. Corolla apopetalous above the hypanthium or absent in 1 species of Ludwigia [Figs. 738, 743]; stamens 2, 4, or usually 8; leaves alternate or opposite, exstipulate . . . . (in part) Onagraceae 20b. Perianth 5-merous or absent altogether (sepals numbering 2 in the Portulacaceae; petals rarely 4 or 6 in Portulaca) 23a. Leaves opposite; flowers with 3 or 4 stamens (5 in Triosteum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Caprifoliaceae 23b. Leaves alternate or all basal (often both alternate and opposite in Portulaca, a prostrate, fleshy-leaved herb); flowers with 5–many stamens 24a. Inflorescence an umbel [Figs. 347, 348]; fruit a schizocarp or, less commonly, a berry; leaves compound (rarely simple), usually with a sheathing petiole base; styles swollen at the base forming a stylopodium; calyx a series of small teeth around the rim of the receptacle or absent (rarely well-developed) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Apiaceae

g ro u p 6   23

24b. Inflorescence otherwise (except in Primula); fruit a capsule, pyxis, or drupe; leaves simple (trifoliate in Menyanthes), mostly without a sheathing petiole base; gynoecium without a stylopodium; calyx of evident sepals (or apparently absent in the Comandraceae) 25a. Leaves compound, with 3 thick, fleshy leaflets; petals conspicuously pubescent on the adaxial (i.e., inner) surface [Fig. 724] . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Menyanthaceae 25b. Leaves simple; petals glabrous on the adaxial surface (with a tuft of hairs near the base in Comandra) 26a. Flowers with ca. 20 stamens; herbage pubescent with barbed hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Loasaceae 26b. Flowers with 5–10 stamens; herbage glabrous or with hairs that lack barbs 27a. Flowers with mostly 6–10 stamens; petals yellow; prostrate weeds . . . . . . . . . . . . . . (in part) Portulacaceae 27b. Flowers with 5 stamens; petals (or tepals) white, red, green-purple, blue, or purple; plants not prostrate weeds 28a. Perianth monochlamydeous; fruit a drupe (dryish in Comandra); leaf blades ± entire . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Comandraceae 28b. Perianth dichlamydeous; fruit a capsule; leaf blades entire or toothed . . . (in part) Campanulaceae

Group 7 1a. Perianth with a nectary spur [Figs. 478, 708, 759] 2a. Plants insectivorous by means of bladders or viscid leaf blades [Figs. 708, 709, 711]; flowers with 2 stamens; placentation free-central . . . . . . . . . . . . . . . . . . . . . Lentibulariaceae 2b. Plants not insectivorous; flowers with 4–6 stamens; placentation axile or parietal 3a. Stamens numerous; both the calyx and corolla spurred, the 2 upper petals prolonged backward into a spur that extends into the upper, spurred sepal; leaf blades palmately divided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Ranunculaceae 3b. Stamens 4–12; either the calyx or corolla spurred, but not both (hypanthium with a spur in Cuphea); leaf blades not palmately divided (palmately lobed in some Viola) 4a. Corolla 6-merous; androecium with 11 or 12 stamens; flowers perigynous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lythraceae 4b. Corolla 3- to 5-merous; androecium with 4–8 stamens; flowers hypogynous 5a. Androecium with 8 stamens; leaf blades with peltate petioles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tropaeolaceae 5b. Androecium with 4–6 stamens; leaf blades with basifixed petioles 6a. Calyx spurred (note: the calyx is, in part, petaloid and can be misinterpreted as part of the corolla) [Fig. 478]; capsule dehiscing elastically; gynoecium with 5 carpels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Balsaminaceae

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6b. Corolla spurred [Figs. 757, 759]; capsule not dehiscing elastically; gynoecium with 2 or 3 carpels 7a. Calyx of 2, often caducous, sepals; flowers with 6 stamens; leaf blades decompound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Papaveraceae 7b. Calyx of 5 sepals; flowers with 4 or 5 stamens; leaf blades variously simple to lobed, but not many times dissected 8a. Corolla apopetalous [Fig. 931]; flowers with 5 stamens; gynoecium with 3 carpels, therefore, the capsule with 3 valves . . . . . . . . . . . Violaceae 8b. Corolla gamopetalous [Fig. 770]; flowers with only 4 functional stamens; gynoecium with 2 carpels, therefore, the capsule with 2 valves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Plantaginaceae 1b. Perianth lacking a nectary spur 9a. Upper sepal much larger than the other 4, petaloid, arched, forming a helmet-shaped structure that conceals the 2 petals; leaf blades palmately divided, with pinnately lobed segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Ranunculaceae 9b. Calyx without a single, larger, helmet-shaped sepal, sepaloid or petaloid; petals 3–8, not concealed by the calyx; leaf blades otherwise 10a. Calyx with 2 sepals; androecium of 3 stamens; fruit a capsule with 1–3 seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Portulacaceae 10b. Calyx with 4–6 sepals; androecium of 2–25 stamens, but not 3; fruit a capsule, silique, legume, schizocarp, or achene 11a. Fruit a legume or schizocarp; gynoecium with 1 carpel; stamens monadelphous or diadelphous (distinct in Baptisia, Chamaecrista, Senna, and Thermopsis); lower 2 petals fused into a keel (only the banner petal present in Amorpha, lower 2 petals distinct in Chamaecrista and Senna) [Figs. 638, 642, 657] . . . . . . . . (in part) Fabaceae 11b. Fruit a capsule, silique, schizocarp, or achene; gynoecium with 2 or more carpels; stamens distinct (monadelphous in Polygala); lower 2 petals variously distinct or connate 12a. Corolla apopetalous; androecium of 6–27 stamens 13a. Leaf blades pinnately compound with 7–13 leaflets, punctate with transparent glands; gynoecium with 5 carpels . . . . . . . . . . . . . (in part) Rutaceae 13b. Leaf blades entire to lobed or compound, when pinnately compound with only 3 leaflets, not glandular-punctate; gynoecium with 2–4 carpels 14a. Petals 4–6, the apical portion bearing a fringe of linear to oblanceolate segments that are ± equal to the length of the petal [Fig. 822]; gynoecium with 3 or 4 carpels; fruit a capsule; leaf blades entire to pinnately lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Resedaceae 14b. Petals 4, the apical portion lacking a fringe; gynoecium with 2 carpels; fruit a silique; leaf blades palmately compound or pinnately trifoliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cleomaceae 12b. Corolla gamopetalous; androecium of 2–8 stamens 15a. Calyx composed of 3 small sepals and 2 large, petaloid sepals [Figs. 783, 784]; flowers with 6 or 8 monadelphous stamens . . . . . . . . . . . . . . Polygalaceae 15b. Calyx composed mostly of 4 or 5 sepaloid sepals, but not as in the Polygalaceae; flowers with 2–5 distinct stamens 16a. Carpels 2, each carpel lobed, often evidently so, therefore, the gynoecium appearing to be composed of 4 carpels, separating at maturity into 4 half-carpellate segments; fruit a schizocarp

g ro u p 7   25

17a. Inflorescence a helicoid cyme [Figs. 490, 496, 503]; leaves alternate; stems terete or angled . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Boraginaceae 17b. Inflorescence not appearing as a coiled raceme [Figs. 695, 700, 927]; leaves opposite; stems usually quadrangular 18a. Style with a conspicuously lobed stigma; carpels weakly lobed; inflorescence indeterminate (e.g., spike, raceme) [Figs. 926, 927]; placentation marginal; plants without aromatic foliage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Verbenaceae 18b. Style usually with an inconspicuous stigma; carpels usually evidently lobed; inflorescence mixed (i.e., thyrsoid), with cymose lateral branches, sometimes the inflorescence condensed in dense verticillasters of flowers, only rarely completely indeterminate [Figs. 695, 700, 701]; ovules attached to lateral partitions and not the carpel margins; plants commonly with mint-, pennyroyal-, or citrus-scented foliage . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lamiaceae 16b. Carpels 2, each carpel unlobed, therefore, the gynoecium appearing to be composed of 2 carpels (appearing unicarpellate in Phryma, but the stigmas 2); fruit a capsule or achene 19a. Fruit an achene; ovary unilocular and uniovulate; calyx tightly reflexed in fruit [Fig. 764] . . . . . . . . . . . . . . . . . . . . . . . . (in part) Phrymaceae 19b. Fruit a capsule; ovary bilocular or unilocular in holoparasitic genera, with 2 or more ovules; calyx not reflexed in fruit 20a. Fruit a drupe-like capsule 10–20 cm long, with a fleshy exocarp that separates from the woody endocarp, prolonged at the apex into a curved beak that splits into 2 halves at maturity; plants pubescent with stipitate glands, the gland composed of 4 or more mucilage-filled cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pedaliaceae 20b. Fruit a capsule without a fleshy exocarp, not terminated by a long, 2-pronged beak; plants stipitate-glandular or not, but never slimy-pubescent 21a. Anthers with 1 locule, dehiscing by a single, transverse, distal slit, not sagittate at the base . . . . . . . . . . (in part) Scrophulariaceae 21b. Anthers with 2 locules, dehiscing by 2 longitudinal slits or the locules connate near the apex and opening by a distal, V-shaped slit, usually sagittate at the base 22a. Plants holoparasitic, lacking chlorophyll; ovary unilocular, with parietal placentation . . . . . . . . . . . . (in part) Orobanchaceae 22b. Plants autotrophic or hemiparasitic, with chlorophyll; ovary bilocular, with axile placentation 23a. Hemiparasitic plants with orobanchin (i.e., usually darkening in drying); corolla galeate (without a galea in Agalinis and Aureolaria) [Figs. 753, 754] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Orobanchaceae 23b. Autotrophic herbs, usually not darkening in drying; corolla not galeate (with a galea in Chelone) 24a. Androecium with 2 stamens, each stamen with 2 pollen sacs positioned perpendicular to one another—the upper pollen sac positioned transversely to the filament, the lower 1 positioned parallel to the filament; fruit an explosively dehiscent, loculicidal capsule . . . Acanthaceae

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24b. Androecium with 1–4 stamens, the stamens with parallel to somewhat divergent pollen sacs (with respect to each other); fruit a loculicidal or septicidal capsule, but not explosively dehiscent 25a. Calyx with a pronounced, tubular, connate portion [Fig. 763]; capsule dehiscence loculicidal; stigma with 2 plate-like lobes that close after contact . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Phrymaceae 25b. Calyx connate at the base, but the connate portion not elongate; capsule dehiscence septicidal; stigmas with 2 lobes that remain open after contact (these platelike in Gratiola) 26a. Lower pair of stamens (which are sterile in our species) with the filament axis bent completely back on themselves and then curved again, therefore, the filaments appearing forked . . . . . . . . . . Linderniaceae 26b. Lower pair of stamens without a conspicuously curved filament axis 27a. Amphibious plants of tidal shores with horizontal stems emitting at the nodes clusters of linear leaves and sometimes also peduncles bearing a solitary flower with a ± actinomorphic corolla . . . . . . . . . . . . . . . (in part) Scrophulariaceae 27b. Terrestrial or non-tidal wetland plants with prostrate to, more commonly, upright stems and variable leaf shapes, though mostly wider than linear, with inflorescences borne on a stem bearing flowers with weakly to conspicuously zygomorphic corollas . . . . . . . . . . . . . . . . (in part) Plantaginaceae

Group 8 1a. Leaves peltate [Figs. 55, 731]; plants aquatic 2a. Leaf blades orbicular, without basal lobes [Fig. 731]; carpels individually sunken in the spongy receptacle; fruit a nut, loosely borne in cavities of the firm, accrescent receptacle [Fig. 731] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nelumbonaceae 2b. Leaf blades elliptic, with basal lobes [Fig. 55]; carpels free from the receptacle; fruit a leathery follicle, not enclosed by the receptacle . . . . . . . . . . . . . . . . (in part) Nymphaeaceae 1b. Petiole, if present, attached at the base of the leaf blade; plants terrestrial or sometimes aquatic in Caltha, Comarum, Mentha, Ranunculus, and Saururus 3a. Inflorescence a cylindrical, usually nodding spike produced on a peduncle opposite the leaves with 175–300 flowers that lack perianth and have 3–5 (usually 4) carpels that are basally connate and mature as rugose mericarps . . . . . . . . . . . . . . . . . . (in part) Saururaceae 3b. Inflorescence not combinging the above characteristics, usually of a different type (e.g., cyme, raceme), with fewer flowers, and/or with perianth 4a. Corolla gamopetalous; carpels 2, each carpel evidently lobed, therefore, the gynoecium appearing to be composed of 4 carpels, separating at maturity into 4 halfcarpellate segments

g ro u p 8   27

5a. Inflorescence a helicoid cyme [Figs. 490, 496, 503]; leaves alternate (opposite in Cryptantha and Plagiobothrys); stems terete or angled . . . . . . (in part) Boraginaceae 5b. Inflorescence mixed (i.e., thyrsoid), with cymose lateral branches, sometimes the inflorescence condensed in dense verticillasters of flowers, only rarely completely indeterminate [Figs. 695, 700, 701]; leaves opposite; stems usually square . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lamiaceae 4b. Corolla apopetalous or absent; carpels 1–many, not schizocarpic (except Malvaceae) 6a. Stamens monadelphous [Fig. 720]; anthers unilocular; fruit a schizocarp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Malvaceae 6b. Stamens distinct; anthers bilocular; fruit otherwise 7a. Flowers lacking a hypanthium 8a. Plants ± succulent with fleshy, simple leaf blades; carpels with a small, scale-like, nectariferous appendage at the base; stamens usually 10 or fewer per flower (up to 80 in Sempervivum); fruit a follicle . . . . . . . . . . . Crassulaceae 8b. Plants not succulent, with thinner, simple, lobed, or compound leaf blades; carpels without a scale-like, nectariferous appendage; stamens usually many per flower; fruit an achene, berry, or follicle 9a. Intrastaminal nectar disk present; calyx sepaloid, with coriaceous, persistent sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paeoniaceae 9b. Intrastaminal nectar disk absent; calyx sepaloid or petaloid, usually membranaceous to herbaceous and deciduous . . . . (in part) Ranunculaceae 7b. Flowers with a hypanthium 10a. Leaves with simple blades, without stipules; flowers with 2 or 4 carpels; fruit a capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Saxifragaceae 10b. Leaves with simple to compound blades, stipulate (without stipules in Aruncus); flowers with 2–many carpels; fruit an achene, drupe, or follicle, either solitary or in an aggregate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Rosaceae

Group 9 1a. Plants parasitic, with haustoria, lacking chlorophyll; stems twining, pink-yellow to orange, with scale-like leaves [Figs. 589, 591] . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Convolvulaceae 1b. Plants autotrophic, without haustoria, with chlorophyll (without chlorophyll in Pterospora); stems, when present, upright or twining, usually green for some period during the growing season, with foliaceous leaves (scale-like in Bartonia and Pterospora) 2a. Depressed, pulvinate [Fig. 603], evergreen, alpine herbs of elevation exceeding 1000 meters; inflorescence a solitary, peduncled flower . . . . . . . . . . . . . . . (in part) Diapensiaceae 2b. Plants of various habit, but commonly upright or vining, with deciduous leaves, of lower elevation habitats; inflorescence various, but commonly composed of more than 1 flower 3a. Leaves all basal (opposite and borne on a stem in Plantago arenaria); calyx and corolla inconspicuous, the corolla scarious, 3- or, more commonly, 4-merous [Figs. 771, 775] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Plantaginaceae 3b. Leaves borne on a stem (all basal in Limosella, Menyanthes, and Primula); calyx and/or corolla evident, at least 1 of which is petaloid, usually 4- or 5-merous

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4a. Plants often with milky latex; carpels connate at the summit only, with a common stigma; fruit a pair of follicles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apocynaceae 4b. Plants with watery sap; carpels connate throughout or only in the basal portion; fruit an achene, berry, schizocarp, capsule, or pyxis 5a. Leaves with sheathing stipules (i.e., ocreae) [Fig. 793]; pedicels of the flowers subtended by sheathing bracteoles (i.e., ocreolae); perianth mostly monochlamydeous, composed of ± petaloid tepals; fruit an achene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Polygonaceae 5b. Leaves without sheathing stipules; pedicels without sheathing bracteoles; perianth dichlamydeous; fruit otherwise 6a. Stems prostrate, emitting at nodes a tuft of 5–10 narrow leaves and 1 or more, 1-flowered peduncles; plants of tidal shores . . . . . . (in part) Phrymaceae 6b. Stems upright or vining, the leaves normally 1 or 2 from a node, sometimes more; inflorescence various, usually with more than 1 flower; plants not of tidal shores (of coastal shorelines in Mertensia maritima and Heliotropium) 7a. Plants aquatic, with densely crowded, dissected leaves; inflorescence an umbel or raceme, the peduncle and axis of each individual raceme inflated and hollow, constricted at the nodes and bearing a whorl of flowers [Fig. 806] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Primulaceae 7b. Plants aquatic or terrestrial, the leaves not both densely crowded and dissected; inflorescence without an inflated peduncle and axis 8a. Foliage leaves opposite or whorled 9a. Carpels 2, each carpel lobed, often evidently so, therefore, the gynoecium appearing to be composed of 4 carpels, separating at maturity into 4 half-carpellate segments; fruit a schizocarp; flowers with 4 stamens (5 in Boraginaceae); stems quadrangular (terete or angled in Boraginaceae) 10a. Inflorescence a helicoid cyme [Figs. 490, 496, 503]; flowers with 5 stamens; stems terete or angled . . . . . (in part) Boraginaceae 10b. Inflorescence not appearing as a coiled raceme [Figs. 695, 700, 927]; flowers with 4 stamens; stems quadrangular 11a. Style usually with an inconspicuous stigma; carpels evidently lobed; inflorescence mixed (i.e., thyrsoid), with cymose lateral branches, these usually arranged in dense verticillasters of flowers, ovules attached to lateral partitions and not the carpel margins [Figs. 926, 927]; plants with mint-, pennyroyal-, or citrusscented foliage . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lamiaceae 11b. Style with a conspicuously lobed stigma; carpels weakly lobed; inflorescence indeterminate (e.g., spike, raceme) [Figs. 926, 927]; placentation marginal; plants without aromatic foliage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Verbenaceae 9b. Carpels 1–3, not separating at maturity; fruit a capsule or pyxis; flowers with 4–7 stamens; stems mostly terete or angled 12a. Stamens antepetalous [Fig. 729]; ovary with 1 stigma; placentation free-central . . . . . . . . . . . . . . . . . . . (in part) Myrsinaceae 12b. Stamens antesepalous; ovary usually with a 2- or 3-lobed stigma; placentation parietal or axile 13a. Gynoecium with 3 carpels and 3 stigmas; placentation axile; calyx with scarious intervals between the green, longitudinal ribs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Polemoniaceae

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13b. Gynoecium with 2 carpels and 2 stigmas; placentation parietal; calyx without scarious intervals between longitudinal ribs 14a. Leaf blades entire; capsule dehiscence septicidal; surface of vegetative organs ± glabrous (though the margins sometimes with cilia) . . . . . . . . . . . . . . . . (in part) Gentianaceae 14b. Leaf blades pinnately lobed; capsule dehiscence loculicidal; surface of at least some vegetative parts pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Boraginaceae 8b. Foliage leaves alternate or all basal (appearing opposite in some Solanaceae due to forking of the stem) 15a. Inflorescence a helicoid cyme [Figs. 490, 496, 503]; fruit a schizocarp (a 2-valved capsule in Hydrophyllum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Boraginaceae 15b. Inflorescence otherwise; fruit a capsule, pyxis, or berry (the berry sometimes dry in the Solanaceae) 16a. Flowers with 10 stamens and 5 carpels; stipitate-glandular, mycoparasitic herbs lacking chlorophyll . . . . . . . . (in part) Ericaceae 16b. Flowers with 4 or 5 stamens and 2 or 3 carpels; autotrophic herbs (except Bartonia), with or without stipitate glands 17a. Leaves reduced to scales on the stem [Fig. 664]; flowers 2.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Gentianaceae 17b. Leaves foliaceous, not scale-like; flowers longer 18a. Plants aquatic; leaves either floating or with 3 fleshy leaflets [Figs. 724, 725] . . . . . . . . . . . . . (in part) Menyanthaceae 18b. Plants terrestrial; the leaves neither floating nor with 3 fleshy leaflets 19a. Stamens antepetalous; placentation free-central 20a. Leaves confined to the base of the stem; corolla ± salverform . . . . . . . . . . . . . . . . . . . . (in part) Primulaceae 20b. Leaves borne on the stem; corolla ± rotate [Fig. 729] . . . . . . . . . . . . . . . . . . . . . . . (in part) Myrsinaceae 19b. Stamens antesepalous; placentation basal, parietal, or axile 21a. Ovary with 4 or 6 ovules; fruit a 4- or 6-seeded capsule; plants climbing or trailing (twining only at the tip in Calystegia spithamaea) . . . . . (in part) Convolvulaceae 21b. Ovary usually with more than 6 ovules; fruit usually a many-seeded capsule, berry, or pyxis; plants upright (climbing or trailing in Solanum dulcamara) 22a. Gynoecium composed of 3 carpels (2 in Navarretia); calyx with scarious intervals between the green, longitudinal ribs (calyx of uniform texture in Collomia); fruit a 3-valved capsule (2-valved in Navarretia) . . . . . . . . . . . . . . . (in part) Polemoniaceae 22b. Gynoecium composed of 2 carpels (3–5 in Nicandra); calyx of ± uniform texture; fruit a berry, 2-valved capsule, or pyxis

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23a. Stamens each flanked by a pair of small scales; fruit a loculicidal capsule dehiscing by 2 valves . . . . . . . . . . . . . . . . . . (in part) Boraginaceae 23b. Stamens without associated scales; fruit usually a septicidal capsule opening by 2–4 valves, berry, or pyxis 24a. Fruit an unarmed capsule; anthers with 1 locule, opening by a single, transverse, distal slit; inflorescence a raceme or panicle . . . . . . . . . . . . . . . . . . . (in part) Scrophulariaceae 24b. Fruit a berry, prickly capsule, or pyxis; anthers with 2 locules, opening by a pair of pores or longitudinal slits; inflorescence of solitary flowers in the axils of leaves or cymose (note: the cymes sometimes appear racemeor panicle-like) . . . . . . . . . . (in part) Solanaceae

Group 10 1a. Plants submerged aquatics, attaching to the substrate by fleshy disks; flowers without perianth, borne singly in the axils of leaves, subtended by a tubular spathe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podostemaceae 1b. Plants aquatic or terrestrial, but not attaching to the substrate by fleshy disks; flowers with at least 1 whorl of perianth (sometimes so in only 1 sex of flower; lacking perianth in Callitriche, Euphorbia, and Saururus), borne variously, not subtended by a tubular spathe 2a. Inflorescence a cylindrical, usually nodding spike produced on a peduncle opposite the leaves with 175–300 flowers that lack perianth and have 3–5 (usually 4) carpels that are basally connate and mature as rugose mericarps . . . . . . . . . . . . . . . . . . (in part) Saururaceae 2b. Inflorescence not combinging the above characteristics, usually of a different type (e.g., cyme, raceme), with fewer flowers, and/or with perianth 3a. Placentation laminar; plants aquatic, with conspicuous elliptic or orbicular floating leaf blades with 2 basal lobes [Fig. 55], arising directly from a rhizome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Nymphaeaceae 3b. Placentation otherwise; plants terrestrial or aquatic; leaf blades various, but not at once floating, basally lobed, and arising from an underwater rhizome 4a. Inflorescence a cyathium, with a single carpellate flower and 2 or more staminate flowers borne in a cupuliform involucre, the margin of the involucre sometimes subtended by petaloid glands, the entire arrangement resembling a single flower; plants often with milky latex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Euphorbiaceae 4b. Inflorescence not pseudanthial; plants with watery sap (milky or colored latex in the Papaveraceae) 5a. Calyx of 2, often caducous, sepals; petals wrinkled in bud; plants with milky or colored latex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Papaveraceae 5b. Calyx of 3 or more sepals (2 sepals in the Portulacaceae and some Elatinaceae), or the perianth monochlamydeous and with 3 or more tepals, or the perianth absent; petals arranged otherwise in bud (wrinkled in the Lythraceae); plants with watery sap 6a. Plants insectivorous by means of viscid hairs or pitcher-shaped leaves [Figs. 604, 902], of organic soils; leaves all basal

g ro u p 10   3 1

7a. Leaf blades with viscid, glandular-hairs, flat, the margins not connate; inflorescence a raceme; styles 3, distinct . . . . . . . . . . . . . . . . . . . . . Droseraceae 7b. Leaf blades without glandular hairs, the margins connate, thereby forming a tube that is retrorsely pubescent on the adaxial (i.e., inside) surface [Fig. 902]; inflorescence a single flower; styles 5, connate, modified into a 5-rayed, umbrella-shaped body . . . . . . . . . . . . . . . . . . . . . . . . . . Sarraceniaceae 6b. Plants not insectivorous, of mineral or organic soils; leaves variously arranged 8a. Leaves opposite or whorled (sometimes the upper alternate in Chrysosplenium) [Figs. 628, 685, 768] 9a. Leaves parapinnate, 1 of each node conspicuously larger than the other; fruit a schizocarp, each mericarp with 2 divergent horns and a central row of tubercles and transversely divided into 3–5 1-seeded compartments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zygophyllaceae 9b. Leaves simple, lobed, or compound, when pinnately compound then imparapinnate, those of a given node ± equal in size; fruit an achene, berry, pyxis, silique, silicle, capsule, or a schizocarp but then lacking horns and tubercles and not transversely divided into compartments 10a. Plants free-floating, rootless, aquatic; leaves 2- to 4-times dichotomously forked [Fig. 51]; perianth composed of 7–15 basally connate sepals; fruit an achene with 2–15 marginal spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratophyllaceae 10b. Plants usually rooted, terrestrial or aquatic; leaves not dichotomously forked; perianth otherwise (consisting of up to 10 basally connate sepals in some Euphorbiaceae); fruit various, but not an achene with marginal spines 11a. Basal leaves peltate, 30–40 cm wide; fruit a yellow berry 40–50 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Berberidaceae 11b. None of the leaves peltate, narrower; fruit an achene, capsule, pyxis, silique, or schizocarp 12a. Leaf blades punctate with pellucid dots, sessile; petals yellow or flesh-colored to pink . . . . . . . . . . (in part) Hypericaceae 12b. Leaf blades not pellucid-punctate, sessile or petiolate; petals variously colored, usually not yellow or flesh-colored (yellow in 1 species of Portulaca and some Linum) or absent 13a. Leaf blades scale-like and sessile; flowers sunken into the fleshy, jointed stem [Fig. 340]; plants coastal halophytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Amaranthaceae 13b. Leaf blades foliaceous (though they may be small), sessile or petiolate; flowers not sunken into a fleshy, jointed stem; plants of various habitats 14a. Plants aquatic (though sometimes stranded after water level decline); androecium with 1–3 stamens 15a. Leaves crowded at the apex forming a floating rosette (not forming a rosette in 2 species) [Fig. 766]; flowers unisexual, without perianth; fruit a schizocarp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Plantaginaceae 15b. Leaves not crowded at the apex; flowers bisexual, with sepals and petals; fruit an achene . . . . Elatinaceae 14b. Plants terrestrial or of wetland habitats; androecium with 4–10 stamens

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16a. Perianth monochlamydeous 17a. Perianth petaloid 18a. Leaves whorled, 3–8 per node; placentation axile . . . . . . . . . . . . . . . . . . . . (in part) Molluginaceae 18b. Leaves opposite; placentation basal or freecentral (axile in Sesuvium) 19a. Sepals with subterminal, slender, horn-like appendages; fruit a pyxis . . . . (in part) Aizoaceae 19b. Sepals lacking subterminal appendages; fruit an achene or capsule 20a. Flowers in a cyme, with a sepaloid involucre subtending 1–5 flowers and a petaloid calyx [Fig. 732]; fruit an achene, enclosed in an indurate calyx (collectively called an anthocarp); cultivated and escaped plants . . . . . . . . . . . . Nyctaginaceae 20b. Flowers axillary, with only a petaloid calyx; fruit a 5-valved capsule; native plants of coastal saltmarshes . . . . . . . . . . . . . . . . . . . . . (in part) Myrsinaceae 17b. Perianth sepaloid or inconspicuous 21a. Flowers with 7–20 or more stamens; style branches bifid or laciniately divided; fruit a capsule-like schizocarp with 2 or 3 mericarps . . . . . . . . . . . . . . . . . . . . . . . . (in part) Euphorbiaceae 21b. Flowers with 1–8 stamens; style branches simple (though the stigmatic surface may be fringed in some fashion); fruit a 2- to 10-valved capsule or utricle 22a. Stamens 4–8, set in the notches of an 8-lobed disk that fills much of the center of the flower; fruit a 2-valved capsule . . . . . . . . . . . . . . . . . . . . . . (in part) Saxifragaceae 22b. Stamens 1–5, not set on a disk; fruit an achene or a 3- to 10-valved capsule 23a. Flowers unisexual; fruit an achene 24a. Ovary with 2 styles; leaf blades palmately lobed or divided; staminate flowers each with 5 stamens . . . . . . . . . . . . . . . . . . (in part) Cannabaceae 24b. Ovary with 1 style; leaf blades simple, without lobes or divisions; staminate flowers each with 4 stamens or 5 in Laportea . . . . . . . . . . . . (in part) Urticaceae 23b. Flowers bisexual; fruit a capsule (a utricle in Scleranthus) 25a. Ovary with 3–5 locules; placentation axile; leaves whorled . . . . . . . . . . . . . . . . . (in part) Molluginaceae

g ro u p 10   3 3

25b. Ovary with a single locule; placentation free-central; leaves opposite . . . . . . . . . . . . . . (in part) Caryophyllaceae 16b. Perianth dichlamydeous 26a. Calyx of 2 sepals [Fig. 805]; stamens antepetalous . . . . . . . . . . . . . . . (in part) Portulacaceae 26b. Calyx of 3 or more sepals [Figs. 556, 572, 712]; stamens antesepalous 27a. Flowers 4-merous [Figs. 504, 527] 28a. Leaf blades palmately lobed or palmately divided; androecium composed of 6 tetradynamous stamens; fruit a silique or silicle . . . . . . . . . . . . . . . (in part) Brassicaceae 28b. Leaf blades simple; androecium composed of 4 or 8 stamens; fruit a capsule 29a. Anthers dehiscing by a single, terminal pore; filaments twisted, thereby bringing all the stamens to 1 side of the flower at anthesis; leaf blades 20–70 mm long . . . . . . . . . . . . . . . . . . . . . . . Melastomataceae 29b. Anthers dehiscing by longitudinal slits; filaments not twisted to 1 side; leaf blades 2–23 (–30) mm long 30a. Sepals (1–) 1.5–3 mm long, entire at the apex; leaf blades linearsubulate; capsules 2–4 mm long . . . . . . . . . . . . . . (in part) Caryophyllaceae 30b. Sepals 0.7– 1 mm long, 3-cleft at the apex; leaf blades ovate to oblong; capsules 0.6–0.9 mm long . . . . . . (in part) Linaceae 27b. Flowers 3-, 5-, or 6-merous [Figs. 557, 611, 671] 31a. Anthers dehiscing by 2 terminal pores; stigmas large, peltate, orbicular, with 4 or 5 lobes . . . . . . . . . . . . . . . . . . . . (in part) Ericaceae 31b. Anthers dehiscing by longitudinal slits; stigmas smaller, linear to capitate, without lobes 32a. Style branches bifid or laciniately divided [Fig. 625]; fruit a capsule-like schizocarp with 3 mericarps . . . . . . . . . . . . . . . . . . (in part) Euphorbiaceae 32b. Style branches simple (though the stigmatic surface may be fringed in some fashion); fruit a 2- to 10-valved capsule or a schizocarp separating into 5 mericarps 33a. Leaf blades palmately lobed or pinnately divided; fruit a schizocarp, with 5 1-seeded mericarps that elastically dehisce from a persistent, central column [Fig. 672] . . . . . . . . . . . . . . . . . . Geraniaceae

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33b. Leaf blades simple and unlobed; fruit a capsule 34a. Flowers perigynous, with intersepalar appendages; ovary 2-locular . . . . . . . . (in part) Lythraceae 34b. Flowers hypogynous (perigynous in Scleranthus), without intersepalar appendages; ovary 1- or 10-locular 35a. Capsule with 10 locules; placentation apical-axile; stamens alternating with small staminodes in some species; calyx aposepalous . . . . . . . . . . . . . . . . . (in part) Linaceae 35b. Capsule with a single locule; placentation free-central; androecium without staminodes; calyx apo- or gamosepalous 36a. Androecium with 6 stamens in 2 whorls, the outer whorl shorter than the inner; flowers sessile; placentation parietal . . . . . . . . . . . . . . . . . Frankeniaceae 36b. Androecium with 1–10 stamens in 1 or 2 whorls, when in 2 whorls the stamens not dissimilar as to length; flowers pedicellate (sessile in Herniaria and Scleranthus); placentation free-cental . . . . . . (in part) Caryophyllaceae 8b. Leaves alternate or all basal [Figs. 336, 715, 790] 37a. Leaf blades compound, trifoliate, with obcordate leaflets [Fig. 756]; petals white with pink stripes or yellow. . . . . . . . . . . . . . . . . . . . Oxalidaceae 37b. Leaf blades simple to compound, but not as above; petals variously colored or absent 38a. Flowers with 4 petals (very rarely 0 or 2); androecium with 6 stamens arranged in tetradynamous fashion (rarely only 2 stamens, numbering 6 but not tretradynamous in Cleome); fruit a silique or silicle 39a. Flowers, except sometimes for the lower ones, ebracteate [Figs. 504, 525, 531]; stamens usually 6 (rarely only 2), the outer pair shorter than the other 4; silique or silicle usually divided into 2 locules by an unvascularized septum . . . . . (in part) Brassicaceae 39b. Flowers subtended by bracts; stamens 6, all elongate; silique unilocular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cleomaceae 38b. Flower with 0–6 petals; androecium with a variable number of stamens, but never 6 and tetradynamous; fruit otherwise 40a. Stamens monadelphous, the filaments forming a conspicuous tube surrounding the style [Fig. 720]; gynoecium composed of 5–40 carpels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Malvaceae

g ro u p 10   3 5

40b. Stamens distinct or connate at only the very base; gynoecium composed of 1–6 (–7) carpels 41a. Leaves with tubular, sheathing stipules (i.e., ocreae); flowers subtended by tubular, sheathing bracteoles (i.e., ocreolae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Polygonaceae 41b. Leaves without tubular, sheathing stipules; flowers not subtended by tubular, sheathing bracteoles 42a. Leaf blades compound 43a. Perianth 3-merous, the sepals not petaloid; flowers solitary on thin peduncles from the axils of leaves; fruit a schizocarp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnanthaceae 43b. Perianth (3–) 4- to 6-merous, the sepals petaloid; flowers borne in cymes or racemes, or borne singly at the summit of the stem or tips of branches, or singly on scapes; fruit a blue seed or an achene, berry, follicle, or capsule 44a. Anthers dehiscing by flaps; stamens as many as the petals and opposite them (i.e., antepetalous); fruit an exposed, blue seed or a capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Berberidaceae 44b. Anthers dehiscing by longitudinal slits; stamens numbering more than the petals; fruit an achene, berry, or follicle 45a. Leaf blades 2- or 3-times divided or dissected into numerous, very narrow segments; fruit a follicle or berry; sepals caducous or not . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Ranunculaceae 45b. Leaf blades 1-time divided ; fruit an achene enclosed in an indurate hypanthium or a follicle; sepals not caducous. . . . . . . . . . . . . (in part) Rosaceae 42b. Leaf blades simple, toothed, or lobed but not divided 46a. Perianth sepaloid 47a. Stamens 10 per flower; fruit a 5- to 7-lobed and locular capsule [Fig. 762] . . . . . . . . . . . . . . Penthoraceae 47b. Stamens 1–5 per flower (as many as 20 or more in some Euphorbiaceae); fruit an achene, utricle, pyxis, berry, multiple, or 3-locular capsule or capsule-like schizocarp 48a. Flowers perigynous, bisexual; leaf blades palmately lobed . . . . . . . . . . . . . . . . . (in part) Rosaceae 48b. Flowers hypogynous, unisexual or sometimes bisexual in the Amaranthaceae; leaf blades simple to pinnately lobed (palmately lobed in Ricinus, which has a peltate attachment of the petiole) 49a. Filaments stout, nearly as wide as the anthers, white, primarily responsible for the flower color; leaf blades thick and evergreen . . . . . . . . Buxaceae 49b. Filaments more slender, not as wide as the anthers, not imparting an evident white color to flowers; leaf blades thinner and deciduous

36   k ey to the fa mi lie s

50a. Ovary 3-locular (1-locular in Crotonopsis); styles 2 or 3, each 1 bifid [Fig. 625]; fruit a capsule-like schizocarp with 2 or 3 mericarps (an achene in Crotonopsis) . . . . . . . . . . . . . . . . . . . . . (in part) Euphorbiaceae 50b. Ovary 1-locular; styles 1–3, without additional branching (though the stigmatic surface may be fringed in some fashion); fruit an achene, utricle, multiple, or pyxis 51a. Sepals usually dry and scarious; filaments distinct or, more commonly, connate at least near the base; ovary with 3 stigmas . . . . . . . . (in part) Amaranthaceae 51b. Sepals membranaceous to herbaceous (hyaline in Axyris); filaments distinct; ovary with usually 1 or 2 stigmas 52a. Proximal portion of first pair of primary lateral veins at the very margin of the leaf blade (i.e., the basiscopic side of the vein exposed for a short distance); stems with minute, uncinate hairs; fruit a multiple of achenes . . . . (in part) Moraceae 52b. First pair of primary lateral veins not at the margin of the leaf blade (i.e., both the basiscopic and acroscopic sides bordered by green leaf tissue) or the leaf blades with 1 nerve and lateral veins lacking; stems variously glabrous to pubescent, but lacking uncinate hairs; fruit an achene, utricle, or pyxis 53a. Ovary with 1 stigma; flowers unisexual; plants sometimes with stinging hairs . . . . . (in part) Urticaceae 53b. Ovary with 2 (–5) stigmas; flowers unisexual or bisexual; plants without stinging hairs . . . . . . . . . . . . (in part) Amaranthaceae 46b. Perianth, at least in part, petaloid (though caducous in Hydrastis) 54a. Plants lacking chlorophyll, the stems white to pink, yellow to brown, or pink to red; leaves reduced and scale-like [Fig. 616] . . . . . . . . . . . . . (in part) Ericaceae 54b. Plants with chlorophyll, the stems and/or leaves green; leaves foliaceous 55a. Anthers dehiscing by 2 apical pores; leaves all basal; fruit a 5-valved capsule . . . . (in part) Ericaceae 55b. Anthers dehiscing by longitudinal slits; leaves and fruits various 56a. Gynoecium with 5–12 (–15) styles; fruit a berry 57a. Flowers with 5 sepals and consistently with 10 styles, arranged in a raceme; berry dark purple; leaf blades unlobed . . . . . . . . . . . Phytolaccaceae

g ro u p 10   37

57b. Flowers with 3 caducous sepals and with a variable number of styles, solitary at the summit of the stem; berry red; leaf blades palmately lobed . . . . . (in part) Ranunculaceae 56b. Gynoecium with 2–7 styles; fruit a capsule, pyxis, or utricle 58a. Flowers perigynous 59a. Perianth 6-merous; sepals alternating with appendages at the rim of the hypanthium; petals crumpled in bud . . . . . . . . . . . . . . . . . . . . . . (in part) Lythraceae 59b. Perianth 5-merous; sepals not alternating with appendages; petals mostly imbricate in bud, not crumpled 60a. Androecium with 5 stamens and 5 trifid, white staminodes; petals 10–18 mm long; flowers with 4 carpels; fruit a 4-locular capsule . . . . . . . . . . . . . . . . . (in part) Parnassiaceae 60b. Androecium with 10 stamens; petals 2–6 mm long; flowers with 2 carpels; fruit a follicle or a 1- or 2-locular capsule . . . . . . . . . (in part) Saxifragaceae 58b. Flowers hypogynous 61a. Calyx definitely gamosepalous; halophytic plants of Atlantic coast shores . . . . . . . . . . . . . . . . . . . . . . . . . Plumbaginaceae 61b. Calyx aposepalous or nearly so, connate at only the very base, if at all; plants not halophytic 62a. Calyx composed of 2 sepals; stems prostrate; fruit a pyxis . . . . . . . . . . . . . . . . . (in part) Portulacaceae 62b. Calyx composed of 5 sepals; stems upright; fruit a capsule 63a. Gynoecium of 5 carpels; capsule 10-valved; leaves borne on a stem . . . . . . . . . . . . . . . . . . . . (in part) Linaceae 63b. Gynoecium of 2 or 4 carpels; capsule 2- or 4-valved; leaves all basal or essentially so 64a. Androecium with 5 stamens and 5 trifid, white staminodes; petals 10–18 mm long; flowers with 4 carpels; fruit a 4-locular capsule . . . . . . . . . . . (in part) Parnassiaceae 64b. Androecium with 10 stamens; petals 2–6 mm long; flowers with 2 carpels; fruit a follicle or a 1- or 2-locular capsule . . . . . . . . . . . . (in part) Saxifragaceae

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39

Lycophytes Huperziaceae References: Wagner and Beitel (1993), Haines (2003a).

Huperzia 1a. Trophophylls dimorphic, those in the apical portion of the shoot noticeably shorter and more ascending than those near the base [Fig. 1]; gemmiphores borne throughout the apical portion of plant; lateral leaves of gemmae 0.5–1.1 (–1.2) mm wide, narrow-acute to acute at the apex; shoots determinate, the entire stem turning yellow on senescence . . . . . . . . H. appressa 1b. Trophophylls nearly monomorphic, those near the apex neither conspicuously smaller nor more ascending than those near the base; gemmiphores borne in a single whorl at the apex of a season’s growth; lateral leaves of gemmae 1.3–2.5 mm wide, obtuse to rounded at apex; shoots indeterminate, the base eventually turning yellow and rotting away 2a. Trophophylls obovate, definitely widest beyond the middle, with 1–8 evident teeth [Fig. 2], lacking stomates on the adaxial surface; shoots with evident winter bud constrictions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. lucidula 2b. Trophophylls lanceolate to narrow-triangular, widest below the middle, entire or rarely with obscure, papilla-like teeth, with abundant stomates on the adaxial surface (i.e., 30–90 per ½ leaf surface); shoots with obscure winter bud constrictions . . . . . . . . . . . . . . H. selago 1. Huperzia appressa (Desv.) A. & D. Löve N Fig. 1 mountain firmoss. Huperzia appalachiana Beitel & Mickel; H. selago (L.) Schrank & Mart. ssp. appressa (Desv.) D. Löve; Lycopodium selago L. var. appressum Desv.; L. selago L. ssp. appressum (Desv.) Hultén; Urostachys selago (L.) Herter var. appressus (Desv.) Herter ex Ness. • CT, MA, ME, NH, VT; southern New England sites are highly disjunct. Exposed, windswept cliffs, ridges, and summits, often at high elevation, though at low elevation on Mount Desert Island and Isle au Haut, ME. Shade forms of this species are sometimes confused with the hybrid Huperzia ×‌josephbeitelii, which is most frequently encountered in alpine areas. ‌1 × 2. Huperzia ×protoporophila A. Haines is a rare hybrid known from MA, ME, NH, VT. It occurs on cliffs and summits and in alpine gullies. In New England, this taxon is abortive-spored (compared with the fertile tetraploid Huperzia porophila (Lloyd & Underwood) Holub of the mid-Atlantic and Midwest states). It has leaf morphology comparable to H. ×‌buttersii but has somewhat dimorphic trophophylls (as to basal and apical; vs. nearly monomorphic), narrower lateral gemmae leaves (1.2–1.6 mm vs. 1.5–2 mm), and gemmae borne in 1 or 2 pseudowhorls at the apex of each season’s growth (vs. always 1 pseudowhorl at the apex of each season’s growth). ‌1 × 3. Huperzia ×josephbeitelii A. Haines is the most common firmoss hybrid in New England and is known from ME, NH. It is found primarily above treeline in alpine areas. Like other Huperzia hybrids, this nothospecies has polymorphic (i.e., abortive) spores. The plants are similar to shade forms of H. appressa but are more stocky (i.e., shoots 7–10 mm wide inclusive of trophophylls vs. 3–7 mm) and have larger gemmae (the lateral leaves 1–1.5 mm wide vs. 0.5–1.1 (–1.2) mm).

Fig. 1  Apical portion of shoot of Huperzia appressa with gemmae.

40  Lycophy tes

2. Huperzia lucidula (Michx.) Trevisan N Fig. 2 shining firmoss. Lycopodium lucidulum Michx.; Urostachys lucidulus (Michx.) Ness. • CT, MA, ME, NH, RI, VT. Mesic to hydric forests, including conifer and broad-leaved types. 2 ‌ × 3. Huperzia ×buttersii (Abbe) Kartesz & Ghandi is a rare firmoss hybrid known from ME, NH, VT. It usually occurs in hydric and/or coniferous forests. It somewhat resembles small forms of H. lucidula but has trophophylls with ± parallel margins, obscure, papilla-like teeth, and a few stomates (i.e., fewer than 30 per ½ adaxial leaf surface. 3. Huperzia selago (L.) Bernh. ex Mart. & Schrank

Fig. 2  Apical portion of shoot of Huperzia lucidula showing gemmiferous branches.

NC

northern firmoss. Lycopodium selago L.; L. selago L. var. patens (Beauv.) Desv.; Plananthus selago (L.) Beauv.; Urostachys selago (L.) Herter • MA, ME, NH, VT; northern and colder counties. Hydric habitats in ± boreal regions, such as in ditches and low fields, also on lake shores and less commonly in forests, at cliff bases, and in mountain gullies. This species is commonly reported from alpine habitats, but those reports are based on H. appressa and hybrids with that species.

Isoetaceae Isoetes The size and ornamentation of mature megaspores, found in the expanded leaf bases, are crucial for confident identification of most species. Collections of Isoetes from most of the growing season will possess only fragile, yellow-white, and largely unornamented megaspores. For most New England taxa, collections from August and later will yield identifiable material. It is recommended that collectors check the condition of the megaspores while in the field to insure that mature spores are available (they should be spherical in orthospecies and bone white upon drying). Hybrids are to be expected where two or more species are found in the same water body. Hybrid quillworts can be detected by their polymorphic megaspores that range in shape from spherical to pyramidal and flattened. References: Eaton (1900), Kott and Britton (1983), Taylor et al. (1993). 1a. Megaspores echinate with thin, sharp spines; leaves deciduous . . . . . . . . . . I. echinospora 1b. Megaspores variously ornamented, but without a spiny texture; leaves persisting more than 1 year 2a. Megaspores 0.4–0.56 mm in diameter, averaging less than 0.5 mm, patterned with an unbroken reticulum [Fig. 4] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. engelmannii 2b. Megaspores 0.4–0.75 mm in diameter, averaging more than 0.5 mm, cristate, rugulate, or reticulate with broken or anastomosing ridges 3a. Leaves with abundant stomates, bright green; plants aquatic to amphibious, occasionally emergent; girdle of megaspores obscure . . . . . . . . . . . . . . . . . . . . . . . I. riparia 3b. Leaves with few or no stomates, dark green to red-green or red-brown; plants aquatic, usually submerged; girdle evident (obscure in I. prototypus) 4a. Megaspores 0.55–0.75 mm in diameter, averaging more than 0.6 mm; leaves abruptly tapering to the tip; plants typically submerged 1–3 (–5) m 5a. Megaspores with a papillate girdle, cristate to reticulate with sharp or roughened crests [Fig. 5] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. lacustris

is o e tac e a e   41

5b. Megaspores with a smooth girdle, reticulate with low, rounded ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. hieroglyphica 4b. Megaspores 0.4–0.65 mm in diameter, averaging less than 0.6 mm; leaves gradually tapering to the tip; plants submerged to 1 (–3) m 6a. Velum covering the entire sporangium [Fig. 6]; leaves very rigid and straight to the tip; sporangium wall unpigmented; girdle of the megaspore obscure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. prototypus 6b. Velum covering less than half the sporangium [Fig. 7]; leaves pliant and curling at the tip; sporangium wall brown-streaked; girdle evident 7a. Megaspores with a papillate girdle, cristate to reticulate with irregular, roughened crests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. tuckermanii 7b. Megaspores with a smooth girdle, reticulate with smooth, rounded crests [Fig. 3] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. acadiensis 1. Isoetes acadiensis Kott

N C Fig. 3

Acadian quillwort. MA, me, nh; eastern New England. Slightly acidic lakes and slowmoving streams. 2. Isoetes echinospora Durieu ssp. muricata (Durieu) A. & D. Löve N spiny-spored quillwort. Isoetes braunii Durieu; I. echinospora Durieu var. braunii (Durieu) Engelm.; I. muricata Durieu • CT, MA, ME, NH, RI, VT; throughout New England and one of the more common quillworts. Shallow, slightly acidic lakes and slow-moving streams. North American plants of Isoetes echinospora differ morphologically from Old World plants in that the leaves bear stomata (those from the Old World do not). This fact, combined with geographic separation, warrants recognition of North American plants as a distinct subspecies.

Fig. 3  Megaspore of Isoetes acadiensis showing the smooth girdle.

‌ × 3. Isoetes ×eatonii Dodge is a relatively rare quillwort hybrid known from CT, MA, 2 NH, RI. It shows megaspore ornamentation similar to I. riparia (i.e., a pattern intermediate between spines and ridges, usually appearing as columns that sometimes merge to form broken ridges). However, I. × ‌eatonii will have polymorphic megaspores rather than the uniform megaspore morphology of I. riparia. 2 ‌ × 7. Isoetes ×dodgei A.A. Eat. is a rare quillwort hybrid known from NH, VT. It shows spiny megaspores, but many of the spines are fused and/or irregular, resulting in a convoluted-echinate surface (i.e., the spines are not as regular in appearance as those in I. echinospora). The well-formed megaspores are usually smaller than 0.54 mm in diameter. ‌ × 8. Isoetes ×echtuckerii D.F. Brunton & D.M. Britt. is known from MA, ME, NH, 2 but should be expected anywhere the two parental species are found together. It has megaspores that show isolated columns and short, broken ridges that become shorter and more congested near the equatorial ridge (i.e., the hybrid megaspores have a somewhat apparent girdle, which is present in I. tuckermanii but lacking in I. echinospora). The well-formed megaspores are mostly 0.46–0.50 mm in diameter. 3. Isoetes engelmannii A. Braun N Fig. 4 Engelmann’s quillwort. CT, MA, NH, RI, VT. Shallow water of lakes and rivers, sometimes emergent. ‌3 × 7. Isoetes ×brittonii D.F. Brunton & W.C. Taylor is documented from only CT in New England. It has megaspores with ornamentation very similar to those of I. engelmannii, except that the ridges forming the reticulum are broken in places and get more congested toward the equatorial ridge (similar to I. riparia). The wellformed megaspores measure 0.47–0.56 mm in diameter. ‌3 × 8. Isoetes ×foveolata A.A. Eat. ex Dodge is a rare quillwort hybrid known from CT, NH; also reported from MA by Magee and Ahles (1999), but specimens are unknown. The

Fig. 4  Megaspore of Isoetes engelmannii, a species that lacks a girdle.

42   Lycophy te s

megaspores show a broken reticulum that is shorter and denser near the equatorial ridge (i.e., the megaspores have a somewhat apparent girdle). The megaspores measure 0.38–0.56 mm in diameter. 4. Isoetes hieroglyphica A.A. Eat.

NC

carved quillwort. Isoetes lacustris L. forma hieroglyphica (A.A. Eat.) N.E. Pfeiff. • ME; most collections from the northern portion of ME. Cold, clear water of lakes and slow-moving streams with gravel and cobble substrate. 5. Isoetes lacustris L. N Fig. 5

Fig. 5  Megaspore of Isoetes lacustris showing the papillate girdle.

lake quillwort. Isoetes macrospora Durieu • MA, ME, NH, VT; also reported from CT and RI by Taylor et al. (1993), but specimens are unknown. Cold, clear water of lakes and slow-moving streams, sometimes at depths greater than 3 m. North American plants have been segregated from Old World plants as Isoetes macrospora. However, there does not appear to be any basis for separation other than geography. Therefore, I. macrospora is not recognized here as a distinct taxon. ‌5 × 8. Isoetes ×harveyi A.A. Eat. is known only from ME within New England. Given that its two parents are very similar in megaspore ornamentation (though the spores differ in size), this hybrid is recognized by possession of megaspore texture similar to that of I. lacustris or I. tuckermanii, but with many polymorphic spores (as in all quillwort hybrids) and the well-formed ones 0.52–0.64 mm in diameter. 6. Isoetes prototypus D.M. Britt.

N C Fig. 6

prototype quillwort. ME. Cold, clear lakes with mineral substrate. 7. Isoetes riparia Engelm. ex A. Braun var. canadensis Engelm. ex N.E. Pfeiff. N

Fig. 6  Velum of Isoetes prototypus that covers the entire sporangium.

Canada shore quillwort. Isoetes canadensis (Engelm.) A.A. Eat. ex Maxon, nom. illeg. • CT, MA, ME, NH, RI, VT. Sandy and muddy margins of streams and lakes, including tidal shorelines. Isoetes riparia var. canadensis is an allotetraploid, formed by chromosome doubling in I. ×‌eatonii, which is the first generation hybrid between I. echinospora and I. engelmannii. See Caplen and Werth (2000) for justification of recognizing northern populations of the I. riparia complex as distinct. Unfortunately, no combination at the specific level has yet been made. ‌ × 8. Isoetes ×novae-angliae D.F. Brunton & D.M. Britt. is a rare quillwort hybrid 7 known from CT, MA, NH. It has polymorphic megaspores, the well-formed ones measuring mostly 0.45–0.53 mm in diameter and having a dense, broken reticulum, showing an obscure girdle of congested reticulum (no spine-like papillae as occur in I. tuckermanii). 8. Isoetes tuckermanii A. Braun ex Engelm. N Fig. 7 Tuckerman’s quillwort. CT, MA, ME, NH, RI, VT; nearly throughout New England. Slightly acidic lakes and slow-moving streams.

Fig. 7  Velum of Isoetes tuckermanii that covers less than half of the sporangium.

Lycopodiaceae References: Wagner and Beitel (1993), Haines (2003a). 1a. Upright shoots unbranched, lacking winter bud constrictions, produced from horizontal shoots that travel over the surface [Fig. 14]; horizontal shoots lacking winter bud constrictions [Fig. 14]; plants primarily of hydric habitats 2a. Sporophylls green, superficially similar to the trophophylls of the upright shoots, arranged in 10 or more vertical ranks, 2.9–9 mm long; strobilus stalks with abundant and imbricate trophophylls; trophophylls of the horizontal shoots monomorphic; plants deciduous, overwintering by a thickened tip of stem (i.e., turion) . . . . . . . . . . . . Lycopodiella

Lyco p o d i ac e a e  43

2b. Sporophylls stramineous at maturity, clearly differentiated from the trophophylls of the upright shoots, arranged in 4–10 vertical ranks, 2.7–3.3 mm long; strobilus stalks with few and remote trophophylls; trophophylls of the horizontal shoots dimorphic, the lateral trophophylls markedly wider than the median trophophylls; plants evergreen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudolycopodiella 1b. Upright shoots branched, with winter bud constrictions, produced from subterranean or superficial horizontal shoots; horizontal shoots with winter bud constrictions (except Dendrolycopodium), these marked by a zone of small, congested trophophylls; plants primarily of mesic to xeric habitats 3a. Branches 1–4.4 mm wide inclusive of the trophophylls, flat to quadrangular, with 4 ranks of trophophylls (terete and with 5 ranks of trophophylls in one species) [Fig. 11]; branching of strobilus stalks, when present, dichotomous . . . . . . . . . . . . . . . . Diphasiastrum 3b. Branches 5–12 mm wide, terete (somewhat compressed in D. obscurum), with 6 or more ranks of trophophylls; branching of strobilus stalks, when present, pseudomonopodial (i.e., falsely appearing to have a main axis from which branches arise) 4a. Strobili borne on stalks; trophophylls of the upright shoot branches arranged in alternating pseudowhorls of 6–10, tipped with a colorless hair 1.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lycopodium 4b. Strobili sessile; trophophylls of the upright shoot branches arranged in alternating pseudowhorls of 3–5, merely pointed or with a minute, but firm, spine-like tip 5a. Upright shoots with an obvious main axis from which repeatedly forked branches arise (i.e., the shoots dendroid), bearing trophophylls that are merely pointed at the apex; horizontal shoots subterranean, lacking winter bud constrictions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dendrolycopodium 5b. Upright shoots with 1–4 subdichotomous branches (i.e., the shoots not dendroid and the main axis usually obscure), bearing trophophylls with a minute, but firm, spinule 0.4–1 mm long at the apex; horizontal shoots at or near the surface, with winter bud constrictions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spinulum

Dendrolycopodium Individuals are sometimes found with conflicting character states (e.g., the phyllotaxy of one species on one branch and the phyllotaxy of another species on a different branch). These plants have been suspected to be of hybrid origin (Hickey 1978). This hypothesis is corroborated by intermediacy in trophophyll morphology and divergence angle. Given the close morphological similarity of New England’s three species, hybrids are rarely reported and likely undercollected. 1a. Trophophylls of the upright shoot’s main axis 0.9–1 mm wide, widely ascending to spreading (i.e., diverging at an angle of 45–90 degrees), firm, prickly to the touch; trophophylls of the lateral branches arranged in 2 upper, 2 lower, and 2 lateral ranks [Fig. 8] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. dendroideum 1b. Trophophylls of the upright shoot’s main axis 0.5–0.7 mm wide, appressed to ascending (i.e., diverging at an angle of less than 30 degrees), pliable; trophophylls of the lateral branches arranged in 1 upper, 1 lower, and 4 lateral ranks [Figs. 9, 10] 2a. Lateral branches somewhat compressed; trophophylls of the lower rank of lateral branches shorter than those of the other ranks, those of the lateral ranks twisted, so that one surface faces up, the other surface faces down [Fig. 10]; trophophyll margins forming an angle of 27–59 degrees at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. obscurum 2b. Lateral branches terete; trophophylls of the lower rank of lateral branches similar in length to those of the other ranks, those of the lateral ranks not twisted, one surface facing

44  Lyco phy tes

the branch axis, the other surface facing outward [Fig. 9]; trophophyll margins forming an angle of 21–36 degrees at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. hickeyi 1. Dendrolycopodium dendroideum (Michx.) A. Haines N Fig. 8

Fig. 8  Cross-section of upright shoot branch of Dendrolycopodium dendroideum.

prickly tree-clubmoss. Lycopodium dendroideum Michx.; L. obscurum L. var. dendroideum (Michx.) D.C. Eat.; L. obscurum L. var. hybridum Farw. • CT, MA, ME, NH, VT; throughout most of the region except southeastern New England. Interior and edges of hardwood and mixed conifer-hardwood forests and woodlands. 1 × 2. This rare tree-clubmoss hybrid is known from CT, ME, VT. 1 × 3. This rare tree-clubmoss hybrid is known from CT, MA, VT. 2. Dendrolycopodium hickeyi (W.H. Wagner, Beitel, & Moran) A. Haines N Fig. 9 Hickey’s tree-clubmoss. Lycopodium hickeyi W.H. Wagner, Beitel, & Moran; L. obscurum L. var. isophyllum Hickey • CT, MA, ME, NH, RI, VT; throughout most of New England. Interior and edges of hardwood and mixed conifer-hardwood forests and woodlands, frequently in more well-drained soils than Dendrolycopodium obscurum. 2 × 3. This rare tree-clubmoss hybrid is known from CT, MA, ME, NH.

Fig. 9  Cross-section of upright shoot branch of Dendrolycopodium hickeyi.

3. Dendrolycopodium obscurum (L.) A. Haines N Fig. 10 flat-branched tree-clubmoss. Lycopodium obscurum L. • CT, MA, ME, NH, RI, VT; throughout. Interior and edges of hardwood and mixed conifer-hardwood forests and woodlands, sometimes in dry, open areas such as powerline rights-of-way.

Diphasiastrum

Fig. 10  Cross-section of upright shoot branch of Dendrolycopodium obscurum.

Diphasiastrum hybrids can be locally common, particularly in mixed-species populations. Some of the hybrid individuals show reduction in proportion of well-formed spores; unfortunately, so do some collections of orthospecies from the region. Of note is that one nothospecies in particular, D. × ‌sabinifolium, may be located without either parent present. Reference: Wilce (1965). 1a. Trophophylls of the lateral branches monomorphic, arranged in 5 ranks, adnate to the terete branches less than 50% of their length, with stomates on both surfaces; strobilus stalks absent (rarely up to 1 cm tall); strobili solitary; plants commonly shorter than 12 cm, compactly branched, without an evident main axis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sitchense 1b. Trophophylls of the lateral branches di- or trimorphic, arranged in 4 ranks, those of the lateral ranks adnate to the flat or quadrangular branches more than 50% of their length, with stomates on the abaxial (i.e., lower) surface only; strobilus stalks 0.5–15 cm long; strobili 1–4 per stalk; plants commonly taller than 12 cm, usually dendroid 2a. Trophophylls of the lower rank 1–2 mm long, similar in length to the trophophylls of the upper rank; branches narrow, quadrangular, 1.2–2 mm wide, blue-green; horizontal shoots subterranean, frequently buried deeper than 5 cm . . . . . . . . . . . . . . . . . . . . . . . D. tristachyum 2b. Trophophylls of the lower rank 0.3–1 (–1.5) mm long, noticeably shorter than the trophophylls of the other ranks; branches wider, flat, 2–4.4 mm wide, green; horizontal shoots superficial, at the surface or immediately below the litter 3a. Lateral branches of upright shoots with winter bud constrictions [Fig. 11], somewhat irregular, more 3-dimensional and not planar; strobili mostly 10–25 mm tall, without sterile tips; strobilus stalks usually becoming stramineous prior to sporangia dehiscence, mostly 0.4–0.7 mm in diameter, the successive branches separate, appearing as distinct dichotomies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. complanatum 3b. Lateral branches of upright shoots without annual constrictions, with regular and planar aspect; strobili mostly 20–40 mm tall, often some with a slender, sterile tip; strobilus stalks remaining green until after sporangia dehiscence, mostly 0.6–1 mm in

Lyco p o d i ac e a e  45

diameter, the successive branches in close proximity, the strobili often falsely appearing in a pseudowhorl of 4 [Fig. 12] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. digitatum 1. Diphasiastrum complanatum (L.) Holub N Fig. 11 northern ground-cedar. Diphasium complanatum (L.) Rothm.; Lycopodium complanatum L.; L. complanatum L. var. canadense Vict. • ME, NH, VT. Dry-mesic to mesic openings, edges, and forests, often in acid soils. This is the more northern sister species of Diphasiastrum digitatum. 1‌ × 2. Diphasiastrum ×verecundum A.V. Gilman is a rare ground-cedar hybrid of northern New England (ME, NH, VT). The upright shoot branches are somewhat irregular (as in D. complanatum), but the stobilus stalk is stouter than in that species and often remains green after sporangium dehiscence. This hybrid may be more common than collections indicate, but the close similarity between its parents makes it difficult to detect. 1‌ × 4. Diphasiastrum ×zeilleri (Rouy) Holub is a rare ground-cedar hybrid known from ME, NH, VT. It is intermediate between its parents in branch width, number of times the branches divide, and relative size of the trophophylls of the lower rank.

Fig. 11  Adaxial (i.e., upper) view of upright shoot branches of Diphasiastrum complanatum showing winter bud constrictions.

1‌ × Diphasiastrum alpinum (L.) Holub. Diphasiastrum ×issleri (Rouy) Holub is a very rare ground-cedar hybrid in New England known only from northern ME. It generally resembles D. complanatum except that its lateral trophophylls usually roll under the branch (i.e., the branch margin is revolute) and the base of its strobilus is not distinct (i.e., there is a gradual transition from strobilus stalk trophophylls to strobilus sporophylls). 2. Diphasiastrum digitatum (Dill. ex A. Braun) Holub N Fig. 12 southern ground-cedar. Diphasium flabelliforme (Fern.) Blanch.; Lycopodium complanatum L. var. flabelliforme Fern.; L. digitatum Dill. ex A. Braun; L. flabelliforme (Fern.) Blanch. • CT, MA, ME, NH, RI, VT; throughout. Upland forests, borders, and fields. ‌ × 4. Diphasiastrum ×habereri (House) Holub is an infrequent ground-cedar hybrid 2 known from MA, ME, NH, VT. It has winter bud constrictions on the lateral branches, similar to D. tristachyum, but is otherwise intermediate in branch width, relative size of trophophylls of the lower rank, and strobilus stalk branching. 3. Diphasiastrum sitchense (Rupr.) Holub

NC

Sitka ground-cedar. Diphasium sitchense (Rupr.) A. & D. Löve; Lycopodium sabinifolium Willd. var. sitchense (Rupr.) Fern.; L. sitchense Rupr. • ME, NH. Alpine areas, frequently on open, stony plains or near the summit of snow bank gullies; also in disturbed habitats in boreal areas, such as fields and borrow pits. 3 ‌ × 4. Diphasiastrum ×sabinifolium (Willd.) Holub is a somewhat rare but widespread ground-cedar hybrid known from ME, NH, VT that frequently occurs in the absence of its parents. It has somewhat compressed lateral branches with 4 ranks of trophophylls that are adnate to the branches 50–60% of their length (most species, except D. sitchense, with trophophylls adnate more than 60% of their length), and shows a gradual transition from stobilus stalk trophophylls to strobilus sporophylls. 4. Diphasiastrum tristachyum (Pursh) Holub N blue ground-cedar. Diphasium tristachyum (Pursh) Rothm.; Lycopodium chamaecyparissus A. Br. ex Mutel; L. tristachyum Pursh • CT, MA, ME, NH, RI, VT; throughout. Dry-mesic to xeric, often acid, soils of woodlands, forests, edges, and openings. Diphasiastrum tristachyum is involved in more hybrid combinations than any other Diphasiastrum in New England.

Lycopodiella Species of Lycopodiella hybridize more frequently than other clubmosses in the region. With the exception of L. inundata, it is uncommon to find populations lacking hybrids, even when only one species is present. Sporophyll orientation is a very important character, but it cannot

Fig. 12  Strobili and strobilus stalk branching of Diphasiastrum digitatum.

46 Lyco phy te s

be assessed until late summer. Developing strobili have ascending sporophylls that eventually spread from the axis at maturity (except in L. appressa). Abortive-spored plants are present in New England, and not all circumstances appear to be explainable by differences in ploidy level. Reference: Bruce (1975). 1a. Horizontal shoots, excluding the trophophylls, very slender, 0.5–0.9 mm thick, each segment producing 1 (rarely 2) upright shoot; trophophylls of the horizontal shoot entire; upright shoots rarely taller than 10 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. inundata 1b. Horizontal shoots thicker, 1.2–3.1 mm thick, each segment producing 1–5 upright shoots; trophophylls of the horizontal shoot toothed; upright shoots usually taller than 10 cm [Figs. 13, 14] 2a. Horizontal shoots prominently arching over the substrate, rooting 7.5–36 cm distal to the most proximal upright shoot [Fig. 13]; sporophylls 5.5–9 mm long, spreading at maturity, with 1–3 slender teeth on each margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. alopecuroides 2b. Horizontal shoots prostrate, rooting 1.5–4.8 cm distal to the most proximal upright shoot [Fig. 14]; sporophylls 2.9–5 (–5.2) mm long, appressed, entire or rarely with a low tooth on one margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. appressa 1. Lycopodiella alopecuroides (L.) Cranfill

N C Fig. 13

foxtail bog-clubmoss. Lepidotis alopecuroides (L.) Rothm.; Lycopodium alopecuroides L. • CT, MA, ME, RI; southern and eastern New England. Wet, sandy and/or peaty soils along the coastal plain, often in disturbed sites such as abandoned borrow pits and powerline rights-of-way. Regionally rare in New England and disjunct in south-central ME. Fig. 13  Arching horizontal shoot of Lycopodiella alopecuroides.

1‌ × 2. Lycopodiella ×copelandii (Eiger) Cranfill is an extremely rare bog-clubmoss hybrid known from the coastal plain of southern New England in CT, MA. It is marked by ascending sporophylls and weakly arching horizontal stems rooting mostly 3.5–10.5 cm distal to the proximal most upright shoot. Interestingly, many of the sporophylls and trophophylls on most specimens are entire in this hybrid. ‌1 × 3. Lycopodiella ×robusta (R.J. Eat.) A. Haines is a very rare bog-clubmoss hybrid in New England that is known from MA, ME. It generally resembles L. alopecuroides but has a relatively taller strobilus (34–52% vs. 6–36% of the total upright shoot height), less conspicuously arching stems, and fewer teeth on the margins of the horizontal shoot trophophylls (0–2 vs. 1–8 per margin). 2. Lycopodiella appressa (Chapman) Cranfill N Fig. 14

Fig. 14  Prostrate horizontal shoot of Lycopodiella appressa.

appressed bog-clubmoss. Lycopodiella bigelovii (Tuckerman) Holub; Lycopodium appressum (Chapman) Lloyd & Underwood; L. inundatum L. var. appressum Chapman; L. inundatum L. var. bigelovii Tuckerman • CT, MA, ME, NH, RI, VT; populations in the northern states are disjunct, most frequent along the coastal plain and in the lower Connecticut River valley. Wet, sandy and/or peaty soils, often in abandoned borrow pits and draw-down pond shore margins. Most reports of this species from ME were based on Lycopodiella × ‌gilmanii. ‌ × 3. Lycopodiella ×gilmanii A. Haines is a rare bog-clubmoss hybrid that has been 2 found nearly throughout the coastal plain of New England (rarely inland) in CT, MA, ME, NH, VT. It most commonly occurs with L. appressa. It is responsible for reports of L. margueritae Bruce, Wagner, & Beitel in New England, a tetraploid species known ‌gilmanii only from the Great Lakes region. Lycopodiella × differs most notably from L. appressa in its longer (mostly 5–6.2 mm), ascending sporophylls with 0–2 slender teeth per margin, and fewer upright shoots per horizontal shoot segment (commonly 1 or 2 vs. 2–5). Further, many specimens of this hybrid in New England have a large fraction of malformed spores. Rare variants of this nothospecies have appressed sporophylls until very late in the season (i.e., the sporophylls tardily spread from the strobilus axis). 3. Lycopodiella inundata (L.) Holub N northern bog-clubmoss. Lepidotis inundata (L.) Opiz; Lycopodium inundatum L. • CT, MA, ME, NH, RI, VT; throughout. Wet, sandy and/or peaty soils, frequently in abandoned borrow

Lyco p o d i ac e a e   47

pits and on boggy pond margins. This is the only species of Lycopodiella in New England that frequently occurs in the absence of other species or hybrids. Its diminutive size, thin horizontal shoots, and entire trophophylls quickly distinguish most populations.

Lycopodium Lycopodium has been split into smaller, homogeneous groups on the basis of sporophyte and gametophyte morphology, chromosome number, and anatomy (Bruce 1975, Øllgaard 1987, Wagner and Beitel 1993). These genera also have different life histories and hybridization patterns that support the recognition of multiple genera (rather than a broadly defined Lycopodium). Hybrids have not been reported from New England Lycopodium; however, intermediate forms are not uncommon at some locations. 1a. Strobilus stalks with a solitary strobilus, sometimes with a second sessile strobilus; upright shoots with 2 or 3 ascending branches; trophophylls 3–5 mm long; sporophylls usually gradually tapering to the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. lagopus 1b. Strobilus stalks with 1–5 strobili, commonly with 2 or more and each with a short stalk-like branch; upright shoots with 3–6 spreading branches; trophophylls 4–6 mm long; sporophylls usually abruptly tapering to the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. clavatum 1. Lycopodium clavatum L. N common clubmoss. Lycopodium clavatum L. var. subremotum Vict. • CT, MA, ME, NH, RI, VT; throughout. Fields, edges, and forests. Care should be taken when collecting specimens of Lycopodium clavatum. This species occasionally, within a population, will produce solitary strobili. If these are the only portion of the plant that is collected, they will be misleading to later researchers. Lycopodium clavatum and L. lagopus sometimes occur together at the same site. In addition to the characters in the identification key, Lycopodium clavatum has trophophylls of lighter color that spread further from the shoot axis, and it releases spores later than L. lagopus. 2. Lycopodium lagopus (Laestad. ex Hartm.) G. Zinserling ex Kuzeneva-Prochorova N one-cone clubmoss. Lycopodium clavatum L. ssp. monostachyon (Hook. & Grev.) Selander; L. clavatum L. var. lagopus Laestad. ex Hartm.; L. clavatum L. var. megastachyon Fern. & Bissell; L. clavatum L. var. monostachyon Grev. & Hook. • CT, MA, ME, NH, VT; primarily of northern New England, becoming rare to the south. Fields and forest openings and borders. Lycopodium lagopus has a more northern distribution that its sister species, L. clavatum. Intermediate forms between these two species do exist, but whether they represent intermediate morphologies or hybrids has not been determined.

Pseudolycopodiella Both diploid and tetraploid forms of this orthospecies exist. Specimens have been collected in the southeastern United States with abortive spores, which represent largely sterile triploid hybrids between the two different ploidy types. Reference: Bruce (1975). 1. Pseudolycopodiella caroliniana (L.) Holub

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slender false bog-clubmoss. Lycopodiella caroliniana (L.) Pichi-Sermolli; Lycopodium carolinianum L. • MA; known only from Hampshire County, MA. Hydric, often sandy, soils of meadows, ditches, and shores. New England’s single occurrence of this species was extirpated by extensive grading of the site.

Spinulum Spinulum is the sister genus to Lycopodium, and the two genera share such features as terete branches, superficial horizontal shoots, and non-dendroid upright shoots. However, they differ in many important features such as phyllotaxis, leaf apex morphology, and branching pattern

48  Lycophy tes

of upright shoots. The trophophylls change actual length and relative length to width through the growing season. Therefore, to enable consistent comparisons, measurements and counts performed on trophophylls should be assessed on those near the middle of seasonal growth unless specified otherwise. 1a. Trophophylls of upright shoots with abundant stomates on the adaxial surface (i.e., 25–53 per ½ surface), 3–5.9 mm long, entire or very obscurely toothed, arranged in alternating pseudowhorls of 4, those immediately above the winter bud constrictions triangular to lanceolate or narrow-ovate (i.e., broadest at or very near the base); strobili 8–17 (–21) mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. canadense 1b. Trophophylls of upright shoots lacking stomates on upper surface (except the trophophylls at and immediately above winter bud constrictions), 5.2–9.8 mm long, obscurely to evidently toothed, arranged in alternating pseudowhorls of 5, those immediately above the winter bud constrictions oblanceolate or sometimes elliptic (i.e., broadest at or above the middle); strobili (15–) 17–43 mm tall. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. annotinum 1. Spinulum annotinum (L.) A. Haines N common interrupted-clubmoss. Lycopodium annotinum L.; L. annotinum L. var. acrifolium Fern. • CT, MA, ME, NH, RI, VT; nearly throughout, but rare in RI. Forests and forest edges, including

boreal and subalpine areas, infrequently above tree line at the edges of krummholz islands. 1 × 2. This uncommon interrupted-clubmoss hybrid is located most frequently where the parent species are sympatric (e.g., on the higher mountains of ME and NH). It can be recognized by intermediate leaf length and dentition between the parental clones and shows 1–15 stomates per ½ adaxial surface on trophophylls from the middle of seasonal growth (compared with 25–53 per ½ adaxial surface in Spinulum canadense). 2. Spinulum canadense (Ness.) A. Haines N northern interrupted-clubmoss. Lycopodium annotinum L. var. montanum Tuckerman; L. annotinum L. ssp. pungens Hultén; L. annotinum L. var. pungens La Pylaie ex Desv.; L. canadense Ness.; L. dubium, auct. non Zoega (1772); L. pungens La Pylaie ex Iljin in Komarov • ME, NH, VT; northern counties in New England. Northern and/or high-elevation areas, including open, alpine plateaus and ridges; rarely at low elevation (coniferous forests and bogs) in eastern and northern ME.

Selaginellaceae Selaginella There exist two distinct and monophyletic lineages within the Selaginellaceae that could be recognized as genera. One lineage includes the species that lack rhizophores and have terete strobili (Selaginella sensu stricto). The other lineage contains species with rhizophores and tetragonous strobili (Lycopodioides). However, several nomenclatural issues stand in the way of dividing Selaginella. Reference: Valdespino (1993). 1a. Trophophylls monomorphic, spirally arranged, not in distinct ranks [Fig. 16]; axillary trophophylls absent at branching points 2a. Trophophylls thin, spinulose-toothed, acuminate, with stomates scattered over the abaxial surface; strobili terete; sporophylls ascending to spreading, mostly 10-ranked [Fig. 16]; rhizophores absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. selaginoides 2b. Trophophylls thicker, ciliolate, bristle-tipped, with stomates restricted to an abaxial groove; strobili tetragonous; sporophylls appressed, 4-ranked; rhizophores present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rupestris

S e l ag i ne l l ac e a e   49

1b. Trophophylls dimorphic, arranged in 4 ranks, the 2 lateral ranks with larger leaves than the 2 median ranks [Fig. 15]; axillary trophophylls present at branching points 3a. Median trophophylls acute to acuminate at apex, rarely attenuate and then often apically keeled and the vein not reaching tip; megaspores 0.29–0.35 (–0.38) mm in diameter, closely reticulate, dull . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. apoda 3b. Median trophophylls with a long-attenuate, often recurved, apex, not keeled, the vein prolonged into the tip; megaspores 0.33–0.4 mm in diameter, more loosely reticulate, shiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. eclipes 1. Selaginella apoda (L.) Spring in Mart. et al. N Fig. 15

Fig. 15  Horizontal shoot and tetragonous strobilus of Selaginella apoda.

meadow spikemoss. Lycopodioides apoda (L.) Kuntze • CT, MA, ME, NH, RI, VT; nearly throughout but absent from most of ME and northern NH. Stream banks, mesic to wet-mesic meadows and lawns. 2. Selaginella eclipes W.R. Buck

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hidden spikemoss. CT, MA; western counties only. Mesic to hydric meadows and swamps, rarely on rock, habitat usually influenced by high-pH bedrock. 3. Selaginella rupestris (L.) Spring N ledge spikemoss. Bryodesma rupestre (L.) Soják; Lycopodium rupestre L. • CT, MA, ME, NH, RI, VT; throughout. Dry, exposed rock of various pH, less frequently on packed sand and gravel. 4. Selaginella selaginoides (L.) Beauv. ex Mart. & Schrank

N C Fig. 16

northern spikemoss. Lycopodium selaginoides L.; Selaginella spinosa Beauv. • ME; northern counties only. Open and forested circumneutral fens, usually palustrine but sometimes riparian. Fig. 16  Upright shoot and terete strobilus of Selaginella selaginoides.

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51

Monilophytes Aspleniaceae Asplenium Reference: Wagner et al. (1993). 1a. Leaf blades simple, frequently rooting at the tip; venation reticulate; sori irregularly scattered over the blade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. rhizophyllum 1b. Leaf blades 1- to 3-times divided [Figs. 17, 18], not rooting at the tip; venation free (i.e., the veins not rejoining); sori in paired rows on the leaflets 2a. At least the larger leaflets divided into leafules [Figs. 17, 18]; leaf blades deltate or deltate-ovate to lanceolate or obovate, with 2–10 pairs of leaflets 3a. Leaf blades usually with 2–5 pairs of leaflets [Fig. 18], dull blue-green in life; leaflets mostly rhombic to obtriangular-obovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ruta‑muraria 3b. Leaf blades usually with (4–) 5–10 pairs of leaflets [Fig. 17], shiny green in life; leaflets triangular-lanceolate to triangular-ovate . . . . . . . . . . . . . . . . . . . . . . . A. montanum 2b. None of the leaflets divided; leaf blades linear to narrow-oblanceolate, with (6–) 15–45 pairs of leaflets 4a. Leaves dimorphic—the fertile erect and tall, the sterile prostrate or arching and short; leaflets with a basal auricle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. platyneuron 4b. Leaves monomorphic; leaflets without a basal auricle 5a. Rachis red-brown to nearly black throughout; leaflets oblong to oval, the middle ones 2.5–4 mm wide (i.e., measured parallel with the leaf axis); petioles 0.15–0.25 times the length of the blade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. trichomanes 5b. Rachis green; leaflets deltate to rhombic, the middle ones 4–5 mm wide; petioles 0.25–0.5 (–1) times the length of the blade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. viride 1. Asplenium montanum Willd.

N C Fig. 17

mountain spleenwort. CT, MA, RI, VT; in MA confined to western portion of state, in VT confined to southern portion of state. Primarily non-calcareous cliffs. 2. Asplenium platyneuron (L.) Britton N ebony spleenwort. Asplenium platyneuron (L.) Britton var. incisum (Howe ex Peck) B.L. Robins. • CT, MA, ME, NH, RI, VT; in ME only the southwestern portion of state. Rocky, circumneutral slopes, usually growing on thin soil, occasionally on rock or mortar. ‌2 × 3. Asplenium ×ebenoides R.R. Scott is a rare spleenwort hybrid known primarily from ‌ebenoides western New England (CT, MA, NH, VT). Given that Asplenium × is chiefly sterile (i.e., produces abortive spores) and occurs with both parental species, it is appropriate to refer to our material as a nothospecies. The Hale County, Alabama, populations that are fertile tetraploids have appropriately (and finally) been provided a separate name—A. tutwilerae B.R. Keener & R.J. Davenport—because those populations represent an allotetraploid species.

Fig. 17  Leaf blade of Asplenium montanum.

5 2   mo n ilop hy te s

3. Asplenium rhizophyllum L. N walking spleenwort. Camptosorus rhizophyllus (L.) Link • CT, MA, ME, NH, RI, VT; infrequent in eastern MA and in ME only in Kennebec County. Usually on shaded, high-pH rocks and cliffs, rarely at tree bases. 4. Asplenium ruta-muraria L.

N C Fig. 18

wall-rue spleenwort. Asplenium cryptolepis Fern.; A. ruta-muraria L. var. cryptolepis (Fern.) Wherry • CT, MA, RI, VT; primarily western New England, the RI occurrence very disjunct. HighpH rocks and cliffs.

Fig. 18  Leaf blade of Asplenium ruta-muraria.

‌4 × 5. Asplenium ×clermontae Syme is an extremely rare spleenwort hybrid known from VT. It resembles A. trichomanes in general outline (i.e., it has once-pinnately compound leaf blades), but the leaflets are lobed (especially the larger ones) and the rachis is green (rather than brown-red to black-brown). 5. Asplenium trichomanes L. N maidenhair spleenwort. CT, MA, ME, NH, RI, VT; nearly throughout. Growing on cliffs, boulders, and talus. Characters for the following key were extracted from Ekrt and Štech (2008). Those listed in Moran (1982), who drew on European studies of this species, do not appear to be useful for New England populations. 1a. Marginal cells of rachis wings ascending, often longer than wide, rounded at the apex; distance between petiolules on same side of rachis near apex of leaf blade 2–4 mm; terminal leaflet 3–7 mm wide; spores mostly 34–43 μm; plants primarily of circumneutral to basic rock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5a. A. trichomanes ssp. quadrivalens D.E. Mey. 1b. Marginal cells of rachis wings nearly perpendicular to plant axis, often as wide or wider than long, truncate at the apex; distance between petiolules as measured in 1a 3–7 mm; terminal leaflet 1.5–4 mm wide; spores mostly 29–36 μm; plants mostly of acidic rock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5b. A. trichomanes ssp. trichomanes Subspecies quadrivalens is known from CT, MA, ME, NH, RI, VT. Subspecies trichomanes is known from CT, MA, ME, NH, VT. 5a × 5b. Asplenium trichomanes L. nssp. lusaticum (D.E. Mey.) Lawalrée is a very rare spleenwort hybrid in New England known from ME, NH, VT. Given the similarity between the parental subspecies, this nothosubspecies is best identified by its abortive spores. 6. Asplenium viride Huds.

NC

green spleenwort. Asplenium trichomanes-ramosum L. • ME, VT. On calcareous and serpentine rocks and ledges.

Azollaceae Azolla Reference: Lumpkin (1993). 1. Azolla caroliniana Willd. E Carolina mosquito fern. CT, MA; also reported from NH by Magee and Ahles (1999), but voucher specimens are unknown. Still or slow-moving water of lakes, pools, swamps, and streams. Reports of this species from RI were based on a collection taken from a pond on the University of Rhode Island campus—Champlin 51 (Champlin Herb.).

Bl ec hn ac e a e   5 3

Blechnaceae Woodwardia Reference: Cranfill (1993). 1a. Leaves monomorphic, with 15–20 pairs of distinct, lobed leaflets; veins forming 1 row of areoles adjacent to the costae or costules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. virginica 1b. Leaves dimorphic, with 7–10 pairs of leaflets, the sterile leaflets more or less confluent and serrulate; veins forming 2 or more rows of areoles adjacent to the costae . . . . . . . . W. areolata 1. Woodwardia areolata (L.) T. Moore N netted chain fern. Lorinseria areolata (L.) K. Presl, nom. illeg.; Woodwardia onocleoides Willd. • CT, MA, ME, NH, RI. Acid soils of swamps, wetland borders, low fields, seeps, and bogs. Several characters set Woodwardia areolata apart from other members of the genus. On such evidence, generic segregation has been proposed (as Lorinseria areolata, an illegitimate name because Lorinseria C. Presl is considered an orthographic variant of Lorinsera Opiz). Sterile specimens are sometimes confused with vegetative leaves of Onoclea sensibilis. They can be separated through the entire or merely undulate leaflets in O. sensibilis that have a narrow, pale, marginal membrane vs. serrulate leaflets in W. areolata that lack a pale, marginal membrane. 2. Woodwardia virginica (L.) Smith N Virginia chain fern. Blechnum virginicum L. • CT, MA, ME, NH, RI, VT. Acid soil of swamps, bogs, marshes, and ditches.

Dennstaedtiaceae The Dennstaedtiaceae, along with the Pteridaceae, comprise leptosporangiate ferns that have a false indusium covering the sorus. The two families differ morphologically in rhizome details and presence/absence of epipetiolar buds. The Dennstaedtiaceae have rhizomes pubescent with jointed hairs and have epipetiolar buds. The Pteridaceae have rhizomes with scales 2 or more cells wide and lack epipetiolar buds. 1a. Sori discrete, spherical, borne in circular or slightly bivalvate cups formed by fusion of the true indusium with the false indusium; false indusium merely a reflexed tooth or lobe; leaf blade narrow-oblong to lanceolate or broad-lanceolate, 12–25 (–30) cm wide; nectaries absent; base of petiole with 1 or 2 vascular bundles . . . . . . . . . . . . . . . . . . . . . . . . . Dennstaedtia 1b. Sori in a marginal band not borne in cup-like structures; false indusium formed by a ± elongate, revolute margin; leaf blade broad-deltate to ovate-deltate, (25–) 40–70 cm wide; nectaries present at base of leaf divisions, these appearing as dark, oval areas in early season; base of petiole with 10 or more vascular bundles . . . . . . . . . . . . . . . . . . . . . . . . Pteridium

Dennstaedtia Reference: Nauman and Evans (1993). 1. Dennstaedtia punctilobula (Michx.) T. Moore N eastern hay-scented fern. Nephrodium punctilobulum Michx. • CT, MA, ME, NH, RI, VT. Fields, forests, roadsides, and rocky slopes.

5 4   mo n ilop hy tes

Pteridium Reference: Jacobs and Peck (1993). 1. Pteridium aquilinum (L.) Kuhn N bracken fern. 1a. Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underwood ex Heller; P. latiusculum (Desv.) Hieron.; 1b. Pteridium aquilinum (L.) Kuhn var. pseudocaudatum (Clute) Heller; P. latiusculum (Desv.) Hieron. var. pseudocaudatum (Clute) Maxon • CT, MA, ME, NH, RI, VT. Fields, forest openings and edges, woodlands, and roadsides. 1a. Terminal segment of leafules 2–4 times as long as wide; margins of ultimate segments and abaxial costae sparsely to densely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. Pteridium aquilinum ssp. latiusculum (Desv.) Hultén 1b. Terminal segment of leafules mostly 6–15 times as long as wide; margins of ultimate segments and abaxial costae sparsely pubescent to glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. Pteridium aquilinum ssp. pseudocaudatum (Clute) Hultén Subspecies latiusculum is known from CT, MA, ME, NH, RI, VT. Subspecies pseudocaudatum is known from CT, MA.

Dryopteridaceae 1a. Indusium peltate (i.e., attached at the center by a stalk); leaflets or leafules with a basal auricle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polystichum 1b. Indusium attached laterally at a distinct sinus; leaflets and leafules lacking a basal auricle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dryopteris

Dryopteris Hybrids in Dryopteris are common in mixed-species populations. Certain hybrid combinations are almost always found when the two parents grow together (e.g., D. ×‌boottii), whereas ‌pittsfordensis) others are very rare (e.g., D. × or unknown in the region. Hybrids can be recognized by abortive spores and intermediate morphology. References: Thorne and Thorne (1989), Montgomery and Wagner (1993). 1a. Leaf blades mostly 6–25 cm long, with aromatic glands, densely scaly abaxially; indusia often overlapping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. fragrans 1b. Leaf blades 15–120 cm long, mostly more than 25 cm, lacking aromatic glands (but sometimes with stipitate glands), with few or no abaxial scales; indusia separate 2a. Leaf blades (2–) 2.5- to 3.5-times divided near base; leaf segments with bristletipped teeth 3a. First basiscopic (i.e., lower) leafule of basal leaflet no longer than adjacent basiscopic leafule and less than 2 times as long as first acroscopic (i.e., upper) leafule of basal leaflet [Fig. 21]; plant axes and indusia with glandular hairs; leaf blades remaining green through the winter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. intermedia 3b. First basiscopic leafule of basal leaflet longer than adjacent basiscopic leafule and 2–5 times as long as first acroscopic leafule of basal leaflet [Figs. 19, 20]; plants without glandular hairs (rarely with a few glandular hairs in D. campyloptera); leaf blades senescing in the fall

Dryo pt e r i dac e a e  5 5

4a. First pair of leafules of basal leaflet nearly opposite, not more than 4 mm apart, the basiscopic leafule 2 (–3) times the length of the acroscopic one [Fig. 20]; blade ovate-lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. carthusiana 4b. First pair of leafules of basal leaflet offset 4–15 mm, the basiscopic leafule (2–) 3–5 times the length of the acroscopic one [Fig. 19]; blade ovate-deltate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. campyloptera 2b. Leaf blades 1.5- to 2-times divided near base; leaf segments with or without bristletipped teeth 5a. Sori borne at or near margins of leaf segments; leaf blades blue-green; petioles with a dense tuft of white-brown scales at the base . . . . . . . . . . . . . . . . . . . . . . . . D. marginalis 5b. Sori borne between margins and midrib of leaf segments; leaf blades green; petioles with scattered light to dark brown scales 6a. Plants with scales of 2 distinct types—one broad, one hair-like; scales abundant on petiole, rachis, and costae; leaves with 20–30 pairs of leaflets; petiole length up to 0.25 times the length of the blade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. filix‑mas 6b. Plants with scales of various sizes, mostly broad, but not of 2 distinct sizes; scales abundant on only the petiole; leaves with 10–25 pairs of leaflets; petiole length 0.25–0.35 times the length of the blade 7a. Scales of the petiole dark brown to black, with a pale border; sori borne close to midrib; basal leaflets ovate-oblong; teeth of leaf segments not bristle-tipped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. goldiana 7b. Scales of the petiole light brown (sometimes dark brown in the center in D. clintoniana); sori borne midway between margin and midrib; basal leaflets triangular to elongate-triangular; teeth of leaf segments bristle-tipped 8a. Fertile leaves deciduous, their leaflets twisted, parallel to the horizon, noticeably larger and more erect than the persistent sterile leaves; basal leaflets triangular to broad-triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cristata 8b. Fertile leaves persistent, their leaflets not twisted, parallel to the leaf axis, only slightly different from the sterile leaves; basal leaflets narrow-triangular to oblong-triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. clintoniana 1. Dryopteris campyloptera (Kunze) Clarkson N Fig. 19 mountain wood fern. Dryopteris austriaca (Jacq.) Woynar ex Schinz & Thellung; D. spinulosa (O.F. Muell.) Watt var. americana (Fischer ex Kunze) Fern. • CT, MA, ME, NH, VT; also reported from RI by Montgomery and Wagner (1993), but voucher specimens are unknown. Cool forests, including north-temperate, boreal, and subalpine communities; increasingly at higher elevation to the south.

Fig. 19  Lowest leaflet of Dryopteris campyloptera.

2. Dryopteris carthusiana (Vill.) H.P. Fuchs N Fig. 20 spinulose wood fern. Dryopteris austriaca (Jacq.) Woynar ex Schinz & Thellung var. spinulosa (O.F. Muell.) Fiori; D. spinulosa (O.F. Muell.) Watt • CT, MA, ME, NH, RI, VT. Mesic to hydric forests and stream shores. ‌2 × 3. Dryopteris ×benedictii Wherry is a rare wood fern hybrid known from CT, MA, NH, VT. It shows leaf blades mostly 2.5 times divided near base with ± parallel margins and an abrupt apical taper, no stipitate glands on the indusia, and sori positioned midway between the midvein and margin. ‌2 × 4. Dryopteris ×uliginosa (A. Braun ex Dowell) Druce is an uncommon wood fern hybrid known from CT, MA, ME, NH, VT. It shows somewhat narrow leaf blades (relative to length) mostly 2.5 times divided at base with triangular lower leaflets, no stipitate glands on the indusia, and sori positioned midway between the midvein and margin. Very similar to D. ×boottii, but that hybrid would show stipitate glands on the indusia.

Fig. 20  Lowest leaflet of Dryopteris carthusiana.

5 6  m o n ilop hy te s

‌ × 7. Dryopteris ×correllii W.H. Wagner is an extremely rare wood fern hybrid 2 known from VT. It has an oblong leaf blade mostly divided 2.5 times near base with an abruptly tapered apex, dark brown scales on the petiole, no glands on the indusia, and sori positioned midway between the midvein and margin. 2 ‌ × 8. Dryopteris ×triploidea Wherry is an uncommon wood fern hybrid known from CT, MA, ME, NH, RI, VT. It shows long, often outward pointed, basiscopic leafules on the lower leaflets and ± forward-curving teeth (similar to D. carthusiana), but the indusia (and often the rachis and costae) have stipitate glands, and the leaf blades are somewhat evergreen. 2 ‌ × 9. Dryopteris ×pittsfordensis Slosson is a very rare wood fern hybrid known from CT, MA, VT. It shows sori positioned closer to the margin than the midrib of the ultimate segments, and it has fairly dense scales at the base of the petiole (as do other D. marginalis hybrids). However, the hybrid is very similar to D. carthusiana in regard to overall leaf blade outline, division of segments, and margin of segments (serrate vs. entire or crenate in D. marginalis). The leaf blade texture is also thicker than in D. carthusiana, lending a semi-evergreen habit to the blade. 3. Dryopteris clintoniana (D.C. Eat.) Dowell N Clinton’s wood fern. Aspidium cristatum (L.) Sw. var. clintonianum D.C. Eat. in Gray; Dryopteris cristata (L.) Gray var. clintoniana (D.C. Eat.) Underwood • CT, MA, ME, NH, RI, VT. Wet-mesic to hydric forests and swamp margins. 3 × 4. This rare hybrid wood fern is known from CT, MA, ME, NH, VT. It may be more common than collections indicate but it is very difficult to determine due to the morphological similarity of its parent taxa. From Dryopteris clintoniana this hybrid differs in its triangular leaflets twisted parallel to the horizon and relatively narrow leaf blade. From D. cristata this hybrid differs in its relatively abrupt apical taper of the leaf blade and often shows a moderately dark brown interior region on the petiole scales. 3 × 5. This very rare wood fern hybrid is known from VT. It generally resembles Dryopteris clintoniana but has scales of two sizes on the petiole and some microscales on the costules. It can be separated from D. filix-mas by the relative paucity of scales on the rachis and costules and the relatively broader leaflets (lanceolate to triangularlanceolate vs. narrow-lanceolate or narrow-triangular-lanceolate in D. filix-mas). ‌ × 7. Dryopteris ×mickelii Peck is a rare, though sometimes locally frequent, 3 wood fern hybrid known from ME, VT. It could be confused with a robust D. clintoniana. However, unlike that species, this hybrid has oblong lower leaflets, petiole scales with dark brown central regions, and often slightly curved leafules (rather than longtriangular to oblong-triangular lower leaflets, petiole scales commonly light brown, and ± straight leafules). 3 ‌ × 8. Dryopteris ×dowellii (Farw.) Wherry is an infrequent wood fern hybrid known from CT, MA, ME, NH, VT. It resembles D. ×boottii (4 × 8) in many features but has broader leaf blades with a more abrupt taper at the apex. ‌ × 9. Dryopteris ×burgessii Boivin is a rare wood fern hybrid known from CT, MA, NH, 3 VT. It is similar to D. ×slossoniae. However, the leaflets of D. ×burgessii are not as widely spaced on the rachis, the leaf blades are relatively wider, and the apex of the blade tapers more abruptly. 4. Dryopteris cristata (L.) Gray N crested wood fern. Aspidium cristatum (L.) Sw.; Polypodium cristatum L. • CT, MA, ME, NH, RI, VT. Shrub swamps, forested swamps, and wetland margins. ‌4 × 8. Dryopteris ×boottii (Tuckerman) Underwood is a relatively common wood fern hybrid known from CT, MA, ME, NH, RI, VT. It shows stipitate glands on the indusia (and often the rachis and costae), mostly 2.5 times divided leaf blades with ± parallel margins, triangular lower leaflets, and sori positioned midway between midvein and margin.

D ryo pt e r i dac e a e   57

‌ × 9. Dryopteris ×slossoniae Wherry ex Lellinger is a somewhat rare wood fern 4 hybrid known from CT, MA, ME, NH, RI, VT. It shows sori positioned closer to margin than the midrib of the ultimate segments and fairly dense scales at the base of the petiole (as with other D. marginalis hybrids). The leaflets are relatively widely spaced on the rachis, the lower ones triangular, and the leaf blades are lanceolate with ± parallel margins (rather than leaflets more crowded on the rachis, the lower ones oblong-triangular, and the blades broad-lanceolate in D. marginalis). 5. Dryopteris filix-mas (L.) Schott ssp. brittonii Fras.-Jenk. & Widen

NC

male wood fern. Polypodium filix-mas L. • ME, NH, VT. Rich, mesic, often rocky, forests. 5 ‌ × 9. Dryopteris ×montgomeryi Fras.-Jenk. & Widen is a very rare wood fern hybrid known from ME, VT. It shows sori positioned closer to the margin than the midrib of the ultimate segments and fairly dense scales at the base of the petiole (as with other D. marginalis hybrids). However, the general aspect of the plant is very similar to D. filix-mas, including the leaf blade tapering to the base and to the apex, numerous pairs of leaflets per blade, and abundant scales on the rachis and costae. 6. Dryopteris fragrans (L.) Schott N fragrant wood fern. Polypodium fragrans L. • ME, NH, VT. Cliffs, often shaded and also often of high-pH rock. 7. Dryopteris goldiana (Hook. ex Goldie) Gray N Goldie’s wood fern. Aspidium goldianum Hook. ex Goldie • CT, MA, ME, NH, VT; also reported from RI by Montgomery and Wagner (1993), but voucher specimens are unknown. Rich, mesic, often rocky, forests. 7 × 8. This rare wood fern hybrid is known from CT, MA, VT. Overall, the nothospecies appears closer to Dryopteris intermedia on superficial examination. However, it has scales on the petiole that are dark chestnut-brown with broad, paler brown margins (rather than lacking a dark central area as in D. intermedia). The hybrid is further characterized by leaf blades that are 2.5- to 3-times pinnately compound and mostly 1.8–2.2 times as long as wide (rather than mostly 1.7–2.2 in D. goldiana and mostly 2.3–2.6 in D. intermedia), sori positioned in 1 or more rows along each side of the leafule or lobe (rather than 1 row on each side in D. goldiana and 2 or more rows on each side in D. intermedia), and indusia with stipitate-glands (glands lacking in D. goldiana and present in D. intermedia). ‌7 × 9. Dryopteris ×neowherryi W.H. Wagner is a rare wood fern hybrid known from CT, MA, ME, VT. It shows sori positioned closer to the margin than the midrib of the ultimate segments and fairly dense scales at the base of the petiole (as with other D. marginalis hybrids). The petiole scales are dark brown, and the leaf blade tapers abruptly at the apex (rather than pale petiole scales and ± gradual taper to apex in D. marginalis). 8. Dryopteris intermedia (Muhl. ex Willd.) Gray N Fig. 21 evergreen wood fern. Aspidium intermedium Muhl. ex Gray; Dryopteris austriaca (Jacq.) Woynar ex Schinz & Thellung var. intermedia (Muhl. ex Willd.) Morton; D. spinulosa (O.F. Muell.) Watt var. intermedia (Muhl. ex Willd.) Underwood • CT, MA, ME, NH, RI, VT. Forests, including rocky and high-elevation types, edges of forested wetlands. Dryopteris intermedia and D. carthusiana often grow together and are sometimes confused. In addition to those characters stated in the key, the two species can often be separated merely by examining the orientation of the marginal teeth. Those of D. intermedia are ± outward pointing, whereas those of D. carthusiana are ± forward pointing. 8 × 9. This is a relatively rare wood fern hybrid known from CT, MA, VT. It shows sori positioned closer to the margin than the midrib of the ultimate segments and fairly dense scales at the base of the petiole (as with other D. marginalis hybrids). The indusia (and often the rachis and costae) have stipitate glands, and the segment margins are serrate (unlike D. marginalis).

Fig. 21  Lowest leaflet of Dryopteris intermedia.

5 8  mo n i lop hy tes

9. Dryopteris marginalis (L.) Gray N marginal wood fern. Polypodium marginale L. • CT, MA, ME, NH, RI, VT. Rocky forests, cliff bases, talus, and steep banks.

Polystichum Reference: Wagner (1993). 1a. Leaves once-divided, the fertile leaflets contracted and reduced in size compared with the sterile leaflets; sori often confluent and nearly covering the abaxial leaflet surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. acrostichoides 1b. Leaves twice-divided in the basal portion, the fertile leaflets like the sterile and not contracted; sori distinct, not covering the entire abaxial leafule surface . . . . . . . . . . . P. braunii 1. Polystichum acrostichoides (Michx.) Schott N Christmas fern. Nephrodium acrostichoides Michx. • CT, MA, ME, NH, RI, VT. Found in a variety of mesic forest types, especially those on rocky slopes. 1‌ × 2. Polystichum ×potteri Barrington is a rare fern hybrid known from MA, ME, VT. Morphologically this nothospecies closely resembles P. braunii (i.e., the leaf blades are twice-divided in the basal portion and leaves are ± monomorphic). It is best identified by its abortive spores and prominent first acroscopic leafules (simulating the acroscopic auricle on the leaflets of P. acrostichoides). Additionally, it differs from P. braunii with its leaflets that do not diminish in size toward the base as much and its sparser petiole scales. 2. Polystichum braunii (Spenner) Fée N Braun’s holly fern. Aspidium braunii Spenner; Polystichum braunii (Spenner) Fée ssp. purshii (Fern.) Calder & Taylor; P. braunii (Spenner) Fée var. purshii Fern. • CT, MA, ME, NH, VT. Mesic forests, especially those that are rich, rocky, and/or northern. North American plants have been segregated from European populations as ssp. purshii on the basis of broader microscales. However, this difference alone appears too slight to warrant recognition as a subspecific taxon.

Equisetaceae Equisetum Equisetum hybrids are rare in New England, with the exception of two nothospecies— E. ×‌litorale and E. ×‌mackaii. Hybrids can be recognized by their white and misshapen spores (rather than green and spherical). Though spore shape is difficult to assess with a hand lens, color is not (i.e., separation of hybrid individuals from fertile parental species is possible without high magnification). Sequence data from cpDNA suggest that the division of horsetails into two genera (Equisetum and Hippochaete) creates a paraphyletic Equisetum because of the basal position of E. bogotense Kunth (Guillon 2004). Therefore, only one genus is recognized here. References: Hauke (1963, 1978, 1993). 1a. Aerial stems monomorphic, unbranched, persisting more than one year (i.e., evergreen); strobilus with an apiculus at its apex [Fig. 25]; stomates sunken below the surface, arranged in single lines on each side of stem grooves (view at 30×) 2a. Stems 14- to 50-ridged; leaves articulated to and deciduous from the sheaths; central cavity greater than 50% of the stem diameter; strobili 10–20 mm tall . . . . . . . . . E. hyemale

Eq u is e tac e a e   5 9

2b. Stems 3- to 12-ridged; leaves not articulated to the sheaths and persistent on them; central cavity up to 35% of the stem diameter; strobili 2–10 mm tall 3a. Stems tortuous, 6-ridged, 0.5–1 mm thick; sheaths with 3 leaves; central cavity absent; vallecular cavities 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. scirpoides 3b. Stems erect and straight, 3- to 12-ridged, 1–4.5 mm thick; leaves numbering the same as the ridges; central cavity evident; vallecular cavities 5–12 . . . . . . E. variegatum 1b. Aerial stems monomorphic or dimorphic (i.e., the vegetative and spore-bearing stems differing in appearance), commonly with regular whorls of branches (except in unbranched forms of E. fluviatile and E. palustre), deciduous at the end of the growing season; strobilus rounded at its apex; stomates on the surface, scattered or in bands on each side of stem grooves 4a. Strobili appearing in summer, borne on firmer, green, photosynthetic stems that persist throughout the season (i.e., stems monomorphic); length of the first internode of each branch shorter than its subtending stem sheath (or sometimes the branches absent) [Figs. 23, 24]; grooves of branches rounded (i.e., U-shaped in cross-section) 5a. Leaves numbering 12–24 per sheath, 1.5–3 mm long, dark throughout or with a narrow, white band [Fig. 23]; stems with 9–25 low ridges; central cavity 75% or more of the stem diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. fluviatile 5b. Leaves numbering 5–10 per sheath, 2–5 mm long, with prominent white margins [Fig. 24]; stems with 5–10 prominent ridges; central cavity less than 35% of the stem diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. palustre 4b. Strobili appearing in spring, borne on soft, pink to light brown stems, the green, photosynthetic stems forming or appearing later (i.e., stems dimorphic); length of the first internode of each branch equal to or exceeding its subtending stem sheath [Fig. 22]; grooves of branches channeled (i.e., C-shaped in cross-section) 6a. Leaves red-brown, cohering in 3 or 4 large groups, papery; branches of the stem compound (i.e., again branched) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. sylvaticum 6b. Leaves largely brown to black, separate or in 4 or more small groups, firm; branches of the stem simple (except sometimes in damaged plants) 7a. Reproductive stems without stomates, promptly senescing; leaves of the main axis dark at the margin or with a very thin, obscure white margin, those of the branches subulate; central cavity ca. 25% of the stem diameter; stem internodes smooth to moderately scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. arvense 7b. Reproductive stems with stomates, becoming green and branched; leaves of the main axis with a prominent white margin, those of the branches triangular; central cavity 35–50% of the stem diameter; stem internodes evidently scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. pratense

Fig. 22  Node of Equisetum arvense showing branch internodes and leaf sheath.

1. Equisetum arvense L. N Fig. 22 field horsetail. Equisetum arvense L. var. boreale (Bong.) Rupr. • CT, MA, ME, NH, RI, VT; throughout. Open ground including fields, roadsides, and lawns, stream banks, forested swamps. 1‌ × 2. Equisetum ×litorale Kuhlewein ex Rupr. is a frequent horsetail hybrid known from CT, MA, ME, NH, RI, VT. It occurs mainly on shorelines, in ditches, and in other low areas. It can be identified by its white and misshapen spores (unique for New England Equisetum), monomorphic aerial stems that usually have branches, central cavity 66–80% of the stem diameter, 7–14 subulate, dark leaves 1–3 mm long, and first internode of branches equal in length to its subtending stem sheath. 2. Equisetum fluviatile L. N Fig. 23 river horsetail. Equisetum limosum L. • CT, MA, ME, NH, RI, VT; throughout. Standing water of lakes, streams, and marshes.

Fig. 23  Node of Equisetum fluviatile showing branch internodes and leaf sheath.

60  mo n ilop hy tes

3. Equisetum hyemale L. ssp. affine (Engelm.) Calder & Taylor N tall scouring-rush. Equisetum hyemale L. var. robustum (A. Braun) A.A. Eat.; Hippochaete hyemalis (L.) Bruhin ssp. affinis (Engelm.) Holub • CT, MA, ME, NH, RI, VT; throughout. Sandy slopes and roadsides, riverbanks, and borrow pits. ‌3 × 8. Equisetum ×mackaii (Newman) Brichan is an infrequent scouring-rush hybrid known from CT, MA, NH, VT. It is found in many locations where the parental taxa are sympatric. It has 7–16 leaves at each node that are persistent or some deciduous, with dark centers and thin white margins (the white margins broad and prominent in E. variegatum). The sheaths have an apical black band or are ± black throughout, only rarely developing an ash-gray medial band (as in E. hyemale). ‌3 × E. laevigatum A. Braun. Equisteum ×ferrissii Clute is a very rare scouring-rush hybrid known from CT, MA. It is much over-reported in the region, and most collections are merely variants of E. hyemale. Like other New England members of the former subgenus Hippochaete (species 3, 6, and 8), it has an apiculate strobilus apex. The leaves number 14–32 at each node. The sheaths often flare upward and have only a thin apical black band below the articulation point of the teeth, rarely developing also a brown basal band (rather than with apical and basal black bands separated by a medial ash-gray band in E. hyemale). The stem internodes of this hybrid have low, blunt scabrules on the ridges (rather than conspicuous, sharp scabrules as in E. hyemale). 4. Equisetum palustre L. N Fig. 24 marsh horsetail. Equisetum palustre L. var. americanum Vict. • CT, MA, ME, NH, VT; scattered in southern New England. Lake and stream shores, marshes, river shore seeps, and pools. Fig. 24  Leaves of Equisetum palustre with prominent white borders.

5. Equisetum pratense Ehrh. N meadow horsetail. CT, MA, ME, NH, VT; not in southeastern New England and scattered in ME. Meadows, woodlands, riparian forests with rich soils. 6. Equisetum scirpoides Michx. N dwarf scouring-rush. Hippochaete scirpoides (Michx.) Farw. • CT, MA, ME, NH, VT; lacking in southeastern New England and extreme southern New England. Forests and swamps, often growing in cool microclimates and/or among bryophytes. 7. Equisetum sylvaticum L. N wood horsetail. Equisetum sylvaticum L. var. multiramosum (Fern.) Wherry • CT, MA, ME, NH, RI, VT; nearly throughout, lacking from the Cape Cod region of MA. Mesic and hydric forests and forest edges. 8. Equisetum variegatum Schleicher ex F. Weber & D. Mohr ssp. variegatum N Fig. 25 variegated scouring-rush. Hippochaete variegata (Schleicher ex F. Weber & D. Mohr) Bruhin • CT, MA, ME, NH, VT; scattered in southern New England.

Fig. 25  Strobilus of Equisetum variegatum with an apiculus at the apex.

Hymenophyllaceae Trichomanes Reference: Farrar (1993). 1. Trichomanes intricatum Farrar

NC

Appalachian bristle fern. CT, MA, NH, VT. Growing on basic and acidic rock in sheltered caves, pockets, and crevices.

Lyg o d i ac e a e  6 1

Lygodiaceae Lygodium The report by Nauman (1993) for ME was erroneous. 1. Lygodium palmatum (Bernh.) Sw. N American climbing fern. Gisopteris palmata Bernh. • CT, MA, NH, RI, VT. Low forests, forest edges, and swamp margins, mainly on peaty, acid soils overlying sand.

Marsileaceae Marsilea Reference: Johnson (1993a). 1. Marsilea quadrifolia L. E European water-clover. CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving water, sometimes stranded on mud.

Onocleaceae 1a. Sterile leaves deeply lobed, divided only in the basal portion if at all; blade deltate, widest at the base; venation reticulate; sori enclosed in spherical leafules . . . . . . . . . . . . . . . . Onoclea 1b. Sterile leaves once-divided; blade elliptic to narrowly oblanceolate, widest above the middle; venation free (i.e., the veins forking and not rejoining); sori enclosed in linear leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Matteuccia

Matteuccia Reference: Johnson (1993b). 1. Matteuccia struthiopteris (L.) Todaro ssp. pensylvanica (Willd.) A. & D. Löve N fiddlehead fern. Matteuccia pensylvanica (Willd.) Raymond; M. struthiopteris (L.) Todaro var. pensylvanica (Willd.) Morton; M. struthiopteris (L.) Todaro var. pubescens (Terry) Clute; Onoclea struthiopteris (L.) Hoffmann, pro parte; Pteretis pensylvanica (Willd.) Fern. • CT, MA, ME, NH, RI, VT. Rich, mesic soil of riparian and upland forests, sometimes bordering drainages.

Onoclea Reference: Johnson (1993c). 1. Onoclea sensibilis L. N sensitive fern. Onoclea sensibilis L. var. obtusilobata (Schkuhr) Torr. • CT, MA, ME, NH, RI, VT. Wet-mesic to hydric soils of fields, borders, swamps, and shorelines.

62  mo nilophy tes

Ophioglossaceae References: Clausen (1938), Wagner and Wagner (1993), Gilman (2002). 1a. Trophophores (i.e., vegetative portion of leaf blades) simple, entire; venation reticulate; sporangia borne on a compact, linear sporophore . . . . . . . . . . . . . . . . . . . . . . . . . . Ophioglossum 1b. Trophophores lobed or compound (rarely simple in B. tenebrosum), the ultimate segments entire or toothed [Figs. 26, 28]; venation forking; sporangia borne on pinnately branched sporophores [Figs. 28, 29] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Botrychium

Botrychium Botrychium is an enigmatic genus with complexes of species that are separated on very subtle characteristics. Shapes and proportions are very important for determinations. Confounding identifications are the sporadic appearance of individuals from year to year and dwarf plants (which do not display morphologies conducive to identification). It is vitally important that collections include multiple individuals in order to capture variability in the population. Analysis of DNA sequence data shows that the subdivision of Botrychium into three genera (Botrychium, Botrypus, and Sceptridium) creates a paraphyletic Botrypus (Hauk et al. 2003). Therefore, only one genus is recognized here. 1a. Trophophore 2- to 4-times ternately compound, mostly 5–25 cm long, usually as wide or wider than long, commonly sterile (i.e., without a sporophore) [Figs. 26, 27]; leaf buds pubescent 2a. Trophophore sessile or nearly so, thin and deciduous, borne high on the common stalk well above the ground; leaf sheath open; leaf bud only partly enclosed by sheathing base of leaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. virginianum 2b. Trophophore long-stalked, thick and evergreen, borne near the ground on the common stalk; leaf sheath closed; leaf bud completely enclosed by sheathing base of leaf 3a. Trophophore highly lacerate-dissected into linear segments . . (in part) B. dissectum 3b. Trophophore moderately divided into lanceolate to ovate segments 4a. Trophophore leafules regularly dissected, divided to the tip; terminal leaflet segment equal to or slightly longer than adjacent, lateral leaf segments; trophophores green in fall and winter 5a. Ultimate segments of trophophore rounded to obtusely pointed, entire to weakly crenulate, mostly 4–8 mm wide at maturity, plane, coriaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. multifidum 5b. Ultimate segments of trophophore angular, dentate, mostly 2–5 mm wide at maturity, somewhat channeled and concave abaxially in life, semiherbaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. rugulosum 4b. Trophophore leafules irregularly dissected, not divided in the apical portion [Fig. 26]; terminal leaflet segment noticeably larger than adjacent, lateral leaf segments; trophophores green or bronze to purple in fall and winter 6a. Trophophore leafules ovate, crenulate to denticulate, rounded to obtusely pointed at apex [Fig. 27]; trophophores green in fall and winter; dried roots 0.5–4 mm thick 1 cm from stem (mean=2 mm), pale gray to tan (rarely darker) and with sparse transverse circular ridges in the proximal 1–5 cm . . . . . . B. oneidense 6b. Trophophore leafules lanceolate to trullate, denticulate to lacerate, acute to acuminate at apex [Fig. 26]; trophophores usually bronze to purple in fall and winter; dried roots 1.5–5.5 mm thick 1 cm from stem (mean=2.5 mm), dark gray-brown and with abundant transverse circular ridges in the proximal 1–5 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) B. dissectum

O p h i o g loss ac e a e   63

1b. Trophophore simple or up to twice pinnately divided, mostly 2–10 cm long, often longer than wide, always accompanied by a sporophore (i.e., spore-bearing portion of leaf) [Figs. 28, 29]; leaf buds glabrous 7a. Trophophore leaflets linear to narrow-ovate, the apex obtuse to acuminate, the margins shallowly to deeply lobed; at least the tip of the trophophore reflexed in bud 8a. Trophophore oblong to ovate, with 1 main axis, usually stalked; sporophore mainly erect in bud, only the very tip curved over . . . . . . . . . . . . . . . . . . . . . . . B. matricariifolium 8b. Trophophore deltate, with 3 axes, usually ± sessile; sporophore reflexed in bud . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. angustisegmentum 7b. Trophophore leaflets nearly orbicular to obovate or flabellate, the apex rounded to truncate, the margins entire or distally few-lobed; trophophore and sporophore erect in bud or only the extreme tip of trophophore slightly inclined 9a. Trophophore very slender and simple to obscurely lobed or with evident lobes, in the latter case usually with a greater distance between the first and second pairs of leaflets than between the second and third pairs of leaflets counting from the base 10a. Lowest pair of trophophore lobes or leaflets, when present, not obviously larger than the next apical pair [Fig. 29]; lobes on well-developed plants usually squareoblong to obovate-oblong; stalk of trophophore inserted high on common stalk, usually at or above mid-height of plant [Fig. 29]; plants primarily of forested wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. tenebrosum 10b. Lowest pair of trophophore lobes or leaflets, when present, often noticeably larger and more complex than the next apical pair [Fig. 28]; lobes varying from oblong to flabellate to reniform; stalk of trophophore usually borne near base of plant (sometimes toward mid-height) [Fig. 28]; plants primarily of open areas such as fields, shores, and banks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) B. simplex 9b. Trophophore usually with evident lobes, the first and second pairs of leaflets separated by a similar or slightly longer distance than the second and third pairs of leaflets 11a. Lowest pair of trophophore leaflets (or lobes in dwarf individuals) obviously larger and more complex in most plants then next apical pair; many trophophores with a characteristically broad, rounded to obstuse, terminal segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) B. simplex 11b. Lowest pair of trophophore leaflets slightly, if at all, larger and more cleft or lobed than next apical pair; trophophores typically with small, narrow-flabellate terminal segments 12a. Plants pale glaucous-green to white-green in life; lower trophophore leaflets frequently cleft into 2 unequal lobes, the upper lobe larger; sporophore 1.5–4 times as long as the trophophore . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. pallidum 12b. Plants in life green or yellow-green to glaucous-green or blue-green; lower trophophore leaflets entire to crenate or lobed, when lobed the lobes usually nearly equal; sporophore 0.8–2 (–2.5) times as long as trophophore 13a. Trophophore leaflets overlapping to approximate, usually broad-flabellate; apical leaf segments greatly reduced in size and of a shape different from those of medial segments; medial leaflets 6–18 mm wide . . . . . . . . . . . . . . . . . B. lunaria 13b. Trophophore leaflets remote, usually cuneate to narrowly flabellate; apical leaf segments slightly reduced and more or less of similar shape as medial segments; medial leaflets 1–9 mm wide 14a. Trophophore conspicuously stalked, the stalk often equal to or longer than the distance between the lower pairs of leaflets; trophophore leaflets entire to crenulate or, less frequently, notched, the lower ones spreading to

64  m o n ilop hy tes

ascending; branches of sporophore spreading to ascending, not or barely overlapping at maturity; mature sporophore with a stalk (0.5–) 1 or more times as long as the trophophore . . . . . . . . . . . . . . . . . . . . . . . . . B. minganense 14b. Trophophore short-stalked, the stalk often shorter than the distance between the lower pairs of leaflets; trophophore leaflets entire to dentate or symmetrically lobed, the lower ones ascending; branches of sporophore ascending to erect (rarely spreading), somewhat to strongly overlapping at maturity; mature sporophore subsessile or short-stalked, the stalk usually shorter than the length of the trophophore 15a. Stalk of sporophore usually 0.25–0.5 (–1) times as long as the trophophore; trophophore green to bright yellow-green in life, not clasping to loosely clasping the sporophore at maturity, with narrowcuneate to oblong leaflets, the lowest pair sometimes with sporangia; rachis of trophophore relatively narrow, up to 0.25 times as wide as the entire trophophore; plants moderately short, commonly 10–15 cm tall, without spherical gemmae at the root bases . . . . . . . . . . . . . B. ascendens 15b. Stalk of sporophore less than 0.25 times as long as the trophophore; trophophore pale green to glaucous-green in life, loosely to tightly clasping the sporophore at maturity, with linear to narrow-cuneate segments, the lowest pair almost never with sporangia; rachis of trophophore often relatively broad, up to 0.35 times as wide as the entire trophophore; plants very short and compact, rarely taller than 6 cm, sometimes with minute, spherical gemmae at the root bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. campestre 1. Botrychium angustisegmentum (Pease & Moore) Fern. N narrow triangle moonwort. Botrychium lanceolatum (Gmel.) Angst. ssp. angustisegmentum (Pease & Moore) Clausen; B. lanceolatum (Gmel.) Angst. var. angustisegmentum Pease & Moore • CT, MA, ME, NH, RI, VT. Forests, meadows, swamp margins, and peaty openings. 2. Botrychium ascendens W.H. Wagner

NC

upswept moonwort. VT; southwestern portion of state. Meadows, open quarries, grassy roadsides. This species sporulates ca. 10–15 days later than Botrychium campestre when both species occur at the same site. 3. Botrychium campestre W.H. Wagner & Farrar

NC

prairie moonwort. VT; southwestern portion of state. Meadows, open quarries, grassy roadsides. This species sporulates ca. 10–15 days earlier than Botrychium ascendens when both species occur at the same site. 4. Botrychium dissectum Spreng. N Fig. 26 dissected grapefern. Botrychium dissectum Spreng. var. obliquum (Muhl. ex Willd.) Clute; B. obliquum Muhl. ex Willd.; Sceptridium dissectum (Spreng.) Lyon • CT, MA, ME, NH, RI, VT. Forest interiors, edges, and openings, sometimes in fields. 5. Botrychium lunaria (L.) Sw.

Fig. 26  Portion of trophophore of Botrychium dissectum showing pointed apex of leafules.

NC

common moonwort. Botrychium lunaria (L.) Sw. var. onondagense (Underwood) House; B. onondagense Underwood; Osmunda lunaria L. • ME; also reported from NH by Magee and Ahles (1999) and from VT by Wagner and Wagner (1993), but specimens are unknown; highly disjunct locations in ME. Forests, swamp margins, meadows. Gilman (2002) discussed the many erroneous reports of Botrychium lunaria from VT. Shade forms of this species are frequently confused with B. minganense. That species usually has elongate trophophore stalks (i.e., equal to or longer than the distance between the lowest pairs of leaflets) and basal leaflets that are nearly of similar size to medial leaflets. Botrychium lunaria, on the other hand, has sessile or short-stalked trophophores (i.e., the trophophore stalk is shorter than the distance between the lowest pairs of leaflets), and the basal leaflets are often noticeably larger than the medial leaflets.

Op h i o g loss ac e a e   6 5

6. Botrychium matricariifolium (Döll) A. Braun ex Koch N daisy-leaved moonwort. Botrychium lunaria (L.) Sw. var. matricariifolium Döll • CT, MA, ME, NH, RI, VT. Deciduous forests, Thuja occidentalis stands, forest edges, fields. Depauperate specimens of Botrychium matricariifolium can be difficult to distinguish from B. tenebrosum. Examining emerging plants can be useful because the tips of the sporophore and trophophore are curved over in bud (the tips are erect in B. tenebrosum and the related B. simplex). 7. Botrychium minganense Victorin

NC

Mingan moonwort. Botrychium lunaria (L.) Sw. var. minganense (Victorin) Dole • ME, VT. Forests. 8. Botrychium multifidum (Gmel.) Rupr. N leathery grapefern. Botrychium multifidum (Gmel.) Rupr. var. intermedium (D.C. Eat.) Farw.; Osmunda multifida Gmel.; Sceptridium multifidum (Gmel.) Nishida ex Tagawa • CT, MA, ME, NH, RI, VT. Chiefly in open areas, sometimes at forest edges or in forest openings. 9. Botrychium oneidense (Gilbert) House

N C Fig. 27

blunt-lobed grapefern. Botrychium dissectum Spreng. var. oneidense (Gilbert) Clute; B. multifidum (Gmel.) Rupr. var. oneidense (Gilbert) Farw.; Sceptridium oneidense (Gilbert) Holub • CT, MA, ME, NH, RI, VT. Mesic to hydric soils of forests and swamps. 10. Botrychium pallidum W.H. Wagner

NC

Fig. 27  Portion of trophophore of Botrychium oneidense showing blunt apex of leafules.

pale moonwort. ME; single location in Washington County, ME. Open fields. 11. Botrychium rugulosum W.H. Wagner

NC

St. Lawrence grapefern. Botrychium multifidum (Gmel.) Rupr. forma dentatum R. Tryon; B. ternatum, auct. non (Thunb.) Sw.; Sceptridium rugulosum (W.H. Wagner) Skoda & Holub • CT, VT. Open fields, second-growth forests, usually in regions of fine sands deposited during glacial lake periods. The report from NH by Wagner and Wagner (1993) was based on a poorly pressed Botrychium multifidum. Botrychium rugulosum emerges later at a given site than does B. multifidum. 12. Botrychium simplex E. Hitchc. N Fig. 28 least moonwort. CT, MA, ME, NH, RI, VT; nearly throughout but rare in southeastern New England. Meadows, shorelines, and open rights-of-way. Rare plants in New England show greatly enlarged and pinnately divided lower trophophore leaflets and are superficially similar to the primarily western Botrychium simplex var. compositum (Lasch) Milde (a varietal epithet that may or may not prove to properly apply to North American plants). 13. Botrychium tenebrosum A.A. Eat.

Fig. 28  Leaf of Botrychium simplex.

N C Fig. 29

swamp moonwort. Botrychium simplex E. Hitch. var. tenebrosum (A.A. Eat.) Clausen • CT, MA, ME, NH, VT. Swamps, streambanks, shaded forests. 14. Botrychium virginianum (L.) Sw. N rattlesnake fern. Botrypus virginianus (L.) Holub; Osmunda virginiana L. • CT, MA, ME, NH, RI, VT. Forests, usually rich, mesic types.

Ophioglossum 1. Ophioglossum pusillum Raf.

NC

northern adder’s-tongue fern. Ophioglossum vulgatum L. var. pseudopodum (Blake) Farw. • CT, MA, ME, NH, RI, VT. Wet-mesic to hydric open areas such as swales, meadows, boggy fields, and ditches, only rarely in shade.

Fig. 29  Leaf of Botrychium tenebrosum.

66  mo n i lop hy tes

Osmundaceae The report of Osmunda claytoniana × Osmundastrum cinnamomeum by Sorrie and Somers (1999) was based on a collection of Osmunda claytoniana—Ahles 83707 (MASS!). Reference: Whetstone and Atkinson (1993). 1a. Fertile leaves with monomorphic leaflets, borne on a blade separate from the vegetative leaves; axil of leaflets on vegetative leaves bearing a tuft of white hairs (these turning light brown with age) on the abaxial side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmundastrum 1b. Fertile leaves with dimorphic leaflets—some leaflets bearing sporangia and some not; axil of vegetative leaflets without a tuft of hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmunda

Osmunda 1a. Fertile leaflets produced at the apex of the blade; leaflets divided . . . . . . . . . . . . . O. regalis 1b. Fertile leaflets produced in the middle of the blade, with sterile leaflets basal and apical; leaflets lobed, but not divided to the costae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. claytoniana 1. Osmunda claytoniana L. N interrupted fern. CT, MA, ME, NH, RI, VT; throughout and common. Mesic to hydric soils of forests, swamps, and shorelines. 1‌ × 2. Osmunda ×ruggii R. Tryon is an extremely rare fern hybrid known from CT (only two populations, CT and VA, reported from the world by Whetstone and Atkinson 1993). Only vegetative leaves have been seen in the wild. These are twice pinnately divided with sessile leafules (the leaflets of O. regalis are short-stalked). The report of ‌ruggii O. × from NH by Thorne and Thorne (1989) requires confirmation (photos are stored at VT). 2. Osmunda regalis L. var. spectabilis (Willd.) Gray N royal fern. CT, MA, ME, NH, RI, VT; throughout and common. Wet-mesic to hydric soils of swamps, riparian forests, and shorelines.

Osmundastrum 1. Osmundastrum cinnamomeum (L.) C. Presl N cinnamon fern. Osmunda cinnamomea L. • CT, MA, ME, NH, RI, VT; throughout and common. Mesic to hydric soils of forests, swamps, and shorelines.

Polypodiaceae Polypodium Hybridization has been an important force in Polypodium. It has led to the formation of several allopolyploid species, including our P. virginianum (the hybrid-derived species from P. appalachianum and P. sibiricum Sipl.). Though hybridization continues to be an important source of confusion, variation within the species is the major hurdle to understanding the identity of our taxa. Collectors are encouraged to gather reproductive specimens so that spore condition can be observed (fertile vs. abortive). The species show monomorphic and slightly lustrous spores compared with the nothospecies, which has highly polymorphic and dull spores. Reference: Haufler et al. (1993b).

Po ly p o d i ac e a e  67

1a. Sporangiasters more than 40 per sorus; scales of rhizome and petiole golden brown; spores smaller than 52 μm in diameter; leaflet apex usually pointed; leaf blades tending to be more triangular, widest near the base [Fig. 30] . . . . . . . . . . . . . . . . . . . . . . . . . P. appalachianum 1b. Sporangiasters fewer than 40 per sorus; scales of rhizome and petiole golden brown with a dark brown central strip; spores larger than 58 μm in diameter; leaflet apex usually rounded; leaf blades tending to be more oblong, widest near the middle [Fig. 31] . . . . . . P. virginianum 1. Polypodium appalachianum Haufler & Windham N Fig. 30 Appalachian polypody. CT, MA, ME, NH, RI, VT. On rock or thin soil over rock of cliffs, boulders, and talus. The name Polypodium virginianum forma acuminatum (Gilbert) Fern. was sometimes applied to this species in regional herbaria. 1‌ × 2. Polypodium ×incognitum Cusik is a rare polypody hybrid in New England known from CT, MA, ME, NH, RI, VT. This nothospecies is an abortive-spored triploid (from the diploid P. appalachianum and the tetraploid P. virginianum). It is best identified by its intermediate morphology (or combination of characters) and spore examination. 2. Polypodium virginianum L. N Fig. 31 rock polypody. Polypodium vulgare L. var. virginianum (L.) D.C. Eat. • CT, MA, ME, NH, RI, VT. On rock or thin soil over rock of cliffs, boulders, and talus.

Fig. 30  Leaf blade of Polypodium appalachianum.

Pteridaceae 1a. Petiole apparently forking into two main divisions; sori borne on the surface of a false indusium, this best viewed through dissection; ultimate segments without a distinct midrib . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adiantum 1b. Petiole not appearing to fork into two divisions; sori borne on the surface of the leaf blade near margin and covered by a false indusium; ultimate segments with a midrib 2a. Leaf blade tissue pubescent with septate hairs; ultimate segments of reproductive leaves 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilanthes 2b. Leaf blade tissue not pubescent (though the petioles and rachises may be pubescent); ultimate segments of reproductive leaves 5–75 mm long 3a. Leaf blades thin, herbaceous to membranaceous, withering by late summer; rhizome scales colorless; leaves dimorphic—the reproductive ones with more elongate and relatively narrower segments compared with the vegetative ones; veins not reaching margins of ultimate segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptogramma 3b. Leaf blades thick, coriaceous, persistent; rhizome scales red-brown; leaves monomorphic to hemidimorphic (e.g., the reproductive leaves somewhat longer and more divided than the vegetative leaves in P. atropurpurea); veins reaching margins of ultimate segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pellaea

Adiantum Adiantum hispidulum Sw. was reported from CT by Paris (1993), but specimens are unknown. References: Paris (1991, 1993). 1a. Ultimate segments of leaves ± oblong, rounded to obtuse at the apex [Fig. 32], in the same plane with the axis of the blade, thin-herbaceous to herbaceous, bright green to yellow-green; leaf blades lax and arching; plants primarily of rich, mesic forests . . . . . . . . . . . . . . . . A. pedatum 1b. Ultimate segments of leaves ± triangular, acute to obtuse at the apex [Fig. 33], twisted out of the plane of the axis of the blade (but usually in plane with the blade axis in shaded

Fig. 31  Leaf blade of Polypodium virginianum.

68  mo n ilop hy te s

plants), firm-herbaceous to chartaceous, medium green to blue-green (often bright green in the shade); leaf blades usually stiff and upright (lax in shaded plants); plants of serpentine rocks and soil 2a. Medial ultimate segments of leaves on stalks 0.3–1.3 mm long, most stalks shorter than 0.9 mm; false indusia 0.8–2.9 (–3.4) mm long; rhizomes frequently branching and congested, with internodes 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. aleuticum 2b. Medial ultimate segments of leaves on stalks (0.4–) 0.6–1.5 (–1.9) mm long, most stalks longer than 0.9 mm; false indusia 2–5 (–10) mm long; rhizomes occasionally branching (rarely congested), with internodes 4–7.5 mm long . . . . . . . . . . . . . . . . . . . A. viridimontanum 1. Adiantum aleuticum (Rupr.) Paris

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western maidenhair fern. Adiantum pedatum L. ssp. aleuticum (Rupr.) Calder & Taylor; A. pedatum L. var. aleuticum Rupr.;  A. pedatum L. ssp. calderi Cody • ME, VT. Serpentine cliffs and talus. Fig. 32  Ultimate segment of leaf of Adiantum pedatum showing false indusium.

2. Adiantum pedatum L. N Fig. 32 northern maidenhair fern. CT, MA, ME, NH, RI, VT. Rich, mesic forests. 3. Adiantum viridimontanum Paris

N C Fig. 33

Green Mountain maidenhair fern. ME, VT. Serpentine cliffs and talus, thin soil of woodlands and forest edges on serpentine bedrock.

Cheilanthes Reference: Windham and Rabe (1993). 1. Cheilanthes lanosa (Michx.) D.C. Eat.

NC

hairy lip fern. Cheilanthes vestita (Spreng.) Sw. • CT; western portion of state. Cliffs and talus.

Fig. 33  Ultimate segment of leaf of Adiantum viridimontanum showing false indusium.

Cryptogramma Reference: Alverson (1993). 1. Cryptogramma stelleri (Gmel.) Prantl N slender rock-brake. CT, MA, ME, NH, VT. High-pH rocks and cliffs.

Pellaea Reference: Windham (1993a). 1a. Petiole, rachis, and abaxial surface of ultimate segments pubescent with short, curly hairs; leaflet stalks emerging at right angles to the rachis; ultimate segments of fertile leaves mostly 10–75 mm long; leaves hemidimorphic—the fertile leaves larger and with longer ultimate segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. atropurpurea 1b. Petiole and rachis glabrous or nearly so, the abaxial surface of ultimate segments glabrous except for occasional hair-like scales near the midrib; leaflet stalks ascending; ultimate segments of fertile leaves mostly 5–20 mm long; leaves essentially monomorphic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. glabella 1. Pellaea atropurpurea (L.) Link N purple cliff-brake. Pteris atropurpurea L. • CT, MA, NH, RI, VT. High-pH cliffs, rarely on mortar of stone walls. 2. Pellaea glabella Mett. ex Kuhn ssp. glabella N slender cliff-brake. CT, MA, VT. High-pH cliffs, sometimes on roadcuts and mortar of stone and brick walls. The MA occurrence was collected from human-created substrate (a brick wall) in an urban setting.

s a lv i n i ac e a e   6 9

Salviniaceae Salvinia 1. Salvinia minima Baker E common watermoss. MA. Still or slow-moving water of ponds, streams, and canals. The report of Salvinia natans (L.) Allioni in Norfolk County, MA, by Angelo and Boufford (1996) was based on collections originally determined as Salvinia rotundifolia Willd. (a misapplied name)— 13 Aug 1941, Coffin s.n. (GH!, NEBC!). The specimens are S. minima, as evidenced by the leaf blades abaxially with hairs 2–3 mm long that have apically distinct branches. A collection from Middlesex County, MA, determined as S. natans, was taken from the Harvard University Greenhouses (i.e., it was taken from cultivated plants)—Jun 1966, Tryon s.n. (GH!).

Thelypteridaceae 1a. Leaf blades ± triangular; some of the basal leaflets connected by a wing of tissue along the rachis; costae not grooved adaxially; indusium absent . . . . . . . . . . . . . . . . . . . . . Phegopteris 1b. Leaf blades narrow-lanceolate to elliptic; none of the basal leaflets connected by a wing of tissue; costae grooved adaxially; indusium present 2a. Abaxial surface of costae without scales; veins of ultimate segments not forking near margin; leaf blades with either red to orange resinous glands (e.g., P. simulata) or greatly reduced lower leaflets (e.g., P. noveboracensis) . . . . . . . . . . . . . . . . . . . . . . . . . Parathelypteris 2b. Abaxial surface of costae with small, tan scales; veins of ultimate segments of vegetative blades usually forking near margin; leaf blades lacking both red to orange glands and greatly reduced lower leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelypteris

Parathelypteris Reference: Smith (1993). 1a. Leaf blades elliptic, gradually reduced in width through the basal half of the blade, lacking red to orange resin glands; indument of rachis, costae, and costules composed of hairs up to 1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. noveboracensis 1b. Leaf blades lanceolate to broad-lanceolate, only somewhat reduced in the basal portion, with numerous, red to orange resin glands; indument of rachis, costae, and costules of hairs 0.2–0.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. simulata 1. Parathelypteris noveboracensis (L.) Ching N New York fern. Dryopteris noveboracensis (L.) Gray; Polypodium noveboracense L.; Thelypteris noveboracensis (L.) Nieuwl. • CT, MA, ME, NH, RI, VT. Mesic to wet-mesic soils of forests, frequently associated with stream channels, seeps, and swamp borders. 2. Parathelypteris simulata (Davenport) Holttum N Massachusetts fern. Dryopteris simulata Davenport; Thelypteris simulata (Davenport) Nieuwl. • CT, MA, ME, NH, RI, VT. Low-pH soil of swamps, peatlands, and mixed shrub-graminoid wetlands, often found in areas with high proportion of bryophyte cover (especially Sphagnum).

70  mon i lophy tes

Phegopteris Reference: Smith (1993). 1a. Basal leaflets mostly 10–20 cm long, connected to the adjacent pair by a wing; scales on costae narrowly lanceolate, white to light brown, usually 3–5 cells wide at the base; larger leaflets with evidently lobed segments; middle leaflet veins forked or pinnate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. hexagonoptera 1b. Basal leaflets mostly 3–10 cm long, not connected to next pair; scales on costae ovatelanceolate, brown, usually 6–12 cells wide at the base; larger leaflets with entire or crenate segments; middle leaflet veins often simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. connectilis 1. Phegopteris connectilis (Michx.) Watt N long beech fern. Dryopteris phegopteris (L.) C. Christens.; Thelypteris phegopteris (L.) Slosson • CT, MA, ME, NH, RI, VT. Forests, stream banks, ravines, cliff bases, and base of talus slopes. Two forms of this species occur in New England. In the typical form (triploid), the lowest pair of leaflets are usually out of plane with the remaining leaflets and typically are projected downward and form an angle of 45–120 degrees between them. In the tall form (tetraploid), the lowest pair of leaflets usually are ± in plane with the remaining leaflets being only slightly projected downward and form an angle nearly or fully 180 degrees between them. When these two forms occur together (as they frequently do), the tall form is more robust with larger blades and thicker petioles. Preliminary studies showed the tall form is not a hybrid of our two species of beech fern (Driscoll et al. 2003). It is known from ME and VT within New England (though likely overlooked and present in other states). 2. Phegopteris hexagonoptera (Michx.) Fée N broad beech fern. Dryopteris hexagonoptera (Michx.) C. Christens.; Thelypteris hexagonoptera (Michx.) Weatherby • CT, MA, ME, NH, RI, VT. Mesic soils of forests and borders of swamps, found in progressively richer soils to the north.

Thelypteris Reference: Smith (1993). 1. Thelypteris palustris Schott var. pubescens (G. Lawson) Fern. N marsh fern. Dryopteris thelypteris (L.) Gray var. pubescens (G. Lawson) Weatherby; Thelypteris palustris Schott var. haleana Fern. • CT, MA, ME, NH, RI, VT. Swamps, peatlands, marshes, shorelines, and wet ditches.

Woodsiaceae 1a. Indusium inferior, composed of lacerate or filamentous segments at maturity; veins not reaching margin of leaf segments, ending in hydathodes near the margin [Fig. 40]; petioles articulated near base, this recognizable as a small, swollen node (not articulated in W. obtusa) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Woodsia 1b. Indusium absent or present, sometimes inconspicuous, superior when present, attached centrally or laterally, not divided; veins reaching margin of leaf segments [Fig. 36]; petioles not articulated near base 2a. Indusium absent; leaf blades triangular to pentagonal [Fig. 39]; rhizome long and slender; leaflets weakly articulated to rachis; basal leaflets noticeably larger than nextapical pair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnocarpium

Wo o ds i ac e a e  71

2b. Indusium present, though inconspicuous in Cystopteris; leaf blades lanceolate to triangular; rhizome relatively short; leaflets not articulated to rachis; basal leaflets not or only slightly larger than next-apical pair 3a. Indusium ovate to lanceolate, short, inconspicuous, attached laterally and arched over sorus; vascular bundles at the base of the petiole round or oblong in cross-section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cystopteris 3b. Indusium linear, hooked at one end, or horseshoe-shaped, elongate, evident; vascular bundles at the base of the petiole lunate in cross-section 4a. Sori hooked at one end, some crossing the veinlets [Fig. 34]; indument of rachis and costae usually of stipitate glands (rarely indument absent) . . . . . . . . . . . Athyrium 4b. Sori straight or slightly falcate, not crossing the veinlets [Fig. 38]; indument of rachis and costae absent or of septate hairs only 5a. Leaf blade 1.5- to 2-times divided; petiole with 2 rows of sharp or blunt marginal teeth on the swollen base (note: this feature absent in many collections but visible in the field) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Deparia 5b. Leaf blade once divided; petiole base lacking marginal teeth . . . . . . Diplazium

Athyrium Reference: Kato (1993a). 1a. Leaf variegated, such that the distal portion of the leaflets (or much of the plant) suffused with gray to silver . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. nipponicum 1b. Leaf not variegated, the blade green 2a. Petiole scales brown to black-brown; leaf blades elliptic, narrowed to the base, broadest near or just below the middle; leaflets sessile or short-stalked; leafules linear to oblong; indusia ciliate with non-glandular hairs (though the sporangial stalks do have glandular hairs); spores yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. angustum 2b. Petiole scales light brown to brown; leaf blades broad-lanceolate to lanceolate, only slightly narrowed to the base, broadest just above the base; leaflets generally stalked; leafules oblong-lanceolate to narrow-triangular; indusia ciliate with glandular or nonglandular hairs; spores dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. asplenioides 1. Athyrium angustum (Willd.) C. Presl N Fig. 34 narrow lady fern. Athyrium filix-femina (L.) Roth ssp. angustum (Willd.) Clausen; A. filix-femina (L.) Roth var. angustum (Willd.) Lawson; A. filix-femina (L.) Roth var. michauxii (Spreng.) Farw.; A. filix-femina (L.) Roth var. rubellum Gilbert • CT, MA, ME, NH, RI, VT. Mesic to hydric forests and wet-mesic open areas. 2. Athyrium asplenioides (Michx.) Desv.

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southern lady fern. Athyrium filix-femina (L.) Roth ssp. asplenioides (Michx.) Hultén; A. filix-femina (L.) Roth var. asplenioides (Michx.) Farw.; Nephrodium asplenioides Michx. • CT, MA, RI; also reported from VT by Seymour (1982), but voucher specimens are unknown. Mesic to hydric forests. 3. Athyrium nipponicum (Mett.) Hance E Japanese painted lady fern. Asplenium nipponicum Mett. • RI. Gardens, walkways. This species has several morphological features in common with Athyrium asplenioides, including relatively light color of petiole scales, shape of leaf blade (broadest near base), and petiolulate leaflets.

Fig. 34  Leafule of Athyrium angustum showing the sori.

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Cystopteris Cystopteris is a difficult genus due to hybridization (past and present) and the occasional presence of fertile, but stunted, plants in stressful habitats. Hybrids are best determined by examination of spores, which are usually abortive. In order to correctly determine stunted plants, specimens should include rhizomes and complete petioles, as well as carefully preserved leaf blades. Further, for species superficially similar to C. fragilis, it is best to have several collections to adequately capture variation within the population. Reference: Haufler et al. (1993a). 1a. Rachises, costae, and indusia sparsely to densely stipitate-glandular; bulbils often present on the abaxial blade surface (these rare and misshapen in C. laurentiana); leaf blades widest at or near base 2a. Leaf blade triangular to long-triangular; bulbils frequently present; leaf rachis, costae, costules, and ultimate segments usually with dense glandular hairs; spores mostly 33–38 μm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bulbifera 2b. Leaf blade ovate to narrow-ovate; bulbils only rarely present, misshapen; leaf rachis, costae, costules, and ultimate segments with sparse glandular hairs (these often absent from herbarium specimens); spores mostly 49–60 μm in diameter . . . . . . . . C. laurentiana 1b. Rachises, costae, and indusia without glandular hairs; plants lacking bulbils; blade lanceolate to elliptic 3a. Leaves clustered 1–4 cm behind the apex of the rhizome; rhizome pubescent, especially toward apex, with yellow hairs; leafules, especially those of the basal medial portion of the blade, borne on evident stalks; petiole green to stramineous, with dark pigmentation only at the very base; spores mostly 28–34 μm in diameter . . . . . . . . . . . . . . . . . . . . . . C. protrusa 3b. Leaves clustered near rhizome apex; rhizome lacking hairs; leafules varying from sessile to short-stalked; petiole with dark brown color not confined to the base, the pigmentation extending apically for a noticeable distance; spores mostly 39–60 μm in diameter 4a. Basal leafules sessile or nearly so, broadly tapering or rounded at the base, approximate, emerging nearly perpendicular to the costa [Fig. 35]; apical portion of blade with deltate to ovate leaflets; margins of leafules serrate; acroscopic margin of leaflet convex due to the fact that the costa of the leaflet is nearly straight; indusia up to 1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. fragilis 4b. Basal leafules short-stalked, mostly cuneate at the base, remote, emerging at an acute angle to the costa (i.e., ascending) [Fig. 37]; apical portion of blade with ovate to narrowly elliptic leaflets; margins of leafules crenate; acroscopic margin of leaflet concave due to the fact that the costa of the leaflet curves slightly upward; indusia up to 0.5 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tenuis Fig. 35  Veins of Cystopteris fragilis, which reach the margin.

1. Cystopteris bulbifera (L.) Bernh. N bulbil fragile fern. Filix bulbifera (L.) Underwood; Polypodium bulbiferum L. • CT, MA, ME, NH, RI, VT. On rock or thin soil over high-pH substrate. 2. Cystopteris fragilis (L.) Bernh. N Fig. 35, 36 fragile fern. Cystopteris dickieana Sim; Polypodium fragile L. • CT, MA, ME, NH, VT. On rock or thin soil over rock. 2 × 5. Recognition of this nothospecies is generally missed during floristic inventories; it is most commonly misidentified as Cystopteris tenuis. It is intermediate between the two parental taxa, though on average larger than either, and possesses abortive spores. 3. Cystopteris laurentiana (Weatherby) Blasdell

Fig. 36  Leaflet of Cystopteris fragilis showing orientation and base of leafules.

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Laurentian fragile fern. Cystopteris fragilis (L.) Bernh. var. laurentiana Weatherby • ct, MA, VT. Cliffs and ledges, usually high-pH substrate.

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4. Cystopteris protrusa (Weatherby) Blasdell

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southern fragile fern. Cystopteris fragilis (L.) Bernh. var. protrusa Weatherby • CT; also reported from MA by Haufler et al. (1993a), but voucher specimens are unknown. On soil in rich, mesic forests. 5. Cystopteris tenuis (Michx.) Desv. N Fig. 37 Mackay’s fragile fern. Cystopteris fragilis (L.) Bernh. var. mackayi Lawson • CT, MA, ME, NH, RI, VT. On boulders and thin soil over rock, occasionally on cliff faces.

Deparia Reference: Kato (1993b). 1. Deparia acrostichoides (Sw.) M. Kato N Fig. 38

Fig. 37  Leaflet of Cystopteris tenuis showing orientation and base of leafules.

silvery false spleenwort. Asplenium acrostichoides Sw.; Athyrium thelypterioides (Michx.) Desv.; Diplazium acrostichoides (Sw.) Butters • CT, MA, ME, NH, RI, VT. Mesic forests, often rich and/or rocky.

Diplazium Reference: Kato (1993c). 1. Diplazium pycnocarpon (Spreng.) M. Broun N narrow-leaved glade fern. Asplenium pycnocarpon Spreng.; Athyrium pycnocarpon (Spreng.) Tidestrom • CT, MA, NH, VT; also reported from RI by Kato (1993c), but specimens are unknown. Rich, mesic forests. Fig. 38  Leafule of Deparia achrostichoides showing the sori.

Gymnocarpium Hybrids in this genus, which are likely underreported, often possess two types of spores— smaller, black, malformed spores and larger, brown, round spores. Hybrids are not confined to areas of parental sympatry. References: Pryer (1993), Pryer and Haufler (1993). 1a. Leaf stipitate-glandular on petiole, rachis, and abaxial blade surface; basal leaflets and basiscopic leafules curving toward apex of plant [Fig. 39] . . . . . . . . . . . . . . . . . . . . . G. jessoense 1b. Leaf essentially glabrous; basal leaflets and basiscopic leafules spreading, not curving toward plant apex (except in some epipetric populations) . . . . . . . . . . . . . . . . . . . . G. dryopteris 1. Gymnocarpium dryopteris (L.) Newman N northern oak fern. Carpogymnia dryopteris (L.) A. & D. Löve; Dryopteris dryopteris (L.) Britt.; Polypodium dryopteris L. • CT, MA, ME, NH, RI, VT. Forests, base of cliffs, talus slopes. 1‌ × G. disjunctum (Rupr.) Ching. Gymnocarpium ×brittonianum (Sarvela) Pryer & Haufler is a rare oak fern hybrid known from CT, MA, ME, NH, VT. Morphologically it most closely resembles G. disjunctum, with the first pair of leafules of the second pair of leaflets markedly unequal in length (the basiscopic leafules much longer than the acroscopic leafules; those of G. dryopteris are usually ± equal or the basiscopic leafules slightly longer). Spore morphology is an important character for diagnosing this hybrid. Both round and abortive spores are present. Distribution of this nothospecies is based on Pryer and Haufler (1993). Given that G. disjunctum is restricted to northwestern North America, it is plausible that plants seen with dimorphic spores are the result of a different hybrid combination or a separate phenomenon altogether. 2. Gymnocarpium jessoense (Koidzumi) Koidzumi ssp. parvulum Sarvela

N C Fig. 39

Nahanni oak fern. Gymnocarpium continentale (Petrov) Pojark • VT. High-pH rock of cliffs and talus slopes.

Fig. 39  Leaf blade of Gymnocarpium jessoense.

74 m on ilophy tes

Woodsia Reference: Windham (1993b). 1a. Petioles not articulated, therefore the persistent petiole bases of various lengths; leafules dentate with pointed teeth; petiole scales bicolored, with a dark central region; leaf blades (8–) 20–60 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. obtusa 1b. Petioles articulated, therefore the persistent petiole bases mostly of equal length, the articulation point noticeable as a slightly swollen node; leafules entire to crenate; petiole scales concolored; leaf blades 2.5–22 (–25) cm long 2a. Plants glabrous, with sessile glands; basal leaflets flabellate; leaf blades 1–1.5 cm wide; petioles green to light brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. glabella 2b. Plants with hairs, scales, and/or stalked glands; basal leaflets narrowly ovate to deltate; leaf blades 1–3 cm wide; petioles brown to dark brown 3a. Leaflets glabrous or nearly so on the abaxial surface (though scales may be present on the rachis and costae), the larger leaflets with 2 or 3 pairs of leafules; indusium composed of few filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. alpina 3b. Leaflets with scales on the abaxial surface, the larger leaflets with 4–7 pairs of leafules; indusium composed of many filaments . . . . . . . . . . . . . . . . . . . . . . . . . . W. ilvensis 1. Woodsia alpina (Bolton) S.F. Gray

NC

northern cliff fern. Acrostichum alpinum Bolton; Woodsia alpina (Bolton) S.F. Gray var. bellii Lawson; W. glabella R. Br. ex Richards. var. bellii (Lawson) Lawson • ME, VT; also reported from NH by Windham (1993b), but specimens are unknown. High-pH rock of cliffs. An allotetraploid derived from Woodsia glabella and W. ilvensis. Forms of this species with relatively few scales and hairs are sometimes confused with W. glabella. They can be separated by their petioles that are red-brown to purple near the base (vs. green or yellow-brown throughout in W. glabella). 1‌ × 3. Woodsia ×gracilis (Lawson) Butters is a rare, abortive-spored hybrid. Compared with W. alpina, this hybrid fern tends to be larger, with more hairs and scales, and with leaflets more divided and with more pairs of leafules (often more than 3). Compared with W. ilvensis, this nothospecies tends to be smaller, with fewer hairs and scales, and with leaflets less divided. 2. Woodsia glabella R. Br. ex Richards.

NC

smooth cliff fern. Woodsia alpina (Bolton) S.F. Gray var. glabella (R. Br. ex Richards.) D.C. Eat.; W. hyperborea R. Br. var. glabella (R. Br. ex Richards.) Watt • MA, ME, NH, VT. High-pH rock of cliffs. Fig. 40  Veins of Woodsia ilvensis, which end in hydathodes.

3. Woodsia ilvensis (L.) R. Br. N Fig. 40 rusty cliff fern. Acrostichum ilvense L. • CT, MA, ME, NH, RI, VT. Cliffs and talus slopes, including xeric substrates. 4. Woodsia obtusa (Spreng.) Torr. ssp. obtusa N blunt-lobed cliff fern. Physematium obtusum (Spreng.) Hook.; Polypodium obtusum Spreng. • CT, MA, ME, NH, RI, VT. Cliffs and rocky slopes, predominantly on high-pH substrate.

75

Gymnosperms Cupressaceae 1a. Seed cones fleshy and resembling a firm berry, the scales not opening [Fig. 43]; seeds not winged; abaxial glands on leaves elliptic to elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . Juniperus 1b. Seed cones woody, the scales opening and exposing the axis; seeds winged; abaxial glands on leaves circular or oval to elliptic 2a. Branchlets terete or tetragonous; alternating pairs of leaves similar; seed cones nearly spherical, with peltate scales; abaxial leaf glands circular [Fig. 41] . . . . . . . . Chamaecyparis 2b. Branchlets flat; alternating pairs of leaves dimorphic—those of the upper and lower rank flat, those of the lateral ranks folded; seed cones ellipsoidal, with basifixed scales; abaxial leaf glands oval to elliptic [Fig. 42] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thuja

Chamaecyparis Reference: Michener (1993). 1. Chamaecyparis thyoides (L.) B.S.P. N Fig. 41 Atlantic white cedar. Cupressus thyoides L. • CT, MA, ME, NH, RI. Acidic swamps and bogs of the Atlantic coastal plain.

Juniperus Report of Juniperus communis × J. virginiana from ME by Angelo and Boufford (1996) was based on specimens of J. virginiana with a high proportion of subulate (i.e., juvenile) leaves. Reference: Adams (1993). 1a. Leaves whorled, not concealing the branchlet, monomorphic; seed cones axillary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. communis 1b. Leaves opposite, closely imbricate and concealing the young branchlets, dimorphic— longer, subulate ones (i.e., juvenile) and shorter, scale-like ones (i.e., adult); seed cones borne terminally 2a. Plants prostrate shrubs with erect branches, up to 0.3 m high; strobilus with a usually curved stalk, 6–8 (–10) mm long, containing 3–5 seeds; apex of scale-like leaves apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. horizontalis 2b. Plants upright shrubs or trees, to 15 m tall (rarely depressed in exposed situations); strobilus with a usually straight stalk, 3–6 (–8) mm long, containing 1 or 2 (–3) seeds; apex of scale-like leaves merely acute or with a very obscure apiculus . . . . . . . . . . . J. virginiana 1. Juniperus communis L. var. depressa Pursh N common juniper. Juniperus canadensis Lodd. ex Burgsd.; J. communis L. ssp. depressa (Pursh) Franco; J. depressa (Pursh) Raf. • CT, MA, ME, NH, RI, VT. Fields, roadsides, abandoned pastures, and rocky slopes, sometimes ascending high into the mountains. Forms of this species occurring above treeline have sometimes been identified as Juniperus communis var. saxatilis Pallas (a taxonomic synonym of J. communis var. montana Ait.). Plants from New England’s

Fig. 41  Nearly circular abaxial leaf glands of Chamaecyparis thyoides.

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higher peaks are not referable to this taxon because the stomatal bands, though somewhat wider than the green bands, are not within the range of var. montana (which is restricted to Greenland in eastern North America). 2. Juniperus horizontalis Moench N creeping juniper. Juniperus prostrata Pers.; J. repens Nutt.; J. virginiana L. var. prostrata (Pers.) Torr.; Sabina horizontalis (Moench) Rydb. • MA, ME, NH, VT. Sandy or rocky fields, coastal headlands and cliffs, mainly along the coastal plain, rare inland. George (1997) reported this species from Jamestown, RI, based on a collection by Richard Champlin. The only specimen identified as such by Champlin seen by me was collected in Newport, but it was Juniperus virginiana—19 Jun 1983, Champlin s.n. (Champlin Herb.). 2 × 3. This rare hybrid juniper is known from ME, NH. It shows intermediacy in discriminating characters, such as habit, seed cone size, peduncle morphology, and number of seeds (see identification key). Fig. 42  Elliptic abaxial leaf glands of Juniperus virginiana.

3. Juniperus virginiana L. var. virginiana N Fig. 42, 43 eastern red cedar. Juniperus virginiana L. ssp. crebra (Fern. & Grisc.) E. Murr.; J. virginiana L. var. crebra Fern. & Grisc.; Sabina virginiana (L.) Antoine • CT, MA, ME, NH, RI, VT. Dry fields and hillsides, woodlands, and forest openings in sandy or rocky soils.

Thuja Reference: Chambers (1993). 1. Thuja occidentalis L. N northern white-cedar. CT, MA, ME, NH, RI, VT; throughout most of northern New England, but becoming rare and local to the south. Shorelines, fens, swamps, and cliffs, also an occasional component of upland forests to the north. Fig. 43  Seed cones of Juniperus virginiana.

Pinaceae 1a. Leaves, at least in part, borne in fascicles or spirally arranged tufts of 2–60 on short shoots 2a. Leaves annually deciduous, those of the short shoots in spirally arranged tufts of 10–60 [Fig. 44]; each scale of a strobilus without an umbo; bracts exserted in basal portion of strobilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Larix 2b. Leaves persisting more than 1 year, those of the short shoots in fascicles of 2–5; each scale of a strobilus with an umbo; bracts hidden by scales . . . . . . . . . . . . . . . . . . . . . . . . . Pinus 1b. Leaves all borne singly 3a. Winter buds very resinous, the individual scales concealed by resin; strobili erect, with deciduous scales; leaves attached directly to branchlet and expanded to a circular base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Abies 3b. Winter buds not, or only slightly, resinous, the individual scales visible; strobili pendant at maturity [Figs. 45, 47], with persistent scales; leaves either attached to peg-like projections (i.e., sterigmata) from branchlet and narrowed to the attachment or attached directly to the branchlet with an expanded elliptic base 4a. Bracts of seed cone conspicuously exserted, 3-lobed at apex, the central lobe longer than the lateral lobes; young bark with resin blisters; leaves attached directly to the branchlet and expanded to an elliptic base . . . . . . . . . . . . . . . . . . . . . . . . Pseudotsuga

p i nac e a e   77

4b. Bracts of seed cone not exserted, not 3-lobed at apex; bark lacking resin blisters; leaves attached to peg-like projections from branchlet and narrowed to the attachment 5a. Leaf blades flat, petiolate, rounded at apex, dimorphic—longer, spreading ones and shorter, often appressed, ones; terminal shoot nodding . . . . . . . . . . . . . . . . . . Tsuga 5b. Leaf blades tetragonous, sessile, pointed at apex, monomorphic; terminal shoot erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Picea

Abies Reference: Hunt (1993). 1a. Branches deeply grooved, especially the 2- and 3-year-old ones . . . . . . . . . . A. homolepis 1b. Branches ungrooved or scarcely grooved 2a. Leaves 1.2–2.5 cm long, dark green and hardly (if at all) glaucous; seed cones 4–7 cm long; resin canals of leaves median, located well away from either epidermis (this character best viewed in cross-section with a 20× hand lens) . . . . . . . . . . . . . . . . . . . . . . . . A. balsamea 2b. Leaves (1.5–) 4–6 cm long, glaucous on both surfaces; seed cones 7–12 cm long; resin canals of leaves located near the abaxial epidermis . . . . . . . . . . . . . . . . . . . . . . . . . A. concolor 1. Abies balsamea (L.) P. Mill. N balsam fir. Abies balsamea (L.) P. Mill. var. phanerolepis Fern.; Pinus balsamea L. • CT, MA, ME, NH, RI, VT. Forests and swamps. 2. Abies concolor (Gord. & Glend.) Lindl. ex Hildebr. E white fir. MA, ME, VT. Forests and thickets, often in areas of human habitation, though sometimes seen in remote areas (e.g., coastal islands of ME). 3. Abies homolepis Sieb. & Zucc. E Nikko fir. RI. Forest edges, swamp margins.

Larix Reference: Parker (1993). 1a. Branchlets spreading; bark of main stem shedding as small, irregular scales; seed cones 1–2 cm long, with 10–15 (–30) scales that are glabrous abaxially [Fig. 44] . . . . . . . . . . L. laricina 1b. Branchlets pendulous; bark of main stem shedding as large, plate-like scales; seed cones 2–3.5 cm long, with 40–50 scales that are pubescent abaxially . . . . . . . . . . . . . . . . . . L. decidua 1. Larix decidua P. Mill. E European larch. CT, MA, ME, NH, RI, VT. Forests and thickets in areas of human habitation. 2. Larix laricina (Du Roi) K. Koch N Fig. 44 American larch. CT, MA, ME, NH, RI, VT. Hydric, organic soils, becoming occasional in wet-mesic or even mesic soils in the northern part of New England.

Picea Reference: Taylor (1993a). 1a. Seed cones 12–16 cm long; scales elliptic to rhombic, widest near the middle; branches conspicuously pendulous; branchlets red-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. abies 1b. Seed cones 1.5–6 cm long; scales flabellate, widest at or very near the apex; branches ascending to spreading; branchlets light brown to pink-brown or yellow-brown

Fig. 44  Leaves and seed cones of Larix laricina.

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2a. Branchlets glabrous and glaucous, light brown to pink-brown; seed cones 3–6 cm long; scale apex entire [Fig. 45]; winter buds rounded at apex . . . . . . . . . . . . . . . . . . . . . . . P. glauca 2b. Branchlets pubescent, not glaucous, yellow-brown; seed cones 1.5–4.5 (–5) cm long; scale apex slightly erose to evidently and irregularly toothed; winter buds acute at apex 3a. Pubescence of branchlets, in part, glandular [Fig. 46]; leaves gray-green, dull, glaucous; winter buds 3–3.9 mm long, gray-brown; seed cones 1.5–2.5 (–3.5) cm long, persisting on plant for many years; scale apex irregularly toothed. . . . . . . . . . . P. mariana 3b. Pubescence of branchlets entirely eglandular; leaves yellow to dark green, lustrous, not glaucous; winter buds 5–8 mm long, red-brown; seed cones 2.3–4.5 (–5) cm long, usually falling from plant by first winter; scale apex erose [Fig. 47] . . . . . . . . . . . P. rubens 1. Picea abies (L.) Karst. E Norway spruce. Picea excelsa (Lamb.) Link • CT, MA, ME, NH, RI, VT. Forests and thickets, persisting after cultivation. Fig. 45  Seed cone of Picea glauca.

2. Picea glauca (Moench) Voss n Fig. 45 white spruce. Picea canadensis (P. Mill.) B.S.P.; Pinus glauca Moench • CT, MA, ME, NH, RI, VT. Mesic forests, sometimes also found in other hydrologic regimes, occasionally a dominant conifer in maritime habitats. This species is naturalized in CT, MA, RI. 3. Picea mariana (P. Mill.) B.S.P. N Fig. 46 black spruce. Picea nigra (Ait.) Link; Pinus mariana P. Mill. • CT, MA, ME, NH, RI, VT. Growing in peat of bogs and folists on high mountain slopes and plateaus.

Fig. 46  Glandular (in part) pubescence of branchlet of Picea mariana.

3 × 4. This nothospecies is frequent in areas of sympatry between the parental taxa. Such areas include the higher mountains of New England and coniferous forests that are adjacent to acidic peatlands. Hybrids are best recognized by intermediate morphology, but some studies suggest that extensive backcrossing has occurred, causing some hybrid-derived individuals to look more similar to one or the other parent. 4. Picea rubens Sarg. N Fig. 47 red spruce. Picea australis Small; P. nigra (Ait.) Link var. rubra (Du Roi) Engelm. • CT, MA, ME, NH, VT. Predominant in boreal and subalpine forests, though sporadically occurring in many mixed forest types and sometimes in wetlands with a well-developed organic soil horizon.

Pinus Reference: Kral (1993). Fig. 47  Seed cone of Picea rubens.

1a. Leaves in fascicles of 5, with 1 fibrovascular bundle in cross-section; bud scales with entire margins; seed cones 8–20 cm long; umbo terminal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. strobus 1b. Leaves in fascicles of 2 or 3, with 2 fibrovascular bundles in cross-section; bud scales fringed on the margin; seed cones 3–10 cm long; umbo dorsal 2a. Leaves in fascicles of 3; apophyses of mature seed cones red-brown (fading on older, fallen cones) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. rigida 2b. Leaves in fascicles of 2; apophyses of mature seed cones cream to light brown or gray, or red-brown in P. resinosa 3a. Leaves (7–) 9–16 cm long, straight or only slightly twisted; seed cones symmetric, with a straight axis and without unequal development of apophyses 4a. Leaves brittle, tending to break when bent; winter buds red-brown; smaller branches orange-brown; seed cones, when falling, leaving some of the basal scales on the branch (i.e., the seed cone hollow-based); umbo unarmed . . . . . . . . P. resinosa 4b. Leaves pliable, tending not to break when bent; winter buds gray; smaller branches brown to gray-brown; seed cones falling with all the scales intact; umbo armed with a slender prickle (though the prickle often deciduous)

p i nac e a e   79

5a. Winter buds resinous; growth habit relatively uniform and often with an unbranched main stem; seed cones sessile, with a ± rounded base . . . . . . . P. nigra 5b. Winter buds not resinous; growth habit irregular, the main stem and branches often forked; seed cones stalked, with a ± truncate base . . . . . . . . . . . P. thunbergii 3b. Leaves 2–7 (–8) cm long, noticeably twisted (relatively straight in the planted P. mugo); seed cones ± asymmetric, with a curved axis and/or obviously unequal development of apophyses on one side 6a. Umbo encircled by a thin, dark gray to nearly black ring; low, multi-stemmed shrubs; leaves relatively straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. mugo 6b. Umbo without a dark ring; upright, single-stemmed, short to tall trees; leaves twisted 7a. Leaves 3–7 cm long, glaucous; larger branches orange-brown; seed cones usually falling after second winter; apophyses elongating toward base of seed cone on one side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sylvestris 7b. Leaves 2–3.5 (–5) cm long, not glaucous; larger branches gray-brown; seed cones serotinous, persisting on plant for many years; apophyses all ± of equal size [Fig. 48] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. banksiana 1. Pinus banksiana Lamb. n Fig. 48 Jack pine. Pinus divaricata (Ait.) Dum.-Cours. • MA, ME, NH, RI, VT. Nutrient-poor, sandy and/or rocky soils, occasionally in peatlands. This plant is introduced to MA and RI and is native (at least in part) to the other states in New England that it occurs in. 2. Pinus mugo Turra E Mugo pine. MA, NH; also reported from ME by Magee and Ahles (1999), but truly naturalized populations are unknown. Lawns and abandoned lots. 3. Pinus nigra Arnott E Austrian pine. MA, ME. Forest edges and fragments, near areas of human habitation. 4. Pinus resinosa Ait. N red pine. CT, MA, ME, NH, RI, VT. Nutrient-poor, sandy and/or rocky soils, occasionally in wetlands with a well-developed organic soil horizon. 5. Pinus rigida P. Mill. N pitch pine. CT, MA, ME, NH, RI, VT. Nutrient-poor, sandy and/or rocky soils, occasionally in wetlands with a well-developed organic soil horizon. 6. Pinus strobus L. N eastern white pine. Strobus strobus (L.) Small • CT, MA, ME, NH, RI, VT. Forests and woodlands, predominantly on mesic to xeric soils, but sometimes found on hydric soils. 7. Pinus sylvestris L. E Scotch pine. CT, MA, ME, NH, RI, VT. Forest edges, thickets, roadsides, persisting long after planting. 8. Pinus thunbergii Parl. E Japanese black pine. Pinus thunbergiana Franco, nom. illeg. • MA. Forest edges, thickets, roadsides, persisting long after planting.

Pseudotsuga 1. Pseudotsuga menziesii (Mirbel) Franco var. menziesii E coast Douglas-fir. Abies menziesii Mirbel; Pseudotsuga taxifolia (Lamb.) Britt. • CT, MA. Forest edges, river banks, areas of habitation.

Fig. 48  Seed cones of Pinus banksiana.

8 0  gy mnospe rms

Tsuga Reference: Taylor (1993b). 1. Tsuga canadensis (L.) Carr. N eastern hemlock. Pinus canadensis L. • CT, MA, ME, NH, RI, VT. Forests and slopes, sometimes growing in hydric soils.

Taxaceae Taxus ‌media The report of Taxus × Rehd. in RI (Tucker 2006) was based on a hybrid with Taxus umbraculifera (Sieb. ex Endl.) Ravencroft var. hicksii (hort. ex Rehd.) Spjut as the seed parent and possibly Taxus biternata Spjut as the pollen parent (i.e., the second parent is not confidently known). The specimen shows leaves arranged in 4 or more ranks and radiating in ± all directions on the young shoots (as in T. umbraculifera). Taxus cuspidata, with which this plant shares some similarities, such as abruptly bent petioles and many rows of stomates, has leaves arranged in only 2 ranks (as to the attachment of the petioles), but the blades all ascending to erect. Reference: Spjut (2007). 1a. Winter bud scales blunt at the apex, slightly keeled; leaf blades gradually tapering to the apex [Fig. 49 and R inset]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. baccata 1b. Winter bud scales sharply pointed to cuspidate, definitely keeled or folded; leaf blades abruptly tapering to the apex [Fig. 49, L inset] 2a. Low shrubs with straggling, diffusely branched habit; leaf blades 1–2 (–2.4) mm wide; stomates in 5–7 (–9) rows in each band; petioles gradually curved or with an abrupt bend near the junction of the branch, not clasping the branchlet; seeds round or somewhat compressed near apex, but not angled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. canadensis 2b. Plants upright; leaf blades 2–3 mm wide; stomates in (7–) 9–14 (–19) rows in each band; petioles with an abrupt bend near the junction of the blade, clasping the branchlet; seeds 4- or 5-angled near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. cuspidata 1. Taxus baccata L. var. baccata E Fig. 49 and R inset English yew. MA. Forest edges, thickets, roadsides. 2. Taxus canadensis Marsh. N Fig. 49, L inset American yew. 2b. Taxus baccata L. var. minor Michx.; T. minor (Michx.) Britt. • CT, MA, ME, NH, RI, VT. Forests, predominantly deciduous and mixed evergreen-deciduous types. Fig. 49  Branch of Taxus baccata with detail of leaf blades—T. canadensis (left) and T. baccata (right).

1a. Leaves mainly spreading and borne in 2 lateral ranks, the blades acute at the margin as seen in cross-section, slightly revolute, acute to acuminate at the apex, and keeled on the abaxial surface due to the raised midrib . . . . . . . . . . . . . . . . 2a. T. canadensis var. canadensis 1b. Leaves ascending to erect, not borne in 2 spreading ranks, the blades round at the margin, plane, obtsuse to acute at the apex, and flat on the abaxial surface (i.e., the midrib not raised from the surface) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. T. canadensis var. minor (Michx.) Spjut Variety canadensis is known from CT, MA, ME, NH, RI, VT. Variety minor is known from ME. 3. Taxus cuspidata Sieb. & Zucc. E Japanese yew. CT, MA, VT. Forest edges, thickets, roadsides, areas of habitation.

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Magnoliids Aristolochiaceae 1a. Plants without aerial stems, herbaceous; calyx actinomorphic, the basal, tubular portion straight; flowers with 12 stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asarum 1b. Plants with aerial stems, woody or herbaceous; calyx zygomorphic, the basal, tubular portion nearly straight or strongly bent [Fig. 50]; flowers with 6 stamens 2a. Leaf blades 1–5 cm wide, pinnately veined; inflorescence originating from base of stem near ground level; capsule globose, 8–20 mm long . . . . . . . . . . . . . . . . . . . . . . Endodeca 2b. Leaf blades 5–35 cm wide, palmately veined; inflorescence originating from the axils of leaves; capsule cylindric to ovoid or ellipsoid (sometimes globose or pyriform in A. clematitis), 20–80 mm long 3a. Plants herbaceous; inflorescence a fascicle of flowers from the axil or, less commonly, a solitary flower; calyx nearly straight, with a single, terminal lobe; annulus absent; capsules 20–50 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aristolochia 3b. Plants lianas (i.e., woody); inflorescence of a solitary, axillary flower; calyx strongly curved, with 3 terminal lobes [Fig. 50]; annulus (i.e., a circular flange at junction of calyx tube and limb) present; capsules 60–80 mm long . . . . . . . . . . . . . . . . . . . . . . . . . Isotrema

Aristolochia Recent phylogenetic analyses using morphology, DNA sequences, and chromosome numbers supported the division of Aristolochia into multiple segregate genera. The system of Kelly and González (2003), where Aristolochia s.l. was subdivided into four genera, is followed here. The results of Ohi-Toma et al. (2006) also provided support for this system; however, they chose to recognize only two genera. In either case, most of our species reside in genera other than Aristolochia. 1. Aristolochia clematitis L. E creeping birthwort. MA. Roadsides and abandoned lots, near areas of human habitation.

Asarum 1. Asarum canadense L. N Canada wild ginger. Asarum acuminatum (Ashe) Bickn.; A. canadense L. var. acuminatum Ashe; A. canadense L. var. ambiguum (Bickn.) Farw.; A. reflexum Bickn. • CT, MA, ME, NH, VT. Rich, sometimes rocky, mesic forests.

Endodeca 1. Endodeca serpentaria (L.) Raf.

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Virginia serpentaria. Aristolochia convolvulacea Small; A. serpentaria L.; A. serpentaria L. var. hastata (Nutt.) Duchartre; A. serpentaria L. var. nashii (Kearney) Ahles; Endodeca serpentaria (L.) Raf. var. hastata (Nutt.) C.F. Reed • CT. Mesic to dry-mesic forests, often underlain by circumneutral bedrock.

8 2  mag noliids

Isotrema Reference: Barringer (1997). 1a. Stems and abaxial leaf surface glabrous to puberulent; peduncle with a bract near the middle; annulus smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. macrophyllum 1b. Stems and abaxial leaf surface tomentose; peduncle without a bract; annulus rugulose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. tomentosum 1. Isotrema macrophyllum (Lam.) C.F. Reed E Fig. 50 large-leaved Dutchman’s pipe. Aristolochia durior Hill; A. macrophylla Lam. • CT, MA, ME, NH, VT. Forests and forest edges. Fig. 50  Curved perianth of Isotrema macrophyllum.

2. Isotrema tomentosum (Sims) Huber E woolly Dutchman’s pipe. Aristolochia tomentosa Sims • MA. Roadsides and abandoned lots, near areas of human habitation. The report of this species in CT and VT by Kartesz (1999) was erroneous.

Calycanthaceae Calycanthus 1. Calycanthus floridus L. E eastern sweetshrub. 1b. Calycanthus fertilis Walt.; C. floridus L. var. laevigatus (Willd.) Torr. & Gray; C. glaucus Willd. • CT, MA. Forest edges, abandoned lots, near areas of human habitation. 1a. Branchlets, petioles, and abaxial leaf blade surface densely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. C. floridus var. floridus 1b. Branchlets, petioles, and abaxial leaf blade surface glabrous or sparsely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. C. floridus var. glaucus (Willd.) Torr. & Gray Variety floridus is known from MA. Variety glaucus is known from CT, MA.

Ceratophyllaceae Ceratophyllum Ceratophyllum muricatum Cham. ssp. australe (Griseb.) D.H. Les was reported from MA by Angelo and Boufford (in press). The voucher specimen is C. echinatum—21 Jul 1994, Hickler s.n. (MASS!). References: Hellquist and Crow (1984), Les (1997). 1a. Leaves mostly once- or twice-forked, serrate, the ultimate segments flat [Fig. 51]; styles 4.5–6 mm long; achenes with 2 basal spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. demersum 1b. Leaves mostly 3- or 4-times forked, sparsely serrate, the ultimate segments capillary; styles 5–10 mm long; achenes with 4–15 spines—2 basal and 2–13 lateral . . . . . C. echinatum Fig. 51  Branch of Ceratophyllum demersum with detail of branch margin.

1. Ceratophyllum demersum L. N Fig. 51 common hornwort. Ceratophyllum apiculatum Cham. • CT, MA, ME, NH, RI, VT; throughout. Lakes and slow-moving streams.

C e r ato p hy l l ac e a e   83

2. Ceratophyllum echinatum Gray N spineless hornwort. Ceratophyllum demersum L. var. echinatum (Gray) Gray • CT, MA, ME, NH, RI, VT. Lakes and slow-moving streams, less common than Ceratophyllum demersum.

Lauraceae 1a. Leaf blades simple or with a lateral lobe on one or both sides; inflorescence peduncled, a raceme or panicle, produced at the tips of the previous year’s branchlets; tepals accrescent; drupe blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sassafras 1b. Leaf blades simple; inflorescence sessile, a pseudoumbel, produced in the axils along the previous year’s branchlets; tepals deciduous; drupe red . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lindera

Lindera Reference: Wofford (1997). 1. Lindera benzoin (L.) Blume N northern spicebush. Benzoin aestivale Nees; Lindera benzoin (L.) Blume var. pubescens (Palmer & Steyermark) Rehd. • CT, MA, ME, NH, RI, VT. Swamps, margins of streams, and low, riparian forests.

Sassafras Reference: van der Werff (1997). 1. Sassafras albidum (Nutt.) Nees N sassafras. Laurus albidus Nutt.; L. sassafras L.; Sassafras albidum (Nutt.) Nees var. molle (Raf.) Fern.; S. variifolium (Salisb.) Kuntze; S. variifolium (Salisb.) Kuntze var. albidum (Nutt.) Fern. • CT, MA, ME, NH, RI, VT. Forests, forest edges, and sometimes regenerating disturbed places.

Magnoliaceae Reference: Meyer (1997). 1a. Leaf blades with 2–10 lateral lobes (usually 4), the apex truncate to shallowly notched; tepals streaked with an orange band near the base; fruit an aggregate of samaras; winter buds with 2 scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liriodendron 1b. Leaf blades simple, the apex obtuse to acuminate; tepals without orange bands [Fig. 52]; fruit an aggregate of follicles, the seed with a brightly colored aril suspended from the dehisced fruit by the slender funiculus; winter buds with 1 scale . . . . . . . . . Magnolia

Liriodendron 1. Liriodendron tulipifera L. N tuliptree. CT, MA, RI, VT. Hardwood forests, often rich and/or rocky.

8 4  mag noliids

Magnolia 1a. Leaf blades coriaceous, abaxially glaucous; seeds ca. 5 mm long; branchlets green, 2–4 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. virginiana 1b. Leaf blades thick-herbaceous, abaxially pale green to white-green; seeds 9–12 mm long; branchlets gray-brown to brown or red-brown, 4–11 mm thick 2a. Leaves crowded near tip of branchlets; flowers somewhat malodorous, with creamwhite tepals 8–12 cm long; winter buds glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. tripetala 2b. Leaves scattered along the branchlets; flowers pleasantly aromatic, with green-white tepals up to 8 cm long (rarely longer); winter buds pubescent . . . . . . . . . . . . . . M. acuminata 1. Magnolia acuminata (L.) L. E cucumber-tree. Magnolia acuminata L. var. cordata (Michx.) Ser.; M. virginiana L. var. acuminata L.; Tulipastrum acuminatum (L.) Small; T. cordatum (Michx.) Small • CT, MA; also reported from ME by Richards et al. (1983), but specimens are unknown. Thickets and forest edges near areas of human habitation. 2. Magnolia tripetala (L.) L. E umbrella-tree. Magnolia virginiana L. var. tripetala L. • CT, MA, RI. Thickets and forest edges near areas of human habitation. 3. Magnolia virginiana L. ssp. virginiana

n C Fig. 52

sweet-bay. CT, MA; coastal plain. Wet-mesic to hydric forests. This species is introduced in CT and native (in part) to MA.

Fig. 52  Flower of Magnolia virginiana.

Nymphaeaceae 1a. Floating leaf blades without a sinus [Fig. 53]; perianth composed of 3 sepals and 3 petals; carpels 4–8, not embedded in the receptacle 2a. Leaves strongly dimorphic—submersed, opposite, highly dissected ones and floating, alternate, simple ones 0.6–3 cm long; flowers with 3–6 stamens; underwater parts of plant barely coated with mucilage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cabomba 2b. Leaves monomorphic, all alternate, floating, simple [Fig. 53], 3.5–13.5 cm long; flowers with 18–36 (–51) stamens; underwater parts of plant heavily coated with mucilage [Fig. 53] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brasenia 1b. Floating leaf blades (or emergent in Nuphar advena) with a prominent, basal sinus [Figs. 54, 55]; perianth composed of numerous tepals that become progressively petaloid in centripetal fashion; carpels numerous, sunken into a large, spongy receptacle 3a. Leaves with angular basal lobes; petioles with 4 large air cavities; flowers white (pink); sepals sepaloid; outer petals petaloid, gradually grading into the stamens toward the center of the flower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nymphaea 3b. Leaves with rounded basal lobes [Figs. 54, 55]; petioles with many small air cavities; flowers yellow; sepals petaloid; petals all stamen-like . . . . . . . . . . . . . . . . . . . . . . . . . . . Nuphar

Brasenia Reference: Wiersema (1997a). 1. Brasenia schreberi J.F. Gmel. N Fig. 53 Fig. 53  Petiole and leaf blade of Brasenia schreberi.

water-shield. Brasenia peltata Pursh • CT, MA, ME, NH, RI, VT. Still or slow-moving water of lakes and streams, usually oligotrophic to mesotrophic.

Ny m p h a e ac e a e   8 5

Cabomba 1. Cabomba caroliniana Gray E Carolina fanwort. Cabomba caroliniana Gray var. pulcherrima Harper; C. pulcherrima (Harper) Fassett • CT, MA, NH, RI. Still or slow-moving water of lakes, streams, and canals, the water of varied pH.

Nuphar The flowers of Nuphar are interesting. The outer whorl of perianth is composed of 5 or 6 petaloid sepals. The inner whorl of perianth is represented by many small, thick structures that are transitional in appearance to the stamens. Beal (1956) placed all our species as infraspecific taxa under Nuphar lutea, a European plant. Subsequent studies have shown that our species are distinct from N. lutea. However, further research is needed to determine the correct placement of N. microphylla, which appears to be closely related to the Eurasian N. pumila (Timm) DC. (and was synonymized under the latter name by Beal). References: Wiersema and Hellquist (1997), Hellquist and Crow (1984). 1a. Sepals 1–2.5 cm long, numbering 5 per flower; anthers predominantly shorter than the filaments, 1–3 mm long; fruit strongly constricted below the red stigmatic disk; petals and stamens promptly deciduous, usually not persisting as remnants at the base of the fruit; basal sinus of leaf blade (42–) 54–90% of the length of the blade midrib [Fig. 54] . . . . N. microphylla 1b. Sepals 2.5–5 cm long, numbering 6 per flower; anthers longer than the filaments, 3–9 mm long; fruit only slightly constricted below the green (rarely red) stigmatic disk; petals and stamens tardily deciduous, usually persisting as remnants around the base of the fruit; basal sinus of leaf blade 30–59 (–62)% of the length of the blade midrib [Fig. 55] 2a. Petioles strongly flattened on the adaxial surface, often wing-margined; leaves with a narrow, closed, basal sinus, floating; sepals red at the base of the abaxial surface; fruit often purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. variegata 2b. Petioles terete to oval in cross-section, not wing-margined; leaves with a broad, open, basal sinus, often emergent; sepals green (rarely red) at the base of the abaxial surface; fruit green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. advena 1. Nuphar advena (Ait.) Ait. f. in Ait. & Ait. f.

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immigrant pond-lily. Nuphar lutea (L.) Sm.; N. macrophylla Small; Nymphaea advena Ait.; Nymphozanthus advena (Ait.) Fern. • CT, ME. Fresh-tidal river shorelines, still or slow-moving water on the coastal plain. 2. Nuphar microphylla (Pers.) Fern. N Fig. 54 small-leaved pond-lily. Nuphar minima (Willd.) Sm.; Nymphaea microphylla Pers. • CT, MA, ME, NH, VT. Circumneutral to basic water of lakes and slow-moving streams. 2 × 3. Nuphar rubrodisca Morong is an infrequent hybrid of circumneutral to basic water. It is known from CT, MA, ME, NH, VT. It has been treated as a species, but it shows morphological intermediacy, poor fruit set, and lower pollen viability than other sympatric species (Padgett et al. 1998). It resembles N. microphylla in some key characteristics (e.g., short sepals, fruit constricted below stigmatic disk, anthers predominantly shorter than filaments). The hybrid is characterized by 5 or 6 sepals per flower, anthers (2–) 3–6 mm long, stigmatic disk with 8–15 stigmatic rays that terminate 0–1.6 mm from the margin, each ray separated by a shallow sinus, the constriction below the stigmatic disk 5–10 mm in diameter, petals and stamens usually persistent as remnants at the base of the fruit, and basal sinus of leaf blade 39–59 (–62)% of the length of the blade midrib (vs. sepals 5 per flower, anthers 1–3 mm long, stigmatic disk with 6–11 stigmatic rays that terminate 0–0.2 mm from the margin, each ray separated by a relatively deep sinus, the constriction below the stigmatic disk 1.5–5 mm in diameter, petals and stamens promptly deciduous, and basal sinus of leaf blade (42–) 54–90% in N. microphylla).

Fig. 54  Leaf blade of Nuphar microphylla with a relatively deep sinus.

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3. Nuphar variegata Dur. N Fig. 55 bullhead pond-lily. Nuphar lutea (L.) Sm. ssp. variegata (Dur.) E.O. Beal; Nymphaea fraterna Mill. & Standl. • CT, MA, ME, NH, RI, VT. Slightly acidic to basic water of lakes, slow-moving streams, backwaters, and pools.

Nymphaea References: Wiersema (1997b), Hellquist and Crow (1984).

Fig. 55  Leaf blade of Nuphar variegata with a relatively shallow sinus.

1a. Flowers 3–7.5 cm wide, with 20–40 stamens; stigmatic disk with 5–12 lines; leaf blades elliptic, (2–) 4–9 (–15) cm wide, with a wide sinus, mottled when young . . . . . . . . . . . N. leibergii 1b. Flowers 6–19 cm wide, with 40–100 stamens; stigmatic disk with 10–25 lines; leaf blades orbicular, (5–) 10–30 (–40) cm wide, with a narrow sinus, not mottled 2a. Petals acute to narrow-rounded at the apex; abaxial surface of the sepals and leaf blades purple or sometimes green; inner filaments narrower than the anthers; seeds 1.5–2.5 mm long; branches of the rhizome not constricted at the base; petioles faintly, if at all, striped; flowers strongly fragrant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. odorata 2b. Petals broad-rounded at the apex; abaxial surface of the sepals and leaf blades green; all the filaments broader than the anthers; seeds 2.8–4.5 mm long; branches of the rhizome constricted at the base, breaking into tuber-like segments [Fig. 56]; petioles striped with brown-purple; flowers weakly or not at all fragrant . . . . . . . . . . . . . . N. tuberosa 1. Nymphaea leibergii Morong

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dwarf water-lily. Castalia leibergii Morong; Nymphaea tetragona Georgi ssp. leibergii (Morong) Porsild; N. tetragona Georgi var. leibergii (Morong) Boivin • ME, VT; northern portion of states. Circumneutral water of lakes and slow-moving streams. 1 × 2. This rare water-lily hybrid is known from ME, VT. It is sterile and intermediate in morphology between the parents. 2. Nymphaea odorata Ait. N white water-lily. Castalia odorata (Ait.) Wood; Nymphaea odorata Ait. var. gigantea Tricker; N. odorata Ait. var. rosea Pursh • CT, MA, ME, NH, RI, VT. Somewhat acidic to basic water of lakes, slow-moving streams, and pools. Though normally with white petals, this species has occasional forms with pink petals. 3. Nymphaea tuberosa Paine n Fig. 56 tuberous water-lily. Castalia tuberosa (Paine) Greene; Nymphaea odorata Ait. ssp. tuberosa (Paine) Weirsema & Hellquist; N. odorata Ait. var. maxima (Conrad) Boivin • CT, MA, ME, NH, VT. Circumneutral water of lakes, slow-moving streams, and embayments. Considered native to Lake Champlain and tributaries, VT, but introduced elsewhere in New England. Fig. 56  Rhizome and tubers of Nymphaea tuberosa.

Saururaceae Saururus Reference: Buddell and Thieret (1997). 1. Saururus cernuus L. lizard’s-tail. CT, MA, RI. Swamps, shorelines, river banks.

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Monocots Acoraceae Acorus Acorus was traditionally placed in the Araceae. This erroneous placement of the genus was partly due to incorrect interpretation of the distal portion of the sympodial leaf—a structure that had been considered to be a spathe. Two species are now recognized to occur in our area. Acorus calamus is a sterile triploid that has been introduced from Europe. Acorus americanus is a fertile diploid native to North America. References: Haines (2000a), Thompson (2000a). 1a. Secondary veins of dried leaf blades up to ca. 0.5 times the width of the midvein (i.e., leaves with 1 prominent midvein and numerous finer, secondary veins); fruits not produced; sympodial leaf often as tall as or shorter than the vegetative leaves; vegetative leaves (5–) 10–20 mm wide, often crisped or undulate; flowers 3–4 mm long . . . . . . . . . . . A. calamus 1b. One or more of the secondary veins of dried leaf blades ca. 0.75–1 times as wide as the midvein (i.e., leaves with 2–6 prominent veins as well as numerous finer, secondary veins); fruit produced, an obpyramidal berry; sympodial leaf usually as tall as or taller than vegetative leaves; vegetative leaves 3–10 (–12) mm wide, not crisped, entire; flowers 2–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. americanus 1. Acorus americanus (Raf.) Raf. N several-veined sweetflag. Acorus calamus L. var. americanus Raf. • CT, MA, ME, NH, RI, VT. Marshes, lake borders, stream shores, and other open, wet areas. 2. Acorus calamus L. E single-veined sweetflag. CT, MA, ME, NH, RI, VT. Marshes, lake borders, stream shores, and other open, wet areas.

Agavaceae Yucca 1. Yucca filamentosa L. E Adam’s-needle. Yucca filamentosa L. var. smalliana (Fern.) Ahles; Y. smalliana Fern. • CT, MA, RI. Sandy soils of roadsides, fields, and disturbed lots.

8 8  MO N OCOTS

Alismataceae References: Hellquist and Crow (1981), Haynes and Hellquist (2000a). 1a. Androecium composed of 6 stamens; gynoecium with a single whorl of carpels on a small, flat receptacle; inflorescence a panicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alisma 1b. Androecium composed of 7–40 stamens; gynoecium with several whorls of carpels on a convex receptacle, forming a subglobose cluster; inflorescence usually a raceme or an umbel 2a. Roots not septate; flowers all bisexual; achenes longitudinally ribbed; inflorescence usually an umbel (i.e., with a single whorl of pedicels) . . . . . . . . . . . . . . . . . . . . . . . Helanthium 2b. Roots septate; at least the lower flowers of the inflorescence unisexual; achenes not longitudinally ribbed, but sometimes with lateral wings; inflorescence usually a raceme, with (1–) 2–10 whorls of pedicels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sagittaria

Alisma 1a. Leaves linear, 0.2–2 (–3) cm wide; abaxial edge of achenes with 2 grooves; style ± coiled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. gramineum 1b. Emersed leaves with narrow-lanceolate to ovate blades 3–12 cm wide; abaxial edge of achenes with a single groove; style merely curved 2a. Achenes 1.8–3 mm long; petals 3.8–4.5 × 3–3.9 mm, noticeably longer than the sepals; styles 0.4–0.6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. triviale 2b. Achenes 1.5–2.2 mm long; petals 1.8–2.5 × 1.4–2 mm, scarcely, if at all, longer than the sepals; styles 0.2–0.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. subcordatum 1. Alisma gramineum Lej. N narrow-leaved water-plantain. Alisma gramineum Lej. var. angustissimum (DC.) Hendricks • VT; primarily Lake Champlain and its tributaries. Circumneutral still or slow-moving water. 2. Alisma subcordatum Raf. N southern water-plantain. Alisma plantago-aquatica L. var. parviflorum (Pursh) Torr.; A. plantago-aquatica L. ssp. subcordatum (Raf.) Hultén • CT, MA, ME, NH, RI, VT. Marshes, muddy shorelines, ditches, and shallow, circumneutral to slightly basic water of lakes and slowmoving streams. 3. Alisma triviale Pursh N northern water-plantain. Alisma brevipes Greene; A. plantago-aquatica L. var. americanum J.A. Schultes • CT, MA, ME, NH, RI, VT; throughout. Marshes, muddy shorelines, ditches, and shallow, circumneutral water of lakes and slow-moving streams.

Helanthium See Lehtonen and Myllys (2008) for rationale of excluding Helanthium from Echinodorus. 1. Helanthium tenellum (Mart.) Britt.

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dwarf burhead. Alisma tenellum Mart.; Echinodorus parvulus Engelm.; E. tenellus (Mart.) Buch.; E. tenellus (Mart.) Buch. var. parvulus (Engelm.) Fassett; Helanthium parvulum (Engelm.) Small • CT, MA; also reported from ME by Perkins (1936) and from VT by Haynes and Hellquist (2000a), but specimens are unknown. Sandy shorelines of ponds and streams. The report from ME by Perkins was eventually retracted (1938), but the retraction was apparently not based on annotation of the original specimens but rather on an inability to find additional specimens (which could indicate the population was extirpated, not that the original collections were incorrectly determined).

A l i s m atac e a e   8 9

Sagittaria Fruits must be mature to show features such as facial wings, facial glands, and beak orientation. Width of the leaf at its midpoint is used in the following key—the measurement should not be taken at the midpoint of the leaf blade but at the midpoint of the leaf; i.e., combined length of the petiole and blade (when a blade is present). 1a. Sepals of carpellate flowers appressed in fruit [Fig. 59]; pedicels recurved (rarely spreading) in fruit [Fig. 59] 2a. Lower flowers bisexual; leaves represented by thick, spongy phyllodia (2–) 2.5–8 (–8.5) mm wide at the midpoint in drying (sometimes with a small, expanded blade at the apex) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. montevidensis 2b. Lower flowers carpellate; leaves represented by phyllodia that are flattened or lenticular in cross-section and 0.7–2.7 mm wide at the midpoint in drying (sometimes with a small, expanded blade at the apex) 3a. Leaves 5–30 cm long; peduncles 5–40 cm long; plants predominantly of brackish, tidal shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. subulata 3b. Leaves 30–90 (–250) cm long; peduncles 10–200 cm long; plants predominantly of fresh, moderately moving rivers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. filiformis 1b. Sepals reflexed in fruit; pedicels ascending to spreading in fruit 4a. Filaments smooth [Fig. 58]; leaves usually with blades that possess basal lobes 5a. Bracts of inflorescence thick and herbaceous, distinct or with margins connate less than 25% of their total length; achenes with 1–3 facial wings; inflorescence with 2–4 whorls of pedicels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. engelmanniana 5b. Bracts of inflorescence thin and scarious, with margins connate 25% or more of their total length; achenes with 0 or 1 facial wings; inflorescence with 2–10 whorls of pedicels 6a. Beak of the achene 0.1–0.5 mm long, erect; petals 7–10 mm long; achenes 1.8–2.6 mm long; floating leaves with sagittate-ovate blades produced on some deepwater plants; emersed leaves with a broad-lanceolate to triangular-ovate central lobe and borne on recurving petioles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. cuneata 6b. Beak of the achene 0.6–1.8 mm long, horizontally spreading; petals 10–20 mm long; achenes 2.5–4 mm long; floating leaves with expanded blades not produced; emersed leaves with highly variable blades, the central lobe ranging from linear to broad-ovate, borne on ascending petioles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. latifolia 4b. Filaments rough, beset with minute hairs [Fig. 57]; leaves either without blades or with blades that usually lack basal lobes 7a. Leaves all phyllodial, elliptic to terete in cross-section, generally tapering from near the base to the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. teres 7b. Leaves phyllodial and with ± parallel margins that taper somewhat abruptly to the apex or with an expanded blade, in either case flattened 8a. Lowest whorl of carpellate flowers borne on pedicels 1–3 cm long; flowering stem straight, without a distinct bend; achenes 1.5–2 (–2.8) mm long, with a short beak 0.1–0.3 mm long; leaf blades (when formed) narrow-lanceolate to broadlanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. graminea 8b. Lowest whorl of carpellate flowers sessile or on short pedicels up to 0.5 cm long; flowering stem often with a conspicuous bend at the lowest whorl of flowers; achenes 2.5–4 mm long, with a beak 0.8–1.4 mm long; leaf blades (when formed) lanceolate to oblong-ovate or elliptic-ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rigida

90  MO N OCOTS

1. Sagittaria cuneata Sheldon N northern arrowhead. Sagittaria arifolia Nutt. ex J.G. Sm. • CT, MA, ME, NH, VT; mainly in northern New England, confined to western half of the southern New England states. Circumneutral to slightly basic water of lakes, slow-moving streams, and pools. Sagittaria cuneata is known to produce three different leaf morphologies—strap-shaped phyllodia on deep-water plants, floating leaf blades on moderate to deep-water plants, and sagittate leaf blades on recurved petioles on emersed plants. 2. Sagittaria engelmanniana J.G. Sm. N Engelmann’s arrowhead. CT, MA, RI, VT. Acid water of marshes, coastal plain ponds, and peatlands. The occurrence in VT (Lake Champlain) is unusual. 3. Sagittaria filiformis J.G. Sm. N

Fig. 57  Rough filaments of Sagittaria graminea.

narrow-leaved arrowhead. Sagittaria stagnorum Small; S. subulata (L.) Buch. var. gracillima (S. Wats.) J.G. Sm. • CT, MA, ME, RI. Slow to moderately moving rivers, including fresh-tidal sections. Sagittaria filiformis is, like many arrowheads, a very plastic species. Field and museum observations suggest that some characters used to distinguish S. filiformis from S. subulata are not reliable. Though most collections of these two taxa are markedly different, it is likely the result of sampling in widely spaced locations. Specimens taken from some tidal rivers, where exposed shoreline and deep-water habitats co-occur, show a morphological cline between short plants with emergent leaves and tall plants with long, flaccid leaves (though only the tall plants in deep water produce flowers and fruits). It is apparent that determinations based on vegetative, short-leaved plants may be inaccurate (though long-leaved plants are certainly S. filiformis). Adams and Godfrey (1961) have discussed morphological confluence of species within this complex in the southeastern United States. 4. Sagittaria graminea Michx. var. graminea N Fig. 57 grass-leaved arrowhead. Sagittaria eatonii J.G. Sm. • CT, MA, ME, NH, RI, VT; throughout. Shorelines and shallow water of lakes, streams, and pools, including fresh-tidal river shores. 5. Sagittaria latifolia Willd. N Fig. 58 common arrowhead. Sagittaria latifolia Willd. var. obtusa (Engelm.) Wieg.; S. latifolia Willd. var. pubescens (Muhl. ex Nutt.) J.G. Sm.; S. longirostra (Micheli) J.G. Sm.; S. obtusa Muhl. ex Willd.; S. planipes Fern.; S. pubescens Muhl. ex Nutt. • CT, MA, ME, NH, RI, VT; throughout. Shorelines and shallow water of lakes, streams, and pools, including fresh-tidal river shores. 6. Sagittaria montevidensis Cham. & Schlecht. ssp. spongiosa (Engelm.) Bogin N Fig. 59

Fig. 58  Smooth filaments of Sagittaria latifolia.

spongy-leaved arrowhead. Lophotocarpus spathulatus J.G. Sm.; L. spongiosus (Engelm.) J.G. Sm.; Sagittaria calycina Engelm. var. spongiosa Engelm.; S. montevidensis Cham. & Schlecht. var. spongiosa (Engelm.) Boivin; S. spathulata (J.G. Sm.) Buch. • CT, MA, ME, NH. Fresh-tidal river shores, typically on mud flats in areas with well-developed marsh floras. 7. Sagittaria rigida Pursh N sessile-fruited arrowhead. CT, MA, ME, NH, VT. Lakes, river shores, backwaters, and pools, including fresh to brackish-tidal rivers. 8. Sagittaria subulata (L.) Buch.

NC

awl-leaved arrowhead. Alisma subulatum L.; Sagittaria lorata (Chapman) Small • CT; also reported from RI by Haynes and Hellquist (2000a), but specimens are unknown. Fresh to brackish-tidal river shores. Reports of this species from MA were based on specimens of Sagittaria graminea and short, emergent, vegetative forms of S. filiformis. 9. Sagittaria teres S. Wats.

NC

quill-leaved arrowhead. Sagittaria graminea Michx. var. teres (S. Wats.) Bogin • CT, MA, NH, RI. Primarily acidic water of sandy, or sometimes muddy, coastal plain ponds.

Fig. 59  Recurved pedicels of Sagittaria montevidensis.

Al l i ac e a e   9 1

Alliaceae Allium Reference: McNeal and Jacobsen (2002). 1a. Leaves flat, (15–) 20–80 mm wide, produced in early season and soon withering, usually not present during anthesis; capsule prominently 3-lobed . . . . . . . . . . . . . . . . . . . . . A. tricoccum 1b. Leaves flat or terete, 1–20 mm wide, not early withering, present during anthesis; capsule only slightly lobed or not produced 2a. Umbels composed entirely of flowers, without bulbils 3a. Pedicels 10–50 mm long, longer than the flowers; scapes 3–20 mm in diameter; bulbs 3–10 cm in diameter; flowers 3–7 mm long; leaf blades semicircular in crosssection, 3–20 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. cepa 3b. Pedicels 2–6 mm long, shorter than to nearly equaling the flowers; scapes 3–5 mm in diameter; bulbs 1.5–2 cm in diameter; flowers 8–12 mm long; leaf blades terete in cross-section, 2–7 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. schoenoprasum 2b. Umbels with most or all of the flowers replaced by bulbils 4a. Bulbs covered with an outer layer of fibrous reticulum [Fig. 60]; leaf-bearing nodes confined to the basal ¼ of scape; inflorescence subtended by 3 or 4 spathiform bracts (note: the bracts are often caducous, so fewer may be present at time of observation) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. canadense 4b. Bulbs covered with a membranaceous-scaly outer layer, fibers (if present) not forming a reticulum; leaf-bearing nodes extending to midpoint of scape or higher; inflorescence subtended by 1 or 2 spathiform bracts 5a. Umbel subtended by 2 persistent, spathiform bracts; flowers 6–8 mm long, on pedicels 15–60 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. oleraceum 5b. Umbel subtended by a solitary, caducous, spathiform bract; flowers 3–5 mm long, on pedicels 10–20 mm long 6a. Leaf blade hollow in the basal half, terete, 2–4 mm in diameter; bulbs 1–2 cm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. vineale 6b. Leaf blade solid, flat, 5–20 mm wide; bulbs (1.5–) 3–8 cm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sativum 1. Allium canadense L. var. canadense N Fig. 60 meadow garlic. Allium canadense L. var. robustum Farw. • CT, MA, ME, NH, RI, VT. Floodplains, riparian forests, often on rich soils of terraces. 2. Allium cepa L. E garden onion. Allium cepa L. var. bulbiferum Bailey; A. cepa L. var. viviparum M.C. Metz • CT, MA, ME, NH, VT. Fields and disturbed soil in areas of cultivation or agriculture. Many different cultivars exist for this plant. Some of them have bulbils in the inflorescence. Leaf blade characteristics will serve to identify such plants. 3. Allium oleraceum L. E field garlic. MA. Fields, roadsides, and other disturbed areas. 4. Allium sativum L. var. sativum E cultivated garlic. MA. Fields, roadsides, and other disturbed areas.

Fig. 60  Bulb of Allium canadense.

92  MO N OCOTS

5. Allium schoenoprasum L. n wild chives. Allium schoenoprasum L. var. laurentianum Fern.; A. schoenoprasum L. ssp. sibiricum (L.) Čelak.; A. schoenoprasum L. var. sibiricum (L.) Hartman; A. sibiricum L. • CT, MA, ME, NH, RI, VT. River banks, shoreline outcrops, meadows (native populations), fields, roadsides, and vacant lots (non-native populations). Allium schoenoprasum includes both native and non-native races in New England. Non-native plants often have ovate to narrow-ovate, merely pointed outer tepals and leaf blades equaling or overtopping the scape, whereas native plants often have lanceolate to narrow-lanceolate and sharply pointed outer tepals and leaf blades shorter than the scape. However, such differences have been shown to be clinal and are not reliable for discriminating taxa (McNeal and Jacobsen 2002). Therefore, infraspecific taxa are not recognized here. 6. Allium tricoccum Ait. N wild leek. 6a. Allium burdickii (Hanes) A.G. Jones; 6b. Validallium tricoccum (Ait.) Small • CT, MA, ME, NH, RI, VT. Rich, mesic forests and high-terrace floodplain forests. Because bulb size changes throughout the spring, measurements should be performed on individuals that lack leaves. 1a. Bulbs 2–4 (–5) × 1–1.5 cm; leaves not or scarcely petiolate, white at the base, with blades (1.5–) 2–4 (–4.5) cm wide; spathiform bracts 1–2 cm long; umbel with (6–) 12–18 (–24) flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6a. A. tricoccum var. burdickii Hanes 1b. Bulbs 4–6 × 1.5–3 cm; leaves petiolate, anthocyanic (or not) at the base, with blades (3–) 5–8 cm wide; spathiform bracts 2–3 cm long; umbel with (15–) 30–50 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6b. A. tricoccum var. tricoccum Variety burdickii is known from ME, NH, VT. Variety tricoccum is known from CT, MA, ME, NH, RI, VT. The varieties of Allium tricoccum differ morphologically and phenologically (though var. burdickii emerges later than var. tricoccum, it flowers ca. 10–20 days earlier at sites where the two taxa are sympatric). Variety burdickii is far less common in New England. It may deserve species status. 7. Allium vineale L. E crow garlic. CT, MA, ME, RI, VT. Fields, disturbed soil, waste places, often near agricultural areas.

Amaryllidaceae 1a. Perianth with a conspicuous cup-like to tubular corona; tepals 15–35 mm long . . . Narcissus 1b. Perianth without a corona; tepals 7–22 (–25) mm long 2a. Inflorescence with 2–7 flowers; tepals of ± equal length; scape hollow . . . . . . Leucojum 2b. Inflorescence with 1 flower; tepals distinctly unequal; scape solid . . . . . . . . . . Galanthus

Galanthus 1. Galanthus nivalis L. ssp. nivalis E snowdrop. MA. Fields, edges of lawns, roadsides, and other human-disturbed areas. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it was not naturalized.

Leucojum 1. Leucojum aestivum L. E summer snowflake. CT, MA, ME. Fields, edges of lawns, roadsides, and other humandisturbed areas.

A m a ryl l i dac e a e   9 3

Narcissus Reference: Straley and Utech (2002). 1a. Corona tubular, 30–35 mm long, yellow, with a flared and ruffled apex; stamens uniseriate, all exserted to near midlength of corona; tepals yellow, 2.5–3.5 cm long . . . . . . N. pseudonarcissus 1b. Corona cup-like, 3–5 mm long, usually yellow with a red, crenulate apex; stamens biseriate—3 shorter ones included in floral tube of tepals, 3 longer ones exserted from the mouth of corona; tepals white, 1.5–2.5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. poeticus 1. Narcissus poeticus L. E poet’s daffodil. CT, MA, ME, VT. Fields, edges of lawns, roadsides, and other humandisturbed areas. 2. Narcissus pseudonarcissus L. E common daffodil. ct, ma, ri. Fields, edges of lawns, roadsides, and other human-disturbed areas.

Araceae References: Landolt (2000), Thompson (2000b). 1a. Plants thalloid, not differentiated into stems and leaves, 0.5–15 mm long, floating on or near the surface of water [Figs. 61, 62] 2a. Thalli without roots, lacking veins; flowers without a spathe; thallus buds produced from a single, terminal pouch or cavity at base of parent thallus 3a. Thalli ovoid to globose or boat-shaped (i.e., flattened above and rounded below), without air spaces [Figs. 61, 62]; flower produced from cavity along median line of upper thallus surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wolffia 3b. Thalli flat, elongate, narrowly sabre-shaped, with air spaces [Fig. 63]; flower produced from a cavity at the side of the median line of upper thallus surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wolffiella 2b. Thalli with roots, with 1 or more veins; flowers with a small, membranous spathe; thallus buds from 2 lateral pouches at parent thallus base 4a. Each thallus with 2–21 roots, the lower surface red-purple, the upper surface with (3–) 5–16 (–21) veins 5a. Thalli with (5–) 7–16 (–21) veins, 1–1.5 times as long as wide; roots numbering 7–21 per thallus, only 1 or 2 perforating the small, membranous scale at thallus base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spirodela 5b. Thalli with (3–) 5–7 veins, 1.5–2 times as long as wide; roots numbering 2–7 (–12), all perforating the small, membranous scale at thallus base . . . . . . . . . . . . . . Landoltia 4b. Each thallus with a single root, the lower surface green (purple in L. turionifera), the upper surface with 1–3 (–5) nerves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lemna 1b. Plants not thalloid, differentiated into stems and leaves, much larger, terrestrial or emergent aquatics 6a. Leaves compound, with 3–13 (–21) leaflets; flowers concealing only the basal portion of the axis of the spadix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arisaema 6b. Leaves simple; flowers concealing most or all of the axis of the spadix 7a. Spathe absent; adaxial surface of leaf blades dark blue-green with a conspicuous velvety sheen created by a waxy epidermal layer . . . . . . . . . . . . . . . . . . . . . . . . . . . Orontium

94  MO N OCOTS

7b. Spathe present; leaf blades otherwise 8a. Spathe adaxially white, open, nearly flat, not concealing the spadix; fruit a cluster of distinct, red berries; plants from a long, creeping rhizome . . . . . . . . . . . . . . . . . Calla 8b. Spathe green, or green to brown and marked with purple, convolute, concealing the basal portion of the spadix; fruit a cluster of green to brown berries or mature carpels submerged in the fleshy spadix; plants from erect rhizomes or fibrous roots 9a. Mature leaf blades rounded or cordate at the base, lacking a prominent vein that extends into the short, basal lobes (when present); spadix subglobose; flowers with 4 tepals; fruit of mature carpels submerged in the fleshy spadix; plants malodorous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symplocarpus 9b. Leaf blades sagittate to hastate, with pronounced basal lobes and a conspicuous vein extending into each lobe; spadix elongate; flowers without perianth; fruit a green to brown berry; plants not malodorous . . . . . . . . . . Peltandra

Arisaema Reference: Treiber (1980). 1a. Leaves with 3 (–5) leaflets; spadix 3–9 cm long, apically blunt, much shorter than the spathe; spathe 6–16 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. triphyllum 1b. Leaves with (5–) 7–13 (–21) leaflets; spadix 6–20 cm long, tapering to a slender point, much longer than the spathe; spathe 3–6 (–12) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . A. dracontium 1. Arisaema dracontium (L.) Schott N green-dragon. Muricauda dracontium (L.) Small • CT, MA, NH, VT. Rich, mesic forests, riparian forests. 1 × 2c. This very rare Jack-in-the-pulpit hybrid is known from MA. It is usually larger than either parent and has 5- to 7-foliate leaf blades with elliptic or broad-elliptic to broad-oblanceolate leaflets (the leaflets of Arisaema dracontium are narrower— narrow-lanceolate or narrow-elliptic to oblanceolate or elliptic). The hybrid has a spadix that shortly exceeds the spathe, the latter of which is green with purple near the apex and is mostly 5.1–10.5 cm long. 2. Arisaema triphyllum (L.) Schott N Jack-in-the-pulpit. 2a. Arisaema atrorubens (Ait.) Blume; 2b. Arisaema acuminatum Small; A. pusillum (Peck) Nash; A. triphyllum (L.) Schott var. pusillum Peck; 2c. Arisaema atrorubens (Ait.) Blume var. stewardsonii (Britt.) Stevens; A. stewardsonii Britt.; A. triphyllum (L.) Schott var. stewardsonii (Britt.) Stevens • CT, MA, ME, NH, RI, VT; throughout. Mesic forests, swamps, riparian forests, peatlands, and wetland edges. Three varieties of Arisaema triphyllum are found in our area. They are morphologically distinct and show some ecological separation. Further, there are ploidy level differences between some of the taxa. Some authors choose not to recognize these varieties because hybridization is known between them and some characters are lost in pressing and drying. However, neither of these is a valid reason for not recognizing tracheophyte taxa. Characters such as presence vs. absence of bloom on the leaf blades, fluting of the spathe tube, and orientation of the spathe flange (all lost or obscured on herbarium specimens) are best noted in the field so they can be recorded on herbarium specimen labels. 1a. Mature leaf blades glaucous abaxially; lateral leaflets very gibbous or even lobed near the base of the basiscopic margin; spathe flanges (2–) 4.5–7 (–9) mm wide, plane or slightly revolute; sterile spadix appendage clavate (rarely cylindric), 4–10 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a. A. triphyllum ssp. triphyllum 1b. Mature leaf blades green abaxially; lateral leaflets slightly to moderately gibbous near the base of the basiscopic margin; spathe flanges 1–3 mm wide, revolute; sterile spadix appendage cylindric (rarely clavate), 2–5 mm in diameter

Ar ac e a e   9 5

2a. Spathe tube not or only weakly fluted; spathe hood wholly green or wholly purple, without white stripes; spathe acuminate (less frequently acute) at the apex; plants occurring in rich, mesic forests, swamps, and peatlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. A. triphyllum ssp. pusillum (Peck) Huttleston 2b. Spathe tube strongly fluted; spathe hood green with white and/or purple stripes; spathe acute at the apex; plants occurring in swamps, marshes, low floodplains, and other hydric communities . . . . . . . . . . . . . . 2c. A. triphyllum ssp. stewardsonii (Britt.) Huttleston Subspecies triphyllum is known from CT, MA, ME, NH, RI, VT. Subspecies pusillum is known from CT, MA, RI, VT. Subspecies stewardsonii is known from CT, MA, ME, NH, RI, VT. It is the most ecologically restricted of the subspecies and is ± confined to hydric soils.

Calla 1. Calla palustris L. N wild calla. CT, MA, ME, NH, RI, VT; throughout. Marshes, laggs, swamps, and lake borders.

Landoltia 1. Landoltia punctata (G.F.W. Mey.) Les & D.J. Crawford E spotted-duck-meal. Lemna punctata G.F.W. Mey.; Spirodela punctata (G.F.W. Mey.) C.H. Thompson • MA. Mesotrophic to eutrophic waters of lakes and rivers.

Lemna Specimens of Lemna are rarely collected with flowers or fruit, L. perpusilla being the only species that reproduces sexually with any frequency. The veins are sometimes difficult to observe on fresh or dried specimens. Therefore, splitting the collection and using both traditional methods (drying and mounting on acid-free paper) along with chemical preservation assist greatly with identification and vouchers the collection. Preserving thalli in 70% ethanol (C₂H₅OH) makes them transparent, which allows veins and aerenchyma tissue to be more easily viewed (place the thallus on a slide to permit light to pass through it). Dried specimens can be boiled in 70% ethanol and then the pigments cleared with 10% sodium hypochlorite (NaOCl). 1a. Thalli (3–) 6–15 mm long, tapering to a long stalk, with distally denticulate margins; lateral thalli remaining attached to the parent thallus by a narrow, green stalk, forming tangled colonies suspended just below the surface of the water . . . . . . . . . . . . . . . . . . . . . . . . . L. trisulca 1b. Thalli 1–6 (–8) mm long, sessile or nearly so, with entire margins; lateral thalli separating from the parent thallus or remaining attached by a thin, white stipe and forming floating rosettes 2a. Mature thalli with a single vein (the vein sometimes obscure and then the thallus appearing unveined) 3a. Vein usually not extending past aerenchyma tissue, extending less than ⅔ of distance from root node to tip of thallus; thalli 1–2.5 (–4) mm long, 1–2 times as long as broad, broad-elliptic to broad-oblong or obovate, usually solitary or paired, symmetrical at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. minuta 3b. Vein usually extending past aerenchyma tissue, extending at least ¾ of the distance from the root node to the tip of the thallus; thalli 2–4 mm long, 1.3–3 times as long as wide, elliptic to lanceolate or obovate, often in clusters of (2–) 4–8 (–10), asymmetrical at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. valdiviana 2b. Mature thalli with 3 (–5) veins

96  MO N OCOTS

4a. Root sheath winged at base; roots up to 3.5 cm long, usually sharply pointed at apex; thalli without anthocyanic pigment, usually with a distinct apical papilla . . . . . L. perpusilla 4b. Root sheath not winged at base; roots to 15 cm long, frequently longer than 3.5 cm, rounded at apex; thalli often with at least some spotting or diffuse coloring with anthocyanic pigment, with or without a distinct apical papilla 5a. Plants sometimes with small green-brown to brown rootless turions, these 0.8–1.6 mm in diameter and sinking to bottom; thalli often with anthocyanic coloration on lower surface, the coloration more intense than that of upper surface, usually with a distinct row of minute papillae along the midvein . . . . . . L. turionifera 5b. Plants without turions; thalli typically green on lower surface, when slightly anthocyanic, the coloration less intense than that of upper surface, with or without a distinct row of minute papillae along the midvein . . . . . . . . . . . . . . . . . . . . . . . L. minor 1. Lemna minor L. N common duckweed. Lemna minima Chev. ex Schleid. • CT, MA, ME, NH, RI, VT; throughout. Mesotrophic to eutrophic waters of lakes, rivers, beaver flowages, and pools. 2. Lemna minuta Kunth E least duckweed. Lemna minima Phil. ex Hegelm.; L. minuscula Herter; L. valdiviana Phil. var. minima Hegelm. • MA. Pools. Lemna minuta growing in shade often has slightly longer thalli that are more transparent than those growing in sun. Further, shade-grown plants sometimes grow in connected clusters of up to four plants, making shade grown plants much harder to differentiate from L. valdiviana. Relative root length is one of the most important characters and appears to be stable (i.e., not environmentally controlled). 3. Lemna perpusilla Torr.

NC

minute duckweed. Lemna perpusilla Torr. var. trinervis Austin • MA, RI, VT; also reported from CT by Hellquist and Crow (1982), but specimens are unknown. Mesotrophic to eutrophic waters of lakes, rivers, beaver flowages, and pools. The report of Lemna perpusilla from ME by Landolt (2000) was in error. The voucher specimen was L. minor (Elias Landolt, personal communication). 4. Lemna trisulca L. N ivy-leaved duckweed. CT, MA, ME, NH, RI, VT; extending north in the Lake Champlain Valley but not found in a large portion of the interior of northern New England. Circumneutral, mesotrophic to eutrophic waters of lakes and rivers. 5. Lemna turionifera Landolt

NC

turion duckweed. CT, MA, RI, VT. Mesotrophic to eutrophic waters of lakes, rivers, beaver flowages, and pools. 6. Lemna valdiviana Phil.

NC

pale duckweed. Lemna torreyi Austin • CT, MA, NH, RI. Mesotrophic to eutrophic waters of lakes, rivers, beaver flowages, and pools.

Orontium 1. Orontium aquaticum L.

NC

golden-club. CT, MA, RI. Shallow water of lakes and rivers, river banks, including freshtidal shorelines.

Peltandra 1. Peltandra virginica (L.) Raf. ex Schott N green arrow-arum. Arum virginicum L. • CT, MA, ME, NH, RI, VT; not northern. Swamps, shallow water of lakes and rivers, and organic soil wetlands.

A r ac e a e   97

Spirodela 1. Spirodela polyrrhiza (L.) Schleid. N common duck-meal. Lemna polyrrhiza L.; Spirodela polyrrhiza (L.) Schleid. var. masonii Daubs • CT, MA, ME, NH, RI, VT. Slightly acidic to circumneutral, mesotrophic to eutrophic waters of lakes, rivers, beaver flowages, and pools.

Symplocarpus 1. Symplocarpus foetidus (L.) Salisb. ex Nutt. N skunk-cabbage. Dracontium foetidum L.; Spathyema foetida (L.) Raf. • CT, MA, ME, NH, RI, VT. Swamps, lake and stream borders, and hydric riparian forests.

Wolffia 1a. Thalli ovoid to globose, 1–1.5 times as deep as wide, without pigment cells in the vegetative tissue, thus the upper surface transparently green in life . . . . . . . . . W. columbiana 1b. Thalli boat-shaped (i.e., flattened above and rounded below) [Figs. 61, 62], 0.3–1 times as deep as wide, with pigment cells in the vegetative tissue, thus the upper surface intensely green in life 2a. Thalli rounded at each apex, with a large, prominent papilla in the center of the upper surface, 1–1.5 times as long as wide [Fig. 62] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. brasiliensis 2b. Thalli with a minute, upward-bent point at each apex, lacking a central papilla, 1.3–2 times as long as wide [Fig. 61] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. borealis 1. Wolffia borealis (Engelm. ex Hegelm.) Landolt ex Landolt & Wildi N Fig. 61 northern water-meal. Wolffia brasiliensis Weddell var. borealis Engelm. ex Hegelm.; W. punctata, auct. non Griseb. • CT, MA, ME, NH, VT; extending north in the Lake Champlain Valley but absent over much of northern New England. Mesotrophic to eutrophic waters of lakes, rivers, and backwaters.

Fig. 61  Thallus of Wolffia borealis.

2. Wolffia brasiliensis Weddell N Fig. 62 Brazilian water-meal. Bruniera punctata (Griseb.) Nieuwl.; Wolffia papulifera C.H. Thompson; W. punctata Griseb. • CT, MA, ME, RI. Mesotrophic to eutrophic waters of lakes, rivers, and backwaters. 3. Wolffia columbiana Karst. N

Fig. 62  Thallus of Wolffia brasiliensis.

Columbian water-meal. Bruniera columbiana (Karst.) Nieuwl. • CT, MA, ME, NH, VT; rare and sporadic in northern New England. Mesotrophic to eutrophic waters of lakes, rivers, and backwaters.

Wolffiella 1. Wolffiella gladiata (Hegelm.) Hegelm.

N C Fig. 63

sword-like bog-mat. Wolffia floridana (J.D. Sm.) J.D. Sm. ex Hegelm.; W. gladiata Hegelm.; W. gladiata Hegelm. var. floridana J.D. Sm.; Wolffiella floridana (J.D. Sm.) C.H. Thompson • MA. Still or slow-moving, mesotrophic water of lakes and streams. Fig. 63  Thalli of Wolffiella gladiata.

98  MO N OCOTS

Asparagaceae Asparagus 1. Asparagus officinalis L. E asparagus. CT, MA, ME, NH, RI, VT. Fields, roadsides, and disturbed areas.

Butomaceae Reference: Haynes (2000a).

Butomus 1. Butomus umbellatus L. E flowering-rush. CT, MA, ME, VT; locally common in the Lake Champlain Valley, rare elsewhere. Marshes, lake and stream borders.

Colchicaceae 1a. Tepals connate in the lower portion, forming an elongate basal tube, pink to purple or white (rarely yellow); scape very short (plants appearing stemless); ovary subterranean; plants from a corm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colchicum 1b. Tepals distinct, yellow; stem evident, bearing leaves and flowers [Fig. 64]; ovary elevated well above the ground on an aerial stem; plants from rhizomes . . . . . . . . . . . . . . . . . . . . Uvularia

Colchicum 1. Colchicum autumnale L. E autumn-crocus. MA, NH, VT. Fields, edges of lawns, and roadsides.

Uvularia 1a. Leaf blades sessile, minutely papillose-denticulate; stems angled in cross-section distally; rhizomes elongate, 10–15 cm long, bearing scattered fibrous roots; stolons present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. sessilifolia 1b. Leaf blades perfoliate [Fig. 64], entire; stems terete in cross-section; rhizomes short, ca. 1 cm long, bearing clustered fleshy roots; stolons absent 2a. Tepals with large, orange papillae on the adaxial surface [Fig. 64]; leaf blades glabrous on the abaxial surface; stems with (2–) 3 or 4 leaves below the lowest branch; each lobe of the capsule tipped by 2 attenuate beaks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. perfoliata 2b. Tepals smooth on the adaxial surface; leaf blades usually pubescent abaxially on the veins; stems usually with a single leaf below the lowest branch; each lobe of the capsule tipped by 2 truncate beaks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. grandiflora

Co lchi c ac e a e   9 9

1. Uvularia grandiflora J.E. Sm. N large-flowered bellwort. CT, MA, NH, VT; western New England. Rich, mesic, deciduous forests, often in regions of high-pH bedrock. 2. Uvularia perfoliata L. N Fig. 64 perfoliate bellwort. CT, MA, NH, RI, VT; becoming rare in northern New England. Dry-mesic to mesic, deciduous forests and openings. A specimen stated to have been collected from Shapleigh, York County, ME, is likely the result of mix-up (see Fernald 1950a for discussion). 3. Uvularia sessilifolia L. N sessile-leaved bellwort. Oakesiella sessilifolia (L.) S. Wats. • CT, MA, ME, NH, RI, VT; throughout. Deciduous and mixed evergreen-deciduous forests, forest openings and edges, woodlands.

Commelinaceae Reference: Faden (2000). 1a. Corolla weakly zygomorphic—one of the petals usually smaller than the other 2; inflorescence subtended by a folded spathe; flowers with 3 fertile stamens and usually 3 sterile stamens with cruciform anthers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Commelina 1b. Corolla actinomorphic; inflorescence subtended by leaves; flowers with 6 fertile stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tradescantia

Commelina 1a. Spathes white-green or pale green, especially in basal portion of blade, with contrasting dark green veins; lower (i.e., smaller) petal paler than upper 2 petals; capsules 2-locular; seeds rugose and pitted, in addition to reticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. communis 1b. Spathes bright green, without contrasting veins; all petals blue; capsules 3-locular; seeds reticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. diffusa 1. Commelina communis L. E Asiatic dayflower. Commelina communis L. var. ludens (Miq.) C.B. Clarke • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest edges, and disturbed lots. Commelina communis var. ludens is said to be recognized by its darker flowers, yellow antherodes with maroon centers, and relatively broader spathes (among other characters). Faden (2000) reported difficulty separating some material into this variety or var. communis. Therefore, var. ludens is considered a synonym of the typical variety. 2. Commelina diffusa Burman f. var. diffusa E climbing dayflower. Commelina longicaulis Jacq. • VT. Fields, roadsides, forest edges, and disturbed lots.

Tradescantia Garden hybrids that are believed to represent Tradescantia ohiensis crossing with T. subaspera Ker-Gawl. × T. virginiana have been collected in MA (Sorrie and Somers 1999). This ‌andersoniana plant has been called T. × W. Ludwig & Rohweder. However, the name is invalid (Faden 2000). Further, the name has been used as a catchall for several different cultivated taxa (i.e., it does not represent a single, genetic entity). 1a. Sepals and pedicels pubescent with a mixture of glandular and eglandular hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. bracteata

Fig. 64  Flower of Uvularia perfoliata.

10 0  MONOCOTS

1b. Sepals and pedicels glabrous or entirely pubescent with eglandular hairs 2a. Pedicels pubescent [Fig. 65]; sepals pubescent [Fig. 65] . . . . . . . . . . . . . . . . . T. virginiana 2b. Pedicels glabrous; sepals glabrous or with an apical tuft of hairs . . . . . . . . . . T. ohiensis 1. Tradescantia bracteata Small E long-bracted spiderwort. MA, VT. Fields, roadsides, disturbed rights-of-way, areas of cultivation. 2. Tradescantia ohiensis Raf. n smooth spiderwort. Tradescantia canaliculata Raf.; T. foliosa Small; T. incarnata Small; T. ohiensis Raf. var. foliosa (Small) MacRoberts; T. reflexa Raf. • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed rights-of-way, areas of cultivation. This species is native in CT and MA and non-native in ME, NH, RI, VT. 2 × 3. A relatively common spiderwort hybrid that is known from CT, MA, ME, NH. It can be identified by the presence of sparse pubescence on the pedicels and/or sepals (not including an apical tuft of hairs) vs. moderate to dense pubescence in Tradescantia virginiana. 2 × Tradescantia subaspera Ker-Gawl. is a very rare spiderwort hybrid likely originating from a garden planting. It is known from MA. It has glandular hairs on the sepals. 3. Tradescantia virginiana L. E Fig. 65 Virginia spiderwort. Tradescantia brevicaulis Raf.; T. virginiana L. var. alba Hook. ex Raf. • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed rights-of-way, and areas of cultivation.

Fig. 65  Eglandular pubescence of pedicels and sepals of Tradescantia virginiana.

Cyperaceae 1a. Perianth at maturity composed of elongate capillary bristles that greatly exceed the subtending floral scales, creating a “woolly” or “cottony” appearance to the individual spikes [Fig. 120] 2a. Perianth bristles strongly bent, curled, and tangled; inflorescences with 50–500 spikes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Scirpus 2b. Perianth bristles relatively straight, silky; inflorescences with 1–10 (–30) spikelets 3a. Perianth bristles numbering 10–25 per flower; spikelets solitary or 2–10 (–30) per inflorescence, 10–50 mm long in fruit, subtended by 1 or more foliaceous bracts or by 10–15 sterile, basal scales; principal leaf blades up to 45 cm long . . . . . Eriophorum 3b. Perianth bristles mostly 6 in each flower; spikelets solitary, 5.4–8 mm tall in fruit, subtended by a solitary involucral bract that resembles an enlarged, basal scale; leaf blades up to 1 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Trichophorum 1b. Perianth absent or present, when present not composed of greatly elongated capillary bristles, mostly contained within the floral scales [Figs. 68, 73, 110] 4a. Carpellate flowers subtended by an enclosing, sac-like scale (i.e., a perigynium) [Figs. 67, 72, 80, 102], the stigmas protruding through an orifice at the top; flowers unisexual, lacking perianth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carex 4b. Carpellate flowers subtended by flat or folded scales, never enclosed in a sac-like scale [Figs. 105, 110, 125]; flowers bisexual or some unisexual (all unisexual in Scleria), with or without perianth 5a. Spikes 1 per stem

Cy p e r ac e a e   10 1

6a. Involucral bract solitary, erect, appearing to be a continuation of the stem, therefore, the inflorescence falsely appearing to emerge from the side of the stem; leaf blades flaccid, submerged or the tips floating . . . . . . . . (in part) Schoenoplectus 6b. Involucral bracts (if present) not resembling a continuation of the stem, therefore, the inflorescence appearing terminal; leaf blades either not produced or firm 7a. Achenes crowned by a persistent tubercle or expanded style base, usually of different color and texture than body of achene [Figs. 112, 116, 118]; leaf sheaths lacking blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eleocharis 7b. Achenes not crowned by a tubercle; at least the upper leaf sheaths bladebearing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Trichophorum 5b. Spikes usually 2 or more per stem 8a. Scales of spikes distichous (i.e., arranged in 2 ranks) [Fig. 105] 9a. Inflorescence partly or wholly axillary; flowers with a perianth composed of bristles; style persistent on the achene as a slender beak; leaves borne on the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dulichium 9b. Inflorescence terminal; flowers lacking perianth; style deciduous from the achene; leaves basally disposed 10a. Spikelets with 3–30 alternate floral scales [Fig. 105, 107]; inflorescence dense to open, commonly with branches (i.e., not all spikes sessile); achenes biconvex to trigonous in cross-section, when biconvex the edge of the achene fitting into a groove in the rachilla . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperus 10b. Spikelets with 2 (rarely 3) subopposite floral scales, the basal scale subtending a bisexual flower, the apical scale subtending a staminate flower; inflorescence densely capitate, with sessile spikes; achenes biconvex, the edge not fitting into a groove in the rachilla . . . . . . . . . . . . . . . . . . . . . . . . Kyllinga 8b. Scales of spikes spirally arranged (i.e., arranged in 3 or more ranks) 11a. Flowers all unisexual; achenes white, gray, or gray-brown, with a hardened disk at the base (i.e., a hypogynium; the hypogynium vestigial in S. verticillata) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scleria 11b. Flowers bisexual or some unisexual; achenes mostly gray-brown or pale brown to dark brown or black, without a hypogynium 12a. Principal involucral bract solitary, erect, appearing to be a continuation of the stem, therefore, the inflorescence falsely appearing to emerge from the side of the stem 13a. Achenes 0.5–0.7 mm long; spikelets 1–3 (–5) mm long; perianth consisting of a single, diminutive scale; floral scales 0.5–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lipocarpha 13b. Achenes 1.3–4 mm long; spikelets 5–15 mm long; perianth consisting of bristles [Fig. 126], these sometimes caducous; floral scales 2–6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Schoenoplectus 12b. Principal involucral bract erect to spreading, flat, folded, or setaceous, accompanied by additional bracts, not seeming a continuation of the stem, the inflorescence appearing terminal 14a. Achenes with an evident, apical tubercle that covers much or all of the achene summit, the tubercle clearly differentiated from the achene body by color and/or texture [Figs. 122, 123, 124] . . . . . . . . . . . . . . . . . . . . Rhynchospora 14b. Achenes without tubercles or with small, relatively inconspicuous style bases [Figs. 66, 121]

102  MONOCOTS

15a. Perianth dimorphic—an outer set of 3 retrorse-barbellate bristles and an inner set of 3 stipitate blades terminating in acuminate or awned apices [Fig. 121]; leaf sheaths pubescent . . . . . . . . . . . . . . . . . . . . . . . . Fuirena 15b. Perianth monomorphic, of smooth to retrorse-barbellate bristles, or absent [Fig. 66]; leaf sheaths glabrous 16a. Plants annual, with soft stem bases and fibrous roots, 5–30 cm tall 17a. Stems capillary; leaves up to 0.5 mm wide, with two lateral tufts of hairs at the junction of the blade and sheath; style base persistent in fruit and forming a tiny tubercle . . . . . . . . . Bulbostylis 17b. Stems flat; larger leaves 1–3 mm wide, none with two lateral tufts, but with a ligule of short hairs; style base not persistent in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fimbristylis 16b. Plants perennial, with hardened stem bases and rhizomes, 30–200 cm tall 18a. Perianth absent; achenes terete in cross-section; each spike with 1–3 empty, basal scales and an apical, bisexual flower, the remaining scales subtending staminate flowers . . . . . . . . . . Cladium 18b. Perianth usually present, caducous in some species (rarely absent); achenes lenticular or biconvex to trigonous in cross-section; spikes comprised entirely of bisexual flowers 19a. Scales pubescent on the abaxial surface; rhizomes with tuberous thickenings; spikes (7–) 10–40 mm long; achenes 2.3–5.5 mm long; leaves without ligules . . . . . . . . Bolboschoenus 19b. Scales glabrous on the abaxial surface; rhizomes without tubers; spikes 2–12 mm long; achenes 0.6–1.6 mm long; leaves with ligules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Scirpus

Bolboschoenus Anther color, scale translucence, and anatomy of the achenes are critically important for identification of Bolboschoenus specimens. Anther color should be assessed on dried specimens (or dehisced anthers on living plants). Scale translucence is best viewed under a dissecting scope at 10–15 power. Placing a narrow probe or similar tool under the floral scale with good lighting is a good method of assessing how much light passes through the scale (based on how readily the probe can be viewed through the scale). Achene anatomy is viewed in cross-section (i.e., the achenes need to be cut in half with a sharp blade). Hybrids are relatively common and can form large colonies. They generally show decreased fertility and produce fewer well-formed achenes. References: Browning et al. (1995), Smith (2002a). 1a. Inflorescence normally with elongate branches; styles mostly trifid; achenes compressed-trigonous to equilaterally trigonous; perianth bristles present on mature achenes [Fig. 66], sometimes a few caducous; achene exocarp layer 0.15–0.5 times as deep as the mesocarp layer; plants of fresh to slightly brackish shores 2a. Widest leaf blades 2–6 mm wide; spikes slender, mostly 3–5 mm in diameter; achenes 2.5–3.3 mm long, with a minute, persistent style beak; veins of leaf sheaths gradually diverging near summit, leaving an obtriangular, veinless area . . . . . . . . . . . . . . . . . B. glaucus 2b. Widest leaf blades 7–22 mm wide; spikes thicker, mostly 5–10 mm in diameter; achenes 3–5.5 mm long, with an evident style beak 0.1–0.8 mm long [Fig. 66]; veins of leaf sheaths nearly reaching summit before abruptly diverging 3a. Achenes 3.8–5.5 mm long, usually equilaterally trigonous, with isodiametric exocarp cells; perianth bristles firmly attached to achene, equaling the length of the achene [Fig. 66]; anthers yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. fluviatilis

Cy p e r ac e a e   103

3b. Achenes 3–4.3 mm long, usually compressed-trigonous, with exocarp cells 1.5–3 times as tall as wide; perianth bristles weakly attached to achene, 0.5–1 times the length of the achene; anthers dark yellow to orange-yellow . . . . . . . . B. novae‑angliae 1b. Inflorescence normally congested and lacking elongate branches; styles bifid or trifid; achenes plano-convex to compressed-trigonous; perianth bristles rarely present on mature achenes; achene exocarp layer ± 2 times as deep as the mesocarp layer; plants of saline to brackish shores 4a. Floral scales opaque, papery-textured, with awns ca. 0.5 mm wide at the base; anthers yellow when fresh, becoming orange-brown to red-brown after dehiscence or in drying; achenes dark brown, often compressed-trigonous; apex of leaf sheath opposite the leaf blade with abruptly diverging veins, green and firm-textured, prolonged . . . . . . B. robustus 4b. Floral scales translucent, membranous-textured, with awns ca. 0.25 mm wide at the base; anthers yellow when fresh, becoming pale yellow or pale yellow-brown after dehiscence or in drying; achenes light brown to dark brown, lenticular or rarely compressed-trigonous; apex of leaf sheath opposite the leaf blade with gradually diverging veins, with a hyaline obtriangular area, not prolonged . . . . . . . . . . . . B. maritimus 1. Bolboschoenus fluviatilis (Torr.) Soják n Fig. 66 river tuber-bulrush. Schoenoplectus fluviatilis (Torr.) M.T. Strong; Scirpus fluviatilis (Torr.) Gray; S. maritimus L. var. fluviatilis Torr. • CT, MA, ME, NH, VT. Marshes, lake shores, river shores, including fresh-tidal sections. This species is native to all states except NH, where the population was introduced in a human-made wetland planting. 2. Bolboschoenus glaucus (Lam.) S.G. Smith E glaucous tuber-bulrush. Schoenoplectus glaucus (Lam.) Kartesz; Scirpus glaucus Lam. • NH. Marshes, shorelines. Unintentionally introduced into a human-made wetland when it was purchased under the name of another tuberous bulrush (i.e., the plant stock was misidentified). 3. Bolboschoenus maritimus (L.) Palla n saltmarsh tuber-bulrush.  3a. Schoenoplectus maritimus (L.) Lye; Scirpus maritimus L. var. maritimus; 3b. Bolboschoenus maritimus (L.) Palla ssp. paludosus (A. Nels.) A. & D. Löve; B. paludosus (A. Nels.) Soó; Scirpus paludosus A. Nels. • CT, MA, ME, NH, RI. Brackish to saline coastal shorelines and marshes. 1a. Styles mostly trifid; achenes medium to dark brown, compressed-trigonous to, less commonly, lenticular; floral scales medium to dark brown . . . . 3a. B. maritimus ssp. maritimus 1b. Styles mostly bifid; achenes white-brown to medium brown (dark brown), usually lenticular; floral scales light yellow-brown to medium brown (rarely dark brown) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. B. maritimus ssp. paludosus (A. Nels.) T. Koyama Subspecies paludosus is native to New England and known from CT, MA, ME, NH, RI. Subspecies maritimus is introduced from Europe and is known only from ME. 3 × 5. This rare tuber-bulrush hybrid is frequently encountered when both parent species are sympatric. It is known from CT, MA, ME, NH, RI. It is similar to both parents but shows dark yellow to orange-yellow anthers and barely translucent scales. 4. Bolboschoenus novae-angliae (Britt.) S.G. Smith

NC

New England tuber-bulrush. Schoenoplectus novae-angliae (Britt.) M.T. Strong; Scirpus cylindricus (Torr.) Britt.; S. novae-angliae Britt. • CT, MA, ME, RI. Brackish-tidal river shores. 5. Bolboschoenus robustus (Pursh) Soják N sea-coast tuber-bulrush. Schoenoplectus robustus (Pursh) M.T. Strong; Scirpus maritimus L. var. macrostachyus Michx.; S. robustus Pursh • CT, MA, ME, NH, RI. Brackish to saline coastal shorelines and marshes.

Fig. 66  Achene of Bolboschoenus fluviatilis.

104 MO NOCOTS

Bulbostylis 1. Bulbostylis capillaris (L.) Kunth ex C.B. Clarke N tufted hair-sedge. Bulbostylis capillaris (L.) Kunth ex C.B. Clarke var. crebra Fern.; Scirpus capillaris L.; Stenophyllus capillaris (L.) Britt. • CT, MA, ME, NH, RI, VT. Well-drained sand and gravel of grasslands, roadsides, borrow pits, and waste places.

Carex Collections of Carex should include basal portions of the stems and underground organs. The basal portion of the plant is important for observing the morphology of the basal leaves (e.g., sheath color, presence/absence of blades, form of sheath upon disintegration). Underground organs are necessary to determine if a species is rhizomatous or cespitose (at the minimum, notes about a plant’s habit should be made on herbarium collections). Collections of Carex that lack mature spikes may not be able to be confidently determined. Some hybridization does occur, particularly within certain sections (e.g., Glareosae, Hymenochlaenae, Phacocystis, Vesicariae). Carex divulsa Stokes was reported from MA by Kartesz (1999), based on Hermann (1954), who reported C. virens Lam. The specimen was stated to be housed at US; however, searches of that museum have not been able to locate a voucher for this report. Carex laevigata Sm. (section Elatae) was thought to have been collected by Greene from Massachusetts near Boston in the early 1800s (Fernald 1911). It is unsure if the plant represents a mix-up in labeling (i.e., the origin was incorrectly thought to be United States) or if this species was truly collected in New England. In either case, no other collections have been reported, and the plant is not included in the following identification keys. Carex comosa × C. lurida was reported from MA by Sorrie and Somers (1999). The voucher specimen is C. lurida—Fernald and Long 18155 (MASS!). Reports of the hybrid Carex crinita × C. scabrata from ma and nh were based on collections of C. gynandra (specimens at GH! and NEBC!). Reports of Carex × ‌sullivantii Boott were based on a specimen cultivated at the ‌aestivaliformis Mackenzie Cambridge Botanic Garden (specimen at NEBC!). Reports of Carex × from MA were based on a collection of C. aestivalis—3 Aug 1916, Churchill s.n. (NEBC!). 1a. Spikes entirely staminate (i.e., no perigynia present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 1 1b. At least some of the spikes partly or entirely carpellate 2a. Spikes 1 per stem; rachilla often present in perigynium . . . . . . . . . . . . . . . . . . . . . . Group 2 2b. Spikes normally 2 or more per stem (except depauperate individuals); rachilla absent from perigynium 3a. Stigmas 2; achenes lenticular to biconvex 4a. Spikes all similar and sessile, bearing staminate flowers at base or apex; bracteoles usually absent from the base of each spike . . . . . . . . . . . . . . . . . . . . . Group 3 4b. Spikes dissimilar and some or all usually peduncled, with staminate flowers borne in separate or mixed spikes; bracteoles usually present at the base of each spike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 4 3b. Stigmas 3; achenes trigonous to terete 5a. Perigynia pubescent or scabrous with hairs or scabrules several times longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 5 5b. Perigynia glabrous (though long-papillose in some species) 6a. Bract of lowest carpellate spike consisting of a blade only, the sheath absent or extremely short, usually not exceeding 4 mm (or the bract absent altogether) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 6 6b. Bract of lowest carpellate spike consisting of a blade and a prolonged, closed sheath, usually longer than 4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 7

Cy p e r ac e a e   10 5

Group 1 1a. Basal leaf sheaths anthocyanic; stigmas trifid 2a. Leaf blades 2.5–6 mm wide, flat; plants colonial from long rhizomes . . . (in part) Paniceae 2b. Leaf blades 1–3 mm wide, V-shaped in cross-section; plants cespitose, with short rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Scirpinae 1b. Basal leaf sheaths not tinged with anthocyanic pigments; stigmas bifid 3a. Spike 1 per stem 4a. Stems 5–20 cm tall, produced singly or few together from long, slender rhizomes; leaf blades 0.3–0.7 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Physoglochin 4b. Stems 12–70 cm tall, cespitose; widest leaf blades 0.8–1.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Stellulatae 3b. Spikes normally 2 or more per stem 5a. Widest leaf blades 4–8 mm wide; anthers 4–6.5 mm long; inflorescences 10–20 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Macrocephalae 5b. Widest leaf blades 1–3 (–3.5) mm wide; anthers (1–) 1.2–3.9 mm long; inflorescences 3–12 (–14) mm wide 6a. Spikes 5–18 (–25) per stem; stems produced singly on long rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Divisae 6b. Spikes 2–8 per stem; stems cespitose 7a. Ligules 0.3–1.7 mm long; anthers (1–) 1.2–2.2 (–2.3­) mm long; staminate scales 2.2–3.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Stellulatae 7b. Ligules 1–3.5 mm long; anthers 1.7–2.9 mm long; staminate scales 2.8–3.9 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Deweyanae

Group 2 1a. Stigmas 2; achenes lenticular to biconvex 2a. Basal sheaths anthocyanic; carpellate scales obtuse at apex; plants of alpine ridges and plateaus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Capituligerae 2b. Basal sheaths not tinged with anthocyanic pigments; carpellate scales acute to acuminate at apex; plants of organic soil wetlands 3a. Stems 5–20 cm tall, produced singly or few together from long, slender rhizomes; leaves 0.3–0.7 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Physoglochin 3b. Stems 12–70 cm tall, cespitose; widest leaves 0.8–1.5 mm wide . (in part) Stellulatae 1b. Stigmas 3; achenes trigonous 4a. Spike unisexual; perigynia pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Scirpinae 4b. Spike bisexual; perigynia glabrous 5a. Spike gynaecandrous; perigynium with a bidentate beak, the body obconic to obovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Squarrosae 5b. Spike androgynous; perigynium beaked or not, the orifice entire to truncate (i.e., without teeth), the body narrow-subulate to globose 6a. Perigynia (5–) 5.9–7.8 mm long, narrow-subulate in outline, spreading to reflexed; style persistent on achene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucoglochin

10 6  MONOCOTS

6b. Perigynia 2.5–5.4 mm long, narrow-ellipsoid to ovoid, ascending to appressed; style deciduous from achene 7a. Lower carpellate scales foliaceous, surpassing and concealing the perigynium; perigynium with a conspicuous beak; leaves 1.3–5.4 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Phyllostachyae 7b. Lower carpellate scales not foliaceous, shorter than to slightly exceeding the perigynium; perigynium beakless; leaves 0.4–1.3 mm wide . . . . . . . Leptocephalae

Group 3 1a. At least the uppermost spike gynecandrous (i.e., the staminate flowers borne at the base of the spike), the others carpellate or gynecandrous 2a. Perigynia with flat, wing-like margins (at least in the apical half), not distended to the edges by achene, without spongy tissue at the base; style persisting on summit of achene as a short apiculus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperoideae 2b. Perigynia with rounded or sharp, unwinged margins, distended to or close to the edges by achene, with spongy tissue at the base; style deciduous from summit of achene 3a. Perigynium rounded or with a rounded edge at the margins; achene nearly filling body of perigynium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glareosae 3b. Perigynium sharp-edged at margins; achene obviously smaller than body of perigynium 4a. At least the lower perigynia of each spike spreading; beak of perigynium 0.4–1.7 mm long; style base not enlarged . . . . . . . . . . . . . . . . . . . . . (in part) Stellulatae 4b. Perigynia erect to ascending; beak of perigynium 1.2–2.2 (–2.8) mm long; style base enlarged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Deweyanae 1b. Spikes androgynous (i.e., the staminate flowers borne at the summit of the spike) 5a. Perigynium unequally biconvex (i.e., plump), with a smooth beak up to 0.25 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dispermae 5b. Perigynium flat or planoconvex, the beak minutely serrulate in most species and 0.3–3.5 mm long 6a. Stems produced singly or few together from elongate rhizomes, stolons, or prostrate stems 7a. Stems leaning or prostrate, producing upright stems from the axils of dried leaves of the previous year; beak of perigynium 0.3–0.6 mm long; plants of organic soil wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chordorrhizae 7b. Stems upright, produced from elongate rhizomes; beak of perigynium 0.7–2.7 mm long; plants of sandy, gravelly, or rocky, frequently well-drained, habitats 8a. Perigynia sharp-margined but not wing-margined, nerveless or inconspicuously nerved on the adaxial surface; plants phyllopodic, with basal tufts of leaves (in part). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Divisae 8b. Perigynia wing-margined, conspicuously nerved on the adaxial surface; plants aphyllopodic, without basal tufts of leaves . . . . . . . . . . . . . . . . . . . . . . Ammoglochin 6b. Stems cespitose, the rhizomes inconspicuous or very short 9a. Leaf sheaths opposite the blade transversely rugose, appearing corrugated 10a. Spikes 3–10 (–15) per inflorescence, occurring singly at the nodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Phaestoglochin

Cy p e r ac e a e   107

10b. Spikes usually more than 15 per inflorescence, 2 or more produced from some nodes (i.e., the inflorescence branched) 11a. Carpellate scales hyaline, with a single vein or 3 veins near the center; culms often wing-angled and easily compressed, usually more than 1 mm wide apically in drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Vulpinae 11b. Carpellate scales yellow or brown, with hyaline margins, with 3 veins near the center; stems usually firm and less than 1 mm wide apically in drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Multiflorae 9b. Leaf sheaths opposite the blade smooth, not rugose 12a. Perigynia truncate to subcordate at base, tapering from base to apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Vulpinae 12b. Perigynia cuneate to rounded at base, widened above the base and then tapering to the apex 13a. Nodes of the inflorescence with a single spike; leaf sheaths opposite the blade neither red-spotted nor copper-colored . . . . . . . (in part) Phaestoglochin 13b. Some nodes of the inflorescence, especially the basal ones, with 2 or more spikes (i.e., the inflorescence branched); leaf sheaths opposite the blade dotted with red and/or strongly copper-colored . . . . . . . . . . . . . . . . . . . . . . Heleoglochin

Group 4 1a. Perigynia pubescent, with an evident, bidentate beak ca. 1 mm long . . . . . . . . . . . . . Gracilis 1b. Perigynia glabrous (though minutely papillose in several species), beakless or with a short, ± entire beak 0.1–0.4 mm long (with an obscurely bidentate beak 0.2–0.8 mm long in Carex saxatilis) 2a. Perigynia white or gold-orange, biconvex in cross-section; lowest bract of carpellate spikes with a prolonged, closed sheath usually longer than 4 mm . . . . . . . . . . . . . . Bicolores 2b. Perigynia green to brown to black, planoconvex to biconvex in cross-section; lowest bract of carpellate spikes lacking a sheath or the sheath shorter than 3 mm 3a. Style deciduous from achene in fruit; leaf sheaths with no, or very few, septate partitions; perigynia often minutely papillose, with a short, inconspicuous beak 0.1–0.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phacocystis 3b. Style persistent on achene in fruit; leaf sheaths with septate partitions; perigynia smooth, with a distinct beak 0.2–0.8 mm long . . . . . . . . . . . . . . . . . . . . (in part) Vesicariae

Group 5 1a. Leaf sheaths and blades glabrous or scabrous, but not pubescent 2a. Perigynia (5.3–) 6–11.5 mm long, the beak terminated by 2 stiff teeth (1–) 1.2–2.3 (–2.8) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Carex 2b. Perigynia 2.2–6.5 mm long, the beak without teeth or with teeth 0.1–0.8 mm long 3a. Lowermost bract of inflorescence (excluding basal spikes) bladeless or with a blade up to 2 mm long, strongly anthocyanic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clandestinae 3b. Lowermost bract of inflorescence with blades more than 3 mm long (usually much longer), not or scarcely anthocyanic 4a. Leaf sheaths with cross-septa connecting the longitudinal veins (sometimes the cross-veins are few and obscure); adaxial surface of leaf blades usually with 2 marginal veins that are more prominent than midvein

10 8  MONOCOTS

5a. Leaf blades adaxially scabrous; beak of perigynium slightly bent, obscurely bidentate at tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anomalae 5b. Leaf blades adaxially smooth or scabrous only along margins and/or near tip; beak of perigynium straight, terminated by two, stiff teeth 0.4–0.8 mm long (with a soft, obscurely toothed tip in C. vestita) . . . . . . . . . . . . . . . . . . . . . (in part) Paludosae 4b. Leaf sheaths without cross-septa connecting the longitudinal veins or these few and obscure; adaxial surface of leaf blades without 2 marginal veins more prominent than midvein 6a. Vegetative leaf blades involute to triangular-channeled, tapering to an elongate, filiform, curling tip; plants of hydric communities, including peatlands, lakeside fens, and graminoid marshes . . . . . . . . . . . . . . . . . . . . . . (in part) Paludosae 6b. Vegetative leaf blades ± flat to V-shaped in cross-section, pointed at the apex, but not long-prolonged into very thin, curling tips; plants primarily of mesic to xeric soils (frequently of wetlands in C. flacca, but that species with arching or drooping lower spikes) 7a. Basal spike of leafy, flowering stems long-peduncled, often arching or even drooping; lateral spikes cylindric, 20–45 mm long, the basalmost with more than 30 perigynia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thuringiaca 7b. Basal spike of leafy, flowering stems sessile or short-peduncled, erect to ascending; lateral spikes subglobose to short-obloid or short-ellipsoid, 2–15 mm long, the basalmost with (1–) 3–22 perigynia 8a. Style base expanded into a circular ring and persistent at summit of achene; beak of perigynium ca. 0.2 mm long; carpellate scales with a scabrous awn up to 2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mitratae 8b. Style base not expanded, deciduous; beak of perigynium 0.4–2 mm long; carpellate scales awnless or with an awn shorter than 1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Acrocystis 1b. Leaf sheaths and often the blades pubescent 9a. Perigynia 4.8–12 mm long, with a beak terminated by 2 stiff teeth 0.8–3 mm long; style persistent at summit of achene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Carex 9b. Perigynia 1.7–5 mm long, beakless or beaked and then terminated by teeth up to 0.7 mm long; style deciduous 10a. Perigynia 1.7–4 mm long, beakless or essentially so; uppermost spike bearing some perigynia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Porocystis 10b. Perigynia 3.5–5 mm long, with a prominent, bidentate beak; uppermost spike entirely staminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hirtifoliae

Group 6 1a. Leaf sheaths, and often the blades, pubescent 2a. Perigynia (6.5–) 7–12 mm long, with a beak 2.1–4 mm long that is terminated by 2 teeth (1.2–) 1.5–3 mm long; style persistent at summit of achene . . . . . . . . . . . . . . . . . (in part) Carex 2b. Perigynia 2–5 mm long, without a beak or with a beak up to 1.5 mm long that is terminated by 2 teeth up to 0.5 mm long; style deciduous from achene 3a. Perigynia beakless or essentially so; adaxial surface of leaf blades without 2 marginal veins more prominent than midvein . . . . . . . . . . . . . . . . . . . (in part) Porocystis 3b. Perigynia with a distinct beak; adaxial surface of leaf blades usually with 2 marginal veins that are more prominent than midvein . . . . . . . . . . . . . . . (in part) Hymenochlaenae

Cy p e r ac e a e   10 9

1b. Leaf sheaths and blades glabrous or scabrous, but not pubescent 4a. Style persistent on achene; sheaths, and often blades, of larger leaves septate-nodulose 5a. Body of perigynium obconic, abruptly tapering to the beak; terminal spike gynecandrous or entirely staminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Squarrosae 5b. Body of perigynium lanceoloid or ellipsoid to ovoid or rhombic-ovoid, gradually to moderately tapering to the beak; terminal spike staminate 6a. Perigynia 10–20 mm long; staminate spikes usually 1 per plant; carpellate spikes compact, globose to ovoid or obovoid . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lupulinae 6b. Perigynia (3.2–) 3.6–12.5 mm long; staminate spikes mostly 1–3 per plant; carpellate spikes compact to elongate, obloid or ovoid to cylindric (sometimes nearly globose in a few species) 7a. Carpellate scales with a long, scabrous awn, the body of the scale often ciliate on the margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Vesicariae 7b. Carpellate scales without prominent scabrous awns (sometimes with a smooth awn in C. striata), without ciliate margins 8a. Perigynia with 5–14 (–15) veins, relatively thin-walled . . (in part) Vesicariae 8b. Perigynia with 14–28 veins, firm, thick-walled . . . . . . . . . (in part) Paludosae 4b. Style deciduous from achene; sheaths and blades not septate-nodulose (except in section Ceratocystis and very rarely others) 9a. Perigynia with a serrulate wing-margin (this often revolute); achenes 4–7.5 mm long; inflorescence usually composed of 20 or more spikes densely aggregated together into ± a single cluster (the lowest spike sometimes removed from the others) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Macrocephalae 9b. Perigynia without a wing-margin; achenes 1–2.7 mm long; inflorescence composed of 1–9 spikes, these not aggregated into a dense cluster (except in C. media) 10a. Uppermost spike gynecandrous 11a. Perigynia minutely papillose; adaxial surface of leaf blades without 2 marginal veins more prominent than midvein; leaf blades V-shaped in cross-section (at least when young); lateral spikes globose to obloid, 5–20 (–25) mm long . . . . . Racemosae 11b. Perigynia without minute papillae; adaxial surface of leaf blades usually with 2 marginal veins that are more prominent than midvein; leaf blades M-shaped cross-section (at least when young), sometimes ± flat at maturity; lateral spikes cylindric, 15–60 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Hymenochlaenae 10b. Uppermost spike entirely staminate or androgynous 12a. Perigynia minutely papillose 13a. Lateral spikes arching to nodding; some roots with a dense, felt-like pubescence of yellow-brown hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limosae 13b. Lateral spikes erect or ascending; roots without dense pubescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Paniceae 12b. Perigynia smooth 14a. Lowest carpellate scales concealing and much surpassing the perigynia, entirely green, leaf-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Phyllostachyae 14b. Lowest carpellate scales not surpassing perigynia, green to yellow-green or brown, with hyaline margins, not leaflike

110 MO NOCOTS

15a. Lower perigynia in each spike horizontally spreading to reflexed; leaf sheaths, and often blades, of larger leaves septate-nodulose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Ceratocystis 15b. Lower perigynia in each spike ascending to spreading-ascending; leaf sheaths and blades not septate-nodulose 16a. Lateral spikes 25–80 mm long; adaxial surface of leaf blades with 2 marginal veins more prominent than midvein; leaf blades M-shaped in cross-section (at least when young), sometimes ± flat at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Hymenochlaenae 16b. Lateral spikes 4.5–11.3 mm long; adaxial surface of leaf blades without 2 marginal veins more prominent than midvein; leaf blades V-shaped in cross-section (at least when young) . . . . (in part) Acrocystis

Group 7 1a. Style persistent on achene; perigynia (2.2–) 3.4–19 mm long 2a. Teeth of perigynium reflexed; achenes narrow-obloid . . . . . . . . . . . . . . . . . . . . . Collinsiae 2b. Teeth of perigynium erect; achenes obloid-ovoid or obloid-obovoid to ellipsoid, rhomboid, or rhombic-ovoid 3a. Body of perigynium obconic, abruptly tapering to the beak; uppermost spike gynecandrous or entirely staminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Squarrosae 3b. Body of perigynium lanceoloid to ovoid or obovoid, moderately to gradually tapering to the beak; uppermost spike staminate 4a. Carpellate scales with a long, scabrous awn, the body of the scale often ciliate on the margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Vesicariae 4b. Carpellate scales either without prominent scabrous awns or with awns much shorter than the body of the scale, without ciliate margins 5a. Perigynia 8.7–12.1 mm long, lanceoloid, 4–7 times as long as wide; basal sheaths yellow-brown to brown, without anthocyanins . . . . . . . . . . . . . . . . Rostrales 5b. Perigynia 11–19 mm long, narrow-ovoid, less than 7 times as long as wide; basal sheaths usually tinged with anthocyanic pigments . . . . . . (in part) Lupulinae 1b. Style deciduous from achene; perigynia 1.5–6.8 (–9.5) mm long 6a. Leaf blades V-shaped or U-shaped in cross-section (at least when young); adaxial surface of leaf blades without 2 marginal veins that are more prominent than midvein 7a. Bracts of inflorescence consisting of only a sheath; perigynia green, maturing glossy, dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Albae 7b. At least the lowest bracts of the inflorescence with a short blade; perigynia variously colored, but not glossy dark brown at maturity 8a. Lower perigynia in each spike horizontally spreading to reflexed; carpellate spikes globose to short-ellipsoid or short-cylindric . . . . . . . . . . . . . . . . . (in part) Ceratocystis 8b. Lower perigynia in each spike ascending to spreading-ascending; carpellate spikes short-cylindric or ovoid to narrow-cylindric 9a. Abaxial leaf blade and surface of perigynia minutely papillose; plants colonial from long rhizomes, with strongly glaucous leaves . . . . . . . . . . . . (in part) Paniceae 9b. Abaxial leaf blade and surface of perigynia without papillae; plants loosely to densely cespitose from short rhizomes (colonial from long rhizomes in C. crawei), with green to moderately glaucous leaves

Cy p e r ac e a e   1 1 1

10a. Perigynia rounded at base, apically with convexly rounded sides, often with red-brown or yellow streaks and spots . . . . . . . . . . . . . (in part) Granulares 10b. Perigynia tapering to base, apically with flat or somewhat convex sides, without streaks or spots 11a. Perigynia with 2 evident marginal veins, the remaining veins obscure, the beak erect; achenes 1.2–1.7 mm long; lateral spikes, especially the lower, usually spreading or drooping (ascending in alpine populations) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chlorostachyae 11b. Perigynia with (22–) 25–32 evident veins, the beak usually curved or bent; achenes 2–2.8 mm long; lateral spikes usually erect or ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Laxiflorae 6b. Leaf blades M-shaped in cross-section (at least when young), sometimes ± flat at maturity; adaxial surface of leaf blades usually with 2 marginal veins that are more prominent than midvein 12a. Perigynium with 2 evident marginal veins, the remaining veins obscure or evident only in the basal portion of the perigynium body 13a. Perigynium abruptly tapered to a beak 1.7–4 mm long, the beak nearly as long as the perigynium body and terminated by 2 hyaline teeth 0.5–1.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Hymenochlaenae 13b. Perigynium beakless or moderately tapering to a beak 0.1–1.8 (–2.2) mm long, the beak definitely shorter than the perigynium body and entire or weakly bidentate with terminal teeth up to 0.1 mm long 14a. Perigynium with a curved or abruptly bent beak; plants cespitose, with short rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Laxiflorae 14b. Perigynium beakless or with a straight beak; plants colonial from long rhizomes 15a. Perigynium neither streaked nor spotted with red-brown; abaxial leaf blade and surface of perigynia densely and minutely papillose (both usually smooth in C. vaginata) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Paniceae 15b. Perigynium often streaked or dotted with red-brown; abaxial leaf blade and surface of perigynia without minute papillae . . . . . . . . (in part) Granulares 12b. Perigynium with 2 evident marginal veins as well as 8–40+ additional distinct, fine veins that travel ± the length of the perigynium body 16a. Perigynia rounded at base, apically with convexly rounded sides 17a. Perigynia with impressed veins (especially in drying), without red-brown or yellow streaks or dots; carpellate scales with an awn 0.2–6 mm long (sometimes awnless in C. glaucodea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Griseae 17b. Perigynia with elevated nerves, often with red-brown or yellow streaks or dots; carpellate scales sharply pointed to shortly awned, the awns up to 0.5 mm long when present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Granulares 16b. Perigynia tapering to base, apically with flat or somewhat convex sides 18a. Plants colonial from long rhizomes; abaxial leaf blade and surface of perigynia minutely papillose (view at 20× or higher magnification; both usually smooth in C. vaginata) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Paniceae 18b. Plants loosely to densely cespitose from short rhizomes; abaxial leaf blade and surface of perigynia without minute papillae 19a. Perigynia with 8–20 veins; beak of perigynium (when present) terminated by 2 teeth 0.2–1 mm long; uppermost spike gynecandrous or entirely staminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Hymenochlaenae

112  MO NOCOTS

19b. Perigynia with (22–) 25–40+ veins; beak of perigynium ± entire, without 2 apical teeth; uppermost spike entirely staminate 20a. Perigynia with sharp angles and flat faces, usually with 40 or more closely spaced veins; plants often with a long-peduncled, carpellate spike borne from near the base of the plant . . . . . . . . . . . . . . . . . . . . . . . . . Careyanae 20b. Perigynia with rounded angles and convex faces, mostly with 22–32 veins; plants usually without basal spikes . . . . . . . . . . . . . . (in part) Laxiflorae

Acrocystis Hybrids between a species that lacks basal spikes (e.g., Carex pensylvanica) and a species that possesses basal spikes (e.g., C. tonsa, C. umbellata) have been observed in New England. They are difficult to confidently identify and are suspected to be sterile. They typically have the basal spikes of one parent but otherwise resemble the other parent. Reference: Crins and Rettig (2002). 1a. All the carpellate spikes of a plant borne well above the base in proximity to a staminate spike on stems with leaf bearing nodes 2a. Perigynium body (i.e., excluding the beak and stipe-like base) subglobose, about as wide as long as thick 3a. Widest leaf blades (2.6–) 3–5 mm wide; plants cespitose, with short rhizomes; remnants of old leaves not or only slightly fibrous; staminate spikes 1–2.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. communis 3b. Leaf blades 0.5–3 (–3.6) mm wide; plants extensively colonial, with long rhizomes; remnants of old leaves slightly to strongly fibrous; staminate spikes (1.3–) 2–4.6 mm wide 4a. Perigynia with a beak 0.5–0.9 mm long and 0.1–0.5 times as long as the body of the perigynium; uppermost stem leaves with a well-developed blade; apical portion of the sheath opposite the leaf blade shallowly concave, the cleft extending 0.2–1.6 mm below the junction with the leaf blade . . . . . . . . . . . . . . . C. pensylvanica 4b. Perigynia with a beak 0.9–1.6 mm long and 0.5–1 times as long as the body of the perigynium [Fig. 67]; uppermost stem leaves with a poorly developed blade; apical portion of the sheath opposite the leaf blade deeply concave, the cleft extending 0.4–6.9 mm below the junction with the leaf blade . . . . . . . . . . . C. lucorum 2b. Perigynium body ellipsoid to obovoid, definitely longer than thick 5a. Lowest carpellate spike short-pedunculate, separated from the upper spikes by a gap of more than 7 mm; lowest bract of inflorescence usually equaling or exceeding total height of inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. novae‑angliae 5b. Lowest carpellate spike sessile, approximate to or overlapping the upper spikes, usually with a gap of less than 7 mm from the upper spikes; lowest bract of inflorescence usually shorter than total height of inflorescence (except in C. deflexa and C. reznicekii) 6a. Carpellate scales ½ to ⅔ as long as the perigynia they subtend, therefore, the perigynia clearly visible within the intact spikes 7a. Reproductive stems 5–31 cm tall, usually equaled or exceeded by the leaves; perigynia 2.3–3.1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. deflexa 7b. Reproductive stems 21–47 cm tall, usually exceeding the leaves; perigynia 3.2–4.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. peckii 6b. Carpellate scales equaling to slightly exceeding the perigynia, therefore, the perigynia largely concealed within the intact spikes

Cy p e r ac e a e   1 13

8a. Peduncle of staminate spike 0.4–9.9 mm long; remnants of old leaves slightly or not at all fibrous; reproductive stems (10–) 20–45 cm tall; achenes 1.2–1.4 (–1.7) mm long; perigynia 2.3–3.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. albicans 8b. Peduncle of staminate spike (0.2–) 0.3–1.2 mm long; remnants of old leaves slightly to strongly fibrous; reproductive stems 1.9–20 (–27) cm tall; achenes 1.4–2 mm long; perigynia (2.5–) 2.8–3.7 (–3.9) mm long 9a. Widest leaf blades (1.9–) 2.3–4.5 mm wide; carpellate scales sometimes with anthocyanic pigment extending from near the margin to the green or brown stripes on either side of the midvein; reproductive stems 8–38 cm tall; longest staminate spike exceeding the carpellate spikes by (0–) 0.5–5.1 (–7.3) mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nigromarginata 9b. Widest leaf blades 1.2–2.2 (–2.5) mm wide; carpellate scales never with anthocyanic pigment extending from near the margin to the green or brown stripes on either side of the midvein; reproductive stems 1.9–9.1 (–13.7) cm tall; longest staminate spike exceeding the carpellate spikes by 0–3.7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. reznicekii 1b. Some of the carpellate spikes solitary and borne on leafless peduncles originating from the plant base without associated staminate spikes (the solitary carpellate spikes sometimes hidden among the leaf bases), the other carpellate spikes borne on leaf-bearing stems with a staminate spike [Fig. 68] 10a. Carpellate scales ½ to ⅔ as long as the perigynia they subtend, therefore, the perigynia clearly visible within the intact spikes; remnants of old leaves slightly or not at all fibrous; lowermost bract of inflorescence usually equaling or exceeding total height of inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. deflexa 10b. Carpellate scales equaling to slightly exceeding the perigynia, therefore, the perigynia largely concealed within the intact spikes; remnants of old leaves weakly to strongly fibrous; lowermost bract of inflorescence usually shorter than total height of inflorescence 11a. Perigynium 3.1–4.7 mm long, with a beak 0.9–2 mm long; leaf blades coriaceous or herbaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tonsa 11b. Perigynium 2.2–3.2 mm long, with a beak 0.4–1 mm long [Fig. 69]; leaf blades herbaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. umbellata 1. Carex albicans Willd. ex Spreng. N white-tinged sedge. 1a. Carex artitecta Mackenzie; C. emmonsii Dewey ex Torr. var. muehlenbergii (Gray) J. Rettig; C. nigromarginata Schwein. var. muehlenbergii (Gray) Gleason; 1b. Carex emmonsii Dewey ex Torr. • CT, MA, ME, NH, RI, VT. Acidic to basic soils of forests, woodlands, rocky slopes, openings, and wetlands with a well-developed organic soil horizon. 1a. Midvein of median staminate scales usually not reaching apex, usually without minute scabrules along the sides; staminate spikes 8.4–11.1 mm tall; reproductive stems usually firm, erect or ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. C. albicans var. albicans 1b. Midvein of median staminate scales extending to apex of scale and often prolonged as a minute cusp, usually with some minute scabrules along the sides; staminate spikes 5–8.5 mm tall; reproductive stems weak, often spreading-ascending to reclining . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. Carex albicans var. emmonsii (Dewey ex Torr.) J. Rettig Variety albicans is known from CT, MA, ME, NH, RI, VT. Variety emmonsii is known from CT, MA, ME, NH, RI, VT. Though var. emmonsii is found in a wide variety of soil types, it sometimes occurs on more acidic sites and sometimes in peaty soils. 2. Carex communis Bailey var. communis N fibrous-rooted sedge. CT, MA, ME, NH, RI, VT. In shade or in openings within deciduous or mixed evergreen-deciduous forests, also on ledges and at bases of cliffs.

114  MO NOCOTS

3. Carex deflexa Hornem. var. deflexa N northern sedge. MA, ME, NH, VT; also reported from CT by Crins and Rettig (2002), but specimens are unknown. Fields, forest openings and edges, and on the interior of a variety of forest and woodland types. 4. Carex lucorum Willd. ex Link ssp. lucorum N Fig. 67 Blue Ridge sedge. Carex pensylvanica Lam. var. distans Peck • CT, MA, ME, NH, RI, VT. Dry, welldrained, often sandy soils of grasslands and oak-, pine-, and/or hickory-dominated woodlands and forests. 5. Carex nigromarginata Schwein. Fig. 67  Perigynium of Carex lucorum.

NC

black-edged sedge. CT; also reported from RI by Crins and Rettig (2002), but specimens are unknown. Rocky forests and woodlands, clearings and edges. Reports from MA are based on a collection of Carex tonsa var. tonsa—Poland 2135 (MASS!). 6. Carex novae-angliae Schwein. N New England sedge. CT, MA, ME, NH, RI, VT. Mesic deciduous forests and infrequently mixed evergreen-deciduous forests. 7. Carex peckii Howe N Peck’s sedge. Carex clivicola Fern. and Weatherby; Carex nigromarginata Schwein. var. elliptica (Boott) Gleason • MA, ME, NH, VT. Dry-mesic to mesic forests, slopes, and outcrops. 8. Carex pensylvanica Lam. N Pennsylvania sedge. Carex marginata Willd. • CT, MA, ME, NH, RI, VT. Dry, well-drained, often sandy, soils of grasslands and oak-, pine-, and/or hickory-dominated woodlands and forests. The report of Carex inops Bailey ssp. heliophila (Mackenzie) Crins by Angelo and Boufford (2007) is based on a collection of C. pensylvanica—18 May 1913, Murdoch, Jr., and Torrey s.n. (NEBC!). 9. Carex reznicekii Werier

NC

Reznicek’s sedge. CT, RI. Dry-mesic to mesic woodlands and forests, often on circumneutral, rocky or ledgy slopes. 10. Carex tonsa (Fern.) Bickn. N Fig. 68

Fig. 68  Solitary carpellate spikes (lower ones) and carpellate spikes of terminal inflorescences (upper ones) of Carex tonsa var. tonsa.

shaved sedge.  10a. Carex rugosperma Mackenzie; 10b. Carex rugosperma Mackenzie var. tonsa (Fern.) E.G. Voss; Carex umbellata Schkuhr ex Willd. var. tonsa Fern. • CT, MA, ME, NH, VT. Sandy grasslands and heathlands, xeric to mesic forests and woodlands, rocky balds. 1a. Leaf blades stiff, coriaceous, smooth or slightly scabrous adaxially, pale green; perigynia nearly glabrous, usually with a few hairs near the base of the beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10a. C. tonsa var. tonsa 1b. Leaf blades soft, herbaceous, scabrous or papillose adaxially, bright green; perigynia pubescent on the body as well as the beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10b. C. tonsa var. rugosperma (Mackenzie) W.J. Crins 11. Carex umbellata Schkuhr ex Willd. N Fig. 69 parasol sedge. Carex abdita Bickn. • ct, mA, ME, NH, RI, VT; Xeric to mesic fields, forests, and outcrops.

Albae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. eburnea Fig. 69  Perigynium of Carex umbellata.

12. Carex eburnea Boott N bristle-leaved sedge. CT, MA, ME, NH, VT. Open woodlands, cliffs, and outcrops, usually on circumneutral or basic rock.

Cy p e r ac e a e   1 15

Ammoglochin only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. siccata 13. Carex siccata Dewey

NC

dry land sedge. Carex foenea, sensu Fernald (1950) • CT, MA, ME, NH, RI, VT. Dry, well-drained, often coarse soils of fields, balds, and oak-pine woodlands. The name Carex foenea has been misapplied to this species in several regional works.

Anomalae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. scabrata 14. Carex scabrata Schwein. N Fig. 70 eastern rough sedge. CT, MA, ME, NH, RI, VT. Low, wet areas in forests, stream shores, and seeps.

Bicolores Reference: Ball (2002a). 1a. Perigynia becoming gold-orange just prior to being shed, smooth or minutely papillose; lateral spikes relatively sparsely flowered, the internodes of the mid-portion (0.5–) 0.7–1.5 mm long; uppermost spike usually entirely staminate; carpellate scales ovate-oblong, divergent in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. aurea

Fig. 70  Perigynium of Carex scabrata.

1b. Perigynia glaucous-white, densely papillose [Fig. 71]; lateral spikes densely flowered, the internodes of the mid-portion 0.2–0.7 mm long; uppermost spike, or at least many in the population, gynecandrous; carpellate scales ovate-orbicular, ascending . . . . . . . . . . . C. garberi 15. Carex aurea Nutt. N golden-fruited sedge. CT, MA, ME, NH, RI, VT. Cobble pavement and seepy outrcrop river shorelines, wet ledges, and borrow pits, in high-pH bedrock and/or till regions. 16. Carex garberi Fern.

N C Fig. 71

elk sedge. Carex garberi Fern. var. bifaria Fern. • ME, NH, VT. River shores in high-pH bedrock and/or till regions, usually on seepy outcrops or cobble pavement.

Capituligerae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. arctogena 17. Carex arctogena H. Sm.

NC

capitate sedge. Carex capitata L. ssp. arctogena (H. Sm.) Hiitonen; C. capitata L. var. arctogena (H. Sm.) Hultén • NH. Alpine slopes and plateaus. See Reinhammer (2001) for justification of using the epithet arctogena at the rank of species.

Carex 1a. Perigynium glabrous; leaf blades minutely papillose abaxially; ligules (6–) 11–45 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. atherodes 1b. Perigynium pubescent [Fig. 72]; leaf blades not minutely papillose abaxially; ligules 2–12.5 mm long 2a. Leaf sheaths, and often the blades, pubescent; staminate scales, and often the carpellate scales, pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hirta 2b. Leaf sheaths and blades glabrous; staminate and carpellate scales glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. trichocarpa 18. Carex atherodes Spreng.

nC

wheat sedge. ME, NH, VT. Marshes and low, open rights-of-way in high-pH bedrock regions. This species is native in ME and VT, non-native in NH.

Fig. 71  Papillose perigynium of Carex garberi.

1 16 MO NOCOTS

19. Carex hirta L. E hammer sedge. CT, MA, ME, VT. Fields, ditches, railroad embankments, roadsides, and other human-disturbed sites on a variety of soil moisture regimes. 20. Carex trichocarpa Muhl. ex Willd. N C Fig. 72 hairy-fruited sedge. CT, MA, NH, VT. Wet meadows, ditches, lake shores, riverside marshes and fields, usually in high-pH bedrock regions.

Careyanae Reference: Bryson and Naczi (2002a). 1a. Sheaths anthocyanic at the base, those borne from the lower half of the flowering stem with blades up to 1.9 cm long [Fig. 74] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. plantaginea Fig. 72  Perigynium of Carex trichocarpa.

1b. Sheaths white to brown at the base, those borne from the lower half of the flowering stem producing blades 2–24 cm long 2a. Foliage strongly glaucous; leaves dimorphic—those of the vegetative shoots 10–25 mm wide, those of the flowering stems 2–6 mm wide, the former 3.8–9 times as wide as the latter; leaves from the lower half of the flowering stem with blades 2–6.2 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. platyphylla 2b. Foliage green or glaucous; leaves slightly dimorphic if at all, 1–12 mm wide, those of the vegetative shoots 1–3.5 times as wide as those of the flowering stem; leaves from the lower half of the flowering stem with blades 4.5–24 cm long 3a. Staminate spikes 0.6–1.4 (–1.6) mm wide; staminate scales obtuse at the apex, the middle ones 2.6–3.6 (–3.8) mm long; vegetative shoots much taller than the flowering stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. abscondita 3b. Staminate spikes (1–) 1.2–2.7 mm wide; staminate scales acute at the apex, the middle ones 3.6–5.5 mm long; vegetative shoots shorter than to slightly taller than the flowering stems 4a. Lower 1–3 scales of the carpellate spikes empty or subtending staminate flowers only [Fig. 73]; leaves of the vegetative shoots 3–11.8 mm wide, glaucous; lower spikes 4–6 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. laxiculmis

Fig. 73  Lateral spike of Carex laxiculmis with staminate flowers at lower 2 nodes.

4b. All scales of the carpellate spikes subtending carpellate flowers; leaves of the vegetative shoots 1–5.3 mm wide, green; lower spikes 3–4 mm thick . . . . . C. digitalis 21. Carex abscondita Mackenzie N thicket sedge. Carex abscondita Mackenzie var. glauca (Chapman) Fern. • CT, MA, RI; also reported from NH by Magee and Ahles (1999), but specimens are unknown. Rich, mesic forests, riparian forests. 22. Carex digitalis Willd. var. digitalis N slender woodland sedge. CT, MA, ME, NH, RI, VT. Mesic to dry-mesic, deciduous and mixed evergreen-deciduous forests. 23. Carex laxiculmis Schwein. var. laxiculmis N Fig. 73 spreading sedge. CT, MA, ME, NH, RI, VT. Deciduous and mixed evergreen-deciduous forests, often adjacent to streams or seepage areas, sometimes in rich, mesic types. Carex laxiculmis var. copulata (Bailey) Fern. was reported from MA by Sorrie and Somers (1999), but specimens are unknown. 24. Carex plantaginea Lam. N Fig. 74 plantain-leaved sedge. CT, MA, ME, NH, VT. Rich, mesic forests and deciduous seepage forests.

Fig. 74  Leaf subtending a carpellate spike of Carex plantaginea showing anthocyanic color and short blade.

25. Carex platyphylla Carey N broad-leaved sedge. CT, MA, ME, NH, RI, VT. Mesic forests and slopes, usually those underlain by high-pH bedrock.

Cy p e r ac e a e   117

Ceratocystis Carex hostiana DC. was reported to occur in Tewksbury, MA (Crins 2002a). However, the extreme disjunction of the plants and occurrence in atypical habitat suggest that the collection is mislabeled. See Fernald (1911) for discussion of this record. Reference: Crins (2002a). 1a. Perigynia 1.8–3.9 (–4.2) mm long, the lower of each spike spreading; perigynium beak straight or bent at an angle of 0–15 (–25) degrees from the perigynium body; ligule of apical leaves obsolete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. viridula 1b. Perigynia 3.5–6.3 mm long, the lower of each spike reflexed; perigynium beak bent at an angle of 13–72 degrees from the perigynium body [Fig. 75]; ligule of apical leaves well formed 2a. Carpellate scales red-brown, contrasting with color of perigynia; perigynia 4–6.3 mm long, with a moderately to strongly bent, scabrous beak offset (15–) 26–72 degrees from the perigynium body [Fig. 75] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. flava 2b. Carpellate scales yellow-green, of similar color as the perigynia; perigynia 3.5–4.8 mm long, with a weakly to moderately bent, smooth beak offset 13–48 degrees from the perigynium body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cryptolepis 26. Carex cryptolepis Mackenzie N northeastern sedge. Carex flava L. var. fertilis, auct. non Peck. • CT, MA, ME, NH, RI, VT. Shorelines and other open, wet areas, usually on sandy or peaty substrate; calcifuge. 27. Carex flava L. N Fig. 75 yellow-green sedge. Carex flava L. var. fertilis Peck; C. flava L. var. gaspensis Fern.; C. laxior (Kükenth.) Mackenzie • CT, MA, ME, NH, RI, VT. Wet meadows, shorelines, and seepy ledges; calciphile. ‌ × 28. Carex ×ruedtii Kneuck. is a rare sedge hybrid in New England known 27 from MA, ME, NH. It is recognized by its perigynia that are mostly 3.1–4 mm long with a smooth or sparsely serrulate, essentially straight beak, carpellate scales that are brown and contrast with the green perigynia, and upper leaves with ligules (lacking ligules in C. viridula). 28. Carex viridula Michx. n little green sedge.  28a. Carex oederi Ehrh.; C. oederi Retz. var. pumila (Cosson & Germain) Fern.; C. oederi Retz. ssp. viridula (Michx.) Hultén; C. oederi Retz. var. viridula (Michx.) Kükenth.; 28b. Carex demissa Hornem. • CT, MA, ME, NH, RI, VT. Open, mesic to hydric, mineral or organic soils, such as river shores, lake shores, and Atlantic coast shores and islands. 1a. Carpellate spikes contiguous or the lowest sometimes distant; foliage and perigynia gray-green to light green; stems erect . . . . . . . . . . . 28a. C. viridula ssp. viridula var. viridula 1b. Lowest carpellate spike remote; foliage and perigynia brown-green to dark green; stems often arcuate or sinuous . . . . . . . . . . . . . 28b. C. viridula ssp. oedocarpa (Anderss.) B. Schmid Subspecies viridula is native and known from CT, MA, ME, NH, RI, VT. Subspecies oedocarpa is non-native and known from CT (southern portion of state).

Chordorrhizae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. chordorrhiza 29. Carex chordorrhiza Ehrh. ex L. f. N rope-root sedge. MA, ME, NH, VT. Fens, lakeside fens, and graminoid marshes with a welldeveloped organic soil horizon, sometimes growing in shallow standing water.

Fig. 75  Perigynium and bent beak of Carex flava.

1 18 MO NOCOTS

Chlorostachyae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. capillaris 30. Carex capillaris L.

NC

hair-like sedge. 30a. Carex fuscidula V. Krecz. ex Egorova; 30b. Carex capillaris L. var. elongata Olney ex Fern.; C. capillaris L. ssp. robustior (Drej. ex Lange) Böcher • ME, NH, VT. Wet cliffs, river shore outcrops, seeps, and wet-mesic alpine lawns. 1a. Staminate spike partly enclosed in sheath of lowest bract of inflorescence, situated at or below the level of the uppermost carpellate spike, borne on a smooth peduncle 1.5–7 mm long; carpellate scales red-brown to orange-brown (the margins and sometimes also the midrib pale), usually acute at the apex; beak of perigynium usually smooth, relatively abruptly tapering from the body, 0.6–1.2 mm long; stem leaf blades 0.5–1.4 (–2) mm wide; lowermost bract of inflorescence with a sheath 1.7–7 (–10) mm long . . . . . . . . . . . . . . . . . . . . . . . . 30a. C. capillaris ssp. fuscidula (V. Krecz. ex Egorova) A. & D. Löve 1b. Staminate spike exserted from sheath of lowest bract of inflorescence, situated at or above the level of the uppermost carpellate spike, borne on a scabrous peduncle (3–) 7–40 mm long; carpellate scales white-brown to pale brown, usually obtuse to nearly truncate at the apex; beak of perigynium scabrous, gradually tapering from the body, 0.3–0.6 mm long; stem leaf blades 1–2.1 (–2.5) mm wide; lowermost bract of inflorescence with a sheath 4–30 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30b. C. capillaris ssp. capillaris Subspecies fuscidula is known from NH and occurs in alpine habitats. Subspecies capillaris is known from ME, VT and occurs on high-pH bedrock at lower elevations (e.g., boreal cliffs, river shore outcrops, and seeps). Both subspecies are of regional conservation concern.

Clandestinae Fig. 76  Perigynium with basal elaiosome of Carex pedunculata.

1a. Perigynia 3.7–6 mm long; carpellate scales retuse to obtuse, cuspidate; leaves dark green, mostly equaling or exceeding the reproductive stems; most carpellate spikes borne on long, often arching, peduncles to 13 cm emerging from basal nodes . . . . . . . . . . . . . . C. pedunculata 1b. Perigynia 2.4–2.9 mm long; carpellate scales acute to obtuse, without a cusp; leaves pale green, mostly shorter than the reproductive stems; carpellate spikes all borne above the base on peduncles to 7 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. richardsonii 31. Carex pedunculata Muhl. ex Willd. ssp. pedunculata N Fig. 76 long-stalked sedge. CT, MA, ME, NH, RI, VT. Mesic to dry-mesic forests, both upland and riparian. 32. Carex richardsonii R. Br.

NC

Richardson’s sedge. VT. Woodlands in high-pH bedrock regions.

Collinsiae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. collinsii 33. Carex collinsii Nutt.

N C Fig. 77

Collins’ sedge. CT, RI. Forested wetlands, frequently on Sphagnum in the shade of Chamaecyparis thyoides.

Fig. 77  Perigynium of Carex collinsii.

Cyperoideae Carex section Cyperoideae (which includes sedges formerly placed in section Ovales) comprises a very difficult section of sedges given the narrow morphological gaps between some taxa. Flowering individuals from later in the season often have short, contracted inflorescences, regardless of their early season form. Such individuals are not accounted for in the key. Mature perigynia are a requisite for keying, given the importance of perigynium and achene dimensions. Perigynia from about ⅓ the spike length (measuring from the base) are best used for the following key. Reference: Mastrogiuseppe et al. (2002).

Cy p e r ac e a e   1 19

1a. Longest bract of inflorescence 5–20 cm long, (3–) 5 or more times as long as the inflorescence; distance froma apex of achene to apex of perigynium 3–5 mm [Fig. 86] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sychnocephala 1b. Longest bract of inflorescence rarely exceeding 5 cm, up to 2 times as long as inflorescence; distance from apex of achene to apex of perigynium 1.2–3.1 (–3.7) mm [Figs. 79, 83, 84] 2a. Carpellate scales nearly equaling to slightly exceeding the length of the perigynia, mostly concealing the beaks of the perigynia in intact spikes [Fig. 78] 3a. Leaves stiff, very glaucous (except in shade), many with rounded auricles at the summit of the sheath; floral scales white or white-brown; plants of Atlantic coast shores and near shore habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(in part) C. silicea 3b. Leaves softer, slightly or not at all glaucous, without auricles; floral scales whitegreen or brown; plants usually not coastal 4a. Carpellate scales about as wide as the perigynia they subtend; achenes 1.6–2 mm wide; lowest bract usually broad and flat at base, sometimes exceeding the spike it subtends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. adusta 4b. Carpellate scales narrower than the perigynia they subtend; achenes 0.7–1.6 mm wide; lowest bract usually setaceous and shorter than the spike it subtends 5a. Beak of perigynium flat, winged, and minutely serrulate to the apex [Fig. 83] 6a. Perigynium body lanceolate, 0.9–1.4 mm wide, with the marginal wing usually conspicuously reduced (or even absent) on the lower half of the body [Fig. 83] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. oronensis 6b. Perigynium body ovate, (1.5–) 1.7–2.5 mm wide, with the marginal wing scarcely and gradually (if at all) reduced on the lower half of the body [Fig. 84] 7a. Perigynia smooth, adaxially without veins or with 4–8 curving veins of unequal length; inflorescence with 3–7 (–11) spikes, the upper ones usually separate; carpellate scales red-brown to green or gold-brown . . . . . C. foenea 7b. Perigynia granular-papillose, adaxially with (4–) 5–8 (–10) ± parallel veins of equal length; inflorescence with (3–) 7–15 spikes, the upper ones usually densely aggregated; carpellate scales pale white-green . . . . . C. argyrantha 5b. Beak of perigynium cylindric, unwinged, and entire in the apical (0.3–) 0.4–1 mm [Fig. 84] 8a. Perigynia 3.4–4.7 (–5.2) mm long; inflorescence usually erect, with approximate spikes, the internode between the second and third spike counting from the base (2.5–) 3–6 mm long; apical, herbaceous portion of leaf sheath smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ovalis 8b. Perigynia (3.7–) 4.5–6 mm long; inflorescence usually open and ± nodding, the internode between the second and third spike (2.5–) 4–10 mm long; apical, herbaceous portion of leaf sheath minutely papillose . . . . . . . . . . . . C. praticola 2b. Carpellate scales, excluding the awns (when present), clearly exceeded by the perigynia, the tip of the perigynia beaks visible in intact spikes [Fig. 85] 9a. Perigynia up to 2 mm wide 10a. Carpellate scales of the basal median portion of the spike tapering to a slender, acuminate point, often prolonged into a short awn tip 11a. Perigynia bodies lanceolate, 0.9–1.3 mm wide; achene 0.6–0.8 mm wide; inflorescence congested, the lowest internode 2–3 (–5) mm long . . . . C. crawfordii 11b. Perigynia bodies lanceolate to ovate, obovate, or nearly orbicular, 1.2–2.9 mm wide; achene 0.7–1.1 mm wide; inflorescence congested to open, the lowest internode 2–18 mm long

120  MO NOCOTS

12a. Perigynia 1.2–2.1 mm wide, 2.6 or more times as long as wide; leaf sheaths opposite the blade with a prolonged, narrow hyaline area . . . . . . . . C. scoparia 12b. Perigynia 1.8–2.8 mm wide, up to 2.5 times as long as wide; leaf sheaths opposite the blade firm and green-veined nearly to the summit, the hyaline area very short 13a. Staminate portion of lateral spikes 2–6 mm long; beak of perigynium spreading, usually 50% or more of the length of the nearly orbicular body; plants of freshwater wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. straminea 13b. Staminate portion of lateral spikes shorter than 2 mm; beak of perigynium ascending, less than 50% of the length of the narrow-ovate to obovate body; plants of saline-influenced habitats . . . . . . . C. hormathodes 10b. Carpellate scales of the basal median portion of the spike obtuse to subacuminate at the apex, without awn tips 14a. Marginal wing of perigynium usually conspicuously reduced (or even absent) in the lower half of the body; leaves with blades 3–7.5 mm wide, with narrow, decurrent wings on the sheath that are continuous with the blade midrib and margins; vegetative shoots conspicuous, with numerous leaves spaced throughout the apical half of the stem 15a. Perigynia in basal portion of spike spreading or recurved; spikes spherical or subspherical; carpellate scales 1.6–2.3 mm long . . . . . . . . . . . . . . C. cristatella 15b. Perigynia in basal portion of spike appressed-ascending to ascending-spreading; spikes subspherical to ovoid or obovoid; carpellate scales 2–3 mm long 16a. Inflorescence erect, with congested and overlapping spikes, each spike with (30–) 40 or more perigynia; perigynia appressed-ascending to ascending; apical portion of the leaf sheath opposite the blade firm and green-veined nearly to the summit, the hyaline area 3–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tribuloides 16b. Inflorescence arching or nodding, with separate spikes in the basal portion (uncommonly the inflorescence ± congested), each spike with 15–30 perigynia; perigynia ascending to spreading; apical portion of the leaf sheath opposite the blade with a prolonged, hyaline area 4–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. projecta 14b. Marginal wing of perigynium scarcely and gradually (if at all) reduced in the lower half of the body; leaves with blades 1–4.5 mm wide (up to 6.5 mm in C. normalis), with inconspicuous, rounded edges on the sheath that are continuous with the blade midrib and margins; vegetative shoots inconspicuous, with few leaves clustered near apex of stem 17a. Body of the perigynium elliptic to orbicular or obovate, broadest at or above the middle [Fig. 79] 18a. At least some spikes with a pronounced, narrow, staminate base longer than 2 mm; inflorescence arching to nodding; achenes ovate in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. festucacea 18b. Spikes with an inconspicuous staminate base shorter than 2 mm; inflorescence ± erect; achenes narrow-oblong to oblong or elliptic in outline 19a. Perigynium body elliptic to suborbicular, broadest near the middle; achenes 0.9–1.3 mm wide; inflorescence congested, the lowest internode 1.5–6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. molesta 19b. Perigynium body obovate [Fig. 79]; achenes 0.7–1 mm wide; inflorescence close, but usually not congested near the base, the lowest internode (1–) 3–14 mm

Cy p e r ac e a e   1 2 1

20a. Carpellate scales obtuse; perigynium beak triangular [Fig. 81], appressed-ascending; styles straight; apical, herbaceous portion of leaf sheath minutely papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. longii 20b. Carpellate scales acute; perigynium beak slender, spreading [Fig. 79]; styles laterally sinuous; apical, herbaceous portion of leaf sheath smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. albolutescens 17b. Body of the perigynium broad-lanceolate to ovate, widest below the middle 21a. Inflorescence congested, with crowded spikes, the lowest internode usually shorter than 6 mm 22a. Perigynia adaxially with 1–3 faint veins near the base; achenes 0.6–0.9 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bebbii 22b. Perigynia adaxially usually with 3–7 conspicuous veins; achenes 0.9–1.2 mm wide 23a. Leaves with blades 1.5–3.5 mm wide, with lower sheaths that are tight, ± evenly colored, and sometimes cross-corrugated; apical, herbaceous portion of leaf sheath minutely papillose; mature perigynia tan to brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. tincta 23b. Leaves with blades 2.2–6 (–6.5) mm wide, with lower sheaths that are loose, green and white-mottled or -striped, and not corrugated; apical, herbaceous portion of leaf sheath smooth; perigynia green (at least apically) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. normalis 21b. Inflorescence elongate, the lower spikes remote, the lowest internode usually longer than 6 mm 24a. Leaves with blades 2.2–6 (–6.5) mm wide, the lower sheaths loose; flowering stems with 3–7 leaves; apical, herbaceous portion of the leaf sheath smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. normalis 24b. Leaves with blades 1.3–2.5 (–3) mm wide, the lower sheaths tight; flowering stems with 2–4 leaves; apical, herbaceous portion of at least some leaf sheaths minutely papillose (smooth in the very rare C. echinodes) 25a. At least some leaf sheaths minutely papillose near the apex on the herbaceous portion, not prominently mottled or striped with white near the base; beaks of perigynia of intact spikes appressed to spreading-ascending, exceeding the subtending scales by 0–0.8 mm; beaks and shoulders of perigynia stramineous to red-brown at maturity; carpellate scales mostly 0.8–1 times as long as the perigynia they subtend . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tenera 25b. Leaf sheaths smooth near the apex, often mottled or striped with white near the base; beaks of perigynia of intact spikes spreading, exceeding the subtending scales by 0.7–1.6 mm; beaks and shoulders of perigynia green to green-brown at maturity; carpellate scales mostly 0.6–0.85 times as long as the perigynia they subtend . . . C. echinodes 9b. Perigynia wider than 2 mm 26a. Leaves stiff, very glaucous (except in shade), many with rounded auricles at the summit of the sheath; floral scales white or white-brown; plants of Atlantic coast shores and near shore habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. silicea 26b. Leaves softer, not or only slightly glaucous, without auricles; scales light green to green or white-brown to brown; plants not of coastal shores (except C. hormathodes)

122  MO NOCOTS

27a. Carpellate scales of the basal median portion of the spike tapering to a slender, acuminate point, often prolonged into a short awn tip 28a. Perigynia 2.5–3.8 mm wide, broad-obovate; leaf blades 2.5–6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. alata 28b. Perigynia 1.9–2.8 mm wide, narrow-ovate to orbicular or obovate; leaf blades 1–3 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . go to couplet 12 27b. Carpellate scales of the basal median portion of the spike obtuse to subacuminate at the apex, without awn tips 29a. Perigynia obovate, widest above the middle; apical portion of the leaf sheath opposite the blade firm and green-veined nearly to the summit 30a. Leaves with blades 3–6 (–8) mm wide, the summit of the sheaths truncate (unless torn); perigynia adaxially with inconspicuous veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cumulata 30b. Leaves with blades 2–4.5 mm wide, the summit of the sheaths concave; perigynia adaxially with 4–many veins . . . . . . . . . . . . . . . . . . . go to couplet 19 29b. Perigynia broad-lanceolate, ovate, elliptic, or orbicular, widest at or below the middle; apical portion of the leaf sheath opposite the blade with a prolonged, hyaline area (except in C. molesta and C. normalis) 31a. Perigynia broad-lanceolate to ovate, green (at least apically); leaves with blades 2.2–6 (–6.5) mm wide, the lower sheaths green and whitemottled or -striped; sheath summits truncate, prolonged up to 2 mm above sheath apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. normalis 31b. Perigynia ovate to broad-ovate or elliptic to orbicular, yellow-brown to tan or brown; leaves with blades 1.5–4 (–5) mm wide, the lower sheaths ± evenly colored; sheath summits usually concave (sometimes truncate and prolonged) 32a. Perigynia bodies broad-elliptic to orbicular, broadest near the middle 33a. At least some spikes with a pronounced, narrow, staminate base longer than 2 mm; inflorescence arching to nodding, the lowest internode 3–18 mm long; achenes ovate in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. festucacea 33b. Spikes with an inconspicuous staminate base shorter than 2 mm; inflorescence ± erect, the lowest internode 1.5–6 mm; achenes narrowoblong to elliptic in outline . . . . . . . . . . . . . . . . . . . . . (in part) C. molesta 32b. Perigynia bodies ovate to broad-ovate 34a. Perigynia (1.4–) 1.6–2.4 mm wide; achenes 0.9–1.2 mm wide; hyaline region of leaf sheath opposite leaf blade sometimes crosscorrugated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. tincta 34b. Perigynia (2–) 2.3–4.2 mm wide; achenes 1.1–1.8 mm wide; hyaline region of leaf sheath not corrugated 35a. Marginal wing of perigynium evenly and minutely serrulate [Fig. 80]; apical, herbaceous portion of leaf sheaths smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. brevior 35b. Marginal wing of perigynium often irregularly erose or toothed, in addition to minutely serrulate [Fig. 82]; apical, herbaceous portion of leaf sheaths minutely papillose 36a. Perigynia 3.3–5 mm long, adaxially veinless or with 1–5 faint veins [Fig. 82], exceeding the carpellate scales by 0.2–1.3 mm; anthers 1.3–2.6 mm long . . . . . . . . . . . . . . . . . . C. merritt‑fernaldii

Cy p e r ac e a e   1 23

36b. Perigynia (4.5–) 5.1–6.7 (–7.1) mm long, adaxially with 4–8 veins, exceeding the carpellate scales by (1–) 1.4–2.3 mm; anthers (2.4–) 2.8–4.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bicknellii 34. Carex adusta Boott

N C Fig. 78

swarthy sedge. ME, NH. Sandy, often well-drained, soils of woodlands, road shoulders, old road beds, and borrow pits. 35. Carex alata Torr. N broad-winged sedge. CT, MA, NH, RI; primarily along the greater coastal plain. Swamps and wetland margins. 36. Carex albolutescens Schwein. N Fig. 79

Fig. 78  Spike of Carex adusta showing carpellate scales that conceal the associated perigynia.

green-white sedge. CT, MA, RI, VT. Acidic swamps and wetland margins. This species is sometimes confused with Carex festucacea. In addition to characters used in the identification key, these species differ in achene width, style shape, and perigynium shape. Carex albolutescens has achenes 0.75–1 mm wide, styles that are laterally bent near the base (i.e., the bend is toward the margins of the achene), and obovate perigynia bodies. Carex festucacea has achenes 1–1.3 mm wide, styles that are straight or adaxially-abaxially bent (i.e., the bend is toward the broad surfaces of the achene), and orbicular to elliptic perigynia bodies. 37. Carex argyrantha Tuckerman N silvery-flowered sedge. CT, MA, ME, NH, RI, VT. Dry-mesic to xeric, often disturbed, sandy soils of fields, woodlands, and roadsides. 38. Carex bebbii Olney ex Fern. N Bebb’s sedge. Carex tribuloides Wahlenb. var. bebbii Bailey • CT, MA, ME, NH, VT. Mesic to hydric, usually circumneutral, soils of swamps, meadows, shorelines, and seeps. 39. Carex bicknellii Britt.

NC

Fig. 79  Adaxial surface of perigynium of Carex albolutescens.

Bicknell’s sedge. Carex straminea Willd. ex Schkuhr var. crawei Boott • CT, MA, ME, RI, VT; also reported from NH by Kartesz (1999), but specimens are unknown. Xeric to hydric, often sandy, soils of woodlands, fields, open banks and slopes, lake shores, river banks, and swamps. 40. Carex brevior (Dewey) Mackenzie ex Lunell N Fig. 80 short-beaked sedge. Carex straminea Willd. ex Schkuhr var. brevior Dewey • CT, MA, ME, NH, RI, VT. Fields, roadsides, woodlands, and other places in ± open, dry soils. 41. Carex crawfordii Fern. N Crawford’s sedge. CT, MA, ME, NH, VT. Open, xeric to hydric soils, often in human-disturbed areas such as fields and roadsides. 42. Carex cristatella Britt. N crested sedge. CT, MA, NH, RI, VT. Mesic to hydric soils of meadows, marshes, open swamps, and stream banks. 43. Carex cumulata (Bailey) Mackenzie N clustered sedge. Carex straminea Willd. ex Schkuhr var. cumulata Bailey • CT, MA, ME, NH, RI, VT. Sandy, usually well-drained, soils of fields, woodlands, cliff bases, and eroding slopes, sometimes also along railroads. 44. Carex echinodes (Fern.) Rothrock, Reznicek, & Hipp

NC

prickly quill sedge. Carex tenera Dewey var. echinodes (Fern.) Wieg.; C. straminea Willd. ex Schkuhr var. echinodes Fern. • VT. Swamps, lacustrine forests. 45. Carex festucacea Schkuhr ex Willd. N fescue sedge. CT, MA, ME, NH, RI, VT. Seasonally or permanently saturated soils of fields, swamps, and wetland edges. 46. Carex foenea Willd. N straw sedge. Carex aenea Fern. • CT, MA, ME, NH, RI, VT. Woodlands, cliffs, sandy fields, and open, disturbed soil.

Fig. 80  Adaxial surface of perigynium of Carex brevior.

124  MO NOCOTS

47. Carex hormathodes Fern. N marsh straw sedge. Carex straminea Willd. ex Schkuhr var. invisa Boott • CT, MA, ME, NH, RI. Rocky or sandy beaches of the Atlantic coast and bays, and saline to fresh-tidal marshes. 48. Carex longii Mackenzie N Fig. 81 Long’s sedge. CT, MA, ME, RI, VT. Saturated or seasonally saturated soils of fields, pond shores, ditches, and peatland margins. 49. Carex merritt-fernaldii Mackenzie N Fig. 82

Fig. 81  Adaxial surface of perigynium of Carex longii.

Merritt Fernald’s sedge. Carex brevior (Dewey) Mackenzie ex Lunell var. pseudofestucacea Farw. • CT, MA, ME, NH, RI, VT. Well-drained sandy and rocky soils of fields, roadsides, cliffs, and woodlands. 50. Carex molesta Mackenzie ex Bright N troublesome sedge. CT, MA, NH, VT. Fields, roadsides, meadows, usually in relatively hydric soils. 51. Carex normalis Mackenzie N greater straw sedge. Carex mirabilis Dewey • CT, MA, ME, NH, RI, VT. Open, usually ± hydric soils of fields, roadsides, ditches, and meadows. 52. Carex oronensis Fern.

N C Fig. 83

Orono sedge. ME; endemic to the state, mainly in the Penobscot River drainage. Wet-mesic to xeric open places such as fields, roadsides, and abandoned paths. 53. Carex ovalis Goodenough E Fig. 84 oval sedge. Carex leporina, auct. non L., C. traceyi Mackenzie • MA, ME, NH. Mesic to hydric soils of open areas such as fields, meadows, and ditches. Fig. 82  Adaxial surface of perigynium of Carex merrittfernaldii.

54. Carex praticola Rydb.

NC

northern meadow sedge. Carex platylepis Mackenzie; C. pratensis Drej. • ME; northern portion of state. Fields, forest openings, meadows. 55. Carex projecta Mackenzie N necklace sedge. CT, MA, ME, NH, RI, VT. Swamps, riparian forests, wetland borders, marshes, forest seeps, shorelines. 56. Carex scoparia Schkuhr ex Willd. N Fig. 85 pointed broom sedge. CT, MA, ME, NH, RI, VT. Xeric to hydric open soils such as fields, roadsides, ditches, and wetland edges.

Fig. 83  Abaxial surface of perigynium of Carex oronensis.

1a. Carpellate scales and apex of perigynium stramineous to gold-brown, of relatively similar color as body of perigynium and not contrasting with it; perigynia 4.2–6.8 mm long, (2.5–) 2.8– 4 times as long as wide; inflorescence with spikes aggregated to separate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56a. C. scoparia var. scoparia 1b. Carpellate scales and apex of perigynium red-brown, dark brown, or nearly black, sharply contrasting with color of perigynium body; perigynia 4.2–5 mm long, 2–2.6 times as long as wide; inflorescence with aggregated spikes . . . . 56b. C. scoparia var. tessellata Fern. & Wieg. Variety scoparia is known from CT, MA, ME, NH, RI, VT. Variety tessellata is known only from eastern ME (within New England). It has narrower leaves and thinner stems than sympatric plants of C. scoparia var. scoparia and should probably be treated as a distinct species. 57. Carex silicea Olney N sea-beach sedge. CT, MA, ME, NH, RI. Sandy or rocky coastal beaches, dunes, and salted road shoulders. 58. Carex straminea Willd. ex Schkuhr N

Fig. 84  Abaxial surface of perigynium of Carex ovalis.

eastern straw sedge. Carex richii (Fern.) Mackenzie • CT, MA, RI. Wet fields, meadows, graminoid marshes, open swamps.

Cy p e r ac e a e   125

59. Carex sychnocephala Carey

N C Fig. 86

many-headed sedge. ME. Open, hydric or seasonally hydric soils, often disturbed sandy or mucky areas and along shorelines. 60. Carex tenera Dewey

NC

delicate quill sedge. CT, MA, ME, NH, RI, VT. Dry-mesic to wet-mesic fields, meadows, and forest openings. Carex tenera sometimes lacks minute papillae on some leaf sheaths when the plants grow in deep shade or under hydric conditions. However, careful searching will usually reveal minute papillae on some leaf sheaths on the plant. 61. Carex tincta (Fern.) Fern.

N

tinged sedge. Carex mirabilis Dewey var. tincta Fern. • MA, ME, NH, VT; also reported from CT by Dowhan (1979), but specimens are unknown. Dry-mesic to wet-mesic fields, meadows, and forest openings.

Fig. 85  Spike of Carex scoparia var. scoparia showing carpellate scales that are shorter than associated perigynia.

62. Carex tribuloides Wahlenb. var. tribuloides N blunt broom sedge. CT, MA, ME, NH, RI, VT. Mesic to hydric soils of fields, marsh edges, ditches, river banks.

Deweyanae 1a. Perigynia 0.8–1.3 mm wide, (3.5–) 4–6.1 (–6.6) times as long as wide, with 4–8 usually prominent veins adaxially; leaf blades 1.3–3.1 mm wide . . . . . . . . . . . . . . . . . . . . . . . C. bromoides 1b. Perigynia 1.3–1.5 (–1.6) mm wide, 2.9–3.8 times as long as wide, veinless or less commonly with 1–5 obscure veins adaxially; widest leaf blades 2.4–4.2 mm wide . . . . . . . . . C. deweyana 63. Carex bromoides Schkuhr ex Willd. ssp. bromoides N brome-like sedge. CT, MA, ME, NH, RI, VT. Riparian forests, deciduous swamps, seepage forests, and stream shores. 64. Carex deweyana Schwein. var. deweyana N round-fruited short-scaled sedge. CT, MA, ME, NH, VT. Deciduous and mixed evergreendeciduous forests and forest edges. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species could be in RI (i.e., the author is unaware of any collections).

Dispermae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. disperma 65. Carex disperma Dewey N soft-leaved sedge. CT, MA, ME, NH, VT. Shaded, forested wetlands, usually growing among bryophytes.

Divisae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. praegracilis 66. Carex praegracilis W. Boott E freeway sedge. Carex camporum Mackenzie • ME, VT. Roadsides, median strips, and ditches, often along stretches that receive heavy salting in winter.

Glareosae Reference: Toivonen (2002). 1a. Perigynia essentially beakless; inflorescence 6–12 mm tall, consisting of 2–4 aggregated and overlapping spikes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tenuiflora

Fig. 86  Perigynium of Carex sychnocephala.

126  MO NOCOTS

1b. Perigynia definitely, although sometimes shortly, beaked; inflorescence mostly 15–100 mm tall, consisting of (1–) 2–15 aggregated to long-remote spikes 2a. Spikes (1–) 2–4 per stem, the lowest widely separated from the upper spikes (when present) and subtended by a prolonged, setaceous bract (2–) 3–10 cm long; spikes composed of 1–5 carpellate flowers; herbaceous portion of leaf sheath white spotted 3a. Leaf blades (0.8–) 1–1.9 mm wide, flat to partially folded; ligules 0.5–1.9 mm long; inflorescence (14–) 23–55 mm long, with (2–) 3 or 4 spikes, the terminal spike with (1–) 2–5 perigynia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. trisperma 3b. Leaf blades 0.3–0.5 (–0.8) mm wide, involute; ligules 0.3–0.8 (–1.2) mm long; inflorescence 14–32 mm long, with 2 or 3 spikes, the terminal spike with 1–3 perigynia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. billingsii 2b. Spikes 3–15, the lowest approximate to or shortly separated from the upper spikes, subtended by a short, setaceous bract mostly shorter than 3 cm; spikes composed of 5–30 carpellate flowers; herbaceous portion of leaf sheath without white spots 4a. Perigynia ovate in outline, widest near base, with a serrulate-margined beak; inflorescence composed of 5–15 spikes that are aggregated together [Fig. 87] . . . . C. arcta 4b. Perigynia elliptic-ovate to elliptic-oblong in outline, widest near the middle, with an entire- or serrulate-margined beak; inflorescence composed of 3–8 (–10) spikes, at the least the lower usually separate [Fig. 88] 5a. Carpellate scales light red-brown, equaling to shortly exceeding and covering the perigynia; stems smooth on the angles; uppermost spike with a prolonged, basal, staminate portion comprising 33–67% of the entire spike; plants of brackish communities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. mackenziei 5b. Carpellate scales gray, gray-green, or light brown, slightly shorter than and not completely covering the perigynia; stems scabrous on the angles; uppermost spike with a short, basal, staminate comprising (0–) 15–54% of the entire spike; plants of freshwater and terrestrial communities 6a. Spikes composed of 5–10 (–14) carpellate flowers [Fig. 88]; perigynia ascending to ascending-spreading, green, becoming brown; abaxial side of the beak with a distinct longitudinal slit; leaf blades 0.5–2.5 mm wide, green to yellow-green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. brunnescens 6b. Spikes composed of (5–) 10–20 (–30) carpellate flowers; perigynia appressedascending, gray-green, becoming yellow-brown; abaxial side of the beak without a slit or with an inconspicuous one; leaf blades (1.5–) 2–4 mm wide, gray-green to pale green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. canescens Fig. 87  Inflorescence of Carex arcta.

67. Carex arcta Boott N Fig. 87 northern cluster sedge. Carex canescens L. var. polystachya Boott • ME, NH, VT; also reported from MA by Toivonen (2002), but specimens are unknown. Wet-mesic to hydric forests, usually at least in part evergreen, sometimes bordering streams. 68. Carex billingsii (Knight) C.D. Kirschbaum N Billings’ sedge. Carex trisperma Dewey var. billingsii Knight • CT, MA, ME, NH, RI, VT. Open bogs, rarely in the shade of trees. Carex billingsii has been treated as a variety of Carex trisperma and the two are, in fact, quite similar in many morphological details. However, they are easily separated by comparison of leaf blades. Further, they usually grow in different plant communities. Carex billingsii is usually found in open, acidic situations, whereas C. trisperma is normally found in shaded situations of varying pH. 69. Carex brunnescens (Pers.) Poir. N Fig. 88

Fig. 88  Inflorescence of Carex brunnescens var. brunnescens.

brownish sedge. 69a. Carex canescens L. var. alpicola Wahlenb.; 69b. Carex brunnescens ssp. sphaerostachya (Tuckerman) Kalela; C. canescens L. var. sphaerostachya Tuckerman • CT, MA,

Cy p e r ac e a e   127

ME, NH, RI, VT. Hydric or seasonally hydric soils of wetland edges, peatlands, and evergreen forests, also ascending high into the mountains and then often associated with disturbed soil. 1a. Leaf blades (1–) 1.5–2.5 mm wide; reproductive stems usually erect (less commonly ascending or arching), usually 15–60 cm tall; uppermost spike without or with an obscure, slender, staminate base . . . . . . . . . . . . . . . . . . . . . . . . . . 69a. C. brunnescens var. brunnescens 1b. Leaf blades (0.5–) 1–1.5 mm wide; reproductive stems ascending to arching, usually 30–90 cm tall; uppermost spike often with a short, but evident, slender, staminate base . . . . . . . . . . . . . . . . . . . . . . . . . 69b. C. brunnescens var. sphaerostachya (Tuckerman) Kükenth. Variety brunnescens is known from ME, NH, VT, usually high in the mountains, sometimes even near treeline. It frequently grows along trail edges and other disturbed places. It has also been reported from CT, MA, RI by Toivonen (2002), but specimens are unknown. Variety sphaerostachya is is known from CT, MA, ME, NH, RI, VT. It occurs over a wider range of plant communities. 70. Carex canescens L. N hoary sedge. 70a. Carex subloliacea (Fern.) Bickn.; 70b. Carex canescens L. var. disjuncta Fern.; C. disjuncta (Fern.) Bickn. • CT, MA, ME, NH, RI, VT. Wet areas, usually associated with bryophytes and/or organic soils, such as peatlands, boggy marsh edges, and wet evergreen forests. 1a. Inflorescence 3–5 (–7) cm tall; most spikes (except for the lowest 1 or 2) approximate to one another or separated by short internodes, the lowest internode mostly 10–20 mm long; reproductive stems 16–60 cm tall . . . . . . . . . . . . . . . . . . . . . 70a. C. canescens ssp. canescens 1b. Inflorescence 6–12 (–15) cm tall; most spikes (except for the upper 1 or 2) remote from one another, the lowest internode mostly 20–50 mm long; reproductive stems 30–90 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70b. C. canescens ssp. disjuncta (Fern.) Toivonen Subspecies canescens is known from CT, MA, ME, NH, RI, VT. Subspecies disjuncta is known from CT, MA, ME, NH, RI, VT. ‌70 × 71. Carex ×pseudohelvola Kihlm. is a rare sedge hybrid that is known from ME. It is similar to C. mackenziei in that the carpellate scales ± equal the perigynia and cover them. However, the hybrid differs from that species in its scabrous upper angles of the stem and the carpellate scales that are weakly tinged with red-brown. 71. Carex mackenziei Krecz. N Mackenzie’s sedge. Carex norvegica Willd. ex Schkuhr • ME. Brackish and saline marshes. 72. Carex tenuiflora Wahlenb.

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sparse-flowered sedge. ME, NH, VT; also reported from MA by Toivonen (2002), but specimens are unknown. Fens, usually in the shade of Thuja occidentalis. ‌ × 73. Carex ×trichina Fern. is a rare sedge hybrid known from ME. Characteristics 72 that distinguish this plant are its short-beaked perigynia and less congested inflorescence (compared with C. tenuiflora). 73. Carex trisperma Dewey N three-seeded sedge. CT, MA, ME, NH, RI, VT. Hydric, usually peaty soils of bogs, fens, and evergreen swamps, usually in shaded situations.

Gracilis Only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. brunnea 74. Carex brunnea Thunb. E greater brown sedge. Carex brunnea Thunb. var. nakiri Ohwi; C. sendaica Franch. var. nakiri (Ohwi) Koyama • MA. Fields, waste areas.

128  MO NOCOTS

Granulares 1a. Staminate spike on a peduncle 1–35 mm long, shorter than to barely exceeding the uppermost carpellate spike; stems cespitose on short, inconspicuous rhizomes; lowermost spike usually arising from upper half of reproductive stem . . . . . . . . . . . . . . . . . . . . C. granularis 1b. Staminate spike on a peduncle (4–) 9–76 (–91) mm long, overtopping the uppermost carpellate spike; stems produced singly or a few together from elongate rhizomes; lowermost spike usually arising from lower half of reproductive stem . . . . . . . . . . . . . . . . . . . . . . . . C. crawei 75. Carex crawei Dewey

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Crawe’s sedge. CT, ME. Gravel and ledgy river shores, lake shores, meadows, wet cliffs, seeps, and open rights-of-way in high-pH bedrock regions. Specimens from New England show some variation in morphology. In particular, those from CT often have a scabrous staminate spike peduncle, whereas those from ME have a smooth staminate peduncle. 76. Carex granularis Muhl. ex Willd. N limestone-meadow sedge. Carex granularis Muhl. ex Willd. var. haleana (Olney) Porter; C. haleana Olney; C. shriveri Britt. • CT, MA, ME, NH, VT. Shorelines, disturbed soil, meadows, often in high-pH bedrock regions.

Griseae Basal leaf sheath color is a critical character, and, therefore, complete specimens are necessary for confident identification (i.e., collections of only the upper stem and inflorescence may be difficult to assign to species). Color is best assessed with bright light and low magnification. Reference: Naczi and Bryson (2002). 1a. Leaf sheaths, and sometimes the blades along the abaxial midrib, hispidulous; beak of perigynium (0.5–) 0.8–1.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hitchcockiana 1b. Leaf sheaths and blades glabrous; beak of perigynium 0–0.6 mm long (up to 1.2 mm long in C. oligocarpa) 2a. Basal leaf sheaths of reproductive stems yellow-brown to dark brown, without anthocyanic pigments; carpellate spikes with (6–) 12–45 (–65) perigynia [Fig. 90] 3a. Peduncles of lateral spikes scaberulous; leaf blades green; carpellate scales with awns (0.2–) 1.2–2.7 mm long (very rarely awnless) [Fig. 89] . . . . . . . . . . . . . . . . C. conoidea 3b. Peduncles of lateral spikes smooth; leaf blades very glaucous; carpellate scales awnless or with an awn up to 1 (–1.5) mm long [Fig. 90] . . . . . . . . . . . . . . . . . C. glaucodea 2b. Basal leaf sheaths of reproductive stems purple-red to dark purple (very rarely dark brown); carpellate spikes with 2–18 perigynia 4a. Lowermost bract of inflorescence with a tightly clasping sheath; widest leaf blades (1.8–) 2.4–4 (–4.6) mm wide; carpellate spikes loosely flowered, many of the perigynia borne in 2 ranks; beak of perigynium 0.4–1.2 mm long . . . . . . . . . . . . . . . . . . C. oligocarpa 4b. Lowermost bract of inflorescence with a loosely clasping sheath; widest leaf blades (3.7–) 4.2–8.2 (–9.1) mm wide; carpellate spikes densely flowered, the perigynia spirally arranged around the spike axis; beak of perigynium (0.2–) 0.3–0.6 mm long 5a. Perigynia terete or nearly so in cross-section, (1.8–) 2–2.6 mm wide, 1.8–2.3 (–2.6) times as long as wide; achenes borne on a stipe (0.2–) 0.3–0.4 (–0.5) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. grisea 5b. Perigynia obtusely trigonous in cross-section, 1.5–1.9 (–2.2) mm wide, (2.2–) 2.5–3.1 times as long as wide; achenes borne on a stipe (0.3–) 0.4–0.6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. amphibola 77. Carex amphibola Steud. N eastern narrow-leaved sedge. Carex grisea Wahlenb. var. amphibola (Steud.) Kükenth. • CT, MA; also reported from NH by Seymour (1982), but specimens are unknown. Rich, deciduous,

Cy p e r ac e a e   1 2 9

upland or riparian forests. Reports of this species in ME, NH, and VT by Magee and Ahles (1999) are based on specimens of Carex conoidea and (mostly) C. grisea. 78. Carex conoidea Schkuhr ex Willd. N Fig. 89 open-field sedge. Carex katahdinensis Fern. • CT, MA, ME, NH, RI, VT. Meadows, river shores, lake shores, and open, wet, disturbed ground. Carex katahdinensis represents dwarfed individuals of C. conoidea that occur on northern ponds with fluctuating water levels. The dwarf stature appears to represent an adaptation for rapid completion of reproduction. Some specimens of C. conoidea with relatively fewer flowers in the spikes, especially those missing the basal leaf sheaths, can be confused with specimens of C. grisea. However, C. conoidea has narrower perigynia—(1.2–) 1.3–1.7 (–1.8) mm wide—and prominently scabrous staminate spike peduncles, whereas the perigynia of C. grisea are wider—(1.8–) 2–2.6 mm wide—and have a smooth or sparsely scabrous staminate spike peduncle. Carex conoidea is the only member of its section that routinely occurs in open habitats. 79. Carex glaucodea Tuckerman ex Olney

N C Fig. 90

Fig. 89  Spike of Carex connoidea.

blue sedge. Carex flaccosperma Dewey var. glaucodea (Tuckerman ex Olney) Kükenth. • CT, MA, NH, VT. Meadows, disturbed open soil, wetland borders, and woodlands, occurrences in nonhuman-disturbed communities are frequently within rich, oak-hickory woodlands on trap rock. 80. Carex grisea Wahlenb. N inflated narrow-leaved sedge. Carex amphibola Steud. var. turgida Fern. • CT, MA, ME, NH, VT. Rich, deciduous, upland or riparian forests, frequently in sandy fluvial soils. 81. Carex hitchcockiana Dewey N Hitchcock’s sedge. Carex oligocarpa Schkuhr ex Willd. var. hitchcockiana (Dewey) Kükenth. • CT, MA, NH, VT. Rich, mesic forests and woodlands in high-pH bedrock regions. 82. Carex oligocarpa Schkuhr ex Willd.

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rich woods sedge. CT, MA, VT. Rich, mesic to dry-mesic, often rocky, forests and woodlands in high-pH bedrock regions.

Heleoglochin 1a. Apical portion of sheath opposite the leaf blade pale and red-dotted on the hyaline surface; perigynia spreading, olive-brown to black, with a pale, depressed stripe on the abaxial surface, not covered by their subtending scales; inflorescence relatively congested, the lowest internode 3–7 (–10) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. diandra 1b. Apical portion of sheath opposite the leaf blade strongly copper-colored and reddotted on the hyaline surface; perigynia appressed, light brown, without a pale, depressed stripe, covered by their subtending scales; inflorescence more open, the lowest internode 7–26 (–33) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. prairea 83. Carex diandra Schrank N lesser tussock sedge. CT, MA, ME, NH, RI, VT. Fens, swamps dominated by Thuja occidentalis, lakeside fens, marshes, lake shores. 84. Carex prairea Dewey ex Wood N prairie sedge. Carex diandra Schrank var. ramosa (Boott) Fern. • CT, MA, ME, VT. High-pH fens, especially in shade or partial shade of Thuja occidentalis (northern New England) or Larix laricina (southern New England).

Hirtifoliae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hirtifolia 85. Carex hirtifolia Mackenzie N pubescent sedge. Carex pubescens Muhl. ex Willd. • CT, MA, ME, NH, RI, VT. Mesic, often rich, upland and riparian, deciduous forests.

Fig. 90  Spike of Carex glaucodea.

13 0 MO NOCOTS

Hymenochlaenae Because color of the basal portion of the plant is important, complete collections are recommended for confident identification. Reference: Waterway (2002). 1a. Uppermost spike usually gynecandrous 2a. Perigynia with 2 prominent, marginal veins, the remaining veins absent or obscure, without red dots; basal leaf sheaths green to green-brown, sometimes with a slight suffusion of anthocyanin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. prasina 2b. Perigynia with 2 prominent, marginal veins and 7–15 additional, distinct veins, often red-dotted; basal leaf sheaths suffused with dark red to purple-brown 3a. Perigynia 4.5–6 mm long, becoming dull orange-brown at maturity; carpellate scales with conspicuous awns 2.5–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. davisii 3b. Perigynia 2–5 mm long, not becoming orange-brown; carpellate scales acute to cuspidate or with a short awn shorter than 1 mm 4a. Lateral spikes gynecandrous, the basal 1 or 2 flowers staminate, borne on elongate peduncles usually exceeding the length of the associated spike, 4–6 mm thick; perigynia 3.5–5 mm long, abruptly tapering to a short beak . . . . . . . . C. formosa 4b. Lateral spikes unisexual and carpellate, usually borne on peduncles that are shorter than or equal to the length of the associated spike, 2–3.5 mm thick; perigynia 2–3.7 mm long, tapering to a beakless apex 5a. Leaf sheaths glabrous; leaf blades 3–9 mm wide . . . . . . . . . . . . . . . C. gracillima 5b. Leaf sheaths pubescent; leaf blades 1.5–3 mm wide . . . . . . . . . . . . C. aestivalis 1b. Uppermost spike usually unisexual and staminate 6a. Basal leaf sheaths brown, those of previous years persisting as abundant, fibrous remains; perigynia abruptly tapered to a long, tubular beak nearly as long as the perigynium body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sprengelii 6b. Basal sheaths red-purple, those of previous years not fibrous or only inconspicuously so; perigynia gradually tapered to a beak obviously shorter than the perigynium body 7a. Leaf blades pubescent; carpellate spikes 8–25 × 4–5 mm; perigynia with 2 prominent lateral veins and 5–7 additional, finer veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. castanea 7b. Leaf blades glabrous; carpellate spikes 25–80 × 2–3 mm; perigynia with 2 prominent lateral veins and 10–20 additional, finer veins 8a. Perigynia fusiform to narrow-ellipsoid, 5–9.5 mm long; achene elevated above base of perigynium on a stipe 0.5–1.5 mm long; leaf blades 2–5 (–7) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. debilis 8b. Perigynia ellipsoid-ovoid, 3–5 mm long; achene sessile at base of perigynium; leaf blades 3–10 (–13) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. arctata 86. Carex aestivalis M.A. Curtis ex Gray N summer sedge. CT, MA, NH, VT. Dry-mesic to mesic, often rocky, deciduous forests. 87. Carex arctata Boott in Hook. N drooping woodland sedge. Carex arctata Boott in Hook. var. faxonii Bailey • CT, MA, ME, NH, RI, VT. Dry-mesic to wet-mesic forests, including deciduous, evergreen, and mixed types.

Cy p e r ac e a e   13 1

‌ × 88. Carex ×knieskernii Dewey is a rare, largely sterile hybrid know from ME, NH. 87 It has long spikes of intermediate width between the two parents (20–45 × 3–5 mm) and sparsely pubescent leaf blades (the hairs most abundant in the basal portion of the blade). 88. Carex castanea Wahlenb. N chestnut sedge. Carex flexilis Rudge • CT, MA, ME, NH, VT. River shores, lake shores, fens, meadows, usually associated with high-pH bedrock. 89. Carex davisii Schwein. & Torr.

NC

Davis’ sedge. CT, MA, VT. Riparian forests, meadows. 90. Carex debilis Michx. N white-edged sedge.  90a. Carex debilis Michx. var. intercursa Fern.; 90b. Carex debilis Michx. var. allegheniensis Mackenzie; C. debilis Michx. var. interjecta Bailey; C. debilis Michx. var. strictior Bailey; C. flexuosa Muhl. ex Willd. • CT, MA, ME, NH, RI, VT. Forest edges and openings, trailsides, meadows, and open riparian banks and forests. 1a. Perigynia 5.5–9.5 mm long, fusiform, very gradually tapering to the apex; carpellate scales blunt to acute at apex (rarely cuspidate), with white or pale, hyaline margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90a. C. debilis var. debilis 1b. Perigynia 5–6.2 mm long, narrow-ellipsoid, relatively more abruptly tapering to apex; carpellate scales acute to cuspidate (to shortly awned), with hyaline margins that are usually streaked or suffused with red-brown . . . . . . . . . . . . . . . . . . . . 90b. C. debilis var. rudgei Bailey Variety rudgei is known from CT, MA, ME, NH, RI, VT. It is the common form of this species found throughout most of New England. Variety debilis is relatively rare and known only from CT, MA, RI. Rare forms of Carex debilis var. rudgei have erect spikes (the “strictior” form; rather than arching or drooping spikes). 90b × 161. This rare sedge hybrid is known from MA. It can be separated from Carex debilis by the sparse pubescence on the perigynia and the ± absence of perigynium beak teeth. It differs from C. virescens in its short perigynium beak (vs. beak absent), sparser perigynium pubescence, longer perigynia, and less densely flowered spikes. 91. Carex formosa Dewey N handsome sedge. CT, MA, VT. Meadows, rich, mesic forests, often upslope of wetlands and sometimes growing in vernal drainages where leaf litter and other vegetation is sparse. 92. Carex gracillima Schwein. N graceful sedge. Carex gracillima Schwein. var. macerrima Fern. & Wieg. • CT, MA, ME, NH, RI, VT. Mesic to hydric forests, margins of swamps, meadows. 92 × 161. This very rare sedge hybrid is known from CT. It generally resembles Carex virescens in the narrow and elongate spikes and pubescent leaf blades. However, the perigynia are very sparsely pubescent (i.e., the hairs are easily missed without careful examination), some leaf blades are glabrous or nearly so, and the two lateral veins of the leaf blades are more prominent than the midvein on the adaxial surface. 93. Carex prasina Wahlenb. N drooping sedge. CT, MA, ME, NH, RI, VT. Deciduous and mixed evergreen-deciduous forests, usually along streams and in other damp or wet places, including seeps and ditches. 94. Carex sprengelii Dewey ex Spreng. N long-beaked sedge. Carex longirostris Torr. • CT, MA, ME, NH, VT. Deciduous forests and openings, usually associated with riparian systems on high-terrace floodplains or sloping river banks.

13 2  MO NOCOTS

Laxiflorae Color of the basal portion of the plant is critical for identification. Therefore, complete collections are a must. Reference: Bryson and Naczi (2002b). 1a. Shoots anthocyanic at the base, ranging from a slight tinging of the brown background to a strong purple coloration 2a. Vegetative leaf blades 10–52 mm wide; blades of upper bracting leaves spathe-like, lanceolate to narrow-lanceolate, 4–16 mm wide; carpellate scales ± truncate and apiculate at the apex, lacking awns; flowering stems 2–2.4 mm wide, distinctly winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. albursina 2b. Vegetative leaf blades 1–12 mm wide; blades of upper bracting leaves not spathe-like, mostly linear, 0.5–6 mm wide; carpellate scales truncate to acuminate at the apex, usually with awns; flowering stems 0.5–2 mm wide, unwinged or with obscure wing angles 3a. Longest carpellate spike 19–33 mm long, the basal internode (i.e., the distance between the lowest and next apical flowers) 3.5–14 mm [Fig. 93]; perigynia varying from remote to loosely imbricate, with a nearly straight to bent beak; upper portion of flowering stem ± smooth on the angles . . . . . . . . . . . . . . . . . . . . . . . . . . C. ormostachya 3b. Longest carpellate spike 11–21 mm long, with a basal internode of 1.1–3.2 (–4.8) mm [Fig. 91]; perigynia closely imbricate, with an abruptly bent beak; upper portion of flowering stem scabrous on the angles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. gracilescens 1b. Shoots lacking anthocyanic pigments at the base, mostly pale brown to brown 4a. Vegetative leaves 10–52 mm wide; blades of upper bracting leaves spathe-like, lanceolate to narrow-lanceolate, 4–16 mm wide; flowering stems 2–2.4 mm wide, distinctly winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. albursina 4b. Vegetative leaves 3–26 mm wide; blades of upper bracting leaves not spathe-like, mostly linear, 0.8–8 mm wide; flowering stems 0.5–2 mm wide, unwinged or with obscure wing angles 5a. Perigynia with only 2 evident lateral veins, the remaing 6–16 veins faint; anthers (1.4–) 1.6–2 (–2.4) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. leptonervia 5b. Perignynia with 22–39 distinct veins; anthers (2–) 2.4–4 (–4.5) mm long (as short as 1.8 mm in C. blanda) 6a. Carpellate spikes 6–20 mm long, with crowded perigynia, the basal internode (i.e., the distance between the lowest and next apical flowera) 0.9–2.4 mm long [Fig. 91]; upper portion of flowering stem scabrous on the angles 7a. Perigynia tapering to a straight or merely outcurved beak 0.5–1.7 mm long; staminate scales 3.2–4.2 × 1–1.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. styloflexa 7b. Perigynia abruptly tapering to a short, bent beak 0.2–0.5 mm long; staminate scales 1.8–3.2 × 1.4–1.8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. blanda 6b. Carpellate spikes 9–62 mm long, the longest usually more than 18 mm, with approximate but not crowded perigynia, the basal internode (2.2–) 2.8–7.5 mm long [Fig. 93]; upper portion of flowering stem smooth on the angles 8a. Perigynia 2.6–3.8 mm long, usually more abruptly tapered in the apical portion than in the basal portion; staminate spikes 12–26 mm long . . . . . . . . . . C. laxiflora 8b. Perigynia 3.4–5 mm long, usually ± evenly tapered in the apical and basal portions; staminate spikes 22–36 mm long . . . . . . . . . . . . . . . . . . . . . . . . . C. striatula 95. Carex albursina Sheldon N white bear sedge. Carex laxiflora Lam. var. latifolia Boott • CT, MA, NH, VT. Rich, mesic forests, often rocky or with ledge outcrops.

Cy p e r ac e a e   13 3

96. Carex blanda Dewey N Fig. 91 eastern woodland sedge. Carex laxiflora Lam. var. blanda (Dewey) Boott • CT, MA, ME, NH, RI, VT. Deciduous or evergreen-deciduous forests. 97. Carex gracilescens Steud.

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slender loose-flowered sedge. Carex laxiflora Lam. var. gracillima (Boott) Boott ex B.L. Robins. & Fern. CT, MA, VT. Forests, woodlands, edges, and wetland margins. 98. Carex laxiflora Lam. N broad loose-flowered sedge. Carex laxiflora Lam. var. serrulata F.J. Herm. • CT, MA, ME, NH, RI, VT. Deciduous and mixed evergreen-deciduous forests, often associated with rocky slopes and ledge outcrops.

Fig. 91  Spikes with crowded perigynia of Carex blanda.

99. Carex leptonervia (Fern.) Fern. N Fig. 92 nerveless woodland sedge. Carex laxiflora Lam. var. leptonervia Fern. • CT, MA, ME, NH, RI, VT. Mesic to wet-mesic forests. 100. Carex ormostachya Wieg. N necklace spike sedge. Carex laxiflora Lam. var. ormostachya (Wieg.) Gleason • CT, MA, ME, NH, VT. Deciduous and mixed evergreen-deciduous forests. 101. Carex striatula Michx.

N C Fig. 93

lined sedge. Carex laxiflora Lam. var. angustifolia Dewey • CT. Rich, mesic forests, often rocky or with ledge outcrops. 102. Carex styloflexa Buckl. N bent sedge. CT, RI. Deciduous forests, often associated with small streams and seeps.

Leptocephalae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. leptalea 103. Carex leptalea Wahlenb. ssp. leptalea N

Fig. 92  Prolonged sheath of lowermost bract of inflorescence of Carex leptonervia.

bristle-stalk sedge. CT, MA, ME, NH, RI, VT. Forests, evergreen swamps, and wooded fens, usually in rich soils and/or areas with circumneutral bedrock.

Leucoglochin only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pauciflora 104. Carex pauciflora Lightf. N few-flowered sedge. CT, MA, ME, NH, VT. Bogs, usually in the open away from deep shade.

Limosae Reference: Ball (2002b). 1a. Carpellate scales longer than the perigynia they subtend [Fig. 94]; lowest bract of inflorescence 3–15 cm long, equaling or exceeding the height of the inflorescence; stems usually phyllopodic, with dead leaf remains at base . . . . . . . . . . . . . . . . . . . . . . . C. magellanica 1b. Carpellate scales shorter than (rarely as long as) the perigynia they subtend [Fig. 95]; lowest bract of inflorescence 0.5–7 cm long, shorter than the height of the inflorescence; stems usually aphyllopodic, without dead leaf remains at base (usually phyllopodic and with dead leaf remains at base in C. rariflora) 2a. Carpellate scales 1.3–1.8 mm wide, narrower than the perigynia they subtend; lateral spikes usually androgynous, (10–) 25–50 mm long; leaf blades 2–5 mm wide . . . . C. barrattii

Fig. 93  Spikes with approximate perigynia of Carex striatula.

13 4  MONOCOTS

2b. Carpellate scales 2–3.4 mm wide, wider than (rarely as wide as) the perigynia they subtend; lateral spikes usually unisexual and carpellate (sometimes androgynous in C. limosa), 6–20 mm long; leaf blades 1–2.5 mm wide 3a. Perigynia acute at apex, with a beak 0.1–0.5 mm long; stems usually aphyllopodic, without dead leaf remains at base; lateral spikes with 8–30 perigynia; leaf blades glaucous-green, involute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. limosa 3b. Perigynia rounded to obtuse at apex, with a beak up to 0.3 mm long; stems usually phyllopodic, with dead leaf remains at base; lateral spikes with 2–10 perigynia [Fig. 95]; leaf blades green, relatively flat or sometimes with involute margins . . . . . . . C. rariflora 105. Carex barrattii Schwein. & Torr.

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Barratt’s sedge. CT. Swamps, graminoid marshes, wet meadows. 106. Carex limosa L. N mud sedge. CT, MA, ME, NH, VT; also reported from RI by Gould et al. (1998), but specimens are unknown. Bogs and fens, in the open and usually found in acidic or somewhat acidic types. 107. Carex magellanica Lam. ssp. irrigua (Wahlenb.) Hiitonen N Fig. 94 Fig. 94  Lateral spike of Carex magellanica.

boreal bog sedge. Carex limosa L. var. irrigua Wahlenb.; C. paupercula Michx.; C. paupercula Michx. var. irrigua (Wahlenb.) Fern. • CT, MA, ME, NH, VT. Bogs and fens, usually in the open, away from deep shade. 108. Carex rariflora (Wahlenb.) Sm.

N C Fig. 95

loose-flowered alpine sedge. Carex limosa L. var. rariflora Wahlenb. • ME. High-elevation heaths and boggy areas; known only from historical collections from Katahdin, Piscataquis County, ME.

Lupulinae Fig. 95  Lateral spike of Carex rariflora.

Reference: Reznicek (2002a). 1a. Carpellate spikes spherical to short-ovoid, 1–4.2 cm long; lowermost bract of inflorescence usually sheathless or with a sheath shorter than 3 mm; achenes ellipsoid to obovoid; beak of perigynium 1.5–4.2 mm long 2a. Carpellate spikes spherical, with 8–35 perigynia radiating in all directions; base of perigynium cuneate; surface of perigynium dull . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. grayi 2b. Carpellate spikes mostly short-ovoid, with 1–12 (–20) usually ascending to spreading perigynia (uncommonly the lower perigynia are reflexed); base of perigynium convexly rounded; surface of perigynium somewhat lustrous . . . . . . . . . . . . . . . . . . . . . C. intumescens 1b. Carpellate spikes ovoid to cylindric, 1.5–8 cm long; lowermost bract of inflorescence with a closed sheath 5–150 mm long; achenes rhomboid; beak of perigynium 6–10 mm long 3a. Achenes (2.2–) 2.4–3.4 mm wide, nearly as wide as long, the angles knobbed with nipple-like points [Fig. 96] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lupuliformis 3b. Achenes 1.7–2.6 (–2.8) mm wide, longer than wide, the angles without knobs or only obscurely knobbed [Fig. 97] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lupulina 109. Carex grayi Carey N Gray’s sedge. Carex asa-grayi Bailey; C. grayi Carey var. hispidula Gray; C. intumescens Rudge var. globularis Gray • CT, MA, VT. Rich, mesic soils and calcareous seepage swamps, usually associated with riparian systems. 110. Carex intumescens Rudge N greater bladder sedge. Carex intumescens Rudge var. fernaldii Bailey • CT, MA, ME, NH, RI, VT. Mesic to hydric forests and wetland edges. 111. Carex lupuliformis Sartwell ex Dewey

Fig. 96  Achene of Carex lupuliformis.

N C Fig. 96

false hop sedge. CT, MA, VT. Swamps, lacustrine forests, stream banks, edges of marshes, vernal pools.

Cy p e r ac e a e   13 5

112. Carex lupulina Muhl. ex Willd. N Fig. 97 hop sedge. Carex lupulina Muhl. ex Willd. var. pedunculata Gray • CT, MA, ME, NH, RI, VT. Marshy lake shores, openings in swamps, graminoid marshes. 112 × 182. This rare sedge hybrid is known from CT, MA, ME, VT. It appears similar to Carex lupulina but with smaller spikes, shorter than average perigynia (9.5–13 mm long vs. 11–19 mm long), and carpellate scales always with a prominent, scabrous awn.

Macrocephalae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. kobomugi 113. Carex kobomugi Ohwi E Japanese sedge. Carex macrocephala Willd. ex Spreng. var. kobomugi (Ohwi) Miyabe & Kudô • MA, RI. Sandy beaches and dunes of the Atlantic coast, borrow pits near the Atlantic coast.

Mitratae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. caryophyllea 114. Carex caryophyllea Lat. E vernal sedge. MA, ME. Fields, pastures, and other human-disturbed or human-manicured areas.

Multiflorae 1a. Leaves shorter than the flowering stems; perigynia gold-brown at maturity, with a beak ca. ⅓ as long as the body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. annectens 1b. Leaves equaling or taller than the flowering stems; perigynia brown at maturity, with a beak ca. ⅓ to ½ as long as the body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. vulpinoidea 115. Carex annectens (Bickn.) Bickn. N yellow-fruited sedge. Carex brachyglossa Mackenzie; C. setacea Dewey var. ambigua (Barratt ex Boott) Fern.; C. xanthocarpa Bickn. var. annectens Bickn. • CT, MA, ME, NH, RI, VT. Mesic to hydric open areas such as low fields, meadows, and ditches. 116. Carex vulpinoidea Michx. N common fox sedge. Carex setacea Dewey • CT, MA, ME, NH, RI, VT. Wet-mesic to hydric soils of meadows, marshes, and ditches.

Paludosae Reference: Reznicek and Catling (2002). 1a. Perigynia glabrous; style persistent on the achene; leaf blades (2.6–) 5.5–21 mm wide 2a. Perigynia 3–4.5 mm long, glaucous; perigynium beak 0.3–0.6 mm long, weakly bidentate, the apical teeth up to 0.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. acutiformis 2b. Perigynia 3.9–7.8 mm long, not glaucous; perigynium beak 0.5–1.6 mm long, terminated by 2 distinct teeth (0.1–) 0.2–0.7 (–0.9) mm long 3a. Leaves with blades (5.5–) 8.5–21 mm wide, the longest ligules 13–40 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lacustris 3b. Leaves with blades 2.6–5 (–6) mm wide, the longest ligules 1.8–12.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. striata 1b. Perigynia pubescent; style articulated with the summit of the achene and deciduous from it at maturity; leaf blades 0.7–8.5 mm wide 4a. Beak of perigynium with soft, hyaline, obscure teeth; staminate spikes 1 or rarely 2, borne on peduncles 2–20 mm long, usually exceeding the height of the lowest bract of the inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. vestita

Fig. 97  Achene of Carex lupulina.

13 6 MONOCOTS

4b. Beak of perigynium with distinct, firm teeth 0.2–0.8 mm long; staminate spikes 1–3, borne on peduncles (8–) 20–90 mm long, usually equaled or overtopped by the lowest bract 5a. Perigynia 4.5–6.5 mm long, the pubescence short and relatively sparse, not concealing the veins or cellular details of the perigynium body; plants of dry, sandy or rocky soil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. houghtoniana 5b. Perigynia 2.4–5 (–5.2) mm long, the pubescence longer and dense, concealing the veins and cellular details of the perigynium body; plants of shorelines and wetlands 6a. Leaf blades M-shaped in cross-section (at least when young), sometimes ± flat at maturity, (2–) 2.2–4.5 (–6) mm wide, not prolonged apically into an elongate, filiform tip, with a prominent midvein that forms a sharp, abaxial keel for much of the lengt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pellita 6b. Leaf blades involute, 0.7–2 (–2.2) mm wide, prolonged at the apex into an elongate, sometimes curled, filiform tip, with a low, rounded midvein forming an inconspicuous abaxial keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lasiocarpa 117. Carex acutiformis Ehrh. E lesser pond sedge. CT, MA. Graminoid meadows, marsh edges, lake shores, and wet fields. 118. Carex houghtoniana Torr. ex Dewey N Houghton’s sedge. ME, NH, VT. Open, often disturbed, soils of fields, roadsides, logged forests, and ledges. 119. Carex lacustris Willd. N lakeside sedge. CT, MA, ME, NH, RI, VT. Shorelines, marshes, wet fields. 120. Carex lasiocarpa Ehrh. ssp. americana (Fern.) D. Löve & Bernard N woolly-fruited sedge. Carex lanuginosa Michx.; C. lanuginosa Michx. var. americana (Fern.) Boivin; C. lasiocarpa Ehrh. var. americana Fern. • CT, MA, ME, NH, RI, VT. Graminoid marshes, fens, lake shores, even forming floating mats out into water bodies. 121. Carex pellita Muhl. N woolly sedge. Carex lanuginosa, auct. non Michx.; C. lasiocarpa Ehrh. var. latifolia (Boeckl.) Gilly • CT, MA, ME, NH, RI, VT. Marshes, roadsides, meadows, and shorelines, most abundant in regions of high-pH bedrock or till. 122. Carex striata Michx.

NC

Walter’s sedge. Carex striata Michx. var. brevis Bailey; C. walteriana Bailey • MA, NH, RI. Peaty shorelines, lakeside fens, and graminoid marshes. 123. Carex vestita Willd. N velvet sedge. CT, MA, ME, NH, RI. Dry-mesic to wet-mesic, open, sandy soils of fields, roadsides, sandplains, and woodlands.

Paniceae Reference: Rothrock and Reznicek (2002). 1a. Perigynium beakless or with a beak shorter than 0.5 mm 2a. Leaves coriaceous, involute, heavily glaucous; perigynia appressed-ascending, tapering to a beakless or short, straight-beaked apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. livida 2b. Leaves herbaceous, flat, folded, or M-shaped in cross-section (at least when young), sometimes ± flat at maturity, not or only slightly glaucous; perigynia ascending to spreading, concavely tapering (at least on one surface) to a short, curved beak

Cy p e r ac e a e   137

3a. Reproductive stems smooth along the angles; lowermost bract of inflorescence short, the blade and sheath mostly 28–58% of the total inflorescence height . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. panicea 3b. Reproductive stems scabrous along the angles (at least near the summit); lowermost bract of inflorescence relatively elongate, the blade and sheath mostly 62–111% of the total inflorescence height 4a. Achenes 1.2–1.6 (–1.8) mm wide; ligules of leaves (0.6–) 1–6 mm long, (0.8–) 1–2 times as long as wide; leaf blades green; carpellate spikes 3–5.8 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tetanica 4b. Achenes 1.7–2.2 (–2.5) mm wide; ligule of leaves 0.4–3.6 mm long, 0.4–1.2 times as long as wide; leaf blades gray-green; carpellate spikes 3.5–8 mm thick . . . . . C. meadii 1b. Perigynium tapering to a distinct beak 0.6–1.8 mm long 5a. Lower sheaths of reproductive stems pale brown; perigynia and abaxial leaf blade surface not, or only sparsely, minutely papillose; lower perigynia of each spike loosely arranged; plants of forested wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. vaginata 5b. Lower sheaths of reproductive stems strongly tinged with anthocyanin; perigynia and abaxial leaf blade surface minutely papillose; lower perigynia of each spike densely arranged; plants of well-drained, sandy soils . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. polymorpha 124. Carex livida (Wahlenb.) Willd.

NC

livid sedge. Carex livida (Wahlenb.) Willd. var. grayana (Dewey) Fern.; C. livida (Wahlenb.) Willd. var. radicaulis Paine • CT, MA, ME, NH, VT. Fens, including acid, circumneutral, and lakeside types. 125. Carex meadii Dewey

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Mead’s sedge. Carex tetanica Schkuhr var. meadii (Dewey) Bailey • RI. Fens and circumneutral meadows. 126. Carex panicea L. E grass-like sedge. CT, MA, ME, NH, RI. Wet-mesic to dry-mesic, often sandy, soils of fields, roadsides, and cleared areas. 127. Carex polymorpha Muhl.

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variable sedge. CT, MA, ME, NH, RI. Sandy soils of woodlands, forest edges, borrow pits, and cleared rights-of-way. 128. Carex tetanica Schkuhr

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rigid sedge. CT, MA, NH. Circumneutral fens, meadows, and graminoid marshes. Reports of Carex woodii Dewey in New England (e.g., Dowhan 1979, Seymour 1982) were based, in large part, on specimens of C. tetanica that showed slight anthocyanic suffusion of the lower, bladeless sheaths and shallow rhizomes. However, C. woodii shows deep anthocyanic suffusion of the lower, bladeless sheaths and shallow, relatively stout rhizomes that emit aerial shoots at ± regular intervals. Further, the specimens were collected from fens (typical for C. tetanica, atypical for C. woodii). 129. Carex vaginata Tausch sheathed sedge. Carex saltuensis Bailey • ME, VT; also reported from NH by Rothrock and Reznicek (2002), but specimens are unknown. Evergreen swamps dominated by Thuja occidentalis, wooded fens.

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13 8 MO NOCOTS

Phacocystis ‌limula Carex × T. Fries was reported from ME by Campbell et al. (1995). The specimens that supposedly vouchered this occurrence were all C. bigelowii (all at MAINE!). References: Haines (2000b), Standley et al. (2002). 1a. At least the lower carpellate scales with the apex prolonged into a conspicuous, usually scabrous, awn [Figs. 99, 100] 2a. Lowest carpellate spike usually drooping on a peduncle (5–) 14–68 mm long; carpellate scales including awns (2.9–) 3.1–20 mm long, pale brown to copper-brown 3a. Plants colonial from elongate rhizomes; lowermost spike 8–20 mm thick; plants of saline and brackish marshes and strands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. paleacea 3b. Plants cespitose from short rhizomes; lowermost spike 5–10 mm thick; plants of non-saline wetlands and shorelines 4a. Lower sheaths glabrous; apex of carpellate scales retuse or less commonly trunctate [Fig. 99]; perigynia obovoid to broad-obovoid (rarely ellipsoid) . . . . C. crinita 4b. Lower sheaths scabrous with short, stiff hairs; apex of carpellate scales acuminate to truncate (sometimes slightly retuse in C. mitchelliana) [Fig. 100]; perigynia ellipsoid to ovoid 5a. Apex of carpellate scales acute to acuminate; perigynia smooth or weakly papillose near the apex; achenes with a constriction . . . . . . . . . . . . . . . C. gynandra 5b. Apex of carpellate scales ± truncate; perigynia densely papillose throughout; achenes without a constriciton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. mitchelliana 2b. Lowest carpellate spike ascending to arching on a peduncle 6–20 mm long; carpellate scales including awns 2.5–9 mm long, brown to red-brown or purple-brown 6a. Perigynia long-papillose, with 2–5 veins on each face; carpellate scales dark brown to purple-brown, with a paler central band 10–33% as wide as the entire scale; achenes dull, with or without folds or constrictions; perigynium beak sometimes with scabrules about the orifice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. vacillans 6b. Perigynia short-papillose, veinless or obscurely veined; carpellate scales bronze to brown, with a central pale band 33–50% as wide as the entire scale; achenes lustrous, with a fold across one face; perigynium beak without scabrules . . . . . . . . . . . . . . . C. recta 1b. Carpellate scales unawned at the apex, at most with a minute, smooth cusp 7a. Lowermost spike arching to drooping; perigynium smooth, with a triangular, flat, often twisted, apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. torta 7b. Lowermost spike erect to ascending; perigynium minutely papillose, with a rounded to acute, only somewhat flattened, plane apex 8a. Lowest bract of inflorescence exceeding the total height of the inflorescence (often conspicuously so) [Fig. 101]; leaf blades amphistomic 9a. Leaves 1.5–3.5 mm wide; perigynia with (3–) 5–7 veins on each surface; achenes dull; plants cespitose from short rhizomes; lateral spike entirely carpellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lenticularis 9b. Leaves 2.5–8 mm wide; perigynia without veins or with 1–4 obscure (rarely distinct) veins on one surface [Fig. 98]; achenes glossy; plants colonial from elongate rhizomes; usually at least some of the lateral spikes androgynous . . . . . . C. aquatilis 8b. Lowest bract of inflorescence shorter than or equaling the total height of the inflorescence; leaf blades hypostomic (except C. nigra, which has stomates on the adaxial surface)

Cy p e r ac e a e   13 9

10a. Carpellate scales purple-brown to black; stems arising from the center of the previous year’s leaf tufts, surrounded by senescing leaves 11a. Perigynia with 3–9 veins on each surface, the beak with scabrules about the orifice [Fig. 102]; leaf blades with stomates on the adaxial surface . . . . . . . . C. nigra 11b. Perigynia without veins or with 1–4 obscure (rarely distinct) veins on one surface, the beak lacking scabrules; leaf blades lacking stomates on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bigelowii 10b. Carpellate scales pale brown to red-brown; stems arising laterally, not surrounded by the previous year’s leaves 12a. Lower sheaths not fibrillose; sheaths of stems leaves convex and prolonged at summit; perigynia not spotted with red, with 3–5 veins on each surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. emoryi 12b. Lower sheaths splitting into an inconspicuous to prominent ladder-network of fibers [Fig. 103]; sheaths of stem leaves concave at summit; perigynia sometimes red-spotted in apical portion, with 0–5 veins on each face 13a. Carpellate scales shorter than the perigynia (varying to as long as), acute at the apex; perigynia ovoid, acute to obtuse at the apex, with 0–5 veins on each surface, ascending; stems commonly ± equal in height to the leaves; lower leaf sheaths scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stricta 13b. Carpellate scales longer than the perigynia, acute to acuminate at the apex; perigynia obovoid, rounded at apex, without veins, divergent; stems commonly surpassing the leaves in height; lower leaf sheaths glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. haydenii 130. Carex aquatilis Wahlenb. ssp. altior (Rydb.) Hultén N Fig. 98 water sedge. Carex aquatilis Wahlenb. var. altior (Rydb.) Fern.; C. aquatilis Wahlenb. var. substricta Kükenth.; C. substricta (Kükenth.) Mackenzie • CT, MA, ME, RI, VT. Marshes, fens, river shores, graminoid meadows. This species was reported from NH by Magee and Ahles (1999); however, the voucher specimens are Carex haydenii and C. stricta (specimens at NHA!) and C. lenticularis (specimen at MASS!). Recent work by Dragon and Barrington (2009) showed that the traditional infraspecific taxonomy of C. aquatilis does not accurately reflect the phylogeny of this species (i.e., some of the varieties do not form distinct lineages). However, both molecular and morphological evidence support a division between North American and European populations (North American plants have thinner stems and broader carpellate scales than do European ones). Therefore, our plants are here treated as a separate subspecies from the European ssp. aquatilis. In New England, two morphological forms occur that show weak ecological differences. The common form, which occurs on relatively higher pH substrate, has pale carpellate scales with a narrow, red-brown margin and a broad, pale central band (the “substricta” form). The less common form (primarily restricted to ME) has red-brown to black carpellate scales that have a narrow, pale margin and midrib (the “aquatilis” form).

Fig. 98  Perigynium of Carex aquatilis.

131. Carex bigelowii Torr. ex Schwein. ssp. bigelowii N Bigelow’s sedge. Carex rigida Goodenough; C. rigida Goodenough var. bigelowii (Torr. ex Schwein.) Tuckerman; C. rigida Goodenough var. concolor (R. Br.) Kükenth. • ME, NH, VT. Alpine ridges and plateaus. 132. Carex crinita Lam.

N C Fig. 99

fringed sedge. 132a. Carex porteri Olney • CT, MA, ME, NH, RI, VT. Marshes, open swamps, shorelines, wet fields, and ditches. Carex crinata var. porteri (Olney) Fern. is a poorly known taxon from ME. Material of this variety was unavailable for study during preparation of this manual. Its identity is unresolved. 1a. Achenes lacking a constriction or indentation; perigynia broad-obovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132a. C. crinita var. brevicrinis Fern.

Fig. 99  Carpellate scale of Carex crinita.

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1b. Achenes with a constriction or indentation on one or both margins; perigynia ellipsoid to obovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132b. C. crinita var. crinita Variety crinita is known from CT, MA, ME, NH, RI, VT. Variety brevicrinis is known from MA and is of regional conservation concern. It is more rare than previously believed as most of the syntypes cited in the original description are not this variety. 132b × 142. This very rare sedge hybrid is known from NH. It has arching spikes, the lowest one on peduncles 11–31 mm long, basal carpellate flowers subtended by a short-awned scale (awns as long as 2.5 mm and scabrous), the apex of some scales truncate to obscurely retuse, and lower leaf sheaths smooth or sparsely scabrous. 133. Carex emoryi Dewey N Emory’s sedge. Carex millegrana T. Holm; C. stricta Lam. var. elongata (Boeckl.) Gleason; C. stricta Lam. var. emoryi (Dewey) Bailey • CT, MA. Marshes and stream shores. 134. Carex gynandra Schwein. N Fig. 100 nodding sedge. Carex crinita Lam. var. gynandra (Schwein.) Schwein. & Torr.; C. crinita Lam. var. simulans Fern.; C. gynandra Schwein. var. simulans (Fern.) Rolland-Germain • CT, MA, ME, NH, RI, VT. Marshes, open swamps, shorelines, wet fields, and ditches. 135. Carex haydenii Dewey N Fig. 100  Carpellate scale of Carex gynandra.

Hayden’s sedge. Carex stricta Lam. var. decora Bailey; C. stricta Lam. var. haydenii (Dewey) Kükenth. • CT, MA, ME, NH, RI, VT. Open areas with seasonally saturated soils such as river shores, graminoid marshes, meadows. 135 × 138. This very rare sedge hybrid is known from MA. It is most similar to Carex nigra due to the dark carpellate scales and perigynia that are usually suffused with dark red-brown. However, the carpellate scales are acute to acuminate and usually at least reach the base of the perigynium beak varying to sometimes shortly surpassing the beak (vs. carpellate scales that are rounded to obtuse (rarely acute) at the apex that usually do not reach the base of the beak). The leaves of the hybrid are generally shorter and mostly don’t reach the base of the inflorescence (vs. commonly reaching the inflorescence to equaling the height of the plant). 136. Carex lenticularis Michx. N Fig. 101 lake shore sedge. Carex lenticularis Michx. var. albimontana Dewey; C. lenticularis Michx. var. blakei Dewey • MA, ME, NH, VT. Shorelines, often those of high mineral content (e.g., sand, gravel, stone), including moderately high elevation tarns. 137. Carex mitchelliana M.A. Curtis

Fig. 101  Inflorescence and overtopping lowermost bract of Carex lenticularis.

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Mitchell’s sedge. Carex crinita Lam. var. mitchelliana (M.A. Curtis) Gleason • MA, RI. Swamps, both deciduous and evergreen types, stream banks, shorelines, and graminoid marshes. 138. Carex nigra (L.) Reichard N Fig. 102 smooth black sedge. Carex acuta L. var. nigra L.; C. nigra (L.) Reichard var. strictiformis  (Bailey) Fern. • CT, MA, ME, NH, RI, VT. Lawns, low fields and meadows, ditches, usually within the coastal region. 139. Carex paleacea Schreb. ex Wahlenb. N chaffy sedge. Carex crinita Lam. var. paleacea (Schreb. ex Wahlenb.) Dewey; C. paleacea Schreb. ex Wahlenb. var. transatlantica Fern. • MA, ME, NH. Saline and brackish marshes.

Fig. 102  Perigynium of Carex nigra.

139 × 141. This rare sedge hybrid is known from MA, ME, NH. It is very similar to Carex recta and C. vacillans (see identification key). It can be separated from both by its leaf blades that usually lack stomates on the adaxial surface (present in both of the aforementioned species) and its relatively consistent production of prominently ladderfibrillose basal leaf sheaths (this feature usually lacking in the aforementioned species). From C. recta it is further distinguished by its nerved perigynia (usually 1–4 ± prominent veins on mature perigynia) and by its nearly complete sterility (C. recta usually produces many mature achenes). From C. vacillans it is further distinguished by its relatively

Cy p e r ac e a e   1 41

broad, pale central band on the carpellate scales, narrower perigynium apex (acute in this hybrid, rounded or nearly so in C. vacillans), and absence of scabrules about the orifice of the perigynia beak. 140. Carex recta Boott

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‌neofilipendula estuary sedge. Carex kattegatensis Fries ex Lindm.; C. × Lepage; C. salina ‌subnigra Lepage • ME; Wahlenb. var. kattegatensis (Fries ex Lindm.) Almquist; Carex × far-eastern portion of state. Saline and brackish marshes, Atlantic coast shorelines, tidal river shores. This extremely rare species is over-reported, and nearly all records from New England are based on Carex vacillans and Carex paleacea × C. stricta (the latter is specifically responsible for the reports in MA and NH by Standley et al. 2002). 141. Carex stricta Lam. N Fig. 103 tussock sedge. Carex stricta Lam. var. strictior (Dewey) Carey; C. strictior Dewey • CT, MA, ME, NH, RI, VT. Swamps, marshes, shorelines, and low fields. Plants from very wet habitats tend to have cespitose stems (sometimes producing conspicuous clumps of stems) and poorly formed rhizomes. Plants from drier habitats tend to be long-rhizomatous and do not produce clumps of stems. These two forms have been named, but they do intergrade. 142. Carex torta Boott ex Tuckerman N twisted sedge. Carex acuta L. var. sparsiflora Dewey; C. torta Boott ex Tuckerman var. composita Porter • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Rocky and gravelly river shores and stream banks. 143. Carex vacillans Drej. ex Hartman

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‌super-goodenoughii swinging sedge. Carex × (Kükenth.) Lepage • MA, ME. Saline and brackish marshes, Atlantic coast shorelines, tidal river shores. Most (and perhaps all) of the recent reports of this species in MA (e.g., Standley et al. 2002) are based on Carex paleacea × C. stricta.

Phaestoglochin Reference: Ball (2002c). 1a. Sheaths loosely clasping the stem, septate-nodulose (i.e., with cross-veins connecting the elongate veins of the sheath), green and white-striped or -spotted; widest leaf blades 4–10 mm wide 2a. Inflorescence elongate, (3–) 4–10 cm long, with remote lower spikes; body of perigynium with a narrow wing 0.1–0.2 mm wide (sometimes narrower in the basal portion of the perigynium) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sparganioides 2b. Inflorescence relatively congested, 2–4 cm long, with crowded lower spikes; body of perigynium with a very narrow wing up to 0.1 mm wide . . . . . . . . . . . . . . . . . . C. cephaloidea 1b. Sheaths tightly clasping the stem, not septate-nodulose, usually not striped or spotted; widest leaf blades 0.9–4 (–5) mm wide 3a. Beak of perigynium entire along the margins . . . . . . . . . . . . . . . . . . . . . . . . . . C. retroflexa 3b. Beak of perigynium minutely serrulate along the margins 4a. Inflorescence open, with well-separated lower spikes; basal internodes of inflorescence 2 or more times as long as the lower spikes 5a. Stigmas straight, reflexed, or partially coiled, 0.03–0.06 mm wide; spongy base of perigynium 1–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. radiata 5b. Stigmas 1- to 3-times coiled, 0.05–0.1 mm wide; spongy base of perigynium 0.5–1.3 mm long 6a. Widest leaves 0.9–1.5 mm wide; base of flowering stems 0.7–1.4 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. appalachica

Fig. 103  Fibrillose basal leaf sheaths of Carex stricta.

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6b. Widest leaves 1.8–2.6 mm wide; base of flowering stems 1.5–2.2 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rosea 4b. Inflorescence dense, with approximate to crowded spikes; basal internodes of inflorescence less than 2 times as long as the lower spikes 7a. Body of carpellate scale 1–1.8 mm long, up to ½ the length of the perigynium it subtends; achenes 1.4–1.5 × 1.3–1.4 mm; inflorescence dense, the individual spikes not distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cephalophora 7b. Body of carpellate scale 2.3–4.5 mm long, more than ⅔ the length of the perigynium it subtends; achenes 1.5–2.3 × 1.3–2.1 mm; inflorescence dense to somewhat elongate, the individual spikes distinct or not 8a. Ligules (3.5–) 4–8 mm long, much longer than broad; perigynia 4–5.5 mm long; basal sheaths usually tinged with anthocyanin . . . . . . . . . . . . . . . . . . . . . . . C. spicata 8b. Ligules up to 3 mm long, usually wider than long; perigynia 2.7–4.2 mm long; basal sheaths green to brown, without anthocyanic pigments 9a. Inflorescence a dense, globose to ovoid cluster 1–1.5 times as long as wide, the individual spikes highly congested and sometimes not even distinct; perigynia weakly veined (if at all) on the abaxial surface; achenes 1.5–1.7 × 1.3–1.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. mesochorea 9b. Inflorescence elongate to rather congested, usually more than 1.5 times as long as wide, at least the lower spikes usually ± distinct; perigynia either with 9–15 veins on the abaxial surface or ± veinless; achenes (1.8–) 1.9–2.2 × (1.8–) 1.9–2.1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. muehlenbergii 144. Carex appalachica J. Webber & P.W. Ball N Appalachian sedge. CT, MA, ME, NH, VT; also reported from RI by Gould et al. (1998), but specimens are unknown. Dry-mesic to mesic, deciduous and mixed evergreen-deciduous forests. The name Carex radiata was misapplied to this species for many years. 145. Carex cephaloidea (Dewey) Dewey N thin-leaved sedge. Carex muricata L. var. cephaloidea Dewey; C. sparganioides Muhl. ex Willd. var. cephaloidea (Dewey) Carey • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Mesic, often rich, soils of upland deciduous forests and riparian forests. 146. Carex cephalophora Muhl. ex Willd. N oval-headed sedge. CT, MA, ME, NH, RI, VT. Dry-mesic to mesic soils of forests, woodlands, and open areas, often occurring on rocky or ledgy slopes and frequently in regions of moderately high to high pH bedrock. 147. Carex mesochorea Mackenzie

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midland sedge. Carex cephalophora Muhl. ex Willd. var. mesochorea (Mackenzie) Gleason • CT, MA. Dry-mesic to xeric open areas such as grasslands, fields, and roadsides. 148. Carex muehlenbergii Schkuhr ex Willd. N Muhlenberg’s sedge.  148b. Carex plana Mackenzie • CT, MA, ME, NH, RI, VT; rare to the north. Dry-mesic to xeric, open areas such as grasslands, sandy fields, woodlands, and forest openings. 1a. Carpellate scales 2–2.5 mm long; perigynia 2.7–3.1 mm long, lacking veins adaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148a. C. muehlenbergii var. enervis Boott 1b. Carpellate scales 2.5–3.6 mm long; perigynia 3–4.2 mm long, with 5–9 veins or veinless adaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148b. C. muehlenbergii var. muehlenbergii Variety muehlenbergii is known from CT, MA, ME, NH, RI, VT. Variety enervis is known from CT, MA, NH, RI, VT.

Cy p e r ac e a e   1 43

149. Carex radiata (Wahlenb.) Small N eastern star sedge. Carex rosea Schkuhr var. radiata (Wahlenb.) Dewey; C. stellulata Goodenough var. radiata Wahlenb.; C. sylvicola J. Webber & P.W. Ball • CT, MA, ME, NH, RI, VT. Mesic to wet-mesic forests, including slopes and seasonally wet areas. Carex radiata has widest leaf blades 1.3–1.9 mm wide, which is intermediate between C. appalachica and C. rosea. In terms of its stem thickness near the base, C. radiata is closer to C. appalachica with a measurement of 0.8–1.5 mm (see identification key for measurements of other species). The name C. rosea has been misapplied to this species for many years. 150. Carex retroflexa Muhl. ex Willd. N reflexed sedge. CT, MA, NH, RI; also reported from ME by Magee and Ahles (1999), but specimens are unknown. Mesic to dry-mesic, deciduous forests, woodlands, clearings, and open areas. The report of this species in ME by Magee and Ahles (1999) is almost certainly erroneous given that they attribute this taxon with southern affinity to Penobcot County, ME (a region of that state with north-temperate to boreal ecosystems). 151. Carex rosea Schkuhr ex Willd. N rosy sedge. Carex convoluta Mackenzie • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic forests and wooded slopes. 152. Carex sparganioides Muhl. ex Willd. N bur-reed sedge. CT, MA, ME, NH, RI, VT. Rich, mesic, often rocky, deciduous forests, shaded ditches. 153. Carex spicata Huds. E prickly sedge. Carex contigua Hoppe • CT, MA, ME, NH, RI, VT. Open, often human-disturbed and/ or early successional habitats such as fields, pastures, and roadsides.

Phyllostachyae 1a. Carpellate scales 1.6–2.2 mm wide, green with a hyaline margin 0.3–0.8 mm wide; uppermost spike with 3–29 staminate flowers; perigynia 1.2–1.7 mm wide; stigmas long, filiform, strongly papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. willdenowii 1b. Carpellate scales (2–) 2.5–6.5 mm wide, green throughout; uppermost spike with 2 or 3 staminate flowers; perigynia 1.9–3.2 mm wide; stigmas short, clavate-thickened, minutely papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. backii 154. Carex backii Boott N Back’s sedge. Carex backii Boott var. subrostrata (Bates) Dorn; C. durifolia Bailey • CT, MA, ME, NH, VT. Woodlands and forests, frequently on slopes or associated with cliff bases, exposed bedrock, and boulders, higher pH substrate to south, more general to north. 155. Carex willdenowii Schkuhr ex Willd.

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Willdenow’s sedge. CT, MA, VT. Woodlands and forests, usually on dry-mesic, rocky or ledgy slopes and ridge lines.

Physoglochin only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. gynocrates 156. Carex gynocrates Wormsk. ex Drej. northern bog sedge. Carex dioica L. ssp. gynocrates (Wormsk. ex Drej.) Hultén; C. dioica L. var. gynocrates (Wormsk. ex Drej.) Ostenf. • ME. Evergreen swamps dominated by Thuja occidentalis, wooded fens.

NC

14 4  MO NOCOTS

Porocystis Reference: Ball (2002d). 1a. Uppermost spike unisexual, entirely staminate . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pallescens 1b. Uppermost spike bisexual, gynecandrous 2a. Perigynia densely pilose; lateral spikes 2–4 mm wide; leaves with ligules longer than wide 3a. Uppermost spike (5–) 11–20 mm long, densely flowered throughout; anthers 0.7–1.3 (–1.6) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. swanii 3b. Uppermost spike 20–35 (–40) mm long, loosely flowered near base; anthers (1–) 1.6–2 (–2.8) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. virescens 2b. Perigynia glabrous or sparsely pilose; lateral spikes (3–) 4–11 mm wide; leaves with ligules shorter than wide to as long as wide (rarely longer than wide) 4a. Carpellate scales awnless or sometimes with a minute mucro; perigynia glabrous, smooth, bluntly triangular in cross-section; anthers 1.3–2.2 mm long . . . . . . C. hirsutella 4b. Carpellate scales with an awn 0.5–2 mm long; perigynia sparsely pubescent, papillose, ± terete in cross-section; anthers 2–3 mm long . . . . . . . . . . . . . . . . . . . . C. bushii 157. Carex bushii Mackenzie

NC

Bush’s sedge. Carex caroliniana Schwein. var. cuspidata (Dewey) Shinners • CT, MA, ME, VT; also reported from ri by Ball (2002d), but specimens are unknown. Mesic to dry-mesic, often sandy, fields, meadows, and open, human-disturbed areas, infrequently of seasonally saturated, lacustrine floodplains. 158. Carex hirsutella Mackenzie N hirsute sedge. Carex complanata Torr. & Hook. var. hirta (Willd.) Gleason; C. hirsuta Willd.; C. triceps, auct. non Michx. • CT, MA, ME, VT. Forests, forest openings, and woodlands, usually on dry-mesic to mesic, circumneutral substrate. 159. Carex pallescens L. E pale sedge. Carex pallescens L. var. neogaea Fern. • CT, MA, ME, NH, RI, VT. Fields, meadows, forest openings and edges, roadsides. 160. Carex swanii (Fern.) Mackenzie N Swan’s sedge. Carex virescens Muhl. ex Willd. var. swanii Fern. • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic forests and woodlands. 161. Carex virescens Muhl. ex Willd. N ribbed sedge. Carex costata Schwein. • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic, deciduous forests.

Racemosae 1a. Lowermost carpellate spikes spreading or drooping on long peduncles 10–40 mm long; perigynia dark brown to dark red-brown at maturity; achene filling basal ½ of perigynium or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. atratiformis 1b. Lowermost carpellate spikes ascending to erect, sessile or on short peduncles less than 10 mm long; perigynia pale green to white-green or pale brown to yellow-brown at maturity; achene nearly filling body of perigynium 2a. Stems aphyllopodic, arising in tufts from long, horizontal rhizomes; perigynia gray-green or white-green, papillose; carpellate scales usually equaling or exceeding the perigynia; carpellate spikes 10–25 mm long, not forming a dense apical cluster . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. buxbaumii

Cy p e r ac e a e   1 45

2b. Stems phyllopodic and cespitose, arising from leafy tufts, without elongate rhizomes; perigynia pale green becoming gold-brown, smooth or inconspicuously papillose; carpellate scales usually shorter than the perigynia; carpellate spikes 5–12 mm long, the upper spikes densely aggregated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. media 162. Carex atratiformis Britt.

NC

scabrous black sedge. Carex atrata L. ssp. atratiformis (Britt.) Kükenth.; C. ovata Rudge • ME, NH, VT. Wet cliffs, river shores, and stream-side outcrops in high-pH bedrock areas, also of stream shores and wet rocks in subalpine communities. 163. Carex buxbaumii Wahlenb. N brown bog sedge. Carex polygama Schkuhr • CT, MA, ME, NH, RI, VT. Fens, wet meadows, river shores, river shore outcrops and seeps. 164. Carex media R. Br.

NC

closed-headed sedge. Carex alpina Liljeblad var. inferalpina Wahlenb.; C. norvegica Retz. var. inferalpina (Wahlenb.) Hultén • ME; single station known in northwestern portion of state. High-pH cliffs in boreal settings.

Rostrales 1a. Widest leaf blades 1.6–3.5 (–4.2) mm wide; upper leaf sheaths and bract sheaths concave at the summit; staminate spikes 6–15 mm long; carpellate scales 33–66% as long as the perigynia, not terminated by an awn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. michauxiana 1b. Widest leaf blades 6–18 (–21) mm wide; upper leaf sheaths and bract sheaths convex at the summit; staminate spikes 12–40 mm long; carpellate scales mostly 66–100% as long as the perigynia (rarely surpassing the perigynia), usually prolonged into an awn . . . . . . C. folliculata 165. Carex folliculata L. N northern long sedge. CT, MA, ME, NH, RI, VT. Swamps, shorelines, marshes. 166. Carex michauxiana Boeckl. N Michaux’s sedge. Carex abacta Bailey • MA, ME, NH, VT. Peatlands, lakeside fens, peaty shores.

Scirpinae only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. scirpoidea 167. Carex scirpoidea Michx. ssp. scirpoidea

NC

Canadian single-spike sedge. Carex scirpiformis Mackenzie • ME, NH, VT. Boreal and subalpine ridges, cliffs, and basins, usually on high-pH substrate.

Squarrosae 1a. Carpellate spikes with widely radiating perigynia, the lower of each spike often reflexed; achenes 1.1–1.4 mm wide, with a sinuous, persistent style; beak of perigynium usually smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. squarrosa 1b. Carpellate spikes with appressed-ascending to ascending perigynia; achenes 1.4–1.7 mm wide, with a partially deciduous style and only the straight basal portion remaining on the achene; beak of perigynium often sparsely scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . C. typhina 168. Carex squarrosa L. N squarrose sedge. CT, MA. Swamps, wetland margins. Though one of the MA population of Carex squarrosa is considered to be introduced through intentional planting (Sorrie and Somers 1999), there are native occurrences from Berkshire and Hampshire Counties.

146  MONOCOTS

169. Carex typhina Michx. N cattail sedge. Carex squarrosa L. var. typhina (Michx.) Nutt. • CT, MA, ME, VT. Forested wetlands, lacustrine and riverine floodplains.

Stellulatae Due to differences in the shape of perigynia within a spike, it is best to examine those near, but not at, the base of the spike. Reference: Reznicek (2002b). 1a. Spikes usually 1 per stem; anthers (2–) 2.2–3.6 mm long; leaf blades involute . . . . . . C. exilis 1b. Spikes 2 or more per stem; anthers 0.6–2.2 (–2.3) mm long; leaf blades flat or folded 2a. Beak of perigynium with smooth margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. seorsa 2b. Beak of perigynium at least sparsely ciliate-serrulate on the margins 3a. Plants subdioecious, the spikes all or nearly all of a single sex on each stem (rarely with entirely carpellate and entirely staminate spikes on the same plant); uppermost spike either entirely staminate or entirely carpellate or mostly carpellate with as many as 3 staminate flowers at the base (i.e., lacking a prolonged, basal staminate portion); anthers (1–) 1.2–2.2 (–2.3) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sterilis 3b. Plants monoecious, at least the uppermost spike clearly bisexual; uppermost spike gynecandrous, never entirely staminate, usually with a prolonged, slender base of 2–21 staminate flowers; anthers 0.6–1.6 (–2) mm long 4a. Widest leaves 2.8–5 mm wide 5a. Perigynia broad-ovate to suborbicular, 1.5–3 mm wide, 1–1.7 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. atlantica 5b. Perigynia ovate to broad-ovate, (1.25–) 1.4–2 (–2.1) mm wide, (1.4–) 1.6–2.5 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. wiegandii 4b. Widest leaves 0.8–2.7 mm wide 6a. Perigynia (2.6–) 2.9–3.6 (–4) mm long, 1.8–3.2 times as long as wide, with a long beak (0.85–) 0.95–2 mm long and (0.4–) 0.45–0.86 times as long as the body of the perigynium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. echinata 6b. Perigynia 1.9–3 (–3.3) mm long, (1.15–) 1.3–2 (–2.2) times as long as wide, with a short beak 0.4–0.95 mm long and 0.18–0.5 (–0.63) times as long as the body of the perigynium 7a. Perigynia convexly tapered in the apical portion, forming shoulders below the beak, veinless or with as many as 6 faint veins adaxially, with many, fine serrations on the beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. interior 7b. Perigynia gradually or concavely tapered in the apical portion (i.e., without shoulders), with 1–10 veins adaxially, with fewer and coarser serrations on the beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. atlantica 170. Carex atlantica Bailey N prickly bog sedge.  170a. Carex incomperta Bickn.; C. stellulata Goodenough var. conferta Chapman; 170b. Carex atlantica Bailey ssp. capillacea (Bailey) Reznicek.; C. howei Mackenzie • CT, MA, ME, NH, RI, VT. Bogs, deciduous and coniferous swamps, wetland margins. 1a. Widest leaf blades 1.6–4 (–4.5) mm wide; perigynia 2.3–3.8 mm long; inflorescences (1.5–) 1.8–5.5 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170a. C. atlantica var. atlantica 1b. Widest leaf blades (0.65–) 0.8–1.6 mm wide; perigynia 1.9–3 mm long; inflorescences 0.8–2 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . 170b. C. atlantica var. capillacea (Bailey) Cronq.

Cy p e r ac e a e   147

Variety atlantica is known from CT, MA, ME, NH, RI, VT. Variety capillacea is known from CT, MA, ME, NH, RI, VT. 171. Carex echinata Murr. var. echinata N star sedge. Carex angustior Mackenzie; C. cephalantha (Bailey) Bickn.; C. josselynii (Fern.) Mackenzie ex Pease; C. muricata L. var. echinata (Carey) Carey ex Gleason • CT, MA, ME, NH, RI, VT. Peatlands, swamps, wet meadows, mucky or peaty shorelines. 172. Carex exilis Dewey N meager sedge. MA, ME, NH, RI, VT. Fens, bogs, hydric meadows, swamps, boggy pond shores. 173. Carex interior Bailey N inland sedge. Carex scirpoides Schkuhr ex Willd. • CT, MA, ME, NH, VT. Fens, swamps, and calcareous seeps. 174. Carex seorsa Howe N weak stellate sedge. CT, MA, NH, RI. Forested swamps, usually on acidic, sandy substrate and under deciduous trees or in the shade of Chamaecyparis. 175. Carex sterilis Willd. NC dioecious sedge. Carex elachycarpa Fern.; C. muricata L. var. sterilis (Fern.) Gleason; Kobresia elachycarpa (Fern.) Fern. CT, MA, ME, VT. Fens, river shore seeps, and wet meadows. 176. Carex wiegandii Mackenzie N Wiegand’s sedge. MA, ME, NH, VT. Bogs, often in shade or partial shade of Picea or Larix, low areas in evergreen forests, and peaty meadows.

Thuringiaca only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. flacca 177. Carex flacca Schreb. E glaucous sedge. Carex glauca Scop. • CT, VT. Rich forests, swamps, hydric meadows, ditches, wetland edges, usually in regions of high-pH bedrock.

Vesicariae Reference: Reznicek and Ford (2002). 1a. Carpellate scales with long, scabrous awns, the awns often equal in length to the body of scale; carpellate spikes ascending to drooping; carpellate scales often ciliate-margined 2a. Stems produced singly or a few together from elongate rhizomes; staminate scales acute to acuminate at the apex (sometimes the lowermost scales with a short awn tip) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. schweinitzii 2b. Stems cespitose, with short rhizomes connecting the individual stems; at least some of the staminate scales with scabrous awns 3a. Perigynia with 5–12 veins that remain separate nearly to the apex of the beak, the bodies broad-ellipsoid to subglobose, (1.8–) 2–4.2 mm wide; achenes papillose 4a. Widest leaves (4–) 4.5–13 mm wide; spikes (12–) 15–22 mm thick; perigynium moderately contracted to a beak 0.7–0.9 (–1) times as long as its body . . . . . C. lurida 4b. Widest leaves 2.4–4 (–5) mm wide; spikes 9–14 (–15) mm thick; perigynium abruptly contracted to a beak 0.7–1.3 times as long as its body . . . . . . . . . . . C. baileyi 3b. Perigynia with 12–15 veins that converge near the base or middle of the beak (except for the 2 lateral ribs, which remain separate), the bodies ellipsoid to narrowovoid, 1–2.2 mm wide; achenes smooth

14 8  MO NOCOTS

5a. Perigynium somewhat inflated, nearly terete to mildly compressed, with herbaceous texture, ascending to spreading, with gaps between many of the veins wider than 3 times the width of the veins; beak of perigynium terminated by 2 straight teeth 0.3–0.9 mm long; basal leaf sheaths usually strongly tinged with anthocyanic pigments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hystericina 5b. Perigynium scarcely inflated, definitely compressed, with coriaceous texture, horizontally spreading to reflexed, with gaps between most veins less than 2 times the width of the veins; beak of perigynium terminated by 2 straight to outwardly arching teeth 0.7–2.1 (–2.8) mm long; basal leaf sheaths usually pale brown, infrequently tinged with faint anthocyanin 6a. Perigynium beak terminated with 2 straight or slightly outcurved teeth 0.7–1.2 (–1.4) mm long; carpellate spikes 9–12 mm thick . . . . . . . C. pseudocyperus 6b. Perigynium beak terminated with 2 strongly outcurved teeth 1.3–2.1 (–2.8) mm long; carpellate spikes 12–18 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. comosa 1b. Carpellate scales awnless or with short, smooth awns shorter than the body of the scale; carpellate spikes erect to ascending; carpellate scales eciliate 7a. Stigmas 2; achenes biconvex; perigynia obscurely veined . . . . . . . . . . . . . . . . C. saxatilis 7b. Stigmas 3, achenes triangular in cross-section; perigynia distinctly veined 8a. Perigynium beak 0.3–0.9 mm long, obscurely bidentate at tip, the terminal teeth up to 0.3 mm long; leaves filiform, with involute margins, 0.5–2.5 mm wide . . . . . C. oligosperma 8b. Perigynium beak (1–) 1.1–4.8 mm long, with 2 distinct apical teeth (0.2–) 0.3–1.9 mm long; leaves ± flat or folded (rarey U-shaped) in cross-section, 1.5–12 (–15) mm wide 9a. Achenes asymmetrical, with an indentation on one of the surfaces; widest perigynia (4–) 4.5–7 mm wide; lowermost bract of carpellate spikes with a blade more than 3 times as long as the inflorescence . . . . . . . . . . . . . . . . . . . . C. tuckermanii 9b. Achenes symmetrical, lacking indentations; widest perigynia (1.5–) 2.5–3.5 (–4.5) mm wide; lowermost bract of carpellate spikes with a blade less than 3 times as long as the inflorescence (except in C. retrorsa, which has an elongate bract blade) 10a. Bract of lowermost carpellate spike with a blade (2.5–) 3–9 times as long as the inflorescence; perigynia horizontally spreading to reflexed; staminate spike often solitary (infrequently 2 or 3), barely elevated above the crowded cluster of carpellate spikes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. retrorsa 10b. Bract of lowermost carpellate spike with a blade up to 2.5 (–3) times as long as the inflorescence; perigynia ascending to spreading; staminate spikes 1–3, usually well elevated above the separate carpellate spikes 11a. Perigynium beak 2.4–4.2 (–4.8) mm long, scabrous in the apical portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bullata 11b. Perigynium beak (1–) 1.2–2.7 mm long, smooth 12a. Plants cespitose, from short rhizomes, with basally firm stems; ligules of leaves longer than wide; leaf sheaths with few cross-septa connecting the longitudinal veins; perigynia ascending, mostly arranged in 6 or 7 vertical ranks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. vesicaria 12b. Plants colonial, from elongate rhizomes, with basally thickened and ± spongy stems; ligules as wide as long; leaf sheaths with many cross-septa connecting the longitudinal veins; perigynia spreading, mostly arranged in 8–12 vertical ranks 13a. Leaf blades U-shaped in cross-section, white-green, 1.5–4.5 mm wide, the adaxial surface minutely papillose . . . . . . . . . . . . . . . . . . . . . . C. rostrata

Cy p e r ac e a e   149

13b. Leaf blades flat to V-shaped in cross-section, pale green to green, (2.5–) 4.5–12 (–15) mm wide, smooth or rarely scabrous (note: the scabrules are easily viewed with low magnification and are more widely spaced than are the minute papillae of C. rostrata) . . . . . . . . . . . . . . . . C. utriculata 178. Carex baileyi Britt. N Bailey’s sedge. CT, MA, ME, NH, VT. Lake shores, stream edges, ditches, meadows, and other low, wet ground. 179. Carex bullata Schkuhr ex Willd. N button sedge. CT, MA, ME, NH, RI. Open fens, bogs, meadows, and lakeshores. ‌179 × 190. Carex ×olneyi Boott is a rare hybrid known from RI. The hybrid can be recognized by its general similarity to Carex bullata, but the perigynia beaks are shorter (1.9–2.6 mm long) and vary in their presence of scabrules (some beak margins are smooth, some have a few scabrules, others have abundant scabrules). There are additional specimens determined to be this hybrid (including collections from CT, MA, and NH). However, those seen by me from CT and NH are true C. bullata, and most of those from MA are also true C. bullata. More study is needed to determine the range of this nothospecies in New England. 180. Carex comosa Boott N bearded sedge. CT, MA, ME, NH, RI, VT. Swamps, stream and lake shores, wet meadows, fens. 181. Carex hystericina Muhl. ex Willd. N porcupine sedge. CT, MA, ME, NH, RI, VT. Lake and stream shores, wet meadows, fens, calcareous seeps, ditches. 181 × 188. This very rare sedge hybrid is known from MA, VT. It generally resembles Carex schweinitzii but has conspicuous anthocyanic coloration on the lower leaf sheaths, shorter rhizomes (these extending as much as 6–8 cm before turning upward to produce a new aerial shoot), perigynia with ca. 10–16 nerves, and staminate scales with some marginal cilia on the body of the scale. In contrast, C. schweinitzii has non-anthocyanic lower sheaths, rhizomes up to ca. 30 cm long (or longer), perigynia with 7–11 nerves, and no marginal cilia on the staminate scales. Carex hystericina has strongly anthocyanic lower sheaths, rhizomes rarely longer than 3 cm, perigynia with 13–21 nerves, and prominently ciliate staminate scales. 182. Carex lurida Wahlenb. N sallow sedge. CT, MA, ME, NH, RI, VT. Wet fields and meadows, shorelines, swamps, marshes, and ditches. 183. Carex oligosperma Michx. N few-seeded sedge. CT, MA, ME, NH, VT. Bogs, acidic fens, and wet meadows. 184. Carex pseudocyperus L. N cypress-like sedge. CT, MA, ME, NH, VT; also reported from RI by Magee and Ahles (1999), but specimens are unknown. Stream shores, lake shores, marshes, openings in swamps, fens. 185. Carex retrorsa Schwein. N retrorse sedge. CT, MA, ME, NH, RI, VT. Stream shores, lake shores, marshes, graminoid meadows, openings in swamps. 186. Carex rostrata Stokes N beaked sedge. ME, NH; northern portion of states. Lake shores, marshes, river shores, river channel sloughs. 187. Carex saxatilis L.

NC

russet sedge. Carex miliaris Michx.; C. rhomalea (Fern.) Mackenzie; C. saxatilis L. var. miliaris (Michx.) Bailey; C. saxatilis L. var. rhomalea Fern. • ME; in vicinity of Katahdin, Piscataquis County. Tarn shores and moist turf on ledges and in gullies.

15 0 MONOCOTS

‌ 187 × 191. Carex ×stenolepis Lessing is a very rare sedge hybrid, known only from tarn shores on the east slope of Katahdin, Piscataquis County, ME. It is recognized by narrow, ± involute leaf blades mostly 2–3 mm wide, ovoid to obloid-ovoid perigynia with a short beak (0.5–1 mm) with very short terminal teeth (ca. 0.25 mm long). It is largely sterile and produces few (if any) mature achenes. 188. Carex schweinitzii Dewey ex Schwein.

NC

Schweinitz’s sedge. CT, MA, VT; western New England. Open wetlands in regions of high-pH bedrock, including fens, graminoid marshes, stream borders, and open swamps. 189. Carex tuckermanii Dewey N Tuckerman’s sedge. CT, MA, ME, NH, VT. Swamps, lake shores, river shores, and vernal pools. 190. Carex utriculata Boott N swollen-beaked sedge. Carex rostrata Stokes var. utriculata (Boott) Bailey • CT, MA, ME, NH, RI, VT. Lakes shores, river shores, marshes, and graminoid fens. 191. Carex vesicaria L. N lesser bladder sedge. Carex monile Tuckerman; C. vesicaria L. var. jejuna Fern.; C. vesicaria L. var. laurentiana Fern.; C. vesicaria L. var. monile (Tuckerman) Fern.; C. vesicaria L. var. raeana (Boott) Fern. • CT, MA, ME, NH, RI, VT. Lake shores, stream shores, marshes, and openings in swamps. North American plants may be distinct from the Eurasian material (as briefly noted by Fernald 1942a). If treated as such, the name Carex monile would be used for our material.

Vulpinae Reference: Standley (2002). 1a. Perigynia rounded and inconspicuously spongy-thickened at the base, adaxially without veins; sheaths red- or brown-dotted near the margins; achenes circular in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. alopecoidea 1b. Perigynia truncate and conspicuously spongy-thickened at the base, with usually 7 veins adaxially; sheaths not dotted; achenes ovate 2a. Leaf sheaths smooth, not corrugated, the summits firm, thickened, yellow, and truncate; leaf blades minutely papillose on the adaxial surface, epistomic; inflorescence 3–6 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. laevivaginata 2b. Leaf sheaths usually corrugated on surface opposite leaf blade (smooth in C. oklahomensis), the summits thin, fragile, colorless, and convex; leaf blades smooth on adaxial surface, not epistomic; inflorescence 5–15 cm long 3a. Lowermost 2 or 3 leaves with highly reduced blades; leaf sheaths smooth, prolonged above the junction of the blade and forming a tube that surrounds the stem; stems firm; ligules obtuse, up to 4 mm long . . . . . . . . . . . . . . . . C. oklahomensis 3b. Lowermost leaves usually with ± well-developed blades; leaf sheaths corrugated, not prolonged above junction of blade; stems weak, easily compressed; ligules acute, up to 10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stipata 192. Carex alopecoidea Tuckerman

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fox-tail sedge. CT, MA, ME, VT. Usually associated with riverine systems, such as riparian forests and stream-side marshes, rarely also in ditches and other low, wet areas associated with highpH bedrock. 193. Carex laevivaginata (Kükenth.) Mackenzie N smooth-sheathed sedge. Carex stipata Muhl. ex Willd. var. laevivaginata Kükenth. • CT, MA, RI, VT; also reported from ME by Magee and Ahles (1999), but specimens are unknown. Marshes, wet meadows, river shores, and low riparian forests.

Cy p e r ac e a e   15 1

194. Carex oklahomensis Mackenzie E Oklahoma sedge. Carex stipata Muhl. ex Willd. var. oklahomensis (Mackenzie) Gleason • ME. Moist roadsides, ditches, marshes, wet meadows.

195. Carex stipata Muhl. ex Willd. var. stipata N awl-fruited sedge. CT, MA, ME, NH, RI, VT. Marshes, ditches, wet meadows, openings in swamps, shorelines.

Cladium 1. Cladium mariscoides (Muhl.) Torr. N smooth saw-sedge. Mariscus mariscoides (Muhl.) Kuntze • CT, MA, ME, NH, RI, VT. Pond shores, frequently in shallow water, rarely in fens or along river shores.

Cyperus Reference: Tucker et al. (2002). 1a. Styles bifid; achenes lenticular in cross-section; edge of achene facing and fitting into a groove in rachilla 2a. Achenes broad-obovate to nearly suborbicular in outline, minutely marked with elongate, superficial cells and irregular, pale transverse lines at maturity; floral scales broad-ovate; flowers with usually 3 stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. flavescens 2b. Achenes narrow-obovate to oblong in outline, unmarked; floral scales narrow-oblong to ovate; flowers with 2 stamens (upper flowers sometimes with 3 stamens) 3a. Floral scales pigmented, in part, with anthocyanins (i.e., red-brown to purple-brown), rarely pale throughout, without a projecting mucro at the apex 4a. Style divided nearly to its base, usually persistent in fruit; anthocyanic pigments most prominent along medial and apical margin of floral scales and along upper edges of midribs, with a conspicuous pale area between the midrib and margin [Fig. 105] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. diandrus 4b. Style connate in the basal ⅓, often deciduous from the fruits; anthocyanic pigments most prominent in basal and medial portion of floral scales, fading toward apex, the pigmented region not outlining a conspicuous pale region on each side of the scale [Fig. 104] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bipartitus 3b. Floral scales lacking anthocyanic pigments, stramineous, gold-brown, to brown (sometimes light chestnut-brown), with a short mucro at the apex 5a. Achenes narrow-obovate in outline; floral scales 2.5–3.5 mm long . . . . C. filicinus 5b. Achenes oblong in outline; floral scales 1.5–2.3 mm long . . . . . . C. polystachyos 1b. Styles trifid; achenes triangular in cross-section; one surface of achene facing and resting flat on rachilla 6a. Plants annual, with soft stem-bases from a fibrous root-system, lacking rhizomes and knotty tubers 7a. Floral scales with 2 keels in the basal 30–60%; flowers with 1 stamen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. acuminatus 7b. Floral scales either with a single keel or merely nerved and lacking keels altogether; flowers with 2 or 3 stamens (only 1 stamen in C. squarrosus) 8a. Floral scales squarrose at apex [Fig. 109]; flowers with 1 stamen; dwarf plants 5–20 cm tall, when fresh with a sweet scent suggesting Melilotus or Ulmus rubra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. squarrosus

15 2 MONOCOTS

8b. Floral scales not outcurved at tip [Fig. 110]; flowers with 2 or 3 stamens; plants predominantly taller, 10–80 cm in height, without a sweet scent 9a. Floral scales 7- to 13-nerved, (1.5–) 2–4.5 (–5) mm long 10a. Floral scales (3–) 3.2–4.5 (–5) mm long; spikelets disarticulating from the base and falling intact; achenes broad-linear to narrow-oblong in outline; stems usually arising from a swollen, corm-like base . . . . . . . . . . . (in part) C. strigosus 10b. Floral scales (1.5–) 2–2.8 (–3.2) mm long; spikelets disarticulating into segments at maturity, each segment consisting of a floral scale and a winged rachilla joint; achenes elliptic to narrow-obovate in outline; stems not arising from a corm-like base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. odoratus 9b. Floral scales 3- to 5-nerved, 0.5–1.7 mm long 11a. Floral scales broad-obovate to suborbicular; achenes 1–1.5 mm long, brown to nearly black at maturity 12a. Spikes ascending to appressed; distance between floral scales on same side of rachis 0.7–0.9 mm; floral scales tipped by a mucro 0.05–0.12 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. iria 12b. Spikes spreading; distance between floral scales on same side of rachis 1.1–1.5 mm; floral scales tipped by a mucro 0.2–0.25 mm long . . . . . C. microiria 11b. Floral scales ovate; achenes 0.7–1 mm long, pale gray-brown to nearly white at maturity 13a. Spikelets radiating from the summit of short axes, forming subcapitate clusters; rachilla lacking a prominent wing-margin; roots brown . . . . C. fuscus 13b. Spikelets serially disposed along elongate axes, forming cylindric clusters; rachilla wing-margined; roots anthocyanic . . . . . . . C. erythrorhizos 6b. Plants perennial, with firm stem-bases arising from rhizomes, knotty tubers, or a swollen corm-like base [Fig. 106] 14a. Spikelets with 3–5 floral scales, crowded into dense, cylindrical to subglobose clusters, mainly only the tips of the spikelets visible in life 15a. Spikelets (3.5–) 4–7 mm long, with membranaceous, yellow-brown to brown floral scales 3.5–4.5 mm long; spikes globose to short-ovoid; achenes 1.5–2.3 mm long; bracts scabridulous along adaxial surface (especially distally); stems growing singly or loosely cespitose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. echinatus 15b. Spikelets 2.2–4 (–4.5) mm long, with coriaceous, brown to yellow-brown or red-brown floral scales 1.8–2.5 mm long; spikes obloid-ovoid; achenes 1.2–1.7 mm long; bracts smooth along adaxial surface, scabridulous only along margins; stems densely cespitose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. retrorsus 14b. Spikelets with 3–20 (–50) floral scales, loosely to moderately crowded, many spikelets visible throughout their length [Fig. 110] 16a. Floral scales with 2 keels in the basal 30–60%; flowers with 1 stamen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pseudovegetus 16b. Floral scales either with a single keel or merely nerved and lacking keels altogether; flowers with 3 stamens 17a. Spikelets flat, serially disposed on elongate axes, forming cylindric clusters [Fig. 110] 18a. Floral scales (3–) 3.5–4.5 (–5) mm long, keeled along their midrib, persistent and falling with the rachis; plants lacking well-developed

Cy p e r ac e a e   15 3

rhizomes, the stems emerging from a tuberous base; achenes broad-linear to narrow-oblong in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. strigosus 18b. Floral scales 2.3–3 mm long, not or scarcely keeled, deciduous from the rachis; plants with long, slender rhizomes ending in tubers; achenes oblong to narrow-obovate in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. esculentus 17b. Spikelets compressed to subterete, radiating from the summit of short axes, forming hemispherical to subglobose clusters [Figs. 107, 108] 19a. Floral scales with 3–5 nerves arranged close to the center of the scale (i.e., the sides largely nerveless); achenes up to 1 mm long; spikelets often proliferated into vegetative propagules . . . . . . . . . . . . . . . . . . . . . . . C. dentatus 19b. Floral scales with 7–13 nerves, these more widely distributed and only the very margin nerveless; achenes 1.4–3 (–3.5) mm long; spikelets not proliferous 20a. Floral scales suborbicular; achenes with 1 or more evidently concave faces, up to 1.5 times as long as wide . . . . . . . . . . . . . . . . . . . . C. houghtonii 20b. Floral scales broad-elliptic to oblong or ovate; achenes with flat or slightly concave faces, 2 or more times as long as wide 21a. Stems often scabrous near apex; floral scales with a short awn 0.5–1 mm long; achenes 2–3 (–3.5) mm long; spikelets ascending [Fig. 108] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. schweinitzii 21b. Stems smooth; floral scales blunt or with a minute mucro up to 0.2 mm long; achenes 1.4–2.2 mm long; basal spikelets of cluster horizontally spreading to reflexed [Fig. 107] 22a. Involucral bracts ascending to sometimes spreadingascending, with smooth to obscurely scabrous margins, numbering 3–7 per cluster; rachilla broadly wing-margined; inflorescence with (2–) 4–10 elongate rays; anthers 0.7–1 mm long . . . . . . . . . . . C. grayi 22b. Involucral bracts mostly spreading to reflexed, with scabrous margins, numbering 2–4 per cluster; rachilla sharp-edged to narrowly winged; inflorescence with a solitary, sessile cluster or also with 1–4 rays [Fig. 107]; anthers 0.3–0.6 mm long . . . . . . C. lupulinus 1. Cyperus acuminatus Torr. & Hook. ex Torr. E taper-tipped flatsedge. NH. Low, wet, often disturbed, sandy soils. 2. Cyperus bipartitus Torr. N Fig. 104 shining flatsedge. Cyperus rivularis Kunth; Pycreus rivularis (Kunth) Palla • CT, MA, ME, NH, RI, VT. Moist to wet, usually sandy, shorelines.

Fig. 104  Spike of Cyperus bipartitus detailing floral scale pigmentation.

3. Cyperus dentatus Torr. N toothed flatsedge. CT, MA, ME, NH, RI, VT. Sandy, gravelly, or peaty shorelines. 4. Cyperus diandrus Torr. N Fig. 105 umbrella flatsedge. Pycreus diander (Torr.) C.B. Clarke • CT, MA, ME, NH, RI, VT. Moist to wet, usually sandy or peaty, shorelines. 5. Cyperus echinatus (L.) Wood E globe flatsedge. Cyperus ovularis (Michx.) Torr. • CT, MA, RI. Open, mesic, often humandisturbed, soils. 6. Cyperus erythrorhizos Muhl. N red-root flatsedge. Cyperus halei Torr. ex Britt. • CT, MA, ME, NH, RI. Mesic to hydric shorelines and wet areas, often on sandy substrate, but occasionally mucky or peaty.

Fig. 105  Spike of Cyperus diandrus detailing floral scale pigmentation.

15 4  MONOCOTS

7. Cyperus esculentus L. var. leptostachyus Boeck. N nut flatsedge. Cyperus esculentus L. var. angustispicatus Britt.; C. tuberosus Pursh • CT, MA, ME, NH, RI, VT. Open, often human-disturbed, soil such as fields, roadsides, and waste areas. 8. Cyperus filicinus Vahl N fern flatsedge. Cyperus polystachyos Rottb. var. filicinus (Vahl) C.B. Clarke • CT, MA, ME, NH, RI. Sandy, Atlantic coast shorelines, brackish marshes, upper edge of saline marshes, interdune wetlands. 9. Cyperus flavescens L. E yellow flatsedge. Cyperus flavescens L. var. poiformis (Pursh) Fern.; Pycreus flavescens (L.) Reichenb. • MA. Wet-mesic, often human-disturbed, soils. 10. Cyperus fuscus L. E brown flatsedge. CT, MA. Shorelines and temporary and/or human-disturbed wet areas. 11. Cyperus grayi Torr. N Fig. 106 Fig. 106  Base of stem of Cyperus grayi.

Gray’s flatsedge. Cyperus filiculmis Vahl. var. oblitus Fern. & Grisc. • CT, MA, NH, RI. Sandy soils near the Atlantic coast, such as dunes, back beaches, and open woodlands. 12. Cyperus houghtonii Torr.

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Houghton’s flatsedge. MA, ME, NH, VT. Dry-mesic to xeric sands and ledges, including roadsides, lake shores, sandplains, and woodlands. 13. Cyperus iria L. E rice-field flatsedge. CT. Wet-mesic to dry-mesic, often human-disturbed, soils, such as fields and waste areas. 14. Cyperus lupulinus (Spreng.) Marcks N Fig. 107 Great Plains flatsedge. 14a. Cyperus filiculmis Vahl; 14b. Cyperus filiculmis Vahl var. macilentus Fern.; C. lupulinus (Spreng.) Marcks ssp. macilentus (Fern.) Marcks • CT, MA, ME, NH, RI, VT. Open, well-drained, usually sandy soils of fields, roadsides, grasslands, and waste places. 1a. Spikelets 8–16 mm long, with 5–22 floral scales that are green-yellow to stramineous and 2.5–4 mm long; anthers 0.6–1 mm long; spikelets 3–4 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14a. C. lupulinus var. lupulinus Fig. 107  Spikelets of Cyperus lupulinus, the lower ones spreading to reflexed.

1b. Spiklets 3–7 mm long, with 3–7 floral scales that are pale green to light red-brown and 1.8–2.5 mm long; anthers 0.3–0.6 mm long; spikelets 2.5–3.5 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14b. C. lupulinus var. macilentus (Fern.) A. Haines Variety lupulinus is known from CT, MA, RI, VT. Variety macilentus is known from CT, MA, ME, NH, RI, VT. ‌14 × 20. Cyperus ×mesochorus Geise is a very rare flatsedge hybrid known from MA. It typically has fewer inflorescence branches than Cyperus schweinitzii (that species usually with 3–5 elongate branches), widely ascending inflorescence bracts (rather than horizontally spreading to reflexed in C. lupulinus and erect to narrowly ascending in C. schweinitzii), and floral scales with an awn tip 0.4–0.5 mm long (rather than 0.05–0.2 mm in C. lupulinus and 0.1–1 mm in C. schweinitzii). 15. Cyperus microiria Steud. E lesser rice-field flatsedge. Cyperus iria L. var. microiria (Steud.) Franch. & Savatier; C. iria L. var. rectangularis Kükenth. • CT, MA. Wet-mesic to dry-mesic, often human disturbed, soils, such as fields and waste areas. The name Cyperus amuricus Maxim. has been misapplied to (i.e., incorrectly used for) this species by some American authors. 16. Cyperus odoratus L.

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rusty flatsedge. Cyperus engelmannii Steud.; C. ferax L.C. Rich; C. ferruginescens Boeckl.; Torulinium odoratum (L.) S. Hooper • CT, MA, NH, RI, VT. Shorelines, tidal marshes, wet sand, muddy depressions, meadows. Immature specimens of Cyperus odoratus can be confused

Cy p e r ac e a e   15 5

with first-year flowering forms of C. strigosus (the latter species is much more common in New England). In addition to scale length, the red-brown spikelets of C. odoratus can be helpful to separate specimens of C. strigosus (which have yellow-brown to brown spikelets). 17. Cyperus polystachyos Rottb. N many-spiked flatsedge. Cyperus polystachyos Rottb. var. texensis (Torr.) Fern.; Pycreus polystachyos (Rottb.) Beauv. • MA, ME, RI. Sandy pond shores, interdunal wetlands, ditches. 18. Cyperus pseudovegetus Steud. E marsh flatsedge. Cyperus virens Michx. var. arenicola (Boeckl.) Shinners • MA. Hydric soils, usually sandy or muddy, and often associated with stream and pond shores. 19. Cyperus retrorsus Chapman

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pine barren flatsedge. Cyperus cylindricus (Ell.) Britt.; Mariscus cylindricus Ell.; M. retrorsus (Chapman) C.B. Clarke • MA; Marthas Vineyard. Sandy soil of woodlands, clearings, and trail edges. 20. Cyperus schweinitzii Torr. N Fig. 108

Fig. 108  Ascending spikelets of Cyperus schweinitzii.

Schweinitz’s flatsedge. Mariscus schweinitzii (Torr.) T. Koyama • MA. Sandy shorelines, sand dunes, open woodlands. 21. Cyperus squarrosus L. N Fig. 109 awned flatsedge. Cyperus aristatus Rottbøll; Cyperus inflexus Muhl. • CT, MA, ME, NH, RI, VT. River and lake shores, usually in sand. 22. Cyperus strigosus L. N Fig. 110 straw-colored flatsedge. Cyperus strigosus L. var. robustior Britt. • CT, MA, ME, NH, RI, VT. Shorelines, river bars, and ditches, often on sandy substrate.

Dulichium 1. Dulichium arundinaceum (L.) Britt. var. arundinaceum N three-way sedge. Cyperus arundinaceus L. • CT, MA, ME, NH, RI, VT. River shores, lake shores, marshes, and other open wetlands. 1 × Rhynchospora capitellata. This very rare intergeneric hybrid is known from only MA within New England. It has been known by the name Cyperus × ‌weatherbianus Fern.; however, this name is incorrect as the plant is not part of the genus Cyperus (see Raymond (1962) for discussion). Further, the name should not have been a genus but rather a nothogenus (formed by combining in a specific manner the generic names of the species involved). The plant is identified by its general resemblance to R. capitellata with elongate, many-flowered (20–40 flowers) spikelets that resemble a species of Cyperus (except that the scales are spirally arranged and are terminated by a short awn). Most flowers are sterile and do not produce mature achenes, but those achenes that develop somewhat show a subulate tubercle at the apex of the achene body. The plant is further characterized by cespitose habit and terminal and axillary glomerules of spikelets (as in Rhynchospora) combined with flowers that lack perianth (as in Dulichium).

Eleocharis In the following key, achene length does not include the apical tubercle, and stem widths are for dried, pressed specimens. Also, the dimensions of tubercles on freshly collected specimens can be very different from those on dried material, particularly during the early stages of fruit maturation. Therefore, fruits should be dry prior to measurement of tubercles. Reference: Smith et al. (2002). 1a. Floral scales longitudinally striate, usually with red-brown to purple spots on the adaxial (i.e., inner) surface; spikes hardly, if at all, thicker than the stem at ⅔ height

Fig. 109  Spikes of Cyperus squarrosus.

Fig. 110  Spikes of Cyperus strigosus.

15 6 MONOCOTS

2a. Reproductive stems triangular in cross-section; rhizomes often producing flaccid, capillary, leaf-like stems; spikes 4- to 8-flowered, 7–20 (–25) mm tall; floral scales with a keeled midrib . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. robbinsii 2b. Reproductive stems terete or quadrangular in cross-section; rhizomes not producing capillary stems; spikes many-flowered, 15–50 (–60) mm tall; floral scales with an obscure central keel 3a. Reproductive stems both terete and septate . . . . . . . . . . . . . . . . . . . . . E. equisetoides 3b. Reproductive stems quadrangular, lacking septa . . . . . . . . . . . . . . . E. quadrangulata 1b. Floral scales without fine longitudinal striae, lacking spots on the adaxial surface; spikes noticeably thicker than upper portion of stem 4a. Tubercle not obviously differentiated from and nearly confluent with the summit of the achene, forming a stylar beak [Fig. 117] 5a. Stems 1–2 mm thick at midpoint, (15–) 40–100 cm tall, often some arching and rooting at the tip; spikes mostly 10- to 20-flowered; caudex short and thick, the plants lacking rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. rostellata 5b. Stems up to 1 mm thick at midpoint, 3–40 cm tall, none arching and tip-rooting; spikes mostly 2- to 9-flowered; rhizomes present, long and slender, often terminated by a thickened bud 6a. Achenes 1.9–2.6 mm long; floral scales 2.5–5 mm long; plants primarily of high-pH wetlands and shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. quinqueflora 6b. Achenes 0.9–1.3 mm long; floral scales 1.5–2 (–2.5) mm long; plants primarily of brackish water flats and shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. parvula 4b. Tubercle obviously differentiated from the achene, set apart by differences in color and/or texture [Figs. 111, 112, 118] 7a. Achenes gray at maturity, longitudinally striate, with numerous, fine horizontal cross-lines, ca. 2 times as tall as wide [Fig. 111]; floral scales all bearing flowers, arranged in 2 or 3 vertical ranks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. acicularis 7b. Achenes not combining the above characteristics (i.e., lacking conspicuous longitudinal striae, usually no more than 1.5 times as tall as wide); floral scales not all subtending flowers, usually the basal 1 (or more) empty, arranged in more than 3 vertical ranks 8a. Achenes planoconvex to biconvex in cross-section; styles bifid (trifid in E. obtusa) 9a. Base of tubercles with a slight to prominent constriction at the junction of achene [Fig. 116]; plants perennial, with rhizomes (these often difficult to detect in E. flavescens) 10a. Stems cespitose, with slender, often inconspicuous, rhizomes; leaf sheaths prolonged at summit into a white, scarious tip; anthers 0.7–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. flavescens 10b. Stems occurring singly or a few together, produced along obvious, firm rhizomes; leaf sheaths firm at summit; anthers 1–3 mm long 11a. Glumes 2 (rarely 3), lanceolate to triangular, each sheathing ± ½ (up to ⅔) of the base of the spike . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. palustris 11b. Glume solitary, suborbicular, nearly encircling the base of the spike and sheathing ¾ or more of it 12a. Achenes minutely but visibly and regularly patterened with cellular reticulations [Fig. 113]; tubercles 0.15–0.3 (–0.5) mm tall; leaf sheaths firm, many with a thickened, coriaceous collar at the summit . . . . E. ambigens

Cy p e r ac e a e   157

12b. Achenes obscurely patterened, nearly smooth [Fig. 116]; tubercles 0.35–0.8 mm tall; leaf sheaths herbaceous at summit, not or only slightly thickened 13a. Floral scales subcoriaceous, lustrous, the lower and middle ones of the spikelet 3–5 mm long; spikes with (5–) 10–20 (–30) flowers; plants of tidal and/or saline-influenced shores . . . . . . . . . E. uniglumis 13b. Floral scales membranaceous, not or scarcely lustrous, the lower and middle ones of the spikelet 1.8–3 mm long; spikes with (15–) 20– 40 (–50) flowers; plants primarily of non-tidal wetlands and shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. erythropoda 9b. Base of tubercle not constricted at junction of achene; plants annual, with fibrous roots [Fig. 112] 14a. Perianth bristles numbering 5–7 per achene, retrorsely barbellate, exceeding the length of the achene body (and often the tubercle as well) 15a. Tubercles 0.5–0.9 mm wide, 0.65–0.97 times as wide as the achene; flowers usually with 3 stamens 16a. Styles usually trifid; tubercles 0.2–0.4 mm tall, 0.29–0.67 times as tall as wide; perianth bristles usually exceeding tubercle in length; spikes commonly ovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) E. obtusa 16b. Styles bifid or sometimes trifid; tubercles 0.12–0.23 mm tall, 0.1–0.38 times as tall as wide; perianth bristles often not exceeding the tubercle; spikes commonly ellipsoid to ellipsoid-cylindric . (in part) E. engelmannii 15b. Tubercles 0.35–0.5 mm wide, 0.46–0.72 times as wide as the achene; flowers usually with 2 stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. ovata 14b. Perianth bristles lacking or, if present, numbering 2–4, smooth, and shorter than the achene body [Fig. 112] 17a. Tubercles 0.46–0.83 mm wide, 0.65–0.97 times the width of the achene; flowers with usually 3 stamens 18a. Tubercles 0.12–0.23 mm tall, 0.1–0.38 times as tall as wide; styles bifid or sometimes trifid; plants occurring on lake borders and in pools and seepages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) E. engelmannii 18b. Tubercles 0.2–0.4 mm tall, 0.29–0.67 times as tall as wide; styles trifid or sometimes bifid; plants occurring on river shores and lake borders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) E. obtusa 17b. Tubercles 0.24–0.45 mm wide, 0.46–0.68 times the width of the achene; flowers with 2 stamens 19a. Floral scales acute and somewhat keeled at the apex; tubercles mostly 0.1–0.2 mm tall; perianth bristles absent or up to 0.15 mm long; plants occurring on sand shores that are not or only minimally tidal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. diandra 19b. Floral scales rounded and not keeled at the apex; tubercles mostly 0.2–0.3 mm tall [Fig. 112]; perianth bristles absent or up to 0.45 mm long; plants occurring on sand, silt, or mud shores that are usually fresh-tidal (rarely on shores of impounded lakes). . . . . . . . . . . . . . . . . . . . . . E. aestuum 8b. Achenes bluntly triangular to terete in cross-section; styles trifid 20a. Stems cespitose, arising from fibrous roots or a caudex (i.e., rhizomes lacking); plants annual (perennial in E. melanocarpa) 21a. Tubercle as wide as the achene; stems compressed

15 8 MO NOCOTS

22a. Achenes dark brown to nearly black at maturity, smooth on the surface, with a low tubercle much shorter than the achene body [Fig. 115]; perianth bristles shorter than the achene; plants perennial from a stout caudex; stems infrequently arching and rooting or proliferating from tip . . . . E. melanocarpa 22b. Achenes brown to brown-green at matuity, with regular, conspicuous excavations on the surface (i.e., resembling honeycomb), with a large tubercle ± as tall as the achene body; perianth bristles longer than the achene; plants annual from fibrous roots; stems upright, not tip rooting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. tuberculosa 21b. Tubercle clearly narrower than achene; stems terete to capillary, but not obviously compressed 23a. Achenes green to brown-green, 1.2–1.5 mm long, topped with an elongate, cone-shaped tubercle; floral scales 1.7–2.5 mm long; longer perianth bristles exceeding combined length of achene and tubercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. intermedia 23b. Achenes pale, nearly white, 0.5–1 mm long, topped by a minute, depressed-conic tubercle; floral scales 1–1.5 mm long; longer perianth bristles up to as long as the combined length of the achene and tubercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. microcarpa 20b. Stems produced singly or a few together along an obvious rhizome; plants perennial 24a. Achenes with prominent keel-like angles; rhizomes firm, 3–6 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. tricostata 24b. Achenes rounded on the angles, without keels; rhizomes flexible, 0.5–3 mm thick 25a. Stems conspicuously compressed, 2–5 times as wide as thick, 0.5–1.8 mm wide; apical portion of floral scales in the basal portion of the spike often bifid, with a hyaline, colorless region 0.6–1.2 mm long that is longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. compressa 25b. Stems angled to somewhat compressed, 1–2 (–3) times as wide as thick, 0.15–0.8 mm wide; apical portion of floral scales in the basal portion of the spike entire to emarginate, with a hyaline, colorless region that is up to 0.7 mm long and is wider than long to as long as wide 26a. Summit of leaf sheath lacking a tooth-like lobe; achenes 0.6–0.7 × 0.45–0.55 mm; anthers 0.3–0.7 mm long; floral scales 1–1.4 mm long; rhizomes 0.5–1 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. nitida 26b. Summit of leaf sheath usually with a tooth-like lobe; achenes 0.7–1.2 × 0.6–0.9 mm; anthers 1.5–2 mm long; floral scales 1.5–3.5 mm long; rhizomes 1–3 mm thick 27a. Achenes 1.1–1.5 × 0.95–1.25 mm, finely patterned with cellular reticulations; stems 0.5–1.5 mm thick at the midpoint . . . . . . . . E. fallax 27b. Achenes 0.6–1.2 × 0.45–0.9 mm, regularly patterned with conspicuous excavations (i.e., resembling honeycomb); stems 0.2–0.8 mm thick at the midpoint 28a. Stems usually 6- to 8-angled; achenes yellow to orangeyellow, somewhat persistent after the falling of the floral scales, with 12–20 horizontal ridges through the length of the achene [Fig. 114] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. elliptica 28b. Stems 4- or 5-angled; achenes green to brown-green (rarely yellow), deciduous with the falling scales, with 6–10 (–14) horizontal ridges through the length of the achene . . . . . . . . . . . . . . . . . E. tenuis

Cy p e r ac e a e   15 9

1. Eleocharis acicularis (L.) Roemer & J.A. Schultes N Fig. 111 needle spikesedge. Eleocharis acicularis (L.) Roemer & J.A. Schultes var. submersa (H.J. Nilss.) Svens. • CT, MA, ME, NH, RI, VT. Stream shores, lake shores, meadows, and open places with wet, bare soil. 2. Eleocharis aestuum A. Haines

N C Fig. 112

tidal spikesedge. Eleocharis ovata (Roth) Roemer & J.A. Schultes var. aphanactis Moore; E. pallidostachys D.M. Hines, nomen nudum • CT, ME, VT. Tidal river shores on sand, silt, and mud substrate, rarely on inland lakes with altered hydrology. Previous reports of Eleocharis diandra from ME were based on this recently described species. 3. Eleocharis ambigens Fern.

N C Fig. 113

Fig. 111  Achene of Eleocharis acicularis.

ambiguous spikesedge. MA, RI. Coastal plain ponds and marshes, ranging in salinity from fresh to brackish. Eleocharis ambigens is often confused with E. fallax. It has bifid stigmas and achenes with planoconvex cross-section that are capped by a depressed tubercle 0.15–0.3 (–0.5) mm tall that is wider than tall (rarely the tubercles are more elongate and as wide as tall, such as with the collections from Dukes County, MA). Eleocharis fallax shows a high proportion of trifid stigmas and has achenes with planoconvex to trigonous crosssection that are capped by a tubercle 0.3–0.5 mm tall that is ± as wide as tall. 4. Eleocharis compressa Sullivant var. compressa

NC

flat-stemmed spikesedge. Eleocharis elliptica Kunth var. compressa (Sullivant) Drapalik & Mohlenbrock • VT. Seasonally wet soils of lake shores and limestone headlands. 5. Eleocharis diandra C. Wright

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Wright’s spikesedge. Eleocharis diandra C. Wright var. depressa Fern. • CT, MA, ME, NH, VT. Wet sand of major rivers and, rarely, lake shores. In New England known only from the Androscoggin, Connecticut, and Merrimack Rivers and Lake Champlain. This species has been frequently reported from fresh-tidal river shores (e.g., Seymour 1982, Gleason and Cronquist 1991). However, those reports are based on Eleocharis aestuum. Eleocharis diandra occurs in non-tidal situations, with the exception of its downstream populations on the Connecticut River in Hartford County, CT, where the river is minimally tidal.

Fig. 112  Achene of Eleocharis aestuum.

6. Eleocharis elliptica Kunth var. elliptica N Fig. 114 elliptic spikesedge. Eleocharis capitata (L.) R. Br. var. borealis Svens.; E. tenuis (Willd.) J.A. Schultes var. borealis (Svens.) Gleason • CT, MA, ME, NH, RI, VT. Shorelines, meadows, open edges of wetlands, and temporary wet areas. 7. Eleocharis engelmannii Steud. N Engelmann’s spikesedge.  7a. Eleocharis monticola Fern. var. leviseta Fern.; E. obtusa (Willd.) J.A. Schultes var. detonsa (Gray) Drapalick & Mohlenbrock; 7b. Eleocharis monticola Fern.; E. obtusa (Willd.) J.A. Schultes var. engelmannii (Steud.) Britt. • CT, MA, ME, RI; also reported from NH by Smith et al. (2002), but specimens are unknown. Pond shores, temporary pools, and open, disturbed, hydric soils.

Fig. 113  Achene of Eleocharis ambigens.

1a. Perianth bristles lacking or, if present, numbering 2–4, smooth, and shorter than the length of the achene body . . . . . . . . . . . . . 7a. E. engelmannii var. detonsa Gray in H. Patters. 1b. Perianth bristles numbering 5–7 per achene, retrorse-barbellate, exceeding the length of the achene body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7b. E. engelmannii var. engelmannii Variety engelmannii is known from CT, MA, ME, RI. Variety detonsa is known from MA, ME. 8. Eleocharis equisetoides (Ell.) Torr.

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horsetail spikesedge. Scirpus equisetoides Ell. • CT, MA, RI. Lake shores, often growing in standing water. 9. Eleocharis erythropoda Steud. N red-footed spikesedge. Eleocharis calva Torr. • CT, MA, ME, NH, VT; also reported from RI by Smith et al. (2002), but specimens are unknown. Lake shores, marshes, lakeside fens, river shores.

Fig. 114  Achene of Eleocharis elliptica.

16 0 MO NOCOTS

10. Eleocharis fallax Weatherby NC creeping spikesedge. MA. Pond shores on the coastal plain. See comments under Eleocharis ambigens concerning identification. 11. Eleocharis flavescens (Poir.) Urban var. olivacea (Torr.) Gleason N yellow spikesedge. Eleocharis olivacea Torr. • CT, MA, ME, NH, RI, VT. Shorelines, marshes, tidal river shores, peatlands. 12. Eleocharis intermedia J.A. Schultes N mudflat spikesedge. CT, MA, ME, NH, VT. Sandy, muddy, or boggy margins of streams and ponds, often associated with areas of high-pH bedrock or till, also frequent in areas of disturbance (e.g., accretion bars along rivers). 13. Eleocharis melanocarpa Torr. N Fig. 115 black-fruited spikesedge. MA, RI. Sandy or peaty shores of ponds and low areas along the coastal plain. 14. Eleocharis microcarpa Torr. var. filiculmis Torr. small-fruited spikesedge. CT, MA. Sandy pond shores and graminoid marshes. Fig. 115  Achene of Eleocharis melanocarpa.

NC

15. Eleocharis nitida Fern. NC quill spikesedge. ME, NH, VT. Boggy pastures, wet meadows, low areas in trails and fields. 16. Eleocharis obtusa (Willd.) J.A. Schultes NC blunt spikesedge. 16a. Eleocharis obtusa (Willd.) J.A. Schultes var. jejuna Fern.; E. ovata (Roth) Roemer & J.A. Schultes var. obtusa (Willd.) Kükenth.; Scirpus obtusus Willd. • CT, MA, ME, NH, RI, VT. Low, wet areas, pond shores, river shores (these rarely tidal), meadows, disturbed wetlands. 1a. Perianth bristles numbering 5–7 per achene, retrorsely barbellate, exceeding the combined length of the achene body and tubercle . . . . . . . . . . . . . . 16a. E. obtusa var. obtusa 1b. Perianth bristles lacking or, if present, numbering 2–4, smooth, and shorter than the achene body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16b. E. obtusa var. peasei Svens. Variety obtusa is known from CT, MA, ME, NH, RI, VT. Variety peasei is known from ME, NH and is of conservation concern. It is usually found on sandy pond shores and tidal river flats (i.e., places where the water level might fluctuate tremendously daily or seasonally). 17. Eleocharis ovata (Roth) Roemer & J.A. Schultes N ovoid spikesedge. Eleocharis obtusa (Willd.) J.A. Schultes var. heuseri Uechtr.; E. obtusa (Willd.) J.A. Schultes var. ovata (Roth) Drapalik & Mohlenbrock; Scirpus ovatus Roth • CT, MA, ME, NH, RI, VT. Pond shores, river shores (these rarely tidal), meadows, often in watercourses with elevated pH. This species is rare over most of New England but is more frequent in northern ME and NH.

Fig. 116  Achene of Eleocharis palustris.

18. Eleocharis palustris (L.) Roemer & J.A. Schultes N Fig. 116 common spikesedge.  18a. Eleocharis smallii Britt.; Scirpus palustris L.; 18b. Eleocharis palustris (L.) Roemer & J.A. Schultes var. major Sonder; E. palustris (L.) Roemer & J.A. Schultes var. vigens L.H. Bailey • CT, MA, ME, NH, RI, VT. Wet soil or shallow water of lakes, rivers, peatlands, and wetland margins. Various names have been provided for the morphological variation displayed by Eleocharis palustris. Some of the variation is continuous and difficult to interpret. However, present evidence does support recognition of two taxa—a variable diploid (ssp. palustris) and a probable tetraploid (ssp. vigens). 1a. Achene body 1.1–1.5 (–1.6) mm tall; floral scales 3–4 mm long; anthers 1–2 mm long; stem stomates mostly 39–48 µ m long . . . . . . . . . . . . . . . . . . . . . . . . . . 18a. E. palustris ssp. palustris 1b. Achene body 1.6–2 mm tall; floral scales 3.5–4.5 mm long; anthers 1.7–3 mm long; stem stomates 52–65 µ m long . . . . . . . . . . . . . . . 18b. E. palustris ssp. vigens (L.H. Bailey) A. Haines Subspecies palustris is known from CT, MA, ME, NH, RI, VT. Subspecies vigens is known from CT, MA, ME, NH, VT. 19. Eleocharis parvula (Roemer & J.A. Schultes) Link ex Bluff, Nees, & Schauer N little-headed spikesedge. Eleocharis parvula (Roemer & J.A. Schultes) Link ex Bluff, Nees, & Schauer var. anachaeta (Britt.) Svens. • CT, MA, ME, NH, RI. Saline to brackish shorelines.

Cy p e r ac e a e   16 1

20. Eleocharis quadrangulata (Michx.) Roemer & J.A. Schultes

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square-stemmed spikesedge. Eleocharis quadrangulata (Michx.) Roemer & J.A. Schultes var. crassior Fern.; Scirpus quadrangulatus Michx. • CT, MA. Shallow water of lakes and slowmoving rivers. 21. Eleocharis quinqueflora (F.X. Hartmann) Schwarz ssp. fernaldii (Svens.) Hultén



N C Fig. 117

few-flowered spikesedge. Eleocharis pauciflora (Lightf.) Link; E. pauciflora (Lightf.) Link var. fernaldii Svens.; Scirpus quinquefolius F.X. Hartmann • MA, ME, NH, VT. Fens, river shore ledges and seeps, wet cliffs, and peaty lake shores in regions of high-pH bedrock or till. See Svenson (1934) for rationale of recognizing ssp. fernaldii apart from typical ssp. quinqueflora of Europe. 22. Eleocharis robbinsii Oakes N Fig. 118 Robbins’ spikesedge. CT, MA, ME, NH, RI, VT. Lakes and slow moving rivers, often with sand or peat-covered sand substrate.

Fig. 117  Achene of Eleocharis quinqueflora.

23. Eleocharis rostellata (Torr.) Torr. N beaked spikesedge. Scirpus rostellatus Torr. • CT, MA, ME, RI; disjunct from the southern New England locations to Sagadahoc County, ME. Brackish marshes. 24. Eleocharis tenuis (Willd.) J.A. Schultes N slender spikesedge.  24a. Eleocharis capitata (L.) Br. var. pseudoptera Weatherby ex Svens.; E. elliptica Kunth var. pseudoptera (Weatherby ex Svens.) L. Harms; 24b. Eleocharis capitata, sensu Blake • CT, MA, ME, NH, RI, VT. Lake shores, stream shores, ditches, meadows, wet fields. 1a. Achenes yellow, with 10–14 horizontal ridges through the length; stems sharply 4-angled, usually deeply sulcate, up to 0.8 mm wide; some or all of the upper leaf sheaths with a stout apical tooth 0.4–0.6 (–0.9) mm tall; tubercles very depressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24a. E. tenuis var. pseudoptera (Weatherby ex Svens.) Svens. 1b. Achenes green to brown-green, with 6–12 horizontal ridges through the length; stems bluntly and shallowly angled, up to 0.5 mm wide; upper leaf sheaths with a slender apical tooth to 0.2 mm tall; tuberlces often as tall as wide (rarely very depressed) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24b. E. tenuis var. tenuis Variety tenuis is known from CT, MA, ME, NH, RI, VT. Variety pseudoptera is known from CT, MA, ME, VT. It is less common than var. tenuis and transitional, in some respects, to Eleocharis elliptica. 25. Eleocharis tricostata Torr.

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three-angled spikesedge. MA, RI. Sandy or peaty pond shores of the coastal plain. 26. Eleocharis tuberculosa (Michx.) Roemer & J.A. Schultes N long-tubercled spikesedge. Scirpus tuberculosa Michx. • CT, MA, ME, NH, RI. Pond shores, meadows, organic soil wetlands. 27. Eleocharis uniglumis (Link) J.A. Schultes N one-glumed spikesedge. Eleocharis halophila (Fern. & Brack.) Fern. & Brack.; E. uniglumis (Link) J.A. Schultes var. halophila Fern. & Brack.; Scirpus uniglumis Link • CT, MA, ME, NH, RI. Tidal river shores, brackish marshes.

Eriophorum Reference: Ball and Wujek (2002). 1a. Spikelets solitary, erect, with 10–15 sterile scales in the basal portion, not subtended by foliaceous involucral bracts [Fig. 120]; plants cespitose . . . . . . . . . . . . . . . . . . . . . . . E. vaginatum 1b. Spikelets 2–many, spreading or drooping on peduncles; scales all fertile; foliaceous involucral bracts present; stems produced singly or a few together from a rhizome

Fig. 118  Achene of Eleocharis robbinsii.

162 MONOCOTS

2a. Involucral bract 1 per inflorescence (sometimes with another very reduced, scale-like bract), erect; leaf blades 1–2 (–3) mm wide, channeled 3a. Uppermost leaf blade 3–25 cm long, as long or longer than its sheath; stems scabrous near the top; peduncles scabrous; involucral bract (8–) 15–60 mm long, green to red-brown at the base; floral scales colored or at least tinged with red-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. tenellum 3b. Uppermost leaf blade 1–4 cm long, shorter than its sheath; stems smooth; peduncles pubescent but not scabrous; involucral bract 6–20 mm long, gray to black at the base; floral scales black or dark gray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. gracile 2b. Involucral bracts (1–) 2–5 per inflorescence, somewhat spreading; leaves (1.5–) 2–6 (–8) mm wide, flat, at least in the basal half 4a. Scales with 3–7 veins, red-brown near the margin; flowers with 1 stamen; perianth bristles at least tinged with red-brown (rarely entirely white) . . . . . . . . . . . . . E. virginicum 4b. Scales with 1 central vein (i.e., the midrib; occasionally with inconspicuous, short lateral veins in E. viridicarinatum), green or gray to black near the margin; flowers with 3 stamens; perianth bristles white 5a. Midrib of scales inconspicuous near the tip; summit of the sheath darkened [Fig. 119]; anthers 2–5 mm long; peduncles smooth or scabrous . . . . . . E. angustifolium 5b. Midrib of scales prominent to the tip and apically dilated; summit of the sheath not darkened; anthers 0.8–2 mm long; peduncles scabrous . . . . . . E. viridicarinatum Fig. 119  Leaf of Eriophorum angustifolium showing dark summit of sheath.

1. Eriophorum angustifolium Honckeny ssp. angustifolium N Fig. 119 tall cottonsedge. ME, NH; also reported from VT by Hultén and Fries (1986), but specimens are unknown. Peatlands, shorelines, peaty soils around temporary pools. 2. Eriophorum gracile W.D.J. Koch N slender cottonsedge. CT, MA, ME, NH, RI, VT. Bogs, acidic fens, meadows. 3. Eriophorum tenellum Nutt. N few-nerved cottonsedge. CT, MA, ME, NH, RI, VT. Bogs, fens, meadows. 4. Eriophorum vaginatum L. ssp. spissum (Fern.) Hultén N Fig. 120 tussock cottonsedge. Eriophorum spissum Fern.; E. vaginatum L. var. spissum (Fern.) Boivin • CT, MA, ME, NH, RI, VT. Bogs, meadows, alpine tundra.

5. Eriophorum virginicum L. N tawny cottonsedge. CT, MA, ME, NH, RI, VT. Bogs, acidic fens, meadows. 6. Eriophorum viridicarinatum (Engelm.) Fern. N Fig. 120  Solitary spike of Eriophorum vaginatum.

green-keeled cottonsedge. Eriophorum latifolium Hoppe var. viridicarinatum Engelm. • CT, MA, ME, NH, RI, VT. Fens and high-pH meadows.

Fimbristylis 1. Fimbristylis autumnalis (L.) Roemer & J.A. Schultes N slender fimbry. Fimbristylis autumnalis (L.) Roemer & J.A. Schultes var. mucronulata (Michx.) Fern.; Scirpus autumnalis L. • CT, MA, ME, NH, RI, VT. Sandy or peaty pond shores, pools, and temporary wet areas, sometimes occurring in human-disturbed habitats.

Fuirena Underground organs are very important for identification of Fuirena and should be included on herbarium specimens. Reference: Kral (2002a).

Cy p e r ac e a e   163

1a. Plants annual from fibrous roots; anthers 0.5–0.7 mm long; flowers with 1–3 stamens; leaf sheaths subglabrous to hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. pumila 1b. Plants perennial from short rhizomes and corm-like offshoots; anthers 0.7–1 mm long; flowers with 3 stamens; leaf sheaths hispid-hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . F. squarrosa 1. Fuirena pumila (Torr.) Spreng. N Fig. 121 dwarf umbrella-sedge. Fuirena squarrosa Michx. var. pumila Torr. • CT, MA, RI. Sandy or peaty pond shores and low areas on the coastal plain. 2. Fuirena squarrosa Michx.

NC

hairy umbrella-sedge. Fuirena squarrosa Michx. var. hispida (Ell.) Chapman • MA; Cape Cod region; also reported from RI by Kral (2002a), but specimens are unknown. Peaty pond shores and other low, wet areas of the coastal plain. Known from a single collection in New England. The name Fuirena squarrosa has been misapplied to collections of F. pumila.

Kyllinga 1. Kyllinga gracillima Miq. E pasture greenhead-sedge. Cyperus brevifolioides Thieret & Delahoussaye • CT. Stream shores, tidal river shores and coves, usually on muddy substrate.

Lipocarpha 1. Lipocarpha micrantha (Vahl) G. Tucker N small-flowered dwarf-bulrush. Hemicarpha micrantha (Vahl) Pax; Scirpus micranthus Vahl • CT, MA, ME, NH, RI. Sandy shorelines, usually of lakes and ponds, rarey on fresh-tidal river shores.

Rhynchospora A single example of an intergeneric hybrid with Dulichium is known from New England and described under D. arundinaceum. Reference: Kral (2002b). 1a. Achenes 15–25 mm long including the 10–20 (–21) mm long tubercles; leaf blades 3–10 (–15) mm wide; style undivided or bifid only at tip 2a. Stems not cespitose, produced from slender rhizomes, achene body 4–5 mm long; achene tubercle 10–15 mm long; spikes solitary or in clusters of 2–6 . . . . . . . . . R. inundata 2b. Stems cespitose, without rhizomes; achene body 5–6 mm long; achene tubercle (15–) 18–20 (–21) mm long; spikes mostly in dense clusters of 10–30 . . . . . R. macrostachya 1b. Achenes 1–3 mm long including the 0.1–1.7 mm long tubercle; leaf blades 0.2–3 (–5) mm wide; style deeply bifid 3a. Plants annual; perianth bristles absent [Fig. 124]; inflorescences open, diffuse, cyme-like or panicle-like; spikes brown to nearly black, stalked 4a. Tubercle depressed-triangular, 0.1–0.3 mm tall; achene body distinctly but irregularly transverse-rugose, with the marginal edges interrupted at the tubercle base . . . . . R. nitens 4b. Tubercle narrow-triangular, 0.5–0.9 mm tall; achene body obscurely transverserugose, with the marginal edges flowing into the tubercle base . . . . . . . . . . R. scirpoides 3b. Plants perennial; perianth bristles present (though sometimes caducous in R. torreyana) [Figs. 122, 123]; inflorescences of remote to congested clusters of spikes; spikes white to pale brown or red-brown to deep brown, usually clustered 5a. Perianth bristles retrorsely barbellate (smooth in rare forms of R. capillacea and R. capitellata, antrorsely barbellate in rare forms of R. capitellata) [Fig. 122]; base of achene narrowed to a stipe-like process

Fig. 121  Achene of Fuirena pumila.

16 4  MONOCOTS

6a. Flowers with 10–12 perianth bristles, these often antrorsely setose near base; floral scales white to light yellow-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. alba 6b. Flowers with 6 perianth bristles, without antrorsely oriented hairs at the base [Fig. 122]; floral scales pale brown to red-brown or deep brown 7a. Leaf blades filiform, 0.2–0.4 mm wide; spikes all ascending, in ovoid to ellipsoid clusters; achene body elliptic in outline, 1.5–2 mm long . . . . . . . . . . . R. capillacea 7b. Leaf blades ± flat, 1.5–3 mm wide; basal spikes of each cluster spreading, the spikes in turbinate to hemispherical clusters; achene body obovate in outline [Fig. 122], 1.2–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. capitellata 5b. Perianth bristles antrorsely barbellate (sometimes caducous, and therefore absent in fruit, in R. torreyana) [Fig. 123]; base of achene cuneate, not narrowed to a stipe-like process 8a. Perianth bristles firmly attached to achene, reaching to base of tubercle or longer [Fig. 124]; tubercle (0.7–­) 1–1.3 (–1.5) mm long; achene bodies very finely transversely rugose (i.e., pattern not evident without high magnification) . . . . . . . . . . . . . . . R. fusca

Fig. 122  Achene of Rhynchospora capitellata.

8b. Perianth bristles weakly attached to achene, sometimes caducous, not extending more than ⅔ the length of achene body; tubercle 0.2–0.3 (–0.5) mm long; achene bodies finely transversely rugose with irregular horizontal ridges separated by intervals of narrow, vertical alveolae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. torreyana 1. Rhynchospora alba (L.) Vahl N white beaksedge. CT, MA, ME, NH, RI, VT. Wet, organic soils of bogs and acidic fens. 2. Rhynchospora capillacea Torr.

NC

needle beaksedge. Rhynchospora capillacea Torr. var. leviseta E.J. Hill • CT, MA, ME, NH, VT; also reported from RI by Kral (2002b), but specimens are unknown. Circumneutral fens, fen-like meadows, river shore seeps on high-pH bedrock. 3. Rhynchospora capitellata (Michx.) Vahl N Fig. 122 brownish beaksedge. Rhynchospora glomerata (L.) Vahl var. capitellata (Michx.) Kükenth. • CT, MA, ME, NH, RI, VT. Organic soil wetlands, meadows, peaty pond shores, seasonally wet

borrow pits. 4. Rhynchospora fusca (L.) Ait. f. N Fig. 123 brown beaksedge. Rhynchospora alba (L.) Vahl var. fusca (L.) Vahl; Schoenus fuscus L. • CT, MA, ME, NH, RI, VT. Peatlands and peaty pond shores. Fig. 123  Achene of Rhynchospora fusca.

5. Rhynchospora inundata (Oakes) Fern.

NC

narrow-fruited beaksedge. Ceratoschoenus macrostachys (Torr. ex Gray) Gray var. inundatus Oakes; Rhynchospora macrostachya Torr. ex Gray var. inundata (Oakes) Fern. • MA, RI. Coastal plain pond shores. 6. Rhynchospora macrostachya Torr. ex Gray N tall beaksedge. CT, MA, ME, RI. Sandy or peaty pond shores, primarily along the coastal plain. 7. Rhynchospora nitens (Vahl) Gray

N C Fig. 124

short-beaked beaksedge. Psilocarya nitens (Vahl) Wood • MA. Coastal plain pond shores. 8. Rhynchospora scirpoides (Torr.) Gray N long-beaked beaksedge. Psilocarya scirpoides Torr. • CT, MA, RI. Sandy or peaty pond shores, primarily of the coastal plain. 9. Rhynchospora torreyana Gray Fig. 124  Achene of Rhynchospora nitens.

NC

Torrey’s beaksedge. MA, RI; also reported from CT by Kral (2002b), but specimens are unknown. Coastal plain pond shores.

Cy p e r ac e a e   16 5

Schoenoplectus Achene measurements in the following key do not include the persistent style beak, the prominence of which varies by species and is taxonomically informative. Hybridization is frequent between some members of this genus, especially within the Schoenoplectus lacustris (L.) Palla complex (e.g., S. acutus, S. heterochaetus, S. tabernaemontani). Hybrid plants typically show reduced pollen viability and fruit set. Reference: Smith (2002b). 1a. Plants submerged aquatics with flaccid leaves, often forming extensive vegetative colonies; spikelets solitary on a stiff, emergent stem . . . . . . . . . . . . . . . . . . . . . . S. subterminalis 1b. Plants rarely submerged, the leaf blades, when present, commonly stiffer, associated with reproductive shoots; spikelets usually 2 or more per inflorescence 2a. Plants annual, with cespitose stems; rhizomes absent or very short (i.e., ca. 1 mm long); anthers 0.4–0.6 mm long; stems ca. 0.5–1 mm thick at the midpoint 3a. Leaf-bearing nodes confined to the very base of the stem; achenes pitted to nearly smooth [Fig. 126], with or without perianth bristles; amphicarpic flowers absent 4a. Achenes noticeably pitted, 1.75–2 mm long, rounded near base, biconvex; floral scales with a distinct midvein, lacking a green midstrip; perianth bristles, when present, stout and distinctly tapered apically; primary involucral bract often divergent [Fig. 125] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. purshianus 4b. Achenes nearly smooth, 1.5–1.8 mm long, wedge-shaped near base, mostly plano-convex; floral scales with an obscure midvein, possessing a broad green midstrip; perianth bristles, when present, slender throughout [Fig. 126]; primary involucral bract usually erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. smithii 3b. Leaf-bearing nodes basal and one on the stem; achenes with prominent, transverse, wavy ridges, almost always lacking perianth bristles; amphicarpic flowers frequently present, enclosed in the basal leaf sheaths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hallii 2b. Plants perennial, often forming extensive colonies from elongate rhizomes; anthers 1.5–3.5 mm long; stems ca. 1–10 mm thick at the midpoint 5a. Stems terete or nearly so in cross-section; basal sheaths lacking elongate blades 6a. Spikelets all or mostly single at the ends of the pedicels; floral scales smooth along the margin; achenes trigonous in cross-section, with a style beak 0.3–1 mm long, with 4 (or 5) perianth bristles, usually 2 much shorter than the others . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. heterochaetus 6b. Spikelets all or mostly in glomerules of 2–7 at the ends of the pedicels; floral scales ciliate, at least near the apex; achenes plano-convex or with a low, abaxial ridge, with a style beak 0.1–0.5 mm long, with (5–) 6 (–7) perianth bristles, generally of subequal length 7a. Stems firm, with 10–12 aerenchymal lacunae in cross-section at ⅔ stem height, each lacuna 0.3–0.5 mm in diameter; spikelets mostly in glomerules of 3–7; floral scales with a gray background, (3–) 3.5–4 mm long, with a contorted (rarely straight) awn 0.5–2 mm long; achenes 2–2.8 mm long, with a style beak 0.2–0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. acutus 7b. Stems soft, with 2–4 aerenchymal lacunae in cross-section at ⅔ stem height, each lacuna 1–2.5 mm in diameter; spikelets mostly in glomerules of 2 or 3; floral scales with a pale orange-brown to dark red-brown background, 2–3.5 mm long, with a usually straight awn 0.2–0.8 mm long; achenes 1.7–2.3 mm long, with a style beak 0.1–0.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tabernaemontani 5b. Stems sharply trigonous or even triquetrous in cross-section; basal sheaths producing elongate blades

16 6 MONOCOTS

8a. Achenes equilaterally trigonous to compressed-trigonous in cross-section, with a prominent style beak 0.5–2 mm long; floral scales essentially entire at apex, with a central nerve and 2–10 faint to evident veins on the sides; leaves often half to fully as tall as the stem; apical appendage of anthers rounded, smooth 9a. Inflorescence congested, lacking elongate branches, usually with 1–4 spikelets; floral scales 4–5 mm long; style beak 0.5–1 mm long . . . . . . . . . . . . . . . . . . S. torreyi 9b. Inflorescence open, with elongate branches, with 3–20 pedicellate spikelets; floral scales 5–7 mm long; style beak 0.7–2 mm long . . . . . . . . . . . S. etuberculatus 8b. Achenes plano-convex to obscurely compressed-trigonous in cross-section, with a tiny style beak shorter than 0.5 mm; floral scales with an apical notch 0.1–1 mm deep from which the awn emerges, with only a single central nerve, the sides lacking veins; leaves often half as tall as stem or shorter; apical appendage of anthers elongate, spinulose along margin 10a. Stems with deeply concave sides and wing-angles; primary involucral bract 1–6 cm long; floral scales with an apical notch 0.1–0.4 mm deep, with an awn 0.2–0.5 mm long; uppermost leaf blade shorter than to about equaling the length of its associated sheath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. americanus 10b. Stems with shallowly concave to flat sides and sharp angles; primary involucral bract 3–20 cm long; floral scales with an apical notch (0.3–) 0.5–1 mm deep, with an awn 0.5–1.5 (–2.5) mm long; uppermost leaf blade usually several times longer than its associated sheath . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pungens 1. Schoenoplectus acutus (Muhl. ex Bigelow) A. & D. Löve var. acutus N hard-stemmed bulrush. Schoenoplectus lacustris (L.) Palla ssp. acutus (Muhl. ex Bigelow) A. & D. Löve; Scirpus acutus Muhl. ex Bigelow; S. occidentalis (S. Wats.) Chase • CT, MA, ME, NH, RI, VT. Shallow water or wet soil of lakes, slow-moving rivers, and fens. ‌1 × 5. Schoenoplectus ×oblongus (T. Koyama) Soják is a rare hybrid in New England known from MA, RI, VT. It typically shows a high proportion of solitary spikelets (30–80% of the spikelets are solitary vs. 0–20% in S. acutus). However, the achenes are weakly trigonous in cross-section and commonly have 4 or 5 perianth bristles. Also, each spikelet usually shows a mixture of bifid and trifid styles. 1 × 10. This uncommon bulrush hybrid is to be expected anywhere both parents are found growing together. Utilizing only reproductive characters, one can find it difficult to distinguish this hybrid without extreme familiarity with the parental taxa. However, the two species are very different in their vegetative anatomy. The hybrid has 5–8 aerenchymal lacunae in cross-section at ⅔ stem height, each airspace mostly 0.6–1.5 mm in diameter (see identification key for parental character states). 2. Schoenoplectus americanus (Pers.) Volk. ex Schinz & R. Keller N chair-maker’s bulrush. Scirpus americanus Pers.; S. olneyi Gray • CT, MA, RI. Brackish marshes. Reports of this species in ME, NH, VT in Smith (2002b) are based on putative hybrids ‌contortus) (Schoenoplectus × and acknowledged as such by the author. ‌2 × 6. Schoenoplectus ×contortus (Eames) S.G. Sm. is a rare hybrid in New England, and one that has been tentatively identified for some states where one of the parents (S. americanus) has not been documented (e.g., ME, VT). The hybrid is morphologically intermediate between the parents. 3. Schoenoplectus etuberculatus (Steud.) Soják

NC

Canby’s bulrush. Scirpus etuberculatus (Steud.) Kuntze • RI. Pond shores and fresh to brackish marshes along the coastal plain. 4. Schoenoplectus hallii (Gray) S.G. Sm.

NC

Hall’s bulrush. Scirpus hallii Gray; S. supinus L. var. hallii (Gray) Gray • MA; historically known from two pond shores in the eastern portion of the state. Sandy or peaty draw-down pond shores.

Cy p e r ac e a e   167

5. Schoenoplectus heterochaetus (Chase) Soják

NC

slender bulrush. Scirpus heterochaetus Chase; S. lacustris L. var. tenuiculmis Sheldon • MA, VT. Shallow water or wet soil of lake and river shores. It is useful to know (for hybrid detection) that Schoenoplectus heterochaetus has 6–9 aerenchymal lacunae in cross-section at ⅔ stem height that are mostly 0.4–0.9 mm in diameter. These characteristics are inherited additively (i.e., are intermediate) in hybrid plants. See the identification key for the character states of S. acutus and S. tabernaemontani. 5 ‌ × 10. Schoenoplectus ×steinmetzii (Fern.) S.G. Sm. is a very rare bulrush hybrid in New England. It shows a high proportion of solitary spikelets (similar to S. heterochaetus); however, the achenes are weakly trigonous in cross-section and have usually 4 or 5 perianth bristles. Also, the stems have 5–7 aerenchymal lacunae in cross-section at ⅔ stem height that are mostly 0.7–1.4 mm in diameter. This hybrid is known only from ME within New England (it is still extant at the type locality). 6. Schoenoplectus pungens (Vahl) Palla var. pungens N three-square bulrush. Scirpus pungens Vahl • CT, MA, ME, NH, RI, VT. Lake shores, fresh to brackish river shores, lakeside fens, marshes. 7. Schoenoplectus purshianus (Fern.) M.T. Strong

N C Fig. 125

weak-stalked bulrush.  7a. Scirpus debilis Pursh; S. purshianus Fern.; 7b. Scirpus juncoides Roxb. var. williamsii (Fern.) T. Koyama; S. purshianus Fern. var. williamsii Fern. • CT, MA, ME, NH, RI, VT. Sandy or peaty pond shores and temporary pools. 1a. Flowers with 6 perianth bristles that equal or exceed the length of the achene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7a. S. purshianus var. purshianus

Fig. 125  Inflorescence of Schoenoplectus purshianus.

1b. Flowers without perianth bristles . . . . . . 7b. S. purshianus var. williamsii (Fern.) S.G. Sm. Variety purshianus is known from CT, MA, ME, NH, RI, VT. Variety williamsii is known from only MA in New England (and then from only the type locality in Norfolk County). It is of regional conservation concern. 8. Schoenoplectus smithii (Gray) Soják N Fig. 126 Smith’s bulrush.  8a. Scirpus smithii Gray var. smithii; 8b. Scirpus smithii Gray var. levisetus Fassett; 8c. Scirpus smithii Gray var. setosus Fern. • CT, MA, ME, NH, RI, VT. Sandy or silty river shores, including fresh-tidal portions, sandy lake shores. 1a. Perianth bristles absent or vestigial . . . . . . . . . . . . . . . . . . . . . . . . . . 8a. S. smithii var. smithii 1b. Perianth bristles present, shorter than to twice the length of the achene 2a. Perianth bristles 1–4, up to as long as achene, smooth or minutely retrorse-barbellate only at tip [Fig. 126] . . . . . . . . . . . . . . . . . . . . . . 8b. S. smithii var. levisetus (Fassett) S.G. Sm. 2b. Perianth bristles 4–6, equaling to twice as long as achene, minutely retrorse-barbellate at least in the apical 50% . . . . . . . . . . . . . . . . . . . . . 8c. S. smithii var. setosus (Fern.) S.G. Sm. Variety smithii is known from CT, MA, RI, VT. Variety levisetus is known from ME. Variety setosus is known from CT, MA, ME, NH, VT. Varieties levisetus and smithii are found in habitats with usually greatly fluctuating water levels (e.g., tidal river shores, drawn-down pond shores), whereas var. setosus is typically found in places where daily or seasonal variation in water volume is less marked. 9. Schoenoplectus subterminalis (Torr.) Soják N water bulrush. Scirpus subterminalis Torr. • CT, MA, ME, NH, RI, VT. Shallow water of lakes and slow-moving rivers, often forming large mats of thin, flaccid, narrow leaves underwater. 10. Schoenoplectus tabernaemontani (K.C. Gmel.) Palla N soft-stemmed bulrush. Schoenoplectus validus (Vahl) A. & D. Löve; Scirpus lacustris L. ssp. creber (Fern.) T. Koyama; S. tabernaemontani K.C. Gmel.; S. validus Vahl; S. validus Vahl var. creber Fern. • CT, MA, ME, NH, RI, VT. Lake shores, river shores, marshes, fresh to brackishtidal marshes, ditches.

Fig. 126  Achene of Schoenoplectus smithii var. levisetus.

16 8  MONOCOTS

11. Schoenoplectus torreyi (Olney) Palla N Torrey’s bulrush. Scirpus torreyi Olney • CT, MA, ME, NH, RI, VT. Shallow water of lakes, rivers, and marshes.

Scirpus Hybridization is not uncommon in this genus. Hybrid plants can be recognized by low fruit set and frequently by their elongated spikelets (it has been hypothesized that the spiklet axis continues to elongate when resources are not devoted to fruit formation). The perianth bristles are usually either retrorsely barbellate (at least near the tip) or highly bent and contorted. These features help to separate the bristles from the filaments, which may be persistent on the fruits. References: Schuyler (1967), Whittemore and Schuyler (2002). 1a. Perianth bristles at most 1.5 times the length of the achene, retrorsely barbed (often smooth in S. georgianus), straight or merely curved or perianth bristles entirely absent 2a. Leaves 14–22 (–26) on each stem; floral scales red-brown, orbicular or nearly so; spikelets mostly broad-ovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. polyphyllus 2b. Leaves 3–12 on each stem; floral scales green-brown or brown to black, elliptic to broad-elliptic; spikelets cylindrical or narrow-ellipsoid to narrow-ovoid or ovoid (rarely broad-ovoid, as sometimes in S. cyperinus) 3a. Perianth bristles barbed in the apical 60–90% with sharp, stout, thick-walled teeth; achenes 0.6–1 mm wide 4a. Lower sheaths green or white-brown to brown; branches of inflorescence arching to drooping; plants cespitose, with short, brown rhizomes; achenes 0.6–0.8 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. ancistrochaetus 4b. Lower sheaths conspicuously anthocyanic; branches of inflorescence ascending to divaricately spreading; plants spreading by long, anthocyanic rhizomes; achenes (0.6–) 0.8–1 mm wide 5a. Styles bifid; achenes lenticular to plano-convex, with firmly attached, persistent perianth bristles; branches of inflorescence scabrous only in the distal portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. microcarpus 5b. Styles trifid; achenes compressed-trigonous, with weakly attached, sometimes caducous, perianth bristles; branches of inflorescence antrorsely scabrous their entire length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. expansus 3b. Perianth bristles smooth or barbed only in the apical 10–60% with rounded, slender teeth; achenes 0.3–0.6 mm wide 6a. Perianth bristles numbering 0–3 per flower, up to 0.7 times as long as the achene, smooth or with barbs only near the very apex . . . . . . . . . . . . . . . S. georgianus 6b. Perianth bristles numbering mostly 5 or 6 per flower, more than 0.6 times as long as the achene, barbed in the apical half 7a. Longer perianth bristles shorter than to equaling the length of the 0.8–1.1-mm- long achenes; floral scales appearing black under low magnification; spikelets 2–3.5 mm long; leaf blades 5–9 (–10) mm wide; basal leaves and sheaths usually with few and inconspicuous cross-septa . . . . . . . . . S. hattorianus 7b. Longer perianth bristles exceeding the length of the 1–1.3-mm-long achenes; floral scales appearing brown under low magnification; spikelets mostly 3.5–5 mm long; leaf blades 7–17 (–18) mm wide; basal leaves and sheaths with many prominent cross-septa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. atrovirens 1b. Perianth bristles greatly exceeding the achenes when extended, smooth, bent and curled 8a. Perianth bristles largely contained within the floral scales, even in fruit, and therefore not obviously visible; achenes 1–1.2 mm long; floral scales with evident green midribs, even at maturity of the fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pendulus

Cy p e r ac e a e   16 9

8b. Perianth bristles exserted beyond the floral scales, in fruit providing a “woolly” aspect to the inflorescence; achenes 0.6–1 mm long; floral scales with pale, brown, or relatively inconspicuously colored midribs 9a. Achenes red-brown; floral scales 2–3.1 mm long; plants spreading by means of elongate rhizomes, often forming large circular colonies; base of involucral bracts glutinous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. longii 9b. Achenes white to pale brown; floral scales 1.1–2.2 mm long; plants with short, branching rhizomes, forming dense tussocks; base of involucral bracts not glutinous 10a. Spikelets mostly or all in glomerules of 2 or more; floral scales mostly shorter than 1.5 mm; achenes maturing in August and September . . . . . . . . . . . . . S. cyperinus 10b. Spikelets mostly or entirely solitary, sessile or pedicellate; floral scales 1.4–1.8 mm long; achenes maturing in late June through July 11a. Involucels and base of involucral bracts very dark brown to black; stems mostly 1–3 mm thick below the inflorescence; leaf blades 2–5 mm wide; floral scales rounded at apex or with a very tiny mucro to 0.05 mm long; achenes maturing late June through early July. . . . . . . . . . . . . . . . . . . . . . . . . . . . S. atrocinctus 11b. Involucels and base of involucral bracts brown, sometimes with a dark edge; stems mostly 2–5 mm thick below the inflorescence; leaf blades 3–10 mm wide; floral scales rounded at the apex to, more commonly, mucronate, the mucro to 0.1 mm long; achenes maturing in mid- through late July . . . . . . . . . S. pedicellatus 1. Scirpus ancistrochaetus Schuyler northeastern bulrush. MA, NH, VT. Beaver flowages, temporary pools, wet depressions, near small ponds.

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2. Scirpus atrocinctus Fern. N black-girdled woolsedge. Scirpus cyperinus (L.) Kunth var. brachypodus (Fern.) Gilly • CT, MA, ME, NH, RI, VT. Wet fields, lake borders, graminoid marshes, temporary pools. 2 × 4. This woolsedge nothospecies somewhat resembles Scirpus atrocinctus, except that it possesses a very different phenology, especially in regard to fruit maturation (often in late July and early August). 2 ‌ × 7. Scirpus ×peckii Britt. is an uncommon bulrush nothospecies that is known from CT, MA, ME, NH, RI, VT. It has individual spikelets that somewhat resemble those of S. atrocinctus; however, the clusters of spikelets are borne on relatively stiff, straight inflorescence branches. The perianth bristles are elongate and somewhat bent but have a few remote barbules along their length. This hybrid appears to be much less frequent in the southern New England states. 3. Scirpus atrovirens Willd. N dark-green bulrush. CT, MA, ME, NH, RI, VT. Wet fields, graminoid marshes, ditches, open, seasonally wet areas. This species is over-reported. Many reports are actually Scirpus hattorianus. 4. Scirpus cyperinus (L.) Kunth N common woolsedge. Scirpus cyperinus (L.) Kunth var. pelius Fern.; S. cyperinus (L.) Kunth var. rubricosus Fern. • CT, MA, ME, NH, RI, VT. Wet fields, lake borders, graminoid marshes, temporary pools. 5. Scirpus expansus Fern. N wood bulrush. CT, MA, ME, NH, RI, VT. Graminoid marshes, wetland borders, wet fields. 6. Scirpus georgianus Harper N Georgia bulrush. Scirpus atrovirens Willd. var. georgianus (Harper) Fern. • CT, MA, ME, NH, RI, VT; rare in the northern portion of its range. Wet fields, graminoid marshes, ditches, open, seasonally wet areas. Early reports of Scirpus georgianus (e.g., Fernald 1950) in New England are ambiguous in that the taxon was considered to include both this species and the common S. hattorianus.

170 MO NOCOTS

7. Scirpus hattorianus Makino N mosquito bulrush. CT, MA, ME, NH, RI, VT. Wet fields, graminoid marshes, ditches, open, seasonally wet areas. 8. Scirpus longii Fern.

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Long’s woolsedge. CT, MA, ME, NH, RI. Graminoid marshes, often adjacent to river channels, acidic fens, river oxbows. 9. Scirpus microcarpus J. & K. Presl N barber-pole bulrush. Scirpus microcarpus J. & K. Presl var. rubrotinctus (Fern.) M.E. Jones; S. rubrotinctus Fern. • CT, MA, ME, NH, RI, VT. Wet fields, graminoid marshes, ditches, open, seasonally wet areas. 10. Scirpus pedicellatus Fern. N stalked woolsedge. Scirpus cyperinus (L.) Kunth var. pedicellatus (Fern.) Schuyler • CT, MA, ME, NH, RI, VT. Wet fields, lake borders, graminoid marshes, temporary pools. 11. Scirpus pendulus Muhl. N rufous bulrush. Scirpus lineatus, auct. non Michx. • CT, MA, ME, NH, VT. Low fields, ditches, meadows, often associated with high-pH bedrock. The name Scirpus lineatus Michx. was long misapplied to this species. 12. Scirpus polyphyllus Vahl

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leafy bulrush. CT, MA, NH, VT. Low, riparian forests, stream and pond borders, swamp edges. Reports of this species from RI are based on a collection of Scirpus georgianus—Champlin 274 (Champlin Herb.).

Scleria Reference: Reznicek et al. (2002). 1a. Hypogynium vestigial, therefore, the achene base ± continuous with the achene body and without a basal zone of different color and texture; leaf blades 0.5–2 mm wide; plants annual, from fibrous roots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. verticillata 1b. Hypogynium present, therefore, the achene with a distinct basal zone of different color and texture; leaf blades 1–9 mm wide; plants perennial, from firm, nodulose rhizomes or annual with fibrous roots in S. reticularis 2a. Achene body lustrous and smooth; hypogynium unlobed and without tubercles, densely beset with minute, white to light brown papillae; leaf blades 3–9 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. triglomerata 2b. Achene body dull to sublustrous, either pitted or papillate-verrucose; hypogynium either lobed or with tubercles, without minute papillae; leaf blades 1–3.5 mm wide 3a. Hypogynium with 3 oblong lobes, without tubercles; achene body reticulatepatterned with polygonal pits; inflorescence a panicle of clusters, with visible branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. reticularis 3b. Hypogynium without lobes, bearing 6 globose tubercles arranged in 3 pairs; achene body papillate-verrucose, these projections sometimes arranged to provide the appearance of irregular transverse ridges; inflorescence a dense terminal cluster, sometimes with 1 or 2 axillary clusters as well . . . . . . . . . . . . . . . . . . . . . . . . . . S. pauciflora 1. Scleria pauciflora Muhl. ex Willd.

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few-flowered nutsedge. 1a. Scleria pauciflora Muhl. ex Willd. var. kansana Fern.; 1b. Scleria ciliata Michx. var. pauciflora (Muhl. ex Willd.) Kükenth. • CT, MA, NH, RI; also reported from VT by Reznicek et al. (2002), but specimens are unknown. Sandplains, oak shrublands, openings in woodlands, and low, seasonally wet, sandy areas.

Cy p e r ac e a e   171

1a. Plants densely pubescent with spreading hairs 0.5–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. S. pauciflora var. caroliniana (Willd.) Wood 1b. Plants glabrous to sparsely pubescent with hairs less than 0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. S. pauciflora var. pauciflora Variety caroliniana is known from CT, MA, NH, RI. Variety pauciflora is known from MA, NH. 2. Scleria reticularis Michx.

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netted nutsedge. CT, MA, NH, RI. Sandy or peaty shores of ponds along the coastal plain. 3. Scleria triglomerata Michx.

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whip nutsedge. Scleria flaccida Steud.; S. nitida Willd. • CT, MA, RI, VT. Openings in woodlands, moist sandy fields, and low, seasonally wet, sandy areas. The voucher for this species from VT is somewhat enigmatic—it is without location (beyond the state), date, or collector, though it is in the hand of Perkins (Arthur Gilman, personal communication; specimen at VT). 4. Scleria verticillata Muhl. ex Willd.

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low nutsedge. CT. Swamps and graminoid marshes in regions of high-pH bedrock.

Trichophorum Reference: Crins (2002b). 1a. Stems terete, smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. cespitosum 1b. Stems trigonous, usually scabrous on the angles 2a. Perianth bristles white, flattened, 3–5 times longer than the inflorescence; spikelets with 9–20 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. alpinum 2b. Perianth bristles pale brown to brown, terete, much shorter than inflorescence and usually included within the subtending floral scales; spikelets with 3–8 flowers 3a. Floral scales obtuse at apex, the upper scales with an evanescent midrib; leaves 0.5–0.8 (–1) mm wide, up to as tall as the stems at flowering, much shorter than the stems at fruiting; plants of high-pH river shore ledges . . . . . . . . . . . . . . . . . . . . . T. clintonii 3b. Floral scales mucronate at apex, all with an excurrent midrib; leaves 0.8–2 mm wide, equal in height or taller than the stems, even at fruiting; plants of dry-mesic to mesic deciduous forests, often with Quercus in the canopy . . . . . . . . . . . . . . . . . . . T. planifolium 1. Trichophorum alpinum (L.) Pers. N alpine clubsedge. Baeothryon alpinum (L.) Egor.; Eriophorella alpina (L.) Holub; Eriophorum alpinum L.; Scirpus hudsonianus (Michx.) Fern. • CT, MA, ME, NH, RI, VT. Fens, wet rocks, stream shores, circumneutral meadows. 2. Trichophorum cespitosum (L.) Hartman ssp. cespitosum N tufted clubsedge. Baeothryon cespitosum (L.) A. Dietr.; Scirpus cespitosus L.; S. cespitosus L. var. callosus Bigelow; S. cespitosus L. var. delicatulus Fern. • ME, NH, VT. Alpine plateaus, fens, bogs, ice-scoured river shores. 3. Trichophorum clintonii (Gray) S.G. Sm. N Clinton’s clubsedge. Baeothryon clintonii (Gray) A. & D. Löve; Scirpus clintonii Gray • ME. Circumneutral river shore outcrops. 4. Trichophorum planifolium (Spreng.) Palla N bashful clubsedge. Baeothryon verecundum (Fern.) A. & D. Löve; Scirpus planifolius Muhl.; S. verecundus Fern. • CT, MA, ME, NH, RI, VT; restricted in ME and NH to the southern portion of states. Mesic to dry-mesic forests and woodlands, usually with Quercus.

172 MO NOCOTS

Dioscoreaceae Dioscorea 1a. Leaves with 3–5 lobes, with bulbils in the axils, usually opposite in the distal portion of the stem; staminate inflorescences in fascicles of (1–) 3–5 in the axils of leaves; plants tuberous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. polystachya 1b. Leaves without lobes, the margin entire to undulate, without bulbils in the axils, alternate in the distal portion of the stem; staminate inflorescences solitary in the axils of leaves; plants rhizomatous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. villosa 1. Dioscorea polystachya Turcz. E Chinese yam. Dioscorea batatas R. Decr. • CT, MA, VT. Waste places, compost heaps, fields. The name Dioscorea oppositifolia L. has been used for this species, but that name correctly refers to a plant that is native to India and not found in New England (Raz 2002). 2. Dioscorea villosa L. N wild yam. Dioscorea hirticaulis Bartlett; D. villosa L. var. hirticaulis (Bartlett) Ahles • CT, MA, RI. Riparian and lacustrine forests, wetland borders. The report of this species from VT by Angelo and Boufford (2000) is based on a specimen (at NEBC) that was later annotated to Dioscorea polystachya.

Eriocaulaceae Eriocaulon Reference: Kral (2000a). 1a. Inflorescence subglobose, pale gray to white, 4–5 (–10) mm wide; bractlets evidently pubescent with clavate hairs; flowering scape with (4–) 5–7 ridges; each rosette of leaves producing usually a single flowering scape; plants of fresh water habitats. . . . . . E. aquaticum 1b. Inflorescence hemispherical, gray to light yellow-brown, 3–4 mm wide; bractlets sparsely pubescent with clavate hairs; flowering scape with 4 or 5 ridges; each rosette of leaves producing 1–4 flowering scapes; plants of tidal shores . . . . . . . . . . . . . . . . . . . . . . . . . . . E. parkeri 1. Eriocaulon aquaticum (Hill) Druce N seven-angled pipewort. Cespa aquatica Hill; Eriocaulon pellucidum Michx. • CT, MA, ME, NH, RI, VT. Wet soil or shallow water of lakes and slow-moving rivers. The name Eriocaulon septangulare Withering, which has been used for this species, is invalid. 2. Eriocaulon parkeri B.L. Robins.

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Parker’s pipewort. CT, MA, ME. Fresh to brackish-tidal river shores, occurring primarily on sand, silt, and mud substrate.

ha e m o d o r ac e a e   173

Haemodoraceae Lachnanthes 1. Lachnanthes caroliniana (Lam.) Dandy N Carolina bloodroot. Dilatris caroliniana Lam.; Gyrotheca tinctoria (J.F. Gmel.) Salisb.; Lachnanthes tinctoria (J.F. Gmel.) Ell. • CT, MA, RI. Sandy or peaty pond shores along the coastal plain.

Hemerocallidaceae Hemerocallis 1a. Flowers red-orange, not fragrant; tepals reticulate-veined; capsules rarely maturing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. fulva 1b. Flowers yellow, fragrant; tepals parallel-veined; capsules maturing (i.e., containing seeds) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. lilioasphodelus 1. Hemerocallis fulva (L.) L. E orange day-lily. Hemerocallis lilioasphodelus L. var. fulvus L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest edges, abandoned house lots. 2. Hemerocallis lilioasphodelus L. E yellow day-lily. Hemerocallis flava (L.) L. • CT, MA, ME, RI, VT. Fields, roadsides, forest edges, abandoned house lots.

Hostaceae Hosta 1a. Leaf blades lanceolate to broad-lanceolate, 10–17 × (3–) 5–7.5 cm, with 5 or 6 pairs of lateral veins; flowers gradually expanding apically, with pale purple tepals; anthers purple; scape mostly 40–50 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. lancifolia 1b. Leaf blades ovate to cordate-ovate, 20–30 × (10–) 15–20 cm, with 7–9 pairs of lateral veins; flowers abruptly expanding apically, with purple-blue tepals; anthers spotted with purple; scape mostly 80-95 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. ventricosa 1. Hosta lancifolia Tratt. E narrow-leaved plantain-lily. Hemerocallis japonica (Thunb.) Thunb.; H. lancifolia Thunb.; Hosta japonica (Houtt.) Voss • CT. Fields, roadsides, forest edges, abandoned house lots. 2. Hosta ventricosa (Salisb.) Stearn E blue plantain-lily. Bryocles ventricosa Salisb. • CT, MA, RI, VT. Fields, roadsides, forest edges, abandoned house lots.

174 MONOCOTS

Hyacinthaceae 1a. Tepals evidently connate for some distance [Fig. 127]; stamens epipetalous 2a. Perianth urceolate to cylindric [Fig. 127]; tepals connate for most of their length; leaf blades 2–8 mm wide; racemes with usually 12–20 flowers, dense [Fig. 127], with sterile flowers near the apex that often show slightly reduced size and somewhat paler color . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Muscari 2b. Perianth open-campanulate; tepals 15–25 mm long, connate for less than ⅓ their length; leaf blades 10–20 mm wide; racemes with usually 4–10 flowers, open, without apical sterile flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chionodoxa 1b. Tepals essentially distinct (i.e., connate for less than 1⁄10 their total length) [Fig. 128]; stamens free or inserted near the base of the tepals 3a. Tepals white, each with a green abaxial stripe, 15–30 mm long; stamens dimorphic as to length, with flat filaments; leaves mostly 20–60 cm long, with a white adaxial stripe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithogalum 3b. Tepals blue (rarely white), 12–16 mm long; stamens monomorphic, the filaments not conspicuously flattened; leaves mostly 10–15 cm long, lacking a white stripe . . . . . Othocallis

Chionodoxa 1a. Flowers numbering 4–10 per inflorescence, erect to spreading; stems 15–30 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. forbesii 1b. Flowers numbering 1–3 per inflorescence, erect to ascending; stems 10–20 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. luciliae 1. Chionodoxa forbesii Baker E Forbes’ glory-of-the-snow. VT. Edges of lawns, fields, roadsides. 2. Chionodoxa luciliae Boiss. E Lucile’s glory-of-the-snow. MA. Edges of lawns, fields, roadsides.

Muscari 1a. Tepals blue, the connate portion of fertile flowers ± globose [Fig. 127]; leaf blades 3–8 mm wide; racemes with usually 12–20 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. botryoides 1b. Tepals very dark blue, the connate portion of fertile flowers obovoid to cylindric; leaf blades 2–4 (–5) mm wide; racemes with usually 20–40 flowers . . . . . . . . . . . . . . . M. neglectum 1. Muscari botryoides (L.) P. Mill. E Fig. 127 common grape-hyacinth. Hyacinthus botryoides L. • CT, MA, NH, VT. Edges of lawns, fields, roadsides. 2. Muscari neglectum Guss. ex Ten. E starch grape-hyacinth. Hyacinthus racemosus L.; Muscari atlanticum Boiss. & Reut. • CT, MA. Edges of lawns, fields, roadsides. Fig. 127  Inflorescence of Muscaria botryoides.

Ornithogalum 1a. Inflorescence racemose, ± cylindric [Fig. 128], with 5–12 (–18) flowers; pedicels nodding, especially after anthesis, the lower up to 1 cm long; filaments with 2 lateral teeth near the apex; leaves 5–10 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. nutans

hyac i n th ac e a e   175

1b. Inflorescence corymbose, flat-topped, with (4–) 8–20 flowers; pedicels ascending to erect, the lower 2–6 cm long; filaments lacking teeth; leaves 3–5 mm wide . . . . . . . . O. umbellatum 1. Ornithogalum nutans L. E Fig. 128 nodding star-of-Bethlehem. CT. Fields, roadsides, forest edges. 2. Ornithogalum umbellatum L. E nap-at-noon. CT, MA, ME, NH, RI, VT. Fields, roadsides, forest edges and openings.

Othocallis 1. Othocallis siberica (Haw. ex Andr.) Speta E Siberian squill. Scilla siberica Haw. ex Andr. • MA. Edges of lawns, fields, roadsides, abandoned gardens.

Hydrocharitaceae Reference: Haynes (2000b). 1a. Leaves opposite; flowers without a perianth; androecium with 1 stamen; gynoecium composed of a single, unilocular, unicarpellate ovary; fruit an achene . . . . . . . . . . . . . . . . . Najas 1b. Leaves whorled (at least in part) or all basal; flowers with a perianth composed of 3 sepals and 3 petals; androecium with usually 2 or 9 stamens; gynoecium composed of 3–6 carpels; fruit usually ± capsule-like or berry-like 2a. Leaves differentiated into blades and petioles, the blades broad-ovate to cordateorbicular, floating on the surface or becoming emergent in dense vegetation, with evidently non-parallel primary lateral veins [Fig. 130] 3a. Primary lateral veins of leaf blade forming a 75- to 90-degree (or greater) angle with the midvein [Fig. 130]; filaments distinct most of their length; petals broad-obovate, 1.5 times or more or long as the sepals; anthers oval in outline; stipules 2, distinct from the petiole; seeds without blunt, cylindrical hairs . . . . . . . . . . . . . . . . . . . . . . . Hydrocharis 3b. Primary lateral veins of leaf blade forming a 30- to 80-degree angle with the midvein; filaments connate more than ½ their length into a column; petals linear to narrow-oblong, up to 1.5 times as long as the sepals; anthers linear in outline; stipule 1, basally connate to the petiole; seeds covered with blunt, cylindrical hairs . . Limnobium 2b. Leaves not differentiated into blades and petioles, the linear to narrow-ovate blades submersed and with ± parallel veins [Figs. 131, 132] 4a. Leaves all basal, 10–110 cm long, with a prominent, broad, central lacunar band [Fig. 132]; staminate flowers with 2 stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vallisneria 4b. Leaves borne on an elongate stem, mostly in whorls of 3–8, 0.4–4 cm long, without a prominent, central lacunar band; staminate flowers with 3–9 stamens 5a. Leaves with prickles on the abaxial midrib; intravaginal squamules (i.e., small, stipule-like scales in the leaf axils) fringed with orange-brown hairs; staminate flowers with 3 stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrilla 5b. Leaves without prickles; intravaginal squamules entire or fringed with clear hairs; staminate flowers with 7–9 stamens

Fig. 128  Inflorescence of Ornithogalum nutans.

176 MO NOCOTS

6a. Leaves numbering 2–4 per node, usually in whorls of 3 [Fig. 129]; petals inconspicuous, up to 4.5 mm long, scarcely, if at all, longer than the sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elodea 6b. Leaves numbering 4–8 per node, usually in whorls of 5 or more; petals more showy, 9–11 mm long, much longer than the sepals . . . . . . . . . . . . . . . . . . . . . . Egeria

Egeria 1. Egeria densa Planch. E Brazilian-waterweed. Anacharis densa (Planch.) Victorin • CT, MA, NH, VT. Shallow, still or slow-moving water of lakes and rivers. This species has been introduced into the United States through the aquarium trade. Only staminate plants are known outside its native range. Vegetative reproduction is likely through plant fragmentation given that no specialized structures for this form of reproduction (e.g., turions, bulbils) have been observed.

Elodea 1a. Leaves 1.1–5 mm wide, averaging 2 mm, relatively firm, blunt at the apex, the upper densely crowded and overlapping; staminate flowers elevated on a long, slender, pedicellike hypanthium 2–30 cm long [Fig. 129], with a spathe 8–13.5 mm long, with sepals 3–4 mm long; carpellate flowers with sepals 2.0–3.5 mm long . . . . . . . . . . . . . . . . E. canadensis 1b. Leaves 0.9–1.7 (–2.4) mm wide, averaging 1.3 mm, weak, pointed at the apex, the upper not densely crowded; staminate flowers sessile, with a spathe 2–4 mm long, with sepals ca. 2 mm long; carpellate flowers with sepals 1–1.8 mm long . . . . . . . . . . . . . . . . . . . . . E. nuttallii 1. Elodea canadensis Michx. N Fig. 129 Fig. 129  Branch and staminate flower of Elodea canadensis.

common waterweed. Anacharis canadensis (Michx.) Planch. • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving, circumneutral to basic waters of lakes and rivers. 2. Elodea nuttallii (Planch.) St. John N free-flowered waterweed. Anacharis nuttallii Planch. • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving, slightly acidic to basic waters of lakes and rivers.

Hydrilla 1. Hydrilla verticillata (L. f.) Royle E water-thyme. Serpicula verticillata L. f. • CT, MA, ME. Lakes and slow-moving rivers, with a broad toleratance of pH, nutrient level, and salinity. This species can vegetatively perennate by stem fragmentation and by production of tuber-like turions (these structures can be produced underwater and beneath the substrate the plant is rooted in).

Hydrocharis 1. Hydrocharis morsus-ranae L. E Fig. 130 European frog’s-bit. VT. Lakes, embayments and side channels of rivers. Fig. 130  Leaf of Hydrocharis morsus-ranae.

Limnobium 1. Limnobium spongiosa (Bosc) Rich. ex Steud. E American spongeplant. Hydrocharis spongiosa Bosc • CT. Pond shores. This species forms two types of leaf blades—floating ones with conspicuous aerenchyma tissue on the abaxial surface and emergent ones lacking the prominent aerenchyma tissue.

H y d ro c ha r i tac e a e   177

Najas References: Hellquist and Crow (1980), Haynes (2000c). 1a. Each margin of the leaf blade with 7–17 spinules [Fig. 131], these either minute and unicellular or larger and multicellular; apex of leaf sheath truncate or auriculate [Fig. 131] 2a. Axis of seeds curved; style situated at center of seed apex; leaf blades usually recurved in age, the margins with multicellular teeth that are often visible to the unaided eye; staminate flowers 1.9–2.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. minor 2b. Axis of seeds straight; style situated off to one side of seed apex; leaf blades ascending to spreading, the margins with minute, unicellular teeth that usually require magnification to resolve; staminate flowers 1.5–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. gracillima 1b. Each margin of the leaf blade with 18–100 minute, unicellular spinules; apex of leaf sheath convexly tapering into base of blade (rarely truncate in N. guadalupensis) 3a. Testa of seeds dull, minutely pitted and appearing so even at low magnification; seeds fusiform, widest at the middle; anthers 4-locular (rarely 1-locular); leaves 0.2–2.1 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. guadalupensis 3b. Testa of seeds glossy, very minutely pitted and appearing ± smooth at low magnification; seeds narrow-obovoid to broad-obovoid, widest above the middle; anthers 1-locular; leaves 0.2–0.6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. flexilis 1. Najas flexilis (Willd.) Rostk. & Schmidt N wavy waternymph. Caulinia flexilis Willd.; Najas caespitosa (Maguire) Reveal • CT, MA, ME, NH, RI, VT; throughout. Still or slow-moving, circumneutral to basic water of lakes and rivers. 2. Najas gracillima (A. Braun ex Engelm.) Magnus N Fig. 131 slender waternymph. Najas indica (Willd.) Cham. var. gracillima A. Braun ex Engelm. • CT, MA, ME, NH, RI, VT. Still or slow-moving, acidid to basic water of lakes and rivers. 3. Najas guadalupensis (Spreng.) Magnus N Guadalupe waternymph.  3a. Caulinia guadalupensis Spreng.; 3b. Najas guadalupensis (Spreng.) Magnus var. olivacea (Rosendahl & Butters) Haynes; N. olivacea Rosendahl & Butters • CT, MA, ME, NH, RI, VT. Circumneutral, fresh to brackish water of lakes and rivers. 1a. Leaf blade with 50–100 unicellular spicules on each margin; stems up to 0.8 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. N. guadalupensis ssp. guadalupensis

Fig. 131  Leaf sheath and blade of Najas gracillima.

1b. Leaf blade with 20–40 unicellular spicules on each margin; stems 1 mm or more in diameter . . . . . 3b. N. guadalupensis ssp. olivacea (Rosendahl & Butters) Haynes & Hellquist Subspecies guadalupensis is known from CT, MA, ME, NH, RI, VT. Subspecies olivacea is known from CT, MA. 4. Najas minor All. E brittle waternymph. Caulinia minor (All.) Coss. & Germ. • ct, mA, NH, VT; primarily western portion of states. Still or slow-moving, circumneutral to basic water of lakes and rivers.

Vallisneria 1. Vallisneria americana Michx. N Fig. 132 tape-grass. Vallisneria spiralis, auct. non L. • CT, MA, ME, NH, RI, VT. Lakes and slow-moving rivers, primarily in circumneutral to basic water.

Fig. 132  Leaf blade of Vallisneria americana showing lacunar bands.

178  MONOCOTS

Hypoxidaceae Hypoxis 1. Hypoxis hirsuta (L.) Coville N common star-grass. Hypoxis carolinensis Michx.; Ornithogalum hirsutum L. • CT, MA, ME, NH, RI, VT. Woodlands, forest edges, sandy fields.

Iridaceae 1a. Styles broad, petaloid, with paired crests at the apex [Fig. 137]; anthers concealed, appressed to abaxial surface of style branches; sepals, petals, stamens, and carpels adnate at the base forming a floral tube that surmounts the ovary . . . . . . . . . . . . . . . . . . . . . . . . (in part) Iris 1b. Styles not petaloid, without crests; anthers visible, not appressed to style branches; sepals, petals, stamens and carpels not adnate near base (though Crocus with basally connate tepals) 2a. Plants without aerial stems; fruit borne at or even below the ground level; tepals with an elongate, basally connate portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crocus 2b. Plants with evident, above-ground stems; fruit borne well above ground level; tepals distinct or with a very short (i.e., discernable only with close observation) basally connate portion 3a. Leaf blades 10–20 mm wide; tepals light orange to red (rarely yellow); filaments distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Iris 3b. Leaf blades 0.8–5 mm wide; tepals white to blue or blue-violet; filaments basally connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sisyrinchium

Crocus 1. Crocus vernus (L.) Hill ssp. vernus E Dutch crocus. CT, MA. Edges of lawns, fields, roadsides, compost heaps.

Iris Older literature sometimes referred to the sepals as “falls” and the petals as “standards”. These are superfluous terms and are abandoned here. Recent phylogenetic analysis showed that Belamcanda is nested within Iris (Wilson 2004). Two specimens have been seen that were determined as Iris prismatica × I. versicolor—9 Jul 1923, Cheever & Knowlton s.n. (NEBC!) was I. versicolor and Sanford 10009 (NEBC!) is doubtfully as determined. Reference: Henderson (2002). 1a. Styles narrow, not petaloid, without crests; anthers visible, not appressed to style branches; seeds black, fully exposed at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. domestica 1b. Styles broad, petaloid, with paired crests at the apex; anthers concealed, appressed to abaxial surface of style branches; seeds ultimately tan to brown, contained within the capsule 2a. Plants actually or nearly acaulescent at anthesis, the stems up to 4.5 cm tall

i r i dac e a e   179

3a. Sepals with a line of multicellular hairs on the midrib; stems absent or up to 1 cm tall; capsules rounded-triangular in cross-section . . . . . . . . . . . . . . . . . . . . . . . . . . . I. pumila 3b. Sepals with 3, parallel, toothed ridges; stems 2.5–4.5 cm tall; capsules sharply triangular in cross-section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. cristata 2b. Plants caulescent, the stems (5–) 10–120 cm tall 4a. Sepals ornamented with toothed ridges; capsule 6-ribbed . . . . . . . . . . . . I. tectorum 4b. Sepals without ridges or crests, though sometimes pubescent; capsule 3-ribbed or 3-edged 5a. Sepals with a line of multicellular hairs on the midrib; seeds oval in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. germanica 5b. Sepals glabrous or with a small patch of fine, unicellular hairs near the base in I. versicolor; seeds D-shaped to pyriform in outline (± circular in I. ensata) 6a. Perianth yellow; pedicels of fruits arching to sometimes pendulous [Fig. 135] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. pseudacorus 6b. Perianth blue to violet (rarely white); pedicels of fruits erect 7a. Adnate floral tube 2–3 mm tall; branches of rhizomes unlike the primary rhizome, bearing scale-like leaves along its length [Fig. 134], expanding at apex to produce green leaves; capsules sharply triangular in cross-section; seeds in 1 row in each locule of capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. prismatica 7b. Adnate floral tube 5–100 mm tall; branches of rhizome similar to the primary rhizome in size and texture; capsules rounded-triangular to terete in cross-section; seeds in 2 rows in each locule of the capsule 8a. Petals inconspicuous, 1–2 cm long, tipped by a bristle, strongly involute in the basal portion, the margins often nearly or fully touching and ± forming a tube; seeds compressed-pyriform in outline [Fig. 133] . . . . . . . . . . I. hookeri 8b. Petals conspicuous, 2–5.5 cm long, without an apical brisle [Fig. 136], weakly involute in the basal portion and merely forming a shallow trough; seeds D-shaped or ± circular in outline [Fig. 137] 9a. Flowers 8–15 cm wide in life; hypanthium ca. 2 mm long; capsules terete or nearly so in cross-section with 6 longitudinal ridges; seeds ± circular in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. ensata 9b. Flowers mostly 6–8 cm wide in life; hypanthium 4–10 mm long; capsules rounded-triangular in cross-section, lacking longitudinal ridges; seeds D-shaped in outline 10a. Leaf blades 4–6 mm wide; stems hollow; petals 12–20 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. sibirica 10b. Leaf blades 10–30 mm wide; stems solid; petals 8–12 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. versicolor 1. Iris cristata Ait. E crested iris. Neubeckia cristata (Soland. ex Ait.) Alef. • MA. Edges of lawns, fields, roadsides, compost heaps. 2. Iris domestica (L.) Goldblatt & Mabberly E blackberry-lily. Belamcanda chinensis (L.) DC.; Epidendrum domesticum L.; Gemmingia chinensis (L.) Kuntze; Ixia chinensis L. • CT. Fields, edges of lawns, roadsides, abandoned gardens. Reports of this species in VT (e.g., Bean et al. 1951) are based on a collection taken from a cultivated plant (specimen at NEBC).

180  MONOCOTS

2. Iris ensata Thunb. E Russian iris. Iris kaempferi Sieb. ex Lemaire • CT, VT. Edges of lawns, fields, roadsides, compost heaps. Iris ensata has larger flowers than Iris versicolor (8–15 cm wide vs. 6–8 cm wide). 3. Iris germanica L. E German iris. CT, MA; also reported from ME by Campbell et al. (1995), but specimens are unknown. Edges of lawns, fields, roadsides, compost heaps. Iris germanica is the natural hybrid of I. pallida Lam. and I. variegata L. Fig. 133  Seed of Iris hookeri.

4. Iris hookeri Penny ex D. Don N Fig. 133 beach-head iris. Iris canadensis (M. Foster) Peckham; I. setosa Pallas ex Link var. canadensis M. Foster • ME; eastern portion of state. Atlantic coast beaches. 5. Iris prismatica Pursh ex Ker-Gawl. N Fig. 134 slender blue iris. CT, MA, ME, NH, RI; also reported from VT by Seymour (1982), but specimens are unknown; mainly in coastal counties. Fresh to saline marshes, meadows, shorelines. This species has narrow leaf blades 1–7 mm wide. 6. Iris pseudacorus L. E Fig. 135 yellow iris. CT, MA, ME, NH, RI, VT. Marshes, river and lake shores.

Fig. 134  Rhizomes of Iris prismatica.

7. Iris pumila L. E dwarf iris. ME. Edges of lawns, fields, roadsides, compost heaps. 8. Iris sibirica L. E Siberian iris. CT, MA, NH, VT. Edges of lawns, fields, roadsides, compost heaps, waste areas. The report of this species from ME by Campbell et al. (1995) is based on a collection of a cultivated plant— Jun 1957, Richards s.n. (MAINE!). 9. Iris tectorum Maxim. E wall iris. NH. Edges of lawns, fields, roadsides, compost heaps. 10. Iris versicolor L. N Fig. 136, 137 blue iris. CT, MA, ME, NH, RI, VT. Marshes, river and lake shores, wet meadows, ditches. Iris versicolor is the natural hybrid of I. hookeri and I. virginica.

Sisyrinchium Fig. 135  Capsules of Iris pseudacorus.

Identification of Sisyrinchium requires examination of multiple ramets because occasional stems from a given genet will show an unusual character state. For example, it is possible to find “branched stems” among a clump of S. montanum, a species that normally displays “unbranched stems.” Reference: Cholewa and Henderson (2002). 1a. Inflorescences, each of which is subtended by 2 opposed spathe bracts, paired at the summit of the stem and sessile (i.e., without peduncles); spathe bracts closely subtended and partly concealed by a bract-like leaf; tepals white or pale violet . . . . . . . . . . . . . . . . . S. albidum 1b. Inflorescences either occurring singly at the summit or numbering 2–5 and then borne on branch-like peduncles; spathe bracts not closely subtended by leaves; tepals blue to purple (rarely pale blue or white) 2a. Inflorescences sessile, usually solitary at the apex of the stem; stems appearing unbranched [Fig. 139] 3a. Stems 0.9–1.5 (–2) mm wide, with scarcely discernible wing margins; spathes usually suffused with purple; pedicels usually spreading to recurving in fruit; tepals rounded to emarginate at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. mucronatum

Fig. 136  Flower of Iris versicolor.

3b. Stems (1.5–) 2–3.7 mm wide, with obvious wing margins; spathes usually green to bronze, rarely suffused with purple; pedicels usually erect to ascending in fruit; tepals emarginate to retuse at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. montanum

i r i dac e a e   18 1

2b. Inflorescences pedunculate, usually 2–5 per stem; stems appearing branched due to long, branch-like peduncles [Fig. 138] 4a. Stems 2.3–5 mm wide, broadly winged, sometimes with minutely denticulate margins, especially near the base; outer bract of the spathe mostly 18–38 mm long; plants dark olive-green to black-green in drying . . . . . . . . . . . . . . . . . . . . S. angustifolium 4b. Stems 0.8–1.9 mm wide, narrowly winged, with entire margins; outer bract of the spathe mostly 12–18 mm long; plants yellow-green to olive-green in drying 5a. Leaf blades persisting as fibrous tufts at base of plant; ovary green, similar in color to foliage; capsule light to medium brown . . . . . . . . . . . . . . . . . . . . . . . S. fuscatum

Fig. 137  Seed of Iris versicolor.

5b. Leaf blades not persisting as fibrous tufts; ovary dark brown to purple-black, in contrast to lighter foliage; capsule dark brown to purple-black . . . . . . . S. atlanticum 1. Sisyrinchium albidum Raf.

NC

white blue-eyed-grass. Sisyrinchium bermudianum L. var. albidum (Raf.) Gray; S. scabrellum Bickn. • ME. Woodlands, sandy fields, roadsides. This species is considered native to New England by most authors. Its true status in the region is questionable given its highly disjunct location. It is known only as a historical plant from York County, ME. 2. Sisyrinchium angustifolium P. Mill. N Fig. 138 narrow-leaved blue-eyed-grass. Sisyrinchium graminoides Bickn. • CT, MA, ME, NH, RI, VT. Fields, meadows, stream shores, wetland edges. 3. Sisyrinchium atlanticum Bickn. N eastern blue-eyed-grass. Sisyrinchium mucronatum Michx. var. atlanticum (Bickn.) H. Ahles • CT, MA, ME, NH, RI, VT. Fields, meadows, marsh edges. 4. Sisyrinchium fuscatum Bickn.

NC

Fig. 138  Inflorescence of Sisyrhinchium angustifolium.

coastal plain blue-eyed-grass. Sisyrinchium arenicola Bickn. • MA, RI; also reported from CT by Eaton (1958), but specimens are unknown. Mesic to xeric, usually sandy, grasslands. 5. Sisyrinchium montanum Greene N strict blue-eyed-grass.  5a. Sisyrinchium bermudianum L. var. crebrum (Fern.) Boivin; S. montanum Greene ssp. crebrum (Fern.) Böcher; 5b. Sisyrinchium strictum Bickn. • CT, MA, ME, NH, RI, VT. Fields, meadows, open shorelines, forest edges. 1a. Outer spathe bract with margins connate (3.5–) 4–5.7 mm at the base; plants drying dark brown or bronze to olive . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5a. S. montanum var. crebrum Fern. 1b. Outer spathe bract with margins connate 1–3.5 (–4) mm at the base; plants drying green to olive . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5b. S. montanum var. montanum Variety crebrum is known from CT, MA, ME, NH, RI, VT. Variety montanum is known from CT, MA, ME, NH, RI, VT. 6. Sisyrinchium mucronatum Michx. N Fig. 139 needle-tipped blue-eyed-grass. Sisyrinchium intermedium Bickn. • CT, MA, ME, NH, RI, VT. Fields, meadows, open shorelines, forest edges.

Juncaceae 1a. Seeds 3 per capsule, with a caruncle (this modified into a tuft of hairs in L. parviflora); plants pubescent with white hairs, especially at the summit of the leaf sheaths and the basal margins of the leaf blades; leaf sheaths closed (i.e., with connate margins) . . . . . . . . . . . Luzula 1b. Seeds many per capsule, lacking a caruncle (though with white appendages at each end in some species); plants glabrous; leaf sheaths open . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Juncus

Fig. 139  Inflorescence of Sisyrhinchium mucronatum.

182 MO NOCOTS

Juncus References: Hämet-Ahti (1980), Brooks and Clemants (2000). 1a. Involucral bract solitary, of similar cross-sectional shape as the stem and appearing to be a continuation of it, therefore, the inflorescence appearing to emerge from the side of the stem (i.e., pseudolateral) [Figs. 146, 150]; leaves bladeless, consisting of sheaths only 2a. Stems cespitose; flowers with 3 stamens (or with 6 stamens in the rare introduction J. inflexus) 3a. Capsules red-brown to dark brown, (2.5–) 3–4 mm long; stamens numbering 6 per flower, with filaments 0.8–1.5 mm long and anthers 0.8–1 mm long; stems prominently glaucous in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. inflexus 3b. Capsules tan to green-tan or brown, 1.5–3.2 mm long; stamens numbering 3 per flower, with filaments 0.5–0.8 mm long and anthers 0.5–0.8 mm long; stems green 4a. Apical portion of stem relatively lustrous, smooth or nearly so below the inflorescence, the (25–) 30–60 longitudinal striations inconspicuous until drying [Fig. 146]; ridges of dried stems capped with dull, low cells . . . . . . . . . . . . . . . J. effusus 4b. Apical portion of stem relatively dull, evidently ridged below the inflorescence with mostly 10–30 longitudinal grooves [Fig. 150]; ridges of dried stems capped with lustrous, papillose cells 5a. Involucral bract swollen at the base of the inflorescence, sometimes somewhat reflexed in fruit; leaf sheaths with red-brown bases, the upper ones 15–23 cm long; inflorescence compact, mostly 10–25 mm in diameter; stems with 12–30 longitudinal ridges; tepals spreading from the base in fruit . . . . . J. conglomeratus 5b. Involucral bract not swollen, erect in fruit; leaf sheaths with dark red-brown to purple-black bases, the upper ones 5–12 cm long; inflorescence relatively open, mostly 15–80 mm in diameter; stems with 10–20 longitudinal ridges [Fig. 150]; tepals ascending or appressed to the capsule in fruit. . . . . . . . . . . . . . . . . . . J. pylaei 2b. Stems produced singly or a few together from an elongate rhizome; flowers with 6 stamens 6a. Anthers 3–5 times as long as the filaments; tepals usually with a dark stripe on each side of the midvein [Fig. 143]; stems smooth in life, irregularly wrinkled when dry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. balticus 6b. Anthers 0.3–0.5 times as long as the filaments; tepals lacking dark stripes; stems finely grooved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. filiformis 1b. Involucral bracts usually 2 or more, differentiated from the stem in some manner, the inflorescence appearing terminal [Fig. 151]; leaves with blades 7a. Flowers occurring singly on pedicels [Fig. 142]; leaves variously flat to terete, never septate; bracteoles present, these a pair of small bracts at the base of the flowers in addition to the pair of bracts at the base of the pedicels 8a. Plants annual with fibrous roots; inflorescence often occupying 33–90% of the total plant height; leaves without auricles 9a. Inner tepals acute to acuminate, exceeding the capsule [Fig. 144]; capsules mostly acute to subacute at apex (rarely truncate); inflorescences relatively open . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. bufonius 9b. Inner tepals rounded to acute at the apex, many equaling or shorter than the capsule [Fig. 141]; capsules mostly truncate at apex; inflorescences relatively dense . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. ambiguus 8b. Plants perennial; inflorescences rarely occupying more than 25% of the total plant height; leaves with auricles

jun c ac e a e   183

10a. Auricles erose; inflorescence of 1–3 flowers; capsules with a beak up to 0.7 mm long; margins of leaf blades minutely serrulate . . . . . . . . . . . . . . . . . . . . . . . . . . J. trifidus 10b. Auricles entire; inflorescence with several to many flowers; capsules usually beakless (rarely shortly beaked; e.g., J. gerardii); margins of leaf blades entire 11a. Leaves terete or caniculate; tepals shorter than and appressed to the capsule; capsules 3-locular, the septa meeting in the middle 12a. Seeds with inconspicuous pale tails at each end less than 0.2 mm long; capsules 2.9–3.5 (–4) mm long, brown to dark brown; inner tepals mostly 1.9–3.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. greenei 12b. Seeds with conspicuous, slender, slightly curved pale tails at each end 0.2–0.4 mm long; capsules (3.3–) 3.8–4.7 mm long, golden brown to light brown; inner tepals mostly 3.2–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. vaseyi 11b. Leaves flat or becoming involute (nearly terete in some J. dichotomus); tepals equal to or longer than the capsule (often shorter in J. compressus), ascending to spreading; capsules partially 3-locular, the septa not meeting in the center or 3-locular in J. gerardii and J. secundus 13a. Leaves not confined to the base of the stem, some borne on the upper ½ of the stem; tepals with a dark-stripe on each side of the midvein, the outer with an obtuse apex; stems produced singly or a few together from elongate rhizomes (varying to subcespitose in J. compressus); plants of saline habitats 14a. Anthers 1.1–1.6 (–1.8) mm long, 2–4 times as long as the filament; capsules usually shorter than to equaling the length of the tepals; primary bract of the inflorescence mostly 1–5 cm long, usually not conspicuously exceeding the inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. gerardii 14b. Anthers 0.6–1 mm long, 1–2 times as long as the filament; capsules usually exceeding the tepals; primary bract of the inflorescence 2–7.5 (–13) cm long, often conspicuously exceeding the inflorescence . . . . . . J. compressus 13b. Leaves confined primarily to the lower ⅓ of the stem; tepals lacking dark stripes, the outer acuminate at the apex; stems cespitose; plants of nonsaline habitats 15a. Auricles scarious, (1–) 1.5–5 mm long; sheath margin pliable, transparent 16a. Monochasial branches of inflorescence erect to divergent, with crowded to remote flowers, the internodes less than 6 mm long [Fig. 151]; ultimate branches of monochasia 1–2 cm long; tepals (2.8–) 3.5–4.5 mm long; capsules more than 75% of the length of the tepals [Fig. 151]; plants usually shorter than 7 dm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. tenuis 16b. Monochasial branches of inflorescence erect to ascending, often with inwardly curving branches, with widely scattered flowers, the internodes 6 mm or longer [Fig. 142]; ultimate branches of monochasia mostly 3–5 cm long; tepals 2.5–3.5 (–4) mm long; capsules less than 75% of the length of the tepals [Fig. 142]; plants mostly 7–9 dm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. anthelatus 15b. Auricles scarious to cartilaginous, 0.2–0.6 mm long; sheath margin of firmer texture, often brittle, transparent to opaque 17a. Capsule 3-locular; placentation axile; leaves relatively short, often not exceeding midpoint of stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. secundus 17b. Capsules only partially 3-locular, the incomplete septa not meeting in the center; placentation parietal; leaves usually extending past midpoint of stem

184  MONOCOTS

18a. Leaves caniculate to nearly terete, rarely flat; inflorescence varying from compact to open and diffuse, usually with 10–35 or more flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. dichotomus 18b. Leaves flat, though often involute in age or drying; inflorescence usually compact and few-flowered, often with 10 or fewer flowers, rarely more than 15 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. dudleyi 7b. Flowers borne in glomerules (sometimes solitary in J. pelocarpus and J. subtilis); leaves both terete and septate (except in J. biflorus, J. ensifolius, J. marginatus, and J. stygius); bracteoles absent, flowers subtended by only a pair of bracts at the base of the pedicel 19a. Leaf blades compressed and equitant (i.e., set edge to stem), imperfectly septate (i.e., the septa inconspicuous and not continuous through the blade width) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. ensifolius 19b. Leaf blades elliptic to terete in cross-section, septate (imperfectly septate in J. stygius) or flat and non-septate, never equitant 20a. Capsules 5.5–7 (–9) mm long; seeds 2.8–3.5 mm long including the white tails; leaves terete to elliptic in cross-section, imperfectly septate . . . . . . . . . . . . . . J. stygius 20b. Capsules 1.5–5.2 mm long; seeds 0.3–1.9 mm long; leaves either flat or both terete and septate 21a. Leaves flat, without septa; capsules subglobose, often with red flecks [Fig. 148] 22a. Widest leaf blades (1.3–) 1.6–2.6 (–3.5) mm wide; anthers (0.2–) 0.3– 0.5 (–0.7) mm long, concealed by the tepals; tallest stems (19.2–) 26–44 (–56.8) cm tall; sheath of lowest leaf (17–) 22–38 (–47) mm long; base of stem (0.4–) 2– 4.4 (–6) mm thick; inflorescence with (2–) 5–15 (–30) glomerules . . . J. marginatus 22b. Widest leaf blades (2.6–) 3.1–4.5 (­­–5.4) mm wide; anthers (0.5–) 0.6– 1 (–1.3) mm long, exserted beyond tepals; tallest stems (27.2–) 50.8–81.2 (–100.7) cm tall; sheath of lowest leaf (32–) 43–78 (–97) mm long; base of stem (3.4–) 5.8– 9.6 (–12) mm thick; inflorescence with 20–100 (–200) glomerules . . . . J. biflorus 21b. Leaves terete and septate; capsules mostly obloid to ellipsoid or ovoid, without red flecks 23a. Seeds 0.7–1.9 mm long including the pale tails at each end [Fig. 145], the body of the seed covered with a translucent, pale white veil (note: view at 20× magnification); flowers usually with 3 stamens 24a. Seeds 1.1–1.9 mm long, each tail 50–80% as long as the seed body [Fig. 145]; glomerules 5- to 50-flowered . . . . . . . . . . . . . . . . . . . . J. canadensis 24b. Seeds 0.7–1.2 mm long, each tail 10–40% as long as the seed body; glomerules 2- to 10-flowered, rarely with as many as 20 flowers in J. subcaudatus 25a. Outer tepals obtuse to subacute at the apex, soft, with conspicuous scarious margins, 1.8­–2.5 mm long; capsules 2.4–3.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. brachycephalus 25b. Outer tepals sharply acute to acuminate, rigid, with narrow scarious margins, 2.1–3.2 mm long; capsules 3–4.8 mm long 26a. Branches of inflorescence erect to strictly ascending; capsules 3.2–4.8 mm long, prominently exserted, extending more than 0.7 mm beyond perianth; glomerules with 2–5 (–7) flowers . . . . . J. brevicaudatus 26b. Branches of inflorescence widely ascending to spreading; capsules 3–3.8 mm long, weakly to prominently exserted, not exceeding tepals by more than 0.6 (–1.3) mm; glomerules with 2–10 (–20) flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. subcaudatus

j un c ac e a e   18 5

23b. Seeds 0.3–0.7 mm long, merely pointed at the ends, the body clear yellow-brown to red-brown; flowers with 3 or 6 stamens 27a. Flowers solitary or paired, sometimes replaced by vegetative bulbils [Fig. 149] 28a. Stems upright; inflorescences many-flowered; outer tepals 1.6–2.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. pelocarpus 28b. Stems creeping or floating; entire plant with only 1–3 flowers; outer tepals (1.8–) 2.2–2.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. subtilis 27b. Flowers in glomerules of 2–100, not replaced by bulbils 29a. Leaves dimorphic—those produced by the rhizome submersed and capillary, those produced by the stem emergent and thicker; lowest stem leaf overtopping inflorescence, 3–6 mm thick at midpoint . . . . J. militaris 29b. Leaves monomorphic, none capillary and produced by the rhizome; lowest stem leaf shorter than height of plant, mostly 0.5–3 mm thick 30a. Glomerules spheric or subspheric, comprised of (6–) 10–100 flowers 31a. Inflorescences composed of 24–60 glomerules; inner tepals obtuse to subacute at the apex . . . . . . . . . . . . . . . . . . . . . . J. pervetus 31b. Inflorescences composed of 1–23 glomerules; inner tepals acuminate at the apex 32a. Plants cespitose, without elongate rhizomes; tepals lanceolate to narrow-ovate. . . . . . . . . . . . . . (in part) J. acuminatus 32b. Plants not cespitose, the stems produced singly or a few together from elongate rhizomes; tepals subulate 33a. Flowers with usually 3 stamens; capsules included within the perianth, the tips of the valves separate after dehiscence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. brachycarpus 33b. Flowers with usually 6 stamens; capsules equaling tepals to exserted, the tips of the valves often cohering after dehiscence 34a. Auricles 0.5–1 (–1.7) mm long, cartilaginous, dark yellow; glomerules composed of 6–25 (–30) flowers; inner and outer tepals of ± similar length . . . . . . . . . . . J. nodusus 34b. Auricles (1–) 2–4 mm long, scarious, pale brown; glomerules composed of 25–100 flowers; inner tepals shorter than outer tepals . . . . . . . . . . . . . . . . . . . . . . . J. torreyi 30b. Glomerules hemispheric to obconic, comprised of 2–20 flowers 35a. Flowers with usually 3 stamens; plants cespitose; capsules light brown, 1-loculuar 36a. Capsule approximately equal in length to tepals; glomerules composed of 5–20 (or more) flowers . . . . (in part) J. acuminatus 36b. Capsule evidently exserted beyond tepals; glomerules composed of 2–10 flowers 37a. Capsules 2.8–3.7 (–4.2) mm long, approximately 1.3–1.5 times as long as the tepals . . . . . . . . . . . . . . . . . . . . . . . . J. debilis 37b. Capsules 4–5.2 mm long, approximately 2 times as long as the tepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. diffusissimus

186  MONOCOTS

35b. Flowers with usually 6 stamens; plants not cespitose, produced singly or a few together from elongate rhizomes (sometimes subcespitose in J. articulatus); capsules brown to dark brown, imperfectly 3-locular (i.e., the septa not extending all the way to the center) 38a. Capsules rounded to obtuse at apex, with a mucronate tip [Fig. 140]; inner tepals acute to obtuse at apex [Fig. 140]; branches of inflorescence strictly ascending . . . . . . . . . J. alpinoarticulatus 38b. Capsules acute (seldom obtuse) at apex; inner tepals acute to acuminate at apex; branches of inflorescence widely ascending to spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . J. articulatus

Fig. 140  Tepals and capsule of Juncus alpinoarticulatus.

1. Juncus acuminatus Michx. N sharp-fruited rush. Juncus pelocarpus E. Mey. var. crassicaudex Engelm. • CT, MA, ME, NH, RI, VT. Fresh to brackish-tidal river shores, marshes. 2. Juncus alpinoarticulatus Chaix ex Vill. ssp. americanus (Farw.) Hämet-Ahti N C Fig. 140 northern green rush. Juncus alpinoarticulatus Vill. ssp. nodulosus (Wahlenb.) Hämet-Ahti, sensu American authors; J. alpinus Vill. var. americanus Farw.; J. alpinus Vill. var. rariflorus Hartman, sensu American authors • ME, NH, VT; also reported from MA by Hämet-Ahti (1986), but specimens are unknown. Lake shores and ice-scoured river shores in regions of highpH bedrock or till, fresh-tidal river marshes, fen-like meadows. Known in other regions to hybridize with Juncus articulatus, J. brevicaudatus, and J. nodosus (among others). Fig. 141  Tepals and capsule of Juncus ambiguus.

3. Juncus ambiguus Guss.

N C Fig. 141

seaside toad rush. Juncus bufonius L. var. halophilus Buch. & Fern. • CT, MA, ME. Upper edges of saline marshes, low areas in fields and other open areas near the Atlantic coast. See Haines (2000c) and references cited therein for rationale for recognition of this species. 4. Juncus anthelatus (Wieg.) R.E. Brooks N Fig. 142 Wiegand’s rush. Juncus tenuis Willd. var. anthelatus Wieg. • CT, MA, ME, NH, RI, VT. Wet fields, meadows, low powerline rights-of-way, open margins of swamps. 4 × 35. This rare hybrid has been collected from ME (Haines 2001a). It resembles Juncus anthelatus in its stature and tall inflorescence with long internodes. However, the internodes of the inflorescence are straight (rather than often curved or arching), and the capsules are more than 75% as long as the tepals (rather than shorter than 75% as long as the tepals). 5. Juncus articulatus L. N Fig. 142  Inflorescence of Juncus anthelatus with detail of tepals and capsule.

joint-leaved rush. Juncus articulatus L. var. obtusatus Engelm. • CT, MA, ME, NH, RI, VT. Shorelines, ditches, wet sand. Known in other regions to hybridize with Juncus alpinoarticulatus and J. brevicaudatus. 6. Juncus balticus Willd. ssp. littoralis (Engelm.) Snogerup N Fig. 143 Baltic rush. Juncus arcticus Willd. ssp. littoralis (Engelm.) Hultén; J. arcticus Willd. var. littoralis (Engelm.) Boivin; J. balticus Willd. var. littoralis Engelm. • MA, ME, NH, VT. Saline marshes, salted roadsides, fens. Juncus balticus is becoming increasingly frequent along roads and freeways as a result of salting for ice and snow melting. This species has been variously treated as part of the variation displayed by J. arcticus Willd. or, in the narrower sense, as part of distinct species. Snogerup et al. (2002) discussed this complex and presents arguments for treating J. balticus as distinct from J. arcticus. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species could be in RI (i.e., the author is unaware of any collections).

Fig. 143  Tepals and capsule of Juncus balticus showing dark stripes on each side of tepal midvein.

7. Juncus biflorus Ell.

NC

large grass-leaved rush. Juncus aristulatus Michx. var. biflorus Small; J. marginatus Rostk. var. biflorus Wood; J. marginatus Rostk. var. odoratus Torr.; J. odoratus (Torr.) Steud. • MA; coastal

jun c ac e a e   187

plain. Wet, sandy or peaty soils of pond shores. This species (and others) is often included in Juncus marginatus by some authors (e.g., Brooks and Clemants 2000). However, recent research by Knapp and Naczi (2008) shows that multiple taxa should be recognized. 8. Juncus brachycarpus Engelm. E white-root rush. CT, MA, RI. Brackish and saline marshes, Atlantic coast shores, low areas near the coast. 9. Juncus brachycephalus (Engelm.) Buch.

NC

small-headed rush. CT, MA, ME, NH, VT. Meadows, lake shores, slides, lakeside fens, and ice-scoured river shores in regions of high-pH bedrock or till. 10. Juncus brevicaudatus (Engelm.) Fern. N short-tailed rush. CT, MA, ME, NH, RI, VT. Lake and river shores, ditches, and other low, wet areas, tolerant of human disturbance. Known in other regions to hybridize with Juncus alpinoarticulatus and J. articulatus.

Fig. 144  Tepals and capsule of Juncus bufonius.

11. Juncus bufonius L. N Fig. 144 toad rush. CT, MA, ME, NH, RI, VT. Roadsides, meadows, edges of lawns, shorelines, coastal beaches, and seasonally wet, open areas. 12. Juncus canadensis J. Gay ex Laharpe N Fig. 145 Canada rush. Juncus canadensis J. Gay ex Laharpe var. sparsiflorus Fern. • CT, MA, ME, NH, RI, VT. Marshes, shorelines, wetland edges, sometimes an emergent in shallow water. 13. Juncus compressus Jacq. E round-fruited rush. ME, VT. Shoulders and ditches of salted roadways, railroad embankments, disturbed saline soil. 14. Juncus conglomeratus L. E bunch-flowered soft rush. Juncus effusus L. var. conglomeratus (L.) Engelm. • CT. Open wetlands. 15. Juncus debilis Gray

Fig. 145  Seed of Juncus canadensis.

NC

weak rush. CT, MA, RI. Low areas in fields, edges of pools, roadside ditches, damp coastal sands. 16. Juncus dichotomus Ell. N forked rush. Juncus platyphyllus (Wieg.) Fern.; J. tenuis Willd. var. dichotomus (Ell.) Wood • CT, MA, ME, NH, RI. Shorelines, ditches, upper border of coastal marshes, usually in mesic to

hydric, sandy soil. 17. Juncus diffusissimus Buckley E slim-fruited rush. CT, MA. Shorelines, backwater canals, sand and silt bars, disturbed wet ground. 18. Juncus dudleyi Wieg. N

Fig. 146  Finely striate upper stem and inflorescence of Juncus effusus ssp. solutus.

Dudley’s rush. Juncus tenuis Willd. var. dudleyi (Wieg.) F.J. Herm. • CT, MA, ME, NH, RI, VT. River shores, meadows, borrow pits, usually in regions of high-pH bedrock or till. 19. Juncus effusus L. n Fig. 146, 147 common soft rush.  19a. Juncus effusus L. var. solutus Fern. & Wieg.; 19b. Juncus effusus L. var. compactus Lej. & Court. • CT, MA, ME, NH, RI, VT. Graminoid marshes, ditches, shorelines, meadows, low areas in fields. Juncus effusus has been treated variously by different authors, most commonly as being comprised of several varieties. However, some recent treatments (Brooks and Clemants 2000) abandon recognition of any infraspecific taxa. Unfortunately, this approach conceals a great deal of variation in (and understanding of) this complex. Treatment of this species complex follows the work of Hämet-Ahti (1980; considered appropriate by Snogerup et al. 2002). See also J. conglomeratus and J. pylaei, traditionally considered as infraspecific taxa of J. effusus (here treated as separate species).

Fig. 147  Tepals and capsule of Juncus effusus ssp. solutus.

188  MONOCOTS

1a. Tepals ascending or appressed to the capsule [Fig. 147]; leaf sheaths on reproductive stems (12–) 15–27 cm long, usually loose and not clasping the stem (sometimes even unrolled), with margins overlapping only in the basal half (if at all), usually lacking a dark, marginal band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19a. J. effusus ssp. solutus (Fern. & Wieg.) Hämet-Ahti 1b. Tepals spreading from the base in fruit; leaf sheaths on reproductive stems 6–14 cm long, usually clasping the stem, with margins overlapping except for the apical 2–4 cm, usually with a dark, marginal band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19b. J. effusus ssp. effusus Subspecies solutus is native to New England and known from CT, MA, ME, NH, RI, VT. Subspecies effusus is introduced and is known from MA, ME, RI. The latter subspecies varies in the congestion of its flowers (i.e., from tightly compact to open inflorescences). 20. Juncus ensifolius Wikstr. var. ensifolius E sword-leaved rush. Juncus xiphioides E. Mey. var. triandrus Engelm. • RI, VT. Shorelines, ditches, marshes. 21. Juncus filiformis L. N thread rush. MA, ME, NH, VT. Low, wet sandy areas, river shores, peaty and gravelly pond shores, rarely on alpine ridges and plateaus. 22. Juncus gerardii Loisel. n saltmarsh rush. Juncus gerardii Loisel. var. pedicellatus Fern. • CT, MA, ME, NH, RI, VT. Saline marshes, shoulders and ditches of salted roadways. Introduced in VT. Fig. 148  Tepals and capsule of Juncus marginatus.

23. Juncus greenei Oakes & Tuckerman N Greene’s rush. CT, MA, ME, NH, RI, VT. Sandplains, dry fields, sandy road shoulders, rock outcrops. 24. Juncus inflexus L. E European meadow rush. MA. Stream banks, ditches, disturbed wet places. 25. Juncus marginatus Rostk. N Fig. 148 grass-leaved rush. CT, MA, ME, NH, RI, VT. Low fields, pond shores, edges of pools, abandoned borrow pits, often on moist to wet sand. 26. Juncus militaris Bigelow N bayonet rush. CT, MA, ME, NH, RI, VT. Shallow water of lakes and slow-moving rivers, on sand, silt, and mucky substrates. 27. Juncus nodosus L. N

Fig. 149  Inflorescence of Juncus pelocarpus showing proliferations.

knotted rush. CT, MA, ME, NH, RI, VT. River shores, especially in seeps and also pools associated with outcrops, lake shores, meadows, usually in regions of high-pH bedrock and till. Known in other regions to hybridize with Juncus alpinoarticulatus. 28. Juncus pelocarpus E. Mey. N Fig. 149 brown-fruited rush. CT, MA, ME, NH, RI, VT. Sandy, muddy, or peaty shorelines and wetlands, including coastal plain pond shores, abandoned borrow pits, and fens. 29. Juncus pervetus Fern.

NC

blunt-flower rush. MA. Brackish marshes. Not collected since 1928 and believed extinct by many authors. Its origin is debated. The possession of abortive ovules in the capsules suggests hybrid origin. 30. Juncus pylaei Laharpe N Fig. 150

Fig. 150  Coarsely striate upper stem and inflorescence of Juncus pylaei.

Pylae’s soft rush. Juncus effusus L. var. decipiens, auct. non Buch.; J. effusus L. var. pylaei (Laharpe) Fern. & Wieg. • CT, MA, ME, NH, RI, VT. Graminoid marshes, ditches, shorelines, meadows, low areas in fields. Juncus pylaei and J. effusus ssp. solutus are the two native members of the J. effusus complex in New England (and are the most common taxa in that complex). In addition to characters provided in the key, they differ in leaf sheath and inflorescence morphology. Juncus pylaei has longest sheaths on reproductive stems 5–12 cm long that are dark red-brown to purple-black (especially dark near the base) and longest

j un c ac e a e   18 9

inflorescence branches (10–) 20–30 (–50) mm long. Juncus effusus ssp. solutus has longest sheaths on reproductive stems (12–) 15–27 cm long that are pale brown to red-brown (rarely dark brown) and longest inflorescence branches (30–) 40–100 mm long. 31. Juncus secundus Beauv. ex Poir. N lopsided rush. CT, MA, ME, NH, RI, VT. Dry-mesic to xeric, often shallow, soils, including ledges, balds, and sandy areas. 32. Juncus stygius L. ssp. americanus (Buch.) Hultén

NC

moor rush. Juncus stygius L. var. americanus Buch. • ME, NH. Circumneutral fens, often adjacent to or in the vicinity of open pools. 33. Juncus subcaudatus (Engelm.) Coville N somewhat-tailed rush. CT, VT. Marshes, openings in swamps, ditches. The species was attributed to ME by Brooks and Clemants (2000) based on an immature specimen (Steven Clemants, personal communication). It is not accepted here given the southern New England distribution of this plant. Reports of this taxon in MA (e.g., Sorrie and Somers 1999) are based on misidentified voucher specimens of J. acuminatus and J. canadensis. 34. Juncus subtilis E. Mey.

NC

greater creeping rush. ME; northern portions of state. Muddy shores of lakes and rivers. 35. Juncus tenuis Willd. N Fig. 151 path rush. Juncus macer S.F. Gray; J. tenuis Willd. var. williamsii Fern. • CT, MA, ME, NH, RI, VT. Wide variety of open sites and edges, ranging in hydrology from xeric to wet-mesic, including fields, disturbed lots, trail margins, ditches, and shorelines. 35 ‌ × 38. Juncus ×oronensis Fern. is a historically occurring hybrid known from two separate locations in ME (Franklin and Penobscot Counties). It has caniculate leaf blades (i.e., terete with a channel on the upper surface), seeds 0.6–0.8 mm long with short white tails (the seeds of J. tenuis are mostly 0.6 mm or shorter and those of J. vaseyi are longer than 0.8 mm), and conspicuously secund flowers that mature as capsules shorter than the sharply pointed tepals. 36. Juncus torreyi Coville

NC

Torrey’s rush. MA, ME, VT; also reported from NH by Brooks and Clemants (2000), but specimens are unknown. Stream shores and ditches in regions of high-pH bedrock, tidal river shores. 37. Juncus trifidus L. N highland rush. Juncus trifidus L. ssp. carolinianus Hämet-Ahti; J. trifidus L. var. monanthos (Jacq.) Bluff ex Fingerhuth • ME, NH, VT. Boreal and alpine cliffs, ridges and plateaus above treeline, rarely weedy along trails just below treeline. 38. Juncus vaseyi Engelm.

NC

Vasey’s rush. Juncus greenei Oakes & Tuckerman var. vaseyi (Engelm.) Boivin • ME, VT. High-pH river shore outcrops, limestone headlands, low boggy fields and ditches, rarely on roadsides and other places with disturbed soil.

Luzula Seed length measurements in the key include the caruncle. Reference: Swab (2000). 1a. Flowers occurring singly (rarely paired) at the tips of branches 2a. Inflorescence an umbel-like cyme, with simple or once-forked branches; seeds with a conspicuous caruncle; plants of low-elevation woodlands, edges, and forest openings, flowering in April through early June . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. acuminata 2b. Inflorescence a branched dichasial cyme, with several-forked branches; seeds with a tuft of hairs; plants primarily of boreal and subalpine habitats, flowering from mid-June through July . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. parviflora

Fig. 151  Inflorescence of Juncus tenuis with detail of tepals and capsule.

19 0 MO NOCOTS

1b. Flowers clustered in glomerules of 3–12 (–20), the glomerules all at the tips of branches or 1 or 2 of the central ones sessile (rarely all congested) [Figs. 153, 154, 155] 3a. Flowers in 3- to 6-flowered glomerules, these arranged in an open-branched, dichasial cyme; tepals white (rarely pink) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. luzuloides 3b. Flowers mostly in 5- to 12-flowered glomerules, these arranged in an umbel-like or spike-like cyme; tepals light brown to dark brown (rarely nearly white) 4a. Leaves involute, pointed at the tip; bracts subtending individual glomerules of flowers with conspicuously long-ciliate margins; caruncles of seeds inconspicuous, up to 0.2 mm long; branches of inflorescence, when present, arching or drooping; plants of alpine and subalpine habitats 5a. Well-developed inflorescences nodding [Fig. 155]; bracts subtending glomerules of flowers with long, slender tips, often projecting beyond the glomerules . . . . . L. spicata 5b. Inflorescences congested or with arching branches, not nodding; bracts subtending glomerules of flowers obtuse to acuminate at apex, not projecting beyond the glomerules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. confusa 4b. Leaves ± flat, usually with a blunt, callous tip (the callous tip obscure in L. echinata); bracts subtending individual glomerules entire or sparingly lacerate-margined; caruncles of seeds mostly conspicuous, 0.2–0.7 mm long; branches of inflorescence ± straight [Figs. 153, 154]; plants of low-elevation habitats 6a. Leaves with a minute point at the tip, the calloused apex narrow and obscure; anther 2–4 times as long as the filament . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. echinata 6b. Leaves with a blunt, rounded tip, the calloused apex evident; anther not more than 2 times as long as filament (except in L. campestris, a rare introduction) 7a. Plants bearing pale, swollen leaf bases that appear as bublets [Fig. 152] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. bulbosa 7b. Plants without pale, swollen leaf bases 8a. Stems solitary or a few together; plants with short but evident rhizomes; anther 2–6 times as long as the filament . . . . . . . . . . . . . . . . . . . . . . L. campestris 8b. Stems loosely to densely cepsitose; plants without evident rhizomes; anther 1–2 times as long as the filament or shorter 9a. Seeds 0.7–1 mm long, with a caruncle 0.2–0.3 mm long; tepals 1.5–2.6 mm long, clear to pale brown; styles 0.2–0.3 mm long . . . . L. pallidula 9b. Seeds 1.1–1.7 mm long, with a caruncle 0.2–0.6 mm long; tepals (2–) 2.5–3.5 mm long, nearly white to dark brown; styles 0.4–0.7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. multiflora 1. Luzula acuminata Raf.

NC

hairy wood rush. 1a. Luzula carolinae S. Wats.; 1b. Juncoides saltuense (Fern.) Heller; Luzula carolinae S. Wats. var. saltuensis (Fern.) Fern. • CT, MA, ME, NH, VT. Deciduous or mixed evergreen-deciduous forests and woodlands, forest edges. 1a. Most or all of the primary branches of the inflorescence branched 1 or more times and bearing 2–4 flowers . . . . . . . . . . . . . . . . . . . . . . . 1a. L. acuminata var. carolinae (S. Wats.) Fern. 1b. Most or all of the primary branches of the inflorescence simple and bearing a single flower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. L. acuminata var. acuminata Variety carolinae is known from only CT and is of regional conservation concern. The record is based on an herbarium specimen examined by Janice Swab (personal communication). Unfortunately, the location of that specimen is now unknown. Variety acuminata is known from CT, MA, ME, NH, VT and is demonstrably secure in New England.

j un c ac e a e   19 1

2. Luzula bulbosa (Wood) Rydb. N Fig. 152 bulbous wood rush. Juncoides bulbosum (Wood) Small; Luzula campestris (L.) DC. var. bulbosa Wood • CT, MA. Sandy soils of woodlands, dry fields, and forest openings. 3. Luzula campestris (L.) DC. E field wood rush. Juncoides campestre (L.) Kuntze • MA. Lawns, roadsides, fields, disturbed places. 4. Luzula confusa Lindeberg

NC

northern wood rush. Juncoides hyperboreum (R. Br.) Sheldon, in part; Luzula hyperborea R. Br., in part • ME, NH. Alpine ridges, cliffs, and slides. 5. Luzula echinata (Small) Hermann

N C Fig. 153

hedgehog wood rush. Juncoides echinatum Small; Luzula campestris (L.) DC. var. echinata (Small) Fern. & Wieg.; L. multiflora (Ehrh.) Lej. var. echinata (Small) Mohlenbrock • CT, MA. Deciduous forests, forested slopes, fields.

Fig. 152  Bulblet-like leaf bases of Luzula bulbosa.

6. Luzula luzuloides (Lam.) Dandy & Wilmott ssp. luzuloides E oak-forest wood rush. Juncoides nemorosum (Pollard) Kuntze • CT, MA, ME, VT. Forests, woodlands, fields, roadsides. 7. Luzula multiflora (Ehrh.) Lej. N Fig. 154 common wood rush.  7a. Luzula campestris (L.) DC. var. multiflora (Ehrh.) Čelak.;  7b. Juncoides intermedium (Thuill.) Rydb.; Luzula campestris (L.) DC. var. frigida Buch.; L. frigida (Buch.) Samuelsson; L. intermedia (Thuill.) A. Nels.; L. multiflora (Ehrh.) Lej. var. fusconigra Čelak. • CT, MA, ME, NH, RI, VT. Lawns, fields, forests, forest openings and edges. 1a. Tepals pale brown to brown near the center, all pointed at the apex; capsules light brown to brown; caruncle of seeds 0.3–0.6 mm long . . . . . . . . . . . . . 7a. L. multiflora ssp. multiflora

Fig. 153  Inflorescence of Luzula echinata.

1b. Tepals dark brown near the center, the outer pointed, the inner truncate-mucronate; capsules dark brown to nearly black; caruncle of seeds 0.2–0.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7b. L. multiflora ssp. frigida (Buch.) Krecz. Subspecies multiflora is known from CT, MA, ME, NH, RI, VT. Subspecies frigida is known from MA, ME, NH, VT. 8. Luzula pallidula J. Kirschner E pale European wood rush. Luzula campestris (L.) DC. var. pallescens Wahlenb.; Luzula pallescens (Wahlenb.) Bess. • VT. Fields, woodlands, forest openings. 9. Luzula parviflora (Ehrh.) Desv. ssp. melanocarpa (Michx.) Hämet-Ahti N small-flowered wood rush. Juncus melanocarpus Michx.; Luzula melanocarpa (Michx.) Desv.; L. parviflora (Ehrh.) Desv. var. melanocarpa (Michx.) Buch. • MA, ME, NH, VT. Stream shores, gullies, mossy areas, and trail sides in boreal and subalpine areas. 10. Luzula spicata (L.) DC.

N C Fig. 155

spiked wood rush. Juncoides spicatum (L.) Kuntze • ME, NH, VT; northern portion of states. Shorelines of tarns and mountain brooks, snowbank communities, alpine lawns.

Fig. 154  Inflorescence of Luzula multiflora.

Fig. 155  Inflorescence of Luzula spicata.

19 2 MO NOCOTS

Juncaginaceae Triglochin Reference: Haynes and Hellquist (2000b). 1a. Leaves usually equaling the height of the flowering stem, arching outward from the sheath [Fig. 156], 0.5–1 mm wide; racemes 2–5 (–7) cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. gaspensis 1b. Leaves usually shorter than the flowering stem, erect, 0.8–1.4 mm wide; racemes 6–45 cm tall 2a. Flowers with 6 carpels and 6 stigmas; mature schizocarps 1.5–2 mm wide, ca. 2 times as long as wide, the cluster of 6 mericarps on a slender, non-wing-angled axis; ligule apically 2-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. maritima 2b. Flowers with 3 carpels and 3 stigmas; mature schizocarps 0.8–1.2 mm wide, ca. 5–7 times as long as wide, the cluster of 3 mericarps on a triquetrous axis; ligule not lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. palustre 1. Triglochin gaspensis Lieth & D. Löve

N C Fig. 156

Gaspe Peninsula arrow-grass. ME; far-eastern portion of state. Saline marshes with low vegetation. This species is sometimes difficult to separate from small plants of Triglochin maritima. During flowering, the length of stigmatic papillae serves as another useful character in museum settings—mostly 0.09–0.23 mm long (mean=0.16 mm) in T. gaspensis and mostly 0.13–0.37 mm long (mean=0.25 mm) in T. maritima (longest papillae measured in both instances). 2. Triglochin maritima L. N Fig. 156  Habit of Triglochin gaspensis showing leaves that arch outward from sheath apex.

saltmarsh arrow-grass. Triglochin debilis (M.E. Jones) A. & D. Löve; T. maritima L. var. elata (Nutt.) Gray • CT, MA, ME, NH, RI, VT. Saline and brackish marshes, fens. 3. Triglochin palustre L. N marsh arrow-grass. ME, RI; also reported from NH by Haynes and Hellquist (2000b), but specimens are unknown. Saline marshes, ice-scoured river shores.

Liliaceae 1a. Tepals white and spotted with red or purple on the adaxial surface, the outer 3 prominently saccate at the base; each style apically bifid, maculate . . . . . . . . . . . . . . . . Tricyrtis 1b. Tepals variously colored, but not white with dark spots, not prominently saccate; styles neither bifid near apex nor spotted 2a. Tepals green-yellow, green-white, or pink (red-purple to purple in the rare hybrid Streptopus × ‌oreopolus), 0.6–1.6 cm long; fruit a berry; stems arising from rhizomes (the rhizomes tuberous in Medeola) 3a. Styles 3, distinct; tepals recurved from near the base; leaves borne on an aerial stem in 1 or 2 whorls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Medeola 3b. Style 1, the stigma obscurely lobed to 3-parted; tepals spreading or recurving from near the middle; leaves all basal or alternate 4a. Flowers in an umbel at the summit of a scape, with green-yellow tepals; leaves all basal; fruit a blue berry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clintonia

l i l i ac e a e   19 3

4b. Flowers solitary or paired in the axils of foliage leaves, with green-white, pink, or red-purple to purple tepals; leaves borne alternately on an aerial stem; fruit a red berry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Streptopus 2b. Tepals yellow, orange, red or white, (1.3–) 1.5–10 cm long; fruit a capsule; stems arising from bulbs or corms 5a. Leaves appearing all basal (actually borne on a sender, mainly subterranean stem), usually mottled with lighter markings; capsules 12–15 mm tall (up to 22 mm tall in the naturalized E. albidum); spring ephemerals, the above-ground portions of the plant usually senescing by July . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erythronium 5b. Leaves borne on the stem (Gagea with both basal and stem leaves), not mottled; capsules 15–77 mm tall (shorter in Gagea, but that species usually with 3–5 flowers per inflorescence); plants persisting through the summer in typical years 6a. Anthers mesifixed; tepals with basal nectaries; stems with usually more than 5 leaves, whorled (alternate in 1 species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lilium 6b. Anthers basifixed; tepals without basal nectaries; stems with 2–5 alternate leaves or 2 opposite leaves 7a. Stigmas sessile; stems with 2–5 alternate leaves; tepals 20–82 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tulipa 7b. Stigmas elevated on a style; stems with 2 opposite leaves; tepals 13–17 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gagea

Clintonia 1. Clintonia borealis (Ait.) Raf. N yellow bluebead-lily. Dracaena borealis Ait. • CT, MA, ME, NH, RI, VT. North-temperate and boreal forests.

Erythronium 1a. Tepals yellow, the inner with small auricles; style 5–11 mm long . . . . . . . . . E. americanum 1b. Tepals white and tinged with pink, blue, or lavender abaxially, lacking auricles; style 15–25 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. albidum 1. Erythronium albidum Nutt. E white trout-lily. CT. Rich, mesic, deciduous forests. Known historically from a single naturalized population along the Farmington River. 2. Erythronium americanum Ker-Gawl. ssp. americanum N American trout-lily. CT, MA, ME, NH, RI, VT. Mesic deciduous forests, often associated with rich slopes and cliff bases, riparian forests.

Gagea 1. Gagea fragifera (Vill.) E. Bayer & G. López E star-of-Bethlehem. Gagea fistulosa (Ram. ex DC.) Ker-Gawl.; G. liotardii Sternb.; Ornigthogalum fistulosum Ram. ex DC., nom. illeg.; O. fragiferum Vill. • VT. Waste areas. The often used name for this species, Gagea fistulosa, is an illegitimate name (Bayer and López González 1989).

19 4  MONOCOTS

Lilium Details of the flower (e.g., tepal color, tepal shape, flower orientation) are often distorted or obscured in pressing. Noting these features on herbarium labels will assist with identification at a later date. Reference: Skinner (2002). 1a. Leaves alternate throughout, with bulbils in the upper axils; mature buds evenly tapering to a ± flat apex with 3 green, apical, rounded processes . . . . . . . . . . . . . . . . . . . . . . L. lancifolium 1b. Leaves mostly or all whorled, without axillary bulbils; mature buds not as above 2a. Flowers erect; tepals with a distinct, abruptly narrowed, basal portion [Fig. 157]; inflorescence with 1–3 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. philadelphicum 2b. Flowers nodding; tepals gradually narrowed in the basal portion; inflorescence with (1–) 2–22 flowers 3a. Style similar in color to the tepals; tepals merely somewhat recurved, the outer tepals lacking abaxial ridges; buds round in cross-section; margins and veins of leaf blades roughened on the abaxial surface with triangular epidermal spicules . . . . . . L. canadense 3b. Style pale green; tepals strongly recurved, forming a “Turk’s-cap” perianth, the outer tepals with 2 faint, abaxial ridges; buds triangular in cross-section; margins and veins of the leaf blades not roughened on the abaxial surface . . . . . . . . . . . . . . . . . . . L. superbum 1. Lilium canadense L. N Canada lily. Lilium canadense L. var. editorum Fern. • CT, MA, ME, NH, RI, VT. Riparian forests, mesic to wet-mesic fields, wetland borders, roadsides. 2. Lilium lancifolium Thunb. E lance-leaved tiger lily. Lilium tigrinum Ker-Gawl. • CT, MA, ME, NH, RI, VT. Fields, roadsides, edges of lawns. 3. Lilium philadelphicum L. N Fig. 157 wood lily. CT, MA, ME, NH, RI, VT. Woodlands, sandplains, dry powerline rights-of-way, grasslands. 4. Lilium superbum L. N Turk’s-cap lily. Lilium canadense L. ssp. superbum (L.) Baker • CT, MA, NH, RI. Forest openings and edges, thickets, borders of swamps, roadsides. Fig. 157  Flower of Lilium philadelphicum.

Medeola 1. Medeola virginiana L. N Indian cucumber root. CT, MA, ME, NH, RI, VT. Mesic deciduous and mixed evergreendeciduous forests.

Streptopus Reference: Fassett (1935). 1a. Leaves sessile, but not clasping, with usually 30–50 marginal cilia per cm; nodes of the stem pubescent; tepals pink, with darker red stripes, outcurved to recurved near the apex (if at all); anthers ca. 2 mm long, 2-pointed at the apex; stigma 3-lobed; pedicels usually pubescent; berry subglobose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lanceolatus 1b. Leaves cordate-clasping, with usually 0–6 marginal cilia per cm; nodes of the stem glabrous; tepals green-yellow, outcurved or recurved from near the middle; anthers 3–3.5 mm long, with a single point at the apex; stigma unlobed; pedicels glabrous; berry ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. amplexifolius

l i l i ac e a e   19 5

1. Streptopus amplexifolius (L.) DC. N clasping-leaved twistedstalk. Streptopus amplexifolius (L.) DC. ssp. americanus (J.A. & J.H. Schultes) A. & D. Löve; S. amplexifolius (L.) DC. var. denticulatus Fassett; S. amplexifolius (L.) DC. var. grandiflorus Fassett; Uvularia amplexifolia L. • CT, MA, ME, NH, VT; absent from the coastal plain of southern New England. Mesic forests and stream banks in northtemperate to subalpine areas, sometimes ascending above treeline in sheltered gullies and snowbank communities. ‌1 × 2. Streptopus ×oreopolis Fern. is a rare, sterile, diploid hybrid. It is known from subalpine and alpine forests and meadows in ME, NH. From S. amplexifolius it can be recognized apart by its leaf margin cilia (22–40 per cm), red-purple to purple tepals, frequently pubescent pedicels, and green (rather than glaucous) abaxial leaf blade surface. From S. lanceolatus it can be recognized apart by its red-purple to purple tepals, its cordate-clasping leaf blades, usually glabrous nodes, more spreading tepal apices, and more elongate fruit (see identification key for character states of the parental species). 2. Streptopus lanceolatus (Ait.) Reveal N lance-leaved twistedstalk. Stretopus roseus Michx.; S. roseus Michx. var. longipes (Fern.) Fassett; S. roseus Michx. var. perspectus Fassett; Uvularia lanceolata Ait. • ct, ma, me, nh, ri, vt. Mesic forests and stream banks in north-temperate to subalpine areas, sometimes ascending above treeline in sheltered gullies and snowbank communities.

Tricyrtis 1. Tricyrtis hirta (Thunb.) Hook. E hairy toad-lily. MA. Roadsides, forest edges, areas of human habitation.

Tulipa 1a. Filaments pubescent near base; buds often nodding; tepals abruptly narrowed at the base, the inner ones 6–26 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. sylvestris 1b. Filaments glabrous; buds erect; tepals not abruptly narrowed at the base, the inner ones 21–41 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. gesneriana 1. Tulipa gesneriana L. E Didier’s tulip. MA. Edges of lawns, fields, roadsides, compost heaps. 2. Tulipa sylvestris L. E wild tulip. MA. Edges of lawns, fields, roadsides, compost heaps.

Melanthiaceae 1a. Sepals and petals clearly different, the sepals green; leaves borne in a single whorl at the summit of the stem, pinnately veined [Fig. 159]; fruit a berry . . . . . . . . . . . . . . . . . . . . . . . . Trillium 1b. Sepals and petals similar [Fig. 158], the sepals green or petaloid; leaves ± all basal or alternate, parallel veined; fruit a capsule 2a. Ovary partly inferior; anthers mesifixed; seeds twisted and flattened at the ends, unwinged; plants arising from bulbs, lacking rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . Anticlea 2b. Ovary superior; anthers basifixed; seeds variously shaped but with marginal angles, wings, or with a wing-like aril; plants with rhizomes (bulbs sometimes also present)

19 6 MONOCOTS

3a. Leaf blades narrow-linear, 20–70 × 0.5–3 cm; bulbs 30–80 mm long . . . . Stenanthium 3b. Leaf blades narrow-oblanceolate or spatulate to elliptic or ovate, 5–55 × 1–20 cm; bulbs absent or present and 6–17 mm long 4a. Tepals 2–4 mm long, white in staminate flowers, green-white in carpellate flowers (fading or drying yellow), lacking basal nectaries; inflorescence with an unbrached axis, ebracteate; capsule dehiscence loculicidal; plants dioecious . . . . Chamaelirium 4b. Tepals 5–12 mm long, green to yellow-green or green-white to light yellow, with 1 or 2 nectary glands at base; inflorescence branched, with bracts; capsule dehiscence septicidal; plants synoecious or polygamous . . . . . . . . . . . . . . . . . Veratrum

Anticlea 1. Anticlea elegans (Pursh) Rydb. ssp. glaucus (Nutt.) A. Haines

Fig. 158  Flower of Anticlea elegans.

N C Fig. 158

mountain death camas. Anticlea elegans (Pursh) Rydb. var. glaucus (Nutt.) Zomlefer & Judd; A. glauca (Nutt.) Kunth; Melanthium glaucum Nutt.; Zigadenus elegans Pursh var. glaucus (Nutt.) Preece ex Cronq. • VT; known only from Lake Champlain. Lakeshore headlands and bluffs composed of high-pH bedrock.

Chamaelirium 1. Chamaelirium luteum (L.) Gray

NC

devil’s bit. Veratrum luteum L. • CT, MA. Dry-mesic to mesic soils of fields, forest openings, and deciduous and evergreen-deciduous woodlands and forests on trap rock, limestone, and other bedrock types. Chamaelirium luteum is a dioecious species in which the staminate plants and the carpellate plants differ in morphology and abundance (e.g., carpellate plants are larger, carpellate plants have ascending vs. spreading flowers, staminate plants usually have fewer leaves, staminate plants are more frequent in a given population).

Stenanthium 1. Stenanthium gramineum (Ker-Gawl.) Morong E eastern featherbells. Helonias graminea Ker-Gawl.; Stenanthium gramineum (Ker-Gawl.) Morong var. micranthum Fern.; S. gramineum (Ker-Gawl.) Morong var. robustum (S. Wats.) Fern.; S. robustum S. Wats. • CT. Forest openings and clearings.

Trillium Reference: Case and Case (1997). 1a. Leaf blades usually mottled; flowers sessile; petals erect or arching inward . . . T. recurvatum 1b. Leaf blades not mottled; flowers borne on a peduncle 1–10 cm long; petals either spreading from near the base or flaring outwardly from above the base 2a. Petals white, with a red-purple crescent-shaped mark near the base; leaves with a short, but distinct, petiole; ovary 3-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. undulatum 2b. Petals white, pink, or red-purple; leaves sessile (sometimes with an obscure, petiolelike base in T. cernuum); ovary 6-angled or 6-winged (3-lobed in T. nivale) 3a. Stems at anthesis 3–5 cm tall, elongating to 9 cm post anthesis; leaf blades 1.5–4.5 cm long; peduncles 10–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. nivale 3b. Stems 10–60 cm tall; leaf blades 5–20 cm long; peduncles (10–) 15–30 mm long

Me l a nt h i ac e a e   197

4a. Petals 4–7.5 cm long, erect in the basal portion and somewhat recurving in the apical portion to produce a funnelform corolla; stigma straight, of uniform diameter throughout its length, usually equaling or exceeding the length of the ovary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. grandiflorum 4b. Petals 1.5–5 cm long, spreading or recurved from near the base; stigma conspicuously recurved, tapering in the apical portion, usually half as long as the ovary or shorter 5a. Petals red-purple (rarely white), 2.5–5 cm long; ovary red-purple to purple; peduncles (1–) 3–8 (–10) cm long, erect to spreading, the flowers only rarely hidden below the leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. erectum 5b. Petals white, 1.5–2.5 cm long; ovary white to pink; peduncles 1–3 (–5) cm long, recurved, hiding the flowers under the leaves [Fig. 159] . . . . . . . . . . . . . . . T. cernuum 1. Trillium cernuum L. N Fig. 159 nodding wakerobin. Trillium cernuum L. var. macranthum Eames & Wieg. • CT, MA, ME, NH, RI, VT. Deciduous and mixed evergreen-deciduous, upland and riparian forests. 2. Trillium erectum L. N red wakerobin. Trillium purpureum Kin ex Elliot • CT, MA, ME, NH, RI, VT; nearly throughout. Mesic, deciduous forests, less frequently in mixed evergreen-deciduous forests. Trillium erectum normally has dark red to red-purple petals. However, rangewide this species shows white, yellow-green, and paler red forms. None of these variants have been named within T. erectum save for var. album, which was named from plants in the mountains of the Carolinas and, at least in part, may represent introgressants with T. flexipes (Case and Case 1997). Because var. album in the Carolinas may possess real genetic differences from typical T. erectum, it may deserve recognition at some rank. However, our New England plants of T. erectum with white petals are simply sporadic color morphs. Given that other color forms are not provided formal recognition, what has been called var. album in New England is here subsumed in the species and regarded as a form without formal rank. 3. Trillium grandiflorum (Michx.) Salisb. N white wakerobin. Trillium rhomboideum Michx. var. grandiflorum Michx. • CT, MA, ME, NH, VT. Mesic, deciduous, often rich, forests. 4. Trillium nivale Riddell E snow wakerobin. CT. Mesic, deciduous, often rich, forests. Noted from a single naturalized population along the Farmington River but not found in recent searches. 5. Trillium recurvatum Beck E prairie wakerobin. Phyllantherum recurvatum (Beck) Nieuwl.; Trillium unguiculatum Raf. • CT. Mesic, deciduous forest. Known from a single naturalized population along the Farmington River. It was also reported to be growing with Erythronium albidum Nutt. and Trillium nivale Riddell., also believed to have been planted. Recent searches of the location found T. recurvatum present in large numbers. 6. Trillium undulatum Willd. N painted wakerobin. Trillium erythrocarpum Michx. • CT, MA, ME, NH, RI, VT; nearly throughout. Mesic, deciduous forests, less frequently in mixed evergreen-deciduous forests.

Veratrum 1a. Tepals abruptly narrowed to an obvious claw, with 2 basal nectary glands; leaf blades narrow-oblanceolate, 1–7.2 cm wide; filaments introrse . . . . . . . . . . . . . . . . . . . . . . . . V. latifolium 1b. Tepals gradually narrowed to an obscure claw, with a single V-shaped nectary gland on the basal margins; leaf blades ovate to elliptic, 8–20 cm wide; filaments extrorse . . . . . . . . V. viride

Fig. 159  Habit and nodding flower of Trillium cernuum.

19 8  MONOCOTS

1. Veratrum latifolium (Desr.) W.B. Zomlefer

NC

slender bunch-flower. Melanthium hybridum Walt.; M. latifolium Desr.; M. racemosum Michx. • CT. Mesic to dry-mesic, often rocky, forest slopes. 2. Veratrum viride Ait. ssp. viride N American false hellebore. CT, MA, ME, NH, RI, VT. Riparian forests, marshes, swamps.

Nartheciaceae Aletris 1. Aletris farinosa L. N Fig. 160 white colic-root. CT, MA, ME, NH, RI. Open, usually sandy, soils of fields, forest openings, and woodlands.

Fig. 160  Flowers of Aletris farinosa.

Orchidaceae 1a. Labellum deeply concave to saccate (at least in the basal portion), forming a small to very evident pouch [Fig. 164] 2a. Labellum saccate in only the basal portion, 1.8–12 mm long; inflorescence a spike or raceme, with 5–72 flowers [Fig. 166] 3a. Principal leaves all basal, usually white-striped or white-reticulated [Fig. 165]; labellum white to white-green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Goodyera 3b. Principal leaves borne on the stem, alternate, without white markings; labellum purple to purple-brown with pink near the apex or dark red . . . . . . . . . . . . . . . . . Epipactis 2b. Labellum saccate ± throughout, 10–67 mm long; inflorescence a solitary flower or a spike-like raceme with 2 or 3 (–4) flowers 4a. Labellum with 2 apical horns and a prominent fringe of yellow bristles; plants with 1 leaf; flowers with 1 fertile anther; stem arising from a corm . . . . . . . . . . . . . . . . . Calypso 4b. Labellum without a pair of apical horns, lacking evident yellow bristles [Figs. 163, 164]; plants with 2 or more leaves; flowers with 2 fertile anthers; stem arising from a short to elongate rhizome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cypripedium 1b. Labellum neither inflated nor saccate, not forming a pouch [Figs. 167, 171, 174] 5a. Labellum basally prolonged into a hollow spur (the spur short and saccate in Coeloglossum) [Figs. 173, 175] 6a. Flowers bilaterally asymmetrical (i.e., appearing lopsided); leaf single, abaxially purple, produced in autumn, withering by early spring, therefore, the flowering plants without well-formed leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tipularia 6b. Flowers bilaterally symmetrical; leaves 2 or more, less commonly single, abaxially green, produced in spring and present during flowering 7a. Labellum white with magenta spots, 3-lobed, the middle lobe notched at the apex; leaf single, basal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amerorchis

o rc h i dac e a e   19 9

7b. Labellum not spotted, variously shaped but never 3-lobed with a notched middle lobe; leaves 1–several, basal or arranged on a stem 8a. Stems conspicuously angled; labellum unlobed, typically white and strongly contrasting in color from the pink-magenta sepals and lateral petals (rarely all the perianth of similar color); viscidia enclosed in a single, 2-lobed bursicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Galearis 8b. Stems terete; labellum unlobed, lobed, or fringed, ± similar in color to the remainder of the perianth (those Platanthera with 2 basal leaves usually have the sepals more strongly tinged with green than the petals); viscidia free, not enclosed in a bursicle 9a. Labellum 3-lobed, the middle lobe much shorter than the lateral 2 [Fig. 161]; viscidia covered by membranes; orifice of spur covered by a membrane with a small opening; floral bracts conspicuous, usually evidently exceeding the flowers . . . . . . . . . . . . . . . . . . . . . . . . . . Coeloglossum 9b. Labellum variously shaped but never 3-lobed with a very reduced middle lobe [Figs. 172, 173, 174]; viscidia not covered by membranes; orifice of spur open; floral bracts inconspicuous to (rarely) shortly exceeding the flowers 10a. Labellum simple, sometimes with apical teeth or a marginal fringe, but without 3 evident lobes . . . . . . . . . . . . . . . . . . . . . (in part) Plantanthera 10b. Labellum clearly lobed into 3 segments 11a. Labellum 5–25 mm long, fringed on the margin [Fig. 174]; leaf blades entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Platanthera 11b. Labellum 3.5–5 mm long, with entire lobes; leaf blades denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnadenia 5b. Labellum lacking a spur (flowers with a small, inconspicuous, adnate spur in Corallorhiza formed by the 2 lateral sepals) [Figs. 167, 170, 177] 12a. Flowers not resupinate, the labellum the uppermost petal; labellum with a conspicuous brush of yellow-tipped hairs; leaves articulated, this appearing as a thin, transverse line near the base of the leaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calopogon 12b. Flowers resupinate, the labellum the lowest petal; labellum with or without hairs or ornamentation of some type; leaves not articulated 13a. Leaves whorled at the summit of the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isotria 13b. Leaves or scales alternate or opposite, borne at base of plant and/or distributed along the stem 14a. Flowers subtended by foliaceous, ovate bracts [Fig. 179]; labellum ornamented with 3 parallel, green, papillose ridges [Fig. 179] . . . . . . . . . . . Triphora 14b. Flowers subtended by rather inconspicuous, usually scale-like bracts; labellum ornamented or not, never with 3 parallel, green ridges 15a. Flowers large, the labellum (12–) 17–35 mm long; flowers usually solitary 16a. Leaves flat, numbering 1–3 on the stem, present at anthesis, the largest 7–25 (–32) mm wide; sepals and lateral petals distinct, the outer spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pogonia 16b. Leaves obscurely plicate, solitary on the stem, not fully formed at anthesis, 2–4 (–12) mm wide; sepals and lateral petals connate at the base, the outer erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arethusa 15b. Flowers smaller, the labellum 1.3–12 mm long; flowers 3–180 per inflorescence

20 0  MO NOCOTS

17a. Principal leaves arranged in a single, opposite pair on the stem; labellum 2-lobed due to a conspicuous apical notch [Figs. 170, 171] . . . . . . . . . . . Neottia 17b. Principal leaves 1–several, borne at base of plant and/or distributed along the stem; labellum entire or 3-lobed, without a single apical notch (with a prominent notch in Malaxis, but then with a small lobe at the base of the central notch) [Figs. 169, 176] 18a. Flowers predominantly white or cream (except for a yellow or green central area on the labellum in some species), sometimes tinged with yellow or green; leaves and stem bracts above them numbering 5 or more; labellum with a pair of basal callosities . . . . . . . . . . . . . . . Spiranthes 18b. Flowers not predominantly white or cream, variously colored and/ or spotted; leaves and stem bracts (when present), numbering 4 or fewer; labellum without a pair of basal callosities 19a. Plants leafless at flowering time, with sheathing bracts only, arising from coral-like rhizomes; plants without chlorophyll, the stems brown to purple, or yellow-green in one species . . . . . . . . Corallorhiza 19b. Plants with leaves at flowering time (Aplectrum sometimes leafless at flowering, but the withered remnant usually present at stem base), arising from corms; plants chlorophyllous, the leaves and stems ± green at anthesis 20a. Leaf solitary, evidently plicate, produced at distal end of corm in late season and persistent through the winter, usually withering by flowering; labellum white with magenta spots (rarely all whiteyellow); capsules pendent . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aplectrum 20b. Leaf solitary or paired, not or only obscurely plicate, produced in spring, present during flowering, withering before winter; labellum green to yellow-green or brown-purple to pale purple, without spots; capsules spreading to erect 21a. Plants with 2 leaves, in a pair near the base of the stem; labellum 4–12 mm long; capsules 9–13 mm long . . . . . . . . . Liparis 21b. Plants with a single leaf (rarely 2 and then not paired), borne on the stem; labellum 1.3–2.3 mm long; capsules mostly 5–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malaxis

Amerorchis 1. Amerorchis rotundifolia (Banks ex Pursh) Hultén

NC

round-leaved orchid. Galearis rotundifolia (Banks ex Pursh) R.M. Bateman; Habenaria rotundifolia (Banks ex Pursh) Richards.; Orchis rotundifolia Banks ex Pursh • ME, NH, VT. High-pH swamps and fens, usually in the shade of Thuja occidentalis.

Aplectrum 1. Aplectrum hyemale (Muhl. ex Willd.) Nutt.

NC

putty-root. Cymbidium hyemale Muhl. ex Willd. • CT, MA, VT. Mesic to wet-mesic, deciduous, sometime rocky, forests.

Arethusa 1. Arethusa bulbosa L. N dragon’s-mouth. CT, MA, ME, NH, RI, VT. Bogs, fens, wet meadows, and openings in conifer swamps.

o rc h i dac e a e   2 0 1

Calopogon 1. Calopogon tuberosus (L.) B.S.P. var. tuberosus N tuberous grass-pink. Calopogon pulchellus (Salisb.) R. Br.; Limodorum tuberosum L. • CT, MA, ME, NH, RI, VT. Bogs, fens, wet meadows, and openings in conifer swamps.

Calypso 1. Calypso bulbosa (L.) Oakes ssp. americana (R. Br. ex Ait. f.) A. Haines

NC

fairy-slipper. Calypso bulbosa (L.) Oakes var. americana (R. Br. ex Ait. f.) Luer; Cypripedium bulbosum L., pro parte; Cytherea bulbosa (L.) House, pro parte • ME, NH, VT. Evergreen swamps dominated by Thuja occidentalis.

Coeloglossum 1. Coeloglossum viride (L.) Hartman N Fig. 161 long-bracted green orchid. Coeloglossum bracteatum (Muhl. ex Willd.) Parl.; C. viride (L.) Hartman var. virescens (Muhl. ex Willd.) Luer; Dactylorhiza viridis (L.) R.M. Bateman, Pridgeon, & M.W. Chase; Habenaria bracteata (Muhl. ex Willd.) R. Br. ex Ait. f.; H. viridis (L.) R. Br. var. bracteata (Muhl. ex Willd.) Reigenb. ex Gray • CT, MA, ME, NH, RI, VT. Mesic to wet-mesic deciduous and evergreen-deciduous forests, fens, swamps, meadows. Fig. 161  Flower of Coeloglossum viride.

Corallorhiza Reference: Magrath and Freudenstein (2002). 1a. Labellum without lateral lobes; lateral sepals up-curved; plants flowering late-August through late September . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. odontorhiza 1b. Labellum with a lateral lobe or tooth on each side [Fig. 162]; lateral sepals spreading to down-curved; plants flowering mid-May through late-August 2a. Labellum (3.5–) 4–9 mm long, white spotted with red-purple (rarely pure white) [Fig. 162]; sepals and lateral petals with 3 (–5) nerves; spur present, though inconspicuous; stem commonly purple to brown; plants flowering in early July through late August; raceme with (6–) 10–30 (–41) flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. maculata 2b. Labellum 2–3.3 (–5) mm long, white; sepals and lateral petals with 1 nerve; spur absent; stem commonly yellow-green; plants flowering in mid-May through mid-June; raceme with 3–12 (–18) flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. trifida 1. Corallorhiza maculata (Raf.) Raf. N Fig. 162 spotted coral-root.  1a. Corallorhiza multiflora Nutt.; 1b. Corallorhiza maculata (Raf.) Raf. var. immaculata M.E. Peck • CT, MA, ME, NH, RI, VT. Mesic to dry-mesic forests, usually with a relatively open understory. 1a. Floral bracts mostly 0.5–1 mm long, usually entire; middle lobe of labellum not or scarcely widened distally, the ratio of the width of the apical portion to the basal portion of the labellum less than 1.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. C. maculata var. maculata 1b. Floral bracts mostly 1–2.8 mm long, often bifid or even trifid at apex; middle lobe of labellum definitely expanded, the ratio of the width of the apical portion to the basal portion of the labellum more than 1.5 [Fig. 162] . . . . . . . . . . 1b. C. maculata var. occidentalis (Lindl.) Cockerell Variety maculata is known from CT, MA, ME, NH, RI, VT. Variety occidentalis is known from CT, MA, ME, NH, VT. The latter variety flowers earlier than the former when the two taxa occur near one another. 2. Corallorhiza odontorhiza (Willd.) Poir.

NC

fall coral-root.  2b. Corallorhiza pringlei Greenman • CT, MA, ME, NH, RI, VT; scattered throughout New England but restricted in ME to the southwestern portion of the state. Dry-mesic to

Fig. 162  Flower of Corallorhiza maculata var. occidentalis.

202 MO NOCOTS

mesic, deciduous and evergreen-deciduous forests, usually with a relatively open understory, also often on sites influenced by circumneutral to basic bedrock. 1a. Perianth often closed; stigmatic surface 0.3–0.5 mm wide; labellum 1.7–2.2 (–3) mm wide; column without or with poorly developed auricles at base on adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a. C. odontorhiza var. odontorhiza 1b. Perianth open; stigmatic surface 0.7–1 mm wide; labellum 2–3.7 mm wide; column with 2 prominent auricles at base on adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. C. odontorhiza var. pringlei (Greenman) Freudenstein Variety odontorhiza is known from CT, MA, ME, NH, RI, VT. Variety pringlei is only known from CT and is of regional conservation concern; also reported from MA, ME, VT by Brown (1997), but specimens are unknown. 3. Corallorhiza trifida Chatelain N early coral-root. Corallorhiza trifida Chatelain var. verna (Nutt.) Fern. • CT, MA, ME, NH, RI, VT. Wet-mesic to hydric forests, under a variety of canopies but often with some component of evergreen trees.

Cypripedium Reference: Sheviak (2002a). 1a. Plants with 2 basal leaves, the stem scapose; labellum with a cleft along its entire length on the adaxial (i.e., upper) surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. acaule 1b. Plants with 2–9 leaves alternately arranged on the stem; labellum with a round opening near the base on the adaxial surface 2a. Labellum 1–1.6 (–2.5) cm long, strongly red-veined, prolonged downward near the apex into a conical pouch; all 3 sepals distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. arietinum 2b. Labellum 1.5–5.4 cm long, neither red-veined nor prolonged downward to form a protruding sac; lateral sepals connate, thereby forming a single blade positioned beneath the labellum 3a. Labellum white (rarely pink) and streaked with pink or purple; lateral petals white, not twisted, rounded to narrow-obtuse at apex; lateral sepals entirely connate . . . . C. reginae 3b. Labellum yellow, usually with purple veins; lateral petals spotted, streaked, or entirely suffused with red-purple or red-brown, rarely unmarked and then ± yellowgreen, twisted, acute to acuminate at apex; lateral sepals connate except at tip, forming a bilobed apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. parviflorum Fig. 163  Flower of Cypripedium parviflorum var. makasin.

1. Cypripedium acaule Ait. N pink lady’s-slipper. CT, MA, ME, NH, RI, VT. Forests and woodlands, typically on acidic soils. 2. Cypripedium arietinum Ait. f.

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ram’s-head lady’s-slipper. CT, MA, ME, NH, VT. Deciduous and mixed evergreen-deciduous forests, often on enriched soils due to bedrock influence or colluvial deposits, also in swamps dominated by Thuja occidentalis. 3. Cypripedium parviflorum Salisb.

N C Fig. 163, 164

yellow lady’s-slipper.  3a. Cypripedium calceolus L. var. pubescens (Willd.) Correll; C. pubescens Willd.;   3b. Cypripedium calceolus L. var. parviflorum (Salisb.) Fern. • CT, MA, ME, NH, ri, VT. Evergreen swamps, deciduous forests, river banks, river shore ledges. 1a. Labellum usually 3–5.4 cm long; lateral petals mostly 5–8 cm long, entirely yellow-green or sparsely to moderately spotted or streaked with red-purple [Fig. 164] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. C. parviflorum var. pubescens (Willd.) Knight Fig. 164  Flower of Cypripedium parviflorum var. pubescens.

1b. Labellum 1.5–3 (–3.5) cm long; lateral petals mostly 3–5 cm long, either densely spotted or evenly suffused with red-purple or red-brown [Fig. 163]

o rc h i dac e a e   2 03

2a. Uppermost sheathing bract at base of stem conspicuously pubescent with short hairs; red-purple color of lateral petals composed of numerous, densely spaced dots; floral scent rose-like or musty; plants of deciduous forests . . . . . 3b. C. parviflorum var. parviflorum 2b. Uppermost sheathing bract glabrous or inconspicuously pubescent; red-purple color of lateral petals due to an even suffusion of pigment; floral scent intensely sweet; plants of high-pH wetlands and shores . . . . . . . . . . . 3c. C. parviflorum var. makasin (Farw.) Sheviak Variety pubescens is known from CT, MA, ME, NH, RI, VT. Variety parviflorum is known from CT, MA, RI, VT. Variety makasin is known from CT, MA, ME, NH, VT. Both var. makasin and var. parviflorum are of regional conservation concern. Sheviak (1993, 1994) divided the small-flowered yellow lady’sslippers into two varieties based on ecology and floral morphology and fragrance—var. makasin and var. parviflorum. Cypripedium parviflorum var. pubescens is a problematic taxon in New England and has been frequently misidentified. This is, in part, due to a belief that it occurred exclusively in deciduous forests. Though this holds true for the majority of occurrences in southern New England, var. pubescens is frequently located in evergreen swamps dominated by Thuja occidentalis (and rarely shaded river shore ledges) in northern New England. These plants of var. pubescens from wetlands show glabrous sheathing bracts, which is in contradiction to the statements of Sheviak (2002a), suggesting this character to be environmentally influenced. Variety pubescens has a musty floral fragrance similar to var. parviflorum. 4. Cypripedium reginae Walt. N showy lady’s-slipper. Cypripedium spectabile Salisb. • CT, MA, ME, NH, VT. Fens, swamps dominated by Thuja occidentalis.

Epipactis 1a. Labellum 5.5–6.5 mm long, the distal part (i.e., non-saccate portion) broad-triangular or transversely elliptic, with rugose basal calli; ovary tomentose; sepals and lateral petals dark red to nearly red-purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. atrorubens 1b. Labellum 9–12 mm long, the distal part triangular-ovate, with non-rugose basal calli; ovary glabrous or sparsely pubescent; sepals and lateral petals pale green, pink, purple, or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. helleborine 1. Epipactis atrorubens (O. Hoffman) Bess. E dark red helleborine. Epipactis helleborine (L.) Crantz ssp. atrorubens (O. Hoffman ex Bernh.) Syme; Serapias atrorubens O. Hoffmann • VT. Early successional forest on serpentine bedrock in an abandoned asbestos quarry. Searches of the supposed location for Epipactis atrorubens have found only plants of E. helleborine with sparsely pubescent ovaries and some dark red pigment spots on the flowers. The occurrence of E. atrorubens in New England requires confirmation. 2. Epipactis helleborine (L.) Crantz E broad-leaved helleborine. Epipactis latifolia (L.) All.; Serapias helleborine L. • CT, MA, ME, NH, RI, VT. Dry-mesic to hydric forests, stream banks.

Galearis 1. Galearis spectabilis (L.) Raf. N showy orchid. Orchis spectabilis L. • CT, MA, ME, NH, RI, VT. Rich, usually deciduous, forests, often in areas influenced by high-pH bedrock or colluvial deposits.

Goodyera Reference: Kallunki (2002). 1a. Spike cylindric, equally dense on all sides, the spiral of flowers hardly discernible; labellum lacking fleshy callosities on the inner surface, with a reflexed tip; rostellum notched; scape with 4–14 bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. pubescens

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1b. Spike usually less dense, with secund flowers or with flowers arranged in an evident spiral; labellum with fleshy callosities on the inner surface, with a spreading or recurved tip; rostellum with a 2-pronged beak; scape with 2–7 bracts 2a. Labellum 4.9–7.9 mm long; leaf blades typically white only on the midvein [Fig. 165]; beak of the rostellum 2.3–3.6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. oblongifolia 2b. Labellum 1.8–5.5 mm long; leaf blades typically white on the primary lateral veins; beak of the rostellum 0.2–1.7 mm long 3a. Spike with usually secund flowers [Fig. 166]; leaf blades with usually 5 nerves; labellum deeply saccate, the pouch as deep as long, with an elongate, recurved apex; beak of the rostellum 0.2–0.6 mm long; anthers blunt with an apiculus at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. repens 3b. Spike with usually flowers in a loose spiral; leaf blades with usually 5–9 nerves; labellum shallowly saccate, the pouch longer than deep, with a short, spreading to recurved apex; beak of the rostellum 0.6–1.7 mm long; anthers usually acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. tesselata 1. Goodyera oblongifolia Raf. Fig. 165  Leaf of Goodyera oblongifolia.

N C Fig. 165

giant rattlesnake-plantain. Goodyera decipiens (Hook.) Piper; Peramium decipiens (Hook.) Piper • ME, VT; northern portion of states. Evergreen or mixed evergreen-deciduous forests, usually with a component of Picea rubens, Thuja occidentalis, and/or Abies balsamea. 2. Goodyera pubescens (Willd.) R. Br. in Ait. & Ait. f. N downy rattlesnake-plantain. Neottia pubescens Willd.; Peramium pubescens (Willd.) MacM. • CT, MA, ME, NH, RI, VT. Mesic to dry-mesic forests, frequently in association with Pinus strobus, Quercus rubra, and/or Tsuga canadensis. 3. Goodyera repens (L.) R. Br. in Ait. & Ait. f. N Fig. 166 dwarf rattlesnake-plantain. Goodyera ophioides (Fern.) Rydb.; G. repens (L.) R. Br. in Ait. & Ait. f. var. ophioides Fern.; Peramium ophioides (Fern.) Rydb.; Satyrium repens L. • CT, MA, ME, NH, VT. Evergreen and mixed evergreen-deciduous forests, usually growing on humus-rich soil and/or with bryophytes. 4. Goodyera tesselata Lodd. N

Fig. 166  Inflorescence of Goodyera repens.

checkered rattlesnake-plantain. Peramium tesselatum (Lodd.) Heller • CT, MA, ME, NH, RI, VT. Mesic to dry-mesic, evergreen and mixed evergreen-deciduous forests. Goodyera tesselata is thought to be an allotetraploid between G. oblongifolia and a form of G. repens with white-reticulate leaves (Kallunki 1976). The description includes both triploid first-generation hybrids and stabilized tetraploid individuals. Some individuals of Goodyera tesselata approach G. repens in morphology and are very difficult to distinguish with confidence.

Gymnadenia 1. Gymnadenia conopsea (L.) R. Br. E fragrant orchid. Habenaria conopsea (L.) Benth. • CT. Fields, roadsides, meadows. Historically known from Litchfield County, CT, but not seen in recent years.

Isotria 1a. Sepals 12–25 mm long, usually green-yellow throughout; lateral petals 13–18 mm long; labellum 10–15 mm long; peduncle 10–15 cm long, shorter than the ovary; non-flowering plants commonly with a white, arrested floral bud 1–2 mm long; plants short-rhizomatous, occurring singly or in small clumps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. medeoloides 1b. Sepals 34–67 mm long, usually green-yellow in the basal portion and red-purple in the apical portion; lateral petals 15–25 mm long [Fig. 167]; labellum 15–25 mm long; peduncle

o rc h i dac e a e   2 0 5

20–55 mm long, at least as long as the ovary; non-flowering plants only rarely with a white, arrested floral bud; plants long-rhizomatous, forming colonies . . . . . . . . . . . . . . . . I. verticillata 1. Isotria medeoloides (Pursh) Raf.

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small whorled pogonia. Arethusa medeoloides Pursh; Pogonia affinis Austin ex Gray • CT, MA, ME, NH, RI, VT. Deciduous or evergreen-deciduous, often acidic, forests, usually relatively young forests with an open understory, frequently associated with mild topography and growing in the vicinity of small streams. Vegetative plants of Isotria medeoloides superficially resemble Medeola virginiana. The former has glabrous, hollow stems that arise from short rhizomes, while the latter has pubescent, solid stems that arise from a white tuber. 2. Isotria verticillata (Muhl. ex Willd.) Raf. N Fig. 167 large whorled pogonia. Arethusa verticillata Muhl. ex Willd.; Pogonia verticillata (Muhl. ex Willd.) Nutt. • CT, MA, ME, NH, RI, VT. Mesic to dry-mesic, often acidic, forests and woodlands.

Liparis 1a. Labellum brown-purple, 8–12 mm long; column 3–4 mm long; fruiting pedicel 11–18 mm long, nearly as long as or slightly longer than the capsule . . . . . . . . . . . . . . . . . . . . . . L. liliifolia 1b. Labellum green to yellow-green or yellow-white, 4–5.5 mm long; column 2–3 mm long; fruiting pedicel 3–7 mm long, shorter than the capsule . . . . . . . . . . . . . . . . . . . . . . . . . . L. loeselii 1. Liparis liliifolia (L.) L.C. Rich ex Lindl.

Fig. 167  Flower of Isotria verticillata.

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lily-leaved wide-lipped orchid. Leptorchis liliifolia (L.) Kuntze; Malaxis liliifolia (L.) Sw.; Ophrys liliifolia L. • CT, MA, RI, VT. Dry-mesic to wet-mesic forests and woodlands, often on soils influenced by middle to high-pH bedrock such as limestone, trap, and sandstone. Reports of this species in NH are based on a fruiting specimen of Liparis loeselii. 2. Liparis loeselii (L.) L.C. Rich. N Loesel’s wide-lipped orchid. Leptorchis loeselii (L.) MacM.; Liparis correana (W. Bart.) Spreng.; Malaxis correana W. Bart.; M. loeselii (L.) Sw.; Orchis loeselii L. • CT, MA, ME, NH, RI, VT. Mesic to hydric, open soils of meadows, fens, shorelines, and disturbed places such as abandoned borrow pits and cleared rights-of-way.

Malaxis The labellum is to be measured from its base to the apex of the central tooth when using the relative length measurements for Malaxis bayardii and M. unifolia. Reference: Catling and Magrath (2002). 1a. Labellum entire, acuminate at the apex; pedicels (1–) 2–4.5 mm long; bracts 1.5–2 mm long; inflorescence less crowded and tapering to a slender apex . . . . . . . . . . . . . . . . M. monophyllos 1b. Labellum with 2 lobes at the apex and with a shorter, central tooth in the sinus between the lobes [Figs. 168, 169]; pedicels (3.4–) 5–10 (–13) mm long; bracts 0.2–1.6 mm long; inflorescence somewhat to highly crowded near the apex, often ± round- to flat-topped 2a. Basal auricles of labellum 0.4–1.1 mm long, less than 0.6 times as long as labellum [Fig. 169]; pedicels (3.8–) 5–10 (–13) mm long; lower flowers usually withered or shed by anthesis of the upper flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. unifolia 2b. Basal auricles of labellum 0.8–1.1 mm long, 0.6 or more times as long as labellum [Fig. 168]; pedicels 3.4–5 (–5.8) mm long; lower flowers still persisting during anthesis of the upper flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. bayardii 1. Malaxis bayardii Fern.

N C Fig. 168

Bayard’s adder’s-mouth. CT, MA, RI. Woodlands with Pinus and Quercus, dry sandy fields, and among heaths in well-drained soils. This species is reported from VT (Catling and Magrath 2002) based on 16 Jul 1918, Smart s.n. (CONN!). The specimen is a late collection with a few-

Fig. 168  Flower of Malaxis bayardii focusing on the labellum.

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flowered inflorescence that has shed its flowers and fruits, leaving only empty pedicels on the herbarium sheet. Given that the specimen is equivocal and is highly disjunct from the, it is not considered a reliable voucher. 2. Malaxis monophyllos (L.) Sw. ssp. brachypoda (Gray) A. & D. Löve

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white adder’s-mouth. Malaxis brachypoda (Gray) Fern.; M. monophyllos (L.) Sw. var. brachypoda (Gray) F. Morris & Eames • CT, MA, ME, NH, VT; widespread (at least historically) in the northern states, but missing from southern and eastern-central New England. Fens and evergreen swamps, usually in the shade of Thuja occidentalis, rarely on open ledges. 3. Malaxis unifolia Michx. N Fig. 169 green adder’s-mouth. Microstylis unifolia (Michx.) B.S.P. • CT, MA, ME, NH, RI, VT. Swamps, wetland borders, forest cuts and openings, cleared rights-of-way.

Fig. 169  Flower of Malaxis unifolia focusing on the labellum.

Neottia Reference: Magrath and Coleman (2002). 1a. Labellum divided ½ or more of its length into 2 pointed, linear segments [Fig. 171]; column 0.5–1 mm tall; floral bracts about 0.7–2.0 mm long; pedunce usually longer than the leaves 2a. Axis of inflorescence glabrous; labellum 3–4 (–5) mm long, yellow-green to green or redpurple; leaves on many individuals contracted to a short, petiole-like base . . . . . . . N. cordata 2b. Axis of inflorescence stipitate-glandular; labellum 6–12 mm long, red-purple; leaves sessile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. bifolia 1b. Labellum divided ca. ⅓ its length into 2 oblong or ovate segments [Fig. 170]; column 2.5–4 mm tall; floral bracts 2–5 mm long; peduncle usually as long as or shorter than the leaves 3a. Labellum narrowed at the base, usually bearing a pair of small teeth at the base [Fig. 170]; axis of the inflorescence, pedicels, and ovary stipitate-glandular [Fig. 170] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. convallarioides 3b. Labellum maintaining its width and not narrowed at the base, bearing a pair of retrorse auricles at the base; pedicels and ovary glabrous, only the axis of the inflorescence stipitate-glandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. auriculata 1. Neottia auriculata (Wieg.) Szlach.

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auricled twayblade. Bifolium auriculatum (Wieg.) Nieuwl.; Listera auriculata Wieg.; Ophrys auriculata (Wieg.) House • ME, NH, VT. Stream banks, riparian forests of small to large rivers, often in alluvium and commonly associated with Alnus incana, rarely in open peatlands. 1‌ × 3. Neottia ×veltmanii (Case) Baumbach is a rare twayblade hybrid known from ME but is to be expected elsewhere in northern New England. It is usually associated with one or both parents (i.e., it is not typically found alone). It has the basally narrowed labellum of N. convallarioides and, like that species, has stipitate glands on the pedicels and ovary. However, the stipitate glands on these parts are shorter and sparser than those of the inflorescence axis (in N. convallarioides, the stipitate glands are similar in length and ± similar in density on the glandular portions of the inflorescence). Further, the hybrid shows a small pair of outward pointing auricles at the very base of the lip (rather than a small pair of teeth distal to the narrow portion of the labellum). 2. Neottia bifolia (Raf.) Baumbach

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southern twayblade. Bifolium australe (Lindl.) Nieuwl.; Diphryllum bifolium Raf.; Listera australis Lindl.; Ophrys australis (Lindl.) House • VT. Growing on Sphagnum in forested and open bogs. Fig. 170  Inflorescence of Neottia convallarioides with detail of pedicel surface showing stipitateglands.

3. Neottia convallarioides (Sw.) Rich. N Fig. 170 broad-leaved twayblade. Bifolium convallarioides (Sw.) Nieuwl.; Epipactis convallarioides Sw.; Listera convallarioides (Sw.) Nutt. ex Ell.; Ophrys convallarioides (Sw.) W. Wight ex House • ME, NH, VT. Swamps and streambanks, often in the shade of Thuja occidentalis.

o rc h i dac e a e   2 07

4. Neottia cordata (L.) Rich. N Fig. 171 heart-leaved twayblade. Bifolium cordatum (L.) Nieuwl.; Listera cordata (L.) R. Br.; Ophrys cordata L. • CT, MA, ME, NH, RI, VT. Swamps and pond shores, often in the shade of Thuja occidentalis or other evergreen trees, glacial basins, subalpine forests, and rarely near high mountain summits.

Platanthera ‌vossii Platanthera × Case was reported from MA, ME by Kartesz (1999) based on personal communication with Paul Brown, but specimens are unknown. Brown (2002) resurrected the genus Gymnadeniopsis for species allied to Platanthera clavellata; however, phylogenetic studies to date (e.g., Hapeman and Inoue 1997) show that Gymnadeniopsis is nested within Platanthera and not monophyletic. Therefore, it is not recognized here. Reference: Sheviak (2002b). 1a. Labellum simple, not divided into 3 portions, the margins entire, toothed, or fringed [Figs. 172, 175] 2a. Principal leaves all basal, the scapose stems without bracts or with 1–6 minute, bract-like leaves; pollen sacs divergent 3a. Spur 3–8 (–10) mm long; scape with 0 (–1) bract-like leaves; basal leaves 0.8–5 cm wide, oblanceolate to obovate; ovary arcuate . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. obtusata 3b. Spur 11–46 mm long [Fig. 175]; scape with 0–6 bract-like leaves; basal leaves 4–22 cm wide, broad-elliptic or broad-obovate to orbicular; ovary straight 4a. Scape with 0 (very rarely 1) bract-like leaves; ovary sessile; flowers green to green-yellow; labellum upcurved; spur tapering to apex . . . . . . . . . . . . . . . . . . P. hookeri 4b. Scape with 1–6 bract-like leaves; ovary on a stipe 5–10 mm long; flowers white to green-white; labellum downward oriented; spur clavate-thickened at apex 5a. Spur mostly 10–26 mm long; pollinaria 3–4.5 (–4.8) mm long; spur length plus twice the pollinarium length less than 38 (note: measurements in mm for calculation) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. orbiculata 5b. Spur 28–46 mm long; pollinaria (4.6–) 4.7–6.8 mm long; spur length plus twice the pollinarium length equal to or greater than 38 . . . . . . . . . . . . . . P. macrophylla 2b. Principal leaves not confined to the base of the plant, the stems with 1–several leaves; pollen sacs mostly parallel 6a. Margin of the labellum entire [Fig. 175] 7a. Flowers white; labellum abruptly widened at the base; distance between anthers greater at apex than at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. dilatata 7b. Flowers green to white-green or yellow-green; labellum gradually widened at the base; distance between anthers greater at base than at apex 8a. Labellum white-green, 5–12 mm long; anthers nearly parallel or slightly diverging basally, separated apically 0.6–1.5 mm; spur 4–12 mm long, cylindric to slender-clavate; pollinia remaining within anther sacs; flowers with a sweet, pungent scent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. huronensis 8b. Labellum green to yellow-green, 2.5–6 mm long; anthers widely diverging basally, touching near apex or separated by a distance of no more than 0.3 mm; spur 2–5 mm long, clavate or, less commonly, cylindric; pollinia free of anther sacs and/or fragmenting and falling onto stigma (i.e., plants self-pollinating); flowers scentless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aquilonis 6b. Margin of the labellum toothed, erose, or conspicuously fringed [Fig. 172] 9a. Labellum conspicuously fringed, (4–) 8–19 mm long; spur (4–) 10–35 mm long

Fig. 171  Flower of Neottia cordata.

20 8  MO NOCOTS

10a. Flowers white; labellum short-fringed, with most or all marginal fringe segments shorter than width of medial, unfringed portion . . . . . P. blephariglottis 10b. Flowers orange to yellow-orange; labellum long-fringed, with marginal fringe segments equal to or longer than width of medial, unfringed portion 11a. Labellum 8–19 mm long; spur 20–35 mm long; orifice of spur keyholeshaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. ciliaris 11b. Labellum 4–8 mm long; spur 4–10 mm long; orifice of spur triangular to keyhole-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cristata 9b. Labellum with basal or apical teeth or lobes, sometimes with crenations or irregular teeth, but definitely not fringed [Fig. 173], 3–7 mm long; spur 4–13 mm long 12a. Labellum without a protuberance, with 3 teeth or very shallow lobes at the apex; stem with 1 principal leaf; spur 7–13 mm long . . . . . . . . . . . . . . . . P. clavellata 12b. Labellum with a prominent, erect protuberance, usually with 2 basal teeth [Fig. 173], often crenate or irregularly toothed over the margin; stem with (1–) 2 or 3 (–4) principal leaves; spur 4–8 (–11) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . P. flava 1b. Labellum deeply 3-lobed, with erose or conspicuously fringed margins [Fig. 174] 13a. Flowers green-white, yellow-white, or white (rarely tinged with bronze or rose); margin of the labellum conspicuously long-fringed, with some fringe segments exceeding ½ the length of the lobes of the labellum; labellum with cuneate to broad-cuneate lobes 14a. Labellum green-white or yellow-white, 10–17 mm long, its segments cuneate; sepals 4.5–7 mm long, the lateral ones deflexed behind the labellum; spur 11–23 mm long; lateral petals entire (rarely slightly lacerate-toothed at apex); viscidia linear . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lacera 14b. Labellum white, 14–22 mm long, its lobes broad-cuneate; sepals 7–13 mm long, the lateral ones merely divergent with the tips curved forward; spur 28–47 mm long; lateral petals lacerate-toothed at apex; viscidia nearly orbicular . . . . . . . . . . . . . . P. leucophaea 13b. Flowers pink to rose-purple (very rarely white); margin of the labellum erose to shortfringed, with fringe segments up to ½ the length of the lobes of the labellum [Fig. 174]; labellum with broad-flabellate lobes 15a. Orifice of spur horizontally rectangular or oblong; rostellum lobes short, rounded, nearly parallel, directed downward; labellum 5–13 mm long; spur 12–22 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. psycodes 15b. Orifice of spur circular [Fig. 174]; rostellum lobes angular, spreading, directed foward; labellum 10–25 mm long; spur 15–35 mm long . . . . . . . . . . . . . . . . . . P. grandiflora 1. Platanthera aquilonis Sheviak N north wind bog-orchid. CT, MA, ME, NH, RI, VT. Stream banks, fens, deciduous and mixed evergreen-deciduous forests, ditches, wet borrow pits, meadows. Most of our material that was called Platanthera hyperborea is referable to this species. 2. Platanthera blephariglottis (Willd.) Lindl. var. blephariglottis N white fringed bog-orchid. Blephariglottis blephariglottis (Willd.) Rydb.; Habenaria blephariglottis (Willd.) Hook. • CT, MA, ME, NH, RI, VT. Open fens, peaty and sandy meadows. 3. Platanthera ciliaris (L.) Lindl.

Fig. 172  Flower of Platanthera ciliaris.

N C Fig. 172

orange fringed bog-orchid. Blephariglottis ciliaris (L.) Rydb.; Habenaria ciliaris (L.) R. Br. ex Ait. f. • CT, MA, RI. Sandy and peaty meadows, wetland borders, lawns, sandy soils of swamps. Reports of this species from VT are likely based on specimens of Platanthera blephariglottis (see Jenkins and Zika 1995 for discussion). Reports of this species from NH are highly suspect. The specimen—Aug 1872, Jesup s.n. (NHA!)—includes an herbarium label with hand-penned location data that is also the location of the Jesup Herbarium. It is unclear if the label data

o rc h i dac e a e   2 0 9

applies to the location of collection or location of the museum. Given that most collections of this species are from the greater coastal plain, it is doubtful that this species would occur in the interior of NH. 4. Platanthera clavellata (Michx.) Luer N little club-spur bog-orchid. Gymnadeniopsis clavellata (Michx.) Rydb.; Habenaria clavellata (Michx.) Spreng. • CT, MA, ME, NH, RI, VT. Fens, meadows, sandy or peaty lake shores, ditches, openings in swamps. 5. Platanthera cristata (Michx.) Lindl.

NC

crested orange bog-orchid. Blephariglottis cristata (Michx.) Raf.; Habenaria cristata (Michx.) R. Br. ex Ait. f. • MA; eastern portion of the state. Sandy and peaty meadows, marshes, wetland borders, sandy soils of swamps. 6. Platanthera dilatata (Pursh) Lindl. ex Beck var. dilatata N white northern bog-orchid. Habenaria dilatata (Pursh) Hook.; Limnorchis dilatata (Pursh) Rydb. • cT, MA, ME, NH, VT. Bogs, fens, wet openings in boreal and subalpine forests, river shores, bases of wet, high-elevation cliffs, ditches. 7. Platanthera flava (L.) Lindl. var. herbiola (R. Br. in Ait. & Ait f.) Luer N Fig. 173 northern tubercled bog-orchid. Habenaria flava (L.) R. Br.; Perularia flava (L.) Farw. • CT, MA, ME, NH, RI, VT. River shores, usually associated with circumneutral seeps, sandy alluvium, or shrub thickets, rarely in tidal marshes, also in wet fields, meadows, and swamps.

Fig. 173  Flower of Platanthera flava.

8. Platanthera grandiflora (Bigelow) Lindl. N Fig. 174 greater purple fringed bog-orchid. Blephariglottis grandiflora (Bigelow) Rydb.; Habenaria fimbriata (Ait.) R. Br. ex Ait. f.; H. grandiflora (Bigelow) Torr.; H. psycodes (L.) Spreng. var. grandiflora (Bigelow) Gray • CT, MA, ME, NH, RI, VT. Swamp edges, stream banks, ditches, trail edges, fields, mesic forests. 8 ‌ × 11. Platanthera ×keenanii P.M. Brown is a very rare bog-orchid hybrid in New England known from NH; also reported from CT, MA, ME, RI, VT by Kartesz (1999), mainly based on personal communication with Paul Brown, but specimens are unknown. On initial examination, it shows more dissected labellum segments than P. grandiflora, but the perianth is usually only tinged with red-purple to purple. The viscidium length/ width ratio is a useful character for critical determinations. The hybrid has a ratio of 1.5–3.2, P. grandiflora has a ratio of 1–1.5, and P. lacera has a ratio of 2.7–4.6. A similar hybrid, P. ×‌andrewsii (11 × 16), has smaller anther sacs (1.4–2.3 mm long vs. 2.5–4.2 mm long in P. ×‌keenanii). 9. Platanthera hookeri (Torr. ex Gray) Lindl. N Hooker’s bog-orchid. Habenaria hookeri Torr. ex Gray; Lysias hookeriana (Torr. ex Gray) Rydb. • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic forests and forest edges. 10. Platanthera huronensis (Nutt.) Lindl. N Lake Huron green bog-orchid. Habenaria huronensis (Nutt.) Spreng.; H. hyperborea var. huronensis (Nutt.) Farw.; Platanthera hyperborea (L.) Lindl. var. huronensis (Nutt.) Luer; P. ×‌media (Rydb.) Luer • CT, MA, ME, NH, RI, VT. Meadows, fens, marshes, shorelines, ditches, seeps. 11. Platanthera lacera (Michx.) G. Don N green fringed bog-orchid. Blephariglottis lacera (Michx.) Farw.; Habenaria lacera (Michx.) R. Br. • CT, MA, ME, NH, RI, VT. Sand plains, fields, marshes, riparian forests, swamps. 11 ‌ × 16. Platanthera ×andrewsii (M. White) Luer is a rare bog-orchid hybrid in New England known from CT, MA, ME, NH, VT. On initial examination, it shows more dissected labellum segments than P. psycodes, but the perianth is usually only tinged with red-purple to purple. The viscidium length/width ratio is a useful character for critical determinations. The hybrid has a ratio of 1.3–3, P. lacera has a ratio of 2.7–4.6, and P. psycodes has a ratio of 1–1.5. A smilar hybrid, P. ×‌keenanii (8 × 11), has larger anther sacs (2.5–4.2 mm long vs. 1.4–2.3 mm long in P. ×‌andrewsii).

Fig. 174  Flower of Platanthera grandiflora.

210  MONOCOTS

12. Platanthera leucophaea (Nutt.) Lindl.

NC

eastern prairie white fringed bog-orchid. Blephariglottis leucophaea (Nutt.) Farw.; Habenaria leucophaea (Nutt.) Gray • ME; northern part of state. Circumneutral fens. 13. Platanthera macrophylla (Goldie) Lindl. N large-leaved bog-orchid. Habenaria macrophylla Goldie; H. orbiculata Pursh var. macrophylla (Goldie) Boivin; Platanthera orbiculata (Pursh) Lindl. var. macrophylla (Goldie) Luer • CT, MA, ME, NH, RI, VT. Mesic to wet-mesic forests. 14. Platanthera obtusata (Banks ex Pursh) Lindl. ssp. obtusata N blunt-leaved bog-orchid. Habenaria obtusata (Banks ex Pursh) Richards.; Lysiella obtusata (Banks ex Pursh) Rydb. • MA, ME, NH, VT. Evergreen swamps dominated by Thuja occidentalis, forested fens, wet meadows, evergreen and mixed evergreen-deciduous forests. 15. Platanthera orbiculata (Pursh) Lindl. N Fig. 175 round-leaved bog-orchid. Habenaria orbiculata (Pursh) Torr.; Lysias orbiculata (Pursh) Rydb. • CT, MA, ME, NH, RI, VT. Mesic to wet-mesic forests. 16. Platanthera psycodes (L.) Lindl. N Fig. 175  Flower of Platanthera orbiculata.

lesser purple fringed bog-orchid. Blephariglottis psycodes (L.) Rydb.; Habenaria psycodes (L.) Spreng. • CT, MA, ME, NH, RI, VT. Swamp edges, stream banks, ditches, trail edges, fields, mesic forests.

Pogonia 1. Pogonia ophioglossoides (L.) Ker-Gawl. N rose pogonia. Pogonia ophioglossoides (L.) Ker-Gawl. var. brachypogon Fern. • CT, MA, ME, NH, RI, VT. Fens, bogs, sandy meadows, ditches.

Spiranthes References: Sheviak (1982), Sheviak and Brown (2002). 1a. Labellum with a deep, subapical constriction at ca. ⅔ distance from the base, prominently veined, the branches of the veins wide-spreading toward the margins . . . . . S. romanzoffiana 1b. Labellum at most slightly constricted near the apex, if at all, with obscure to prominent veins that have parallel to somewhat diverging branches (S. lucida with short, widely diverging vein branches) 2a. Viscidium oval, 2–2.5 times as long as wide; perianth consisting of white sepals and lateral petals, the labellum with a large, bright yellow or orange-yellow area with green veins near the center; plants flowering in June through July, the lustrous basal leaves persisting long after anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lucida 2b. Viscidium lanceolate to linear, 3–12 times as long as wide; perianth not combining the above characteristics; plants flowering July through early October, the basal leaves senescing prior to or soon after anthesis 3a. Rachis of inflorescence glabrous 4a. Labellum white; root solitary (rarely divided at the tip), 4–12 mm wide; flowers flaring open from near the middle, the basal tubular portion shorter than 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tuberosa 4b. Labellum white with a central green or yellow-green spot; roots few to several, up to 7.5 mm wide; flowers flaring open at ca. ⅔ their length, the basal tubular portion longer than 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. lacera 3b. Rachis of inflorescence pubescent

o rc h i dac e a e   2 1 1

5a. Rachis of inflorescence mostly or entirely pubescent with non-glandular hairs [Fig. 178]; glands of rachis, if present, narrower or equal to width of stipe, therefore, the hairs pointed or cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. vernalis 5b. Rachis of inflorescence pubescent, in part, with glandular hairs [Fig. 176]; glands of rachis wider than stipe, therefore, the hairs capitate at their apex 6a. Inflorescence with 5 or more flowers per cycle of spiral, therefore, the spiral of flowers open and easily discernible; labellum 3.5–8 mm long 7a. Leaves narrow-ovate to obovate, 1–3.5 times as long as wide, strictly basal (sometimes withering prior to anthesis); labellum with a central green or yellow-green spot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. lacera 7b. Leaves narrow-lanceolate to oblanceolate, mostly 5–10 times as long as wide, basal and also on lower stem; labellum cream-white to green-white, sometimes yellow near center . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. casei 6b. Inflorescence with 3 or 4 flowers per cycle of spiral, therefore, the spiral obscure and the inflorescence dense, with flowers appearing arranged in vertical ranks; labellum (6–) 7.5–12 mm long 8a. Blade of labellum strongly curving donward from claw, the interior angle (i.e., measured on the abaxial face) 120–155 degrees [Fig. 177]; claw of labellum 0.8–1.5 mm long; maximum separation of lateral sepals from dorsal sepal mostly 0.5–1.1 mm [Fig. 177]; basal calli of labellum (0.8–) 1–1.6 (–2) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. ochroleuca 8b. Blade of labellum weakly curving downward from claw, the interior angle greater than 150 degrees [Fig. 176]; claw of labellum 0.3–0.8 mm long; maximum separation of lateral sepals from dorsal sepal mostly 0.1–0.5 mm [Fig. 176]; basal calli of labellum (0.5–) 0.7–1.1 (–1.2) mm long . . . . . . . . S. cernua 1. Spiranthes casei Catling & Cruise var. casei

NC

‌intermedia, Case’s ladies’-tresses. Spiranthes × auct. non Ames • ME, NH, VT; northern portion of states. Fields, low meadows, banks, disintegrating roadside ledges. 2. Spiranthes cernua (L.) L.C. Rich. N Fig. 176 nodding ladies’-tresses. Ibidium cernuum (L.) House; Ophrys cernua L. • CT, MA, ME, NH, RI, VT. Open, xeric to hydric sites, including roadsides, borrow pits, woodland openings, fields, and sandy, acid wetlands with a shallow horizon of peat. Both Spiranthes cernua and S. ochroleuca can rarely have open inflorescences of spirally secund flowers, similar to S. casei. They both differ from S. casei in having a rounded or acutely pointed labellum and larger flowers (labellum usually longer than 7.5 mm). Spiranthes casei has an apically truncate labellum usually shorter than 7.5 mm. 3. Spiranthes lacera (Raf.) Raf. N slender ladies’-tresses.  3a. Gyrostachys gracilis (Bigelow) Kuntze; Ibidium gracile (Bigelow) House; Neottia gracilis Bigelow; Spiranthes gracilis (Bigelow) Beck; 3b. Neottia lacera Raf. • CT, MA, ME, NH, RI, VT. Xeric to mesic meadows, fields, woodlands, and roadsides. 1a. Flowers relatively densely arranged on the spike, loosely to tightly spiraled, the ratio of spike length in mm to flower number less than 2.3; basal leaves usually absent at anthesis; rachis of inflorescence glabrous . . . . . . . . . . . . . . . . . . 3a. S. lacera var. gracilis (Bigelow) Luer 1b. Flowers relatively loosely arranged on the spike, secund to loosely spiraled, the ratio of spike length in mm to flower number equal to or greater than 2.3; basal leaves usually persisting through anthesis; rachis of inflorescence sparsely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. S. lacera var. lacera Variety gracilis is known from CT, MA, ME, NH, RI, VT. Vareity lacera is known from CT, MA, ME, NH, RI, VT. Variety gracilis is more restricted and is limited in ME to the southern portion of the state.

Fig. 176  Flowers of Spiranthes cernua with detail of stipitateglands on rachis.

21 2  MO NOCOTS

‌ × 7. Spiranthes ×eamesii P.M. Brown is a rare ladies’-tresses hybrid in New England 3a that is known from CT (it has also been reported from other southern New England states, but specimens are unknown), where it usually grows with both parents. It has a white labellum with green-yellow tinting on the labellum (those of S. tuberosa are pure white), usually 2 white, narrow-conical roots, and lacks leaves during anthesis (similar to both parents). The flowers are intermediate in size between the parents (4–7.5 mm long is S. lacera var. gracilis and 3–5.5 mm long in S. tuberosa).

Fig. 177  Flowers of Spiranthes ochroleuca.

‌3a × 8. Spiranthes ×intermedia Ames is a very rare ladies’-tresses hybrid in New England that is known from southeastern MA. It is similar to Spiranthes vernalis except that the pubescence of the inflorescence is slightly expanded apically into a slender gland (vs. pointed at apex) and the lateral petals show two primary veins, one of which usually branching to create three veins distally (vs. usually three mains originating from the base of each lateral petal). Spiranthes lacera var. gracilis shows stipitate-glands in the inflorescence and usually two unbranched veins on the lateral petals. Spiranthes × ‌intermedia is further characterized by yellow-green coloration along the center of the labellum that may become more yellow near the apex and a stipitate column (similar to S. vernalis). 4. Spiranthes lucida (H.H. Eat.) Ames N shining ladies’-tresses. Ibidium plantagineum Raf.; Neottia lucida H.H. Eat. • CT, MA, ME, NH, RI, VT. River and lake shores, most prevalent in areas influenced by high-pH bedrock, also in seeps and meadows. 5. Spiranthes ochroleuca (Rydb.) Rydb. N Fig. 177 yellow ladies’-tresses. Gyrostachys ochroleuca Rydb. in Britt.; Spiranthes cernua (L.) L.C. Rich. var. ochroleuca (Rydb.) Ames • CT, MA, ME, NH, RI, VT. Open, xeric to hydric sites, including roadsides, borrowpits, woodland openings, fields, and sandy, acid wetlands with a shallow horizon of peat. Rarely this orchid can have a relatively open, spirally secund inflorescence (see discussion under S. cernua). 6. Spiranthes romanzoffiana Cham. N

Fig. 178  Flowers of Spiranthes vernalis with detail of eglandular hairs on ovary.

hooded ladies’-tresses. Gyrostachys stricta Rydb.; Ibidium strictum (Rydb.) House; Spiranthes stricta (Rydb.) A. Nels. • CT, MA, ME, NH, VT. Mesic to hydric open sites including shorelines, meadows, and wetland edges. 7. Spiranthes tuberosa Raf. N little ladies’-tresses. Gyrostachys beckii (Lindl.) W. Stone, sensu Eames; Spiranthes tuberosa Raf. var. grayi (Ames) Fern. • CT, MA, RI. Fields, roadsides, open woodlands, lawns. 8. Spiranthes vernalis Engelm. & Gray N Fig. 178 spring ladies’-tresses. Ibidium vernale (Engelm. & Gray) House • CT, MA, NH, RI. Fields, roadsides, open woodlands, lawns.

Tipularia 1. Tipularia discolor (Pursh) Nutt.

NC

cranefly orchid. Tipularia unifolia B.S.P. • MA; Cape Cod region. Deciduous forests and woodlands on sandy soils, often near upland edge of swamps and ponds.

Triphora 1. Triphora trianthophora (Sw.) Rydb. ssp. trianthophora Fig. 179  Inflorescence of Triphora trianthophora.

N C Fig. 179

three-birds orchid. Arethusa trianthophora Sw.; Pogonia trianthophora B.S.P. • CT, MA, ME, NH, VT. Deciduous forests, usually in association with Fagus grandifolia on slopes and benches in hilly terrain.

P oac e a e   2 13

Poaceae Hesperostipa comata (Trin. & Rupr.) Barkworth was reported from RI by George (1997), but specimens are unknown. 1a. Spikelets appearing spiny, either enclosed in a bur-like fascicle of reduced branches or with an enlarged, uncinate-prickly glume [Figs. 204, 279] . . . . . . . . . . . . . . . . . . . . . . . . . . Group 1 1b. Spikelets not spiny, although pubescence or bristles may be present [Figs. 181, 227, 269] 2a. Spikelets with 2 or more evident florets [Figs. 232, 238, 282] 3a. Inflorescence an open to dense panicle [Figs. 180, 183, 198] 4a. Spikelets dimorphic with the 2 types paired—the lower spikelet of the pair sterile, flabellate, consisting of 2 glumes and several narrow, acuminate lemmas, concealing the upper, fertile spikelets with 2–4 florets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 2 4b. Spikelets of a single plant monomorphic 5a. At least 1 of the glumes (often both) nearly equaling to surpassing the body of the most apical lemma of each spikelet, partially or completely concealing the florets [Figs. 208, 282] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 3 5b. Both glumes shorter than the body of the most apical lemma of the spikelet, the florets largely visible [Figs. 227, 232, 283] . . . . . . . . . . . . . . . . . . . . . . . . . . Group 4 3b. Inflorescence a solitary, terminal spike or raceme, or a panicle with 2 or more secund, spike-like branches [Figs. 205, 224, 239] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 5 2b. Spikelets with only 1 evident floret (members of the subfamily Panicoideae key under this lead) [Figs. 181, 269, 273] 6a. Glumes or lemmas or both folded (i.e., V-shaped in cross-section), sometimes with a low to pronounced keel 7a. Inflorescence a panicle with secund, spike-like branches [Fig. 193] . . . . . . . Group 6 7b. Inflorescence an open to dense panicle, the branches not both secund and spikelike (often neither) or a secund spike [Figs. 182, 277] . . . . . . . . . . . . . . . . . . . . . . . Group 7 6b. Glumes and lemmas flat, arched, or rounded over the abaxial surface, lacking a keel 8a. Inflorescence composed of a single, terminal spike, raceme, or spike-like panicle, the branches, when present, scarcely discernible [Figs. 193, 236, 268] . . . . . . . Group 8 8b. Inflorescence a congested to open panicle with visible branching, or composed of 2 or more spikes, racemes, or rames 9a. Glumes often as long as or longer than the lemmas and concealing the florets; spikelets usually arranged in obvious pairs or triplets with 1 spikelet sessile or shortly pedicellate and the remaining pedicellate (sometimes the pedicellate spikelet reduced and represented by only a pedicel; not organized in pairs or triplets in Milium) [Fig. 240] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 9 9b. One (i.e., the lower) or both glumes shorter than the lemmas or the glumes absent; spikelets not organized in pairs or triplets (sometimes in groups of 2 or 3 in Digitaria and Paspalum) [Fig. 269] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 10

Group 1 1a. Bur formed from concrescent branchlets, surrounding and nearly concealing 1–4 spikelets [Fig. 204]; glumes without prickles; upper lemma and palea indurate . . . . . . . . . . . . . . Cenchrus 1b. Bur formed from an enlarged glume [Fig. 279], each subtending a single floret; glumes with 5–7 rows of uncinate prickles; lemma and palea not indurate . . . . . . . . . . . . . . . . . . . Tragus

21 4  MO NOCOTS

Group 2 only 1 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cynosurus

Group 3 1a. Spikelets, excluding the awns, 19–26 mm long; glumes 5- to 11-veined . . . . . . . . . . . . Avena 1b. Spikelets, excluding the awns, up to 18 mm long; glumes 1- to 5-veined 2a. Rachilla long-pubescent, the longer hairs (2–) 3–7 mm long; ligule of upper stem leaves 5–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Avenula 2b. Rachilla glabrous or pubescent with hairs up to 2 mm long; ligule of upper stem leaves as long as 5 (–6) mm (3–12 mm long in Deschampsia cespitosa and 1–8 mm long in Ventenata) 3a. Spikelets with 2 florets [Fig. 234] 4a. Spikelets 9–15 mm long; upper lemma with an awn 10–16 mm long, terminated by 2 setaceous teeth 1–2 mm long; upper glume 3- to 9-veined . . . . . . (in part) Ventenata 4b. Spikelets 1.7–11 mm long; upper lemma without an awn or with an awn (0.5–) 1–9 mm long, entire or bifid at the apex; upper glume 1- to 3-veined 5a. Lower lemma of spikelet (or both lemmas) lacking an awn 6a. Neither lemma of spikelet with an awn; reproductive stems glabrous or minutely scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Graphephorum 6b. Upper lemma of spikelet with a bent or hooked awn [Fig. 234]; reproductive stems pilose on at least the nodes (glabrous or sparsely pubescent in Holcus mollis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holcus 5b. Lower lemma of spikelet (and usually both lemmas) with an awn 7a. Florets of a given spikelet dimorphic—the lower unisexual, the upper bisexual; lemma awns dimorphic—that of the lower lemma 10–20 mm long and bent near the middle, that of the upper to 6 mm long and straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arrhenatherum 7b. Florets of a given spikelet monomorphic, both bisexual; lemma awns relatively monomorphic, similar in shape, the upper slightly shorter if at all 8a. Lemma awns composed of 2 distinctly unlike members—the basal portion brown and spiraled, the apical portion white and distally thickened— the two members separated by a ciliate joint . . . . . . . . . . . . . . Corynephorus 8b. Lemma awns tapering to the apex, not consisting of 2 articulate members separated by a ring of cilia 9a. Awn (when present) attached above the middle of the lemma [Fig. 281]; lemma apex bifid 10a. Lemmas with awns 3–9 mm long, the awns conspicuously surpassing the apex of the lemma [Fig. 281]; callus of lemma pubescent with hairs up to 1 mm long . . . . . . . . . . . . . . . . . . . . . . . . (in part) Trisetum 10b. Lemmas unawned or rarely with rudimentary awns shorter than 2 mm, the awns, when present, not or scarcely surpassing the apex of the lemma; callus of lemma pubescent with hairs 1.5–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Graphephorum 9b. Awn attached below the middle of the lemma [Fig. 208]; lemma apex entire to bifid 11a. Apex of lemma terminated by 2 setaceous teeth (caution: the terminal teeth are sometimes intertwined and difficult to observe);

Poac e a e   2 15

glumes 2.3–3.3 mm long; rachilla not prolonged beyond upper floret . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aira 11b. Apex of lemma entire or erose, but without terminal teeth; longer glumes 2.7–6.1 mm long; rachilla prolonged beyond upper floret 12a. Glumes slightly shorter than to slightly exceeding the lemmas [Fig. 208]; anthers 1.3–3 mm long; stems with prominent tufts of basal leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Deschampsia 12b. Glumes much exceeding lemmas, approximately twice as long [Fig. 282]; anthers 0.4–0.8 mm long; stems without prominent tufts of basal leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vahlodea 3b. Spikelets with 3 or more florets [Fig. 281] 13a. Lemma apex dissected into numerous, spreading awns . . . . . . . . . . . Pappophorum 13b. Lemma without awns or with a single awn 14a. Upper glume 3- to 9-nerved; upper lemma with an awn 10–16 mm long arising from the abaxial keel, terminated by 2 setaceous teeth 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Ventenata 14b. Upper glume 1- to 3-veined (up to 5 (–7)-veined in Danthonia, but that genus usually with more than 3 florets per spikelet); upper lemma without an awn or with an awn to 10.5 mm long, with or without apical teeth, the awn (when present) emerging from between the apical teeth or from the abaxial keel 15a. Spikelets with 3–12 florets; lemmas with an awn arising from between 2 terminal teeth; spikelets 7–16 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . Danthonia 15b. Spikelets with 3 or 4 florets; lemmas with or without awns, the awns, when present, not arising from between apical teeth; spikelets 3.2–9 mm long 16a. Upper floret bisexual, of different form from the 2 lower, unisexual and staminate florets; 2 lower lemmas evidently pubescent over the surfaces, at least near the apex (note: the hairs visible at low magnification) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anthoxanthum 16b. Florets all bisexual, progressively reduced in size upward; lemmas minutely and/or rather sparsely scabrous-hirtellous over their surfaces (note: the callus and rachilla segments may be evidently pubescent; lemmas pilose in some forms of Trisetum spicatum) 17a. Panicle branches densely pubescent; apex of lemma entire; rachilla segments glabrous or pubescent with hairs shorter than 1 mm . . . . Koeleria 17b. Panicle branches antrorsely scabrous; apex of lemma bifid; rachilla segments pubescent with hairs 0.4–2 mm long, these usually longer than 1 mm in the distal portion of the segment 18a. Lemmas with awns 3–9 mm long, the awns conspicuously surpassing the apex of the lemma [Fig. 281]; callus of lemma pubescent with hairs up to 1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Trisetum 18b. Lemmas unawned or with awns shorter than 2 mm, the awns, when present, not or scarcely surpassing the apex of the lemma; callus of lemma pubescent with hairs 1.5–2 mm long . . . . (in part) Graphephorum

Group 4 1a. Leaf blades narrowed to a short pseudopetiole 2a. Branches from midpoint of stem solitary; reproductive stems terete; spikelets pedicellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudosasa

216 MO NOCOTS

2b. Branches from midpoint of stem in pairs, strongly unequal; reproductive stems grooved on one side; spikelets sessile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllostachys 1b. Leaf blades not narrowed to a petiole-like structure 3a. Plants robust, typically 1.5–4 m tall, forming extensive colonies by stout rhizomes; inflorescence appearing plumose due to long-pubescent rachilla segments . . . . . Phragmites 3b. Plants of short to moderate stature, mostly shorter than 2 m, colonial or not; inflorescence not appearing plumose 4a. Lemmas with 3 veins 5a. Callus and/or nerves of the lemma pubescent 6a. Lemmas entire at the apex, without awns [Fig. 260], arachnoid-pubescent on the callus in most species [Fig. 261]; ligule membranaceous . . . . . . . . . (in part) Poa 6b. Lemmas bifid at the apex, with an awn emerging from between the lobes (note, the lateral lobes may terminate in long awns), pubescent, but not with arachnoid hairs; ligule a band of hairs 7a. Lemmas with 3 slender, elongate awns—1 terminal awn and 2 lateral awns; upper glume bifid at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triraphis 7b. Lemmas with 1 slender awn or 3 excurrent mucros; upper glume entire at the apex 8a. Inflorescence terminal, long-exserted from upper leaf sheaths; paleas glabrous or inconspicuously short-pubescent; lateral veins of lemma shortly excurrent as short awns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tridens 8b. Inflorescence terminal and axillary, usually partly included in the leaf sheaths; paleas densely villous on the apical portion of the 2 keels; lateral veins of lemma not excurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triplasis 5b. Callus and nerves of the lemma glabrous 9a. Lemmas keeled on the abaxial surface (except E. intermedia), longer than the paleas; anthers 0.2–1.2 mm long; rachilla typically not disarticulating, the floral scales falling from it . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eragrostis 9b. Lemmas rounded on the abaxial surface, shorter than the paleas; anthers 1.5–3 mm long; rachilla disarticulating . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Molinia 4b. Lemmas with 5 or more veins (the intermediate veins sometimes inconspicuous) 10a. Panicle composed of dense, somewhat secund, glomerules of spikelets on a few, stiff branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dactylis 10b. Panicle composed of pedicellate spikelets on spirally arranged branches 11a. Plants dioecious (i.e., the flowers of a given plant unisexual and all of one type), of coastal saltmarshes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Distichlis 11b. Plants synoecious (i.e., the flowers of a given plant bisexual), mostly not of saline habitats (except Puccinellia) 12a. Leaf sheath fused (i.e., the sheath closed with united edges) at least in the basal half 13a. Lemmas entire at the apex, without awns, with parallel veins that do not converge near the apex [Fig. 265] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glyceria 13b. Lemmas bifid at the apex, awned in most species [Fig. 196], with arching veins that converge near the apex 14a. Callus of the lemma pubescent; styles borne close together from the summit of the ovary; caryopsis free from the palea . . . . . . . . . Schizachne

P oac e a e   2 17

14b. Callus of the lemma glabrous; styles separated and borne below the summit of the ovary; caryopsis adnate to the palea . . . . . . . . . . . . . Bromus 12b. Leaf sheath with distinct, overlapping edges except at the very base 15a. Glumes dissimilar in shape—the first (i.e., lower) oblong or tapering to the apex, the second obovate and dilated upwards [Figs. 274, 275] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sphenopholis 15b. Glumes relatively similar in shape, though sometimes differing in size, both with parallel or apically tapering margins [Figs. 208, 281] 16a. Lemmas awned, the awn attached to the abaxial surface of the lemma well below the apex [Fig. 208] 17a. Spikelets 9–10 mm long; awn of upper lemmas of spikelet 10–16 mm long; upper glume 3- to 9-veined . . . . . . (in part) Ventenata 17b. Spikelets 2.3–7.5 mm long; awn of lemmas up to 6.5 mm long; upper glume 1- to 3-veined 18a. Awn emerging below the middle of the lemma keel; lemma apex entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Deschampsia 18b. Awn emerging above the middle of the lemma keel; lemma apex bifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Trisetum 16b. Lemmas unawned or with an awn attached at the apex [Fig. 265] 19a. Florets nearly perpendicular to the axis of spikelet; lemmas cordate at their base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Briza 19b. Florets ascending to erect relative to the spikelet axis; lemmas cuneate to rounded or truncate at their base 20a. Lemmas usually blunt at the apex, with parallel veins that do not converge near the tip [Figs. 264, 265] 21a. Lemmas prominently veined; plants of freshwater habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Torreyochloa 21b. Lemmas inconspicuously veined; plants of saline habitats (e.g., saline to brackish marshes, coastal strands, heavily salted roadways) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Puccinellia 20b. Lemmas usually pointed or awned or both at the apex, with arching veins that converge near the tip 22a. Rachilla segments with hairs 1.3–2 mm long; callus of lemma pubescent with straight hairs . . . . . . . . . (in part) Graphephorum 22b. Rachilla segments glabrous or minutely scabrous; callus of lemma glabrous or with arachnoid hairs 23a. Lemmas prominently keeled, often with arachnoid pubescence on the callus [Fig. 261], without awns; leaf tips cucullate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Poa 23b. Lemmas rounded, or keeled only at the apex, with a glabrous callus, usually awned; leaf tips flat 24a. Panicles with short, ± secund branches; lemmas obtuse at the apex, unawned; florets disarticulating with a short rachilla segment such that the fallen florets appear stipitate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Desmazeria

218 MO NOCOTS

24b. Panicles with short to long branches, not secund; lemmas acute to acuminate (obtuse), awned in most species [Fig. 283]; florets disarticulating without a short rachilla segment or with 1 but then the floret not appearing stipitate 25a. Plants annual; flowers remaining closed (i.e., cleistogamous; anthers usually 1 per floret, up to 1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vulpia 25b. Plants perennial; flowers open-pollinated (i.e., chasmogamous) or proliferating in one species; anthers 3 per floret, (0.5–) 0.7–4 mm long 26a. Auricles present at summit of leaf sheath [Fig. 266]; apical half of the lemma with a prominent, scarious margin ca. 0.5 mm wide; leaf blades flat, (2–) 3–18 mm wide . . . . . . . . . . . . . . . . . . . Schedonorus 26b. Auricles absent; lemmas with firm margins or very narrow, inconspicuous, scarious margins; leaf blades involute, conduplicate, or flat, 0.2–3 mm wide or up to 10 mm in F. subverticillata . . . . . Festuca

Group 5 1a. Lemmas with 2, 3, or 5 conspicuous lobes or teeth at the apex 2a. Lemmas 3- or 5-lobed at the apex and with 3 awns; spikelets not sunken into cavities of the inflorescence rachis; inflorescence a panicle consisting of an axis and spike-like branches; disarticulation occurring at the base of the short, spike-like branches, these falling from the panicle axis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Bouteloua 2b. Lemmas with 2 or 3 tooth-like lobes at the apex, usually only 1 of these prolonged into an evident awn (note: the awns usually longest on spikelets in the apical portion of the inflorescence); spikelets appearing sunken into obvious cavities in the thickened rachis; inflorescence a cylindrical spike; disarticulation at the nodes of the spike . . . . . . . . Aegilops 1b. Lemmas entire or inconspicuously bilobed at the apex 3a. Inflorescence a solitary spike or raceme, the spikelets produced from nodes that alternate of each side of the rachis [Figs. 224, 239] 4a. All but the uppermost spikelets borne on short pedicels (i.e., the inflorescence a raceme) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachypodium 4b. All the spikelets sessile (i.e., the inflorescence a spike) [Figs. 224, 239] 5a. Spikelets solitary at each node of the inflorescence [Fig. 239] 6a. Spikelets arranged with the narrow side toward the rachis; only the glume away from the rachis developed, the other obsolete . . . . . . . . . . . . . . . . . . . . . Lolium 6b. Spikelets arranged with the broad side toward the rachis; both glumes developed 7a. Plants annual, short persisting crop species; glumes 1- or 3-veined 8a. Glumes with 1 vein, linear-subulate; keel of lemmas ciliate . . . . . . . Secale 8b. Glumes with 3 veins, ovate; keel of lemmas glabrous . . . . . . . . . . Triticum 7b. Plants perennial, native or introduced species; glumes 5- or 7-veined 9a. Spikelets crowded on short internodes 0.5–2 (–3) mm long, ascending to spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agropyron

Poac e a e   2 19

9b. Spikelets less crowded, at least the lower internodes mostly longer than 4 mm, appressed to appressed-ascending 10a. Glumes gradually tapering from near midlength and ± imperceptively passing into a short awn-tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pascopyrum 10b. Glumes tapering from the apical third, unawned or somewhat abruptly tapering to an awn 11a. Glumes acute at the apex, unawned or merely with a tiny mucro up to 0.5 (–0.7) mm long [Fig. 278] . . . . . . . . . . . . . . . . . . . . . Thinopyrum 11b. Glumes long-acute at the apex, usually with an awn 0.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Elymus 5b. Spikelets numbering 2–4 at each node, although only 1 may be fertile [Figs. 220, 224] 12a. Spikelets consistently 3 at each node, the outer pair usually pedicellate and staminate or sterile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Hordeum 12b. Spikelets 2–4 at each node, commonly 2, all bisexual and fertile 13a. Plants annual; lemmas with awns 50–100 cm long . . . . . . . Taeniatherum 13b. Plants cespitose or rhizomatous perennials; lemmas awnless or with awns up to 40 mm long 14a. Plants mostly self-pollinating, with small anthers 1–3 mm long, cespitose, lacking rhizomes or with short rhizomes; lemmas mostly awned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Elymus 14b. Plants mostly outcrossing, with large anthers mostly 4–9 mm long, not cespitose, with elongate, stout rhizomes; lemmas mostly without awns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leymus 3b. Inflorescence a panicle with 2 or more secund, spike-like branches [Fig. 205] 15a. Panicle with branches digitately arranged (i.e., all or nearly all originating from the same point) [Fig. 205] or with branches in 1–3 closely spaced whorls 16a. Rachis of branches extending beyond distal most flowers 1–5 mm; upper glume with an awn 1–2.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dactyloctenium 16b. Rachis of branches ending in a functional or rudimentary spikelet, not extending as a bristle-tip; glumes unawned or the upper one with a short awn to 0.3 mm 17a. Lemmas awned, pubescent on the marginal veins and callus; spikelets with 2 or 3 (–5) florets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chloris 17b. Lemmas unawned, glabrous; spikelets with 5–7 florets . . . . . . . . . . . . . Eleusine 15b. Panicle with branches racemosely arranged along an elongate axis 18a. Spikelets nearly circular in outline, with 1 or 2 florets, the upper, when present, sterile; glumes obovate, cross-wrinkled, the tips crossing. . . . . . (in part) Beckmannia 18b. Spikelets not circular in outline, either clearly longer than wide or ± obovate, with 2–20 bisexual florets; glumes linear to elliptic to ovate, not cross-winkled, the tips separate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptochloa

Group 6 1a. Spikelets nearly circular in outline; glumes obovate, cross-wrinkled, the tips crossing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Beckmannia 1b. Spikelets not at all circular in outline; glumes linear to lanceolate, not cross-winkled, the tips separate

22 0  MO NOCOTS

2a. Panicle with (2–) 4–6 (–9) digitately arranged branches, lacking an elongate, central axis; plants mat-forming by stolons (and often also rhizomes) . . . . . . . . . . . . . . . . . . Cynodon 2b. Panicle with (1–) 2–80 branches variously attached to an elongate axis; plants without evident stolons 3a. Lemmas entire or bilobed at the apex, unawned or merely apiculate, exceeded by the paleas [Figs. 272, 273]; spikelets with a solitary, bisexual floret; ligules 0.3–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spartina 3b. Lemmas 3-lobed or 5-lobed at the apex, with 3 awns, exceeding the paleas; spikelets usually with a reduced, sterile floret distal to the bisexual floret; ligules 0.1–0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Bouteloua

Group 7 1a. Inflorescence a solitary, terminal spike; spikelets arising from shallow cavities on the rachis, with a single, minute glume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nardus 1b. Inflorescence a panicle or raceme (i.e., the spikelets at least shortly pedicellate), usually branched (though the branches sometimes difficult to discern); spikelets not arising from cavities on the rachis, with 2 glumes (glumes essentially absent in Leersia) 2a. Inflorescence a dense, cylindrical, spike-like panicle, the branches virtually absent 3a. Inflorescence overtopped by the upper leaves, usually partly sheathed or included in the subtending, apically inflated leaf sheath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crypsis 3b. Inflorescence exceeding the leaves, usually well exserted and not included in the upper leaf sheaths 4a. Glumes without awns, connate at the base; paleas wanting . . . . . . . . Alopecurus 4b. Glumes with awns, distinct at the base; paleas present, subequal in length to the lemmas 5a. Glumes shorter than and not concealing the floret, the first glume with 2 (–3) awns, the second with a single awn . . . . . . . . . . . . . . . . . . . . . (in part) Muhlenbergia 5b. Glumes exceeding the length of the floret and concealing it [Fig. 251], glumes awnless or each glume with a single awn 6a. Glumes tapering to a long point, but not awned, slightly swollen at the base; lemmas pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gastridium 6b. Glumes awned [Fig. 251], not swollen at the base; lemmas glabrous over the surface 7a. Awn of glumes 3–8 mm long; lemmas with awns; glumes hispidulous, bilobed at the apex, the awn emerging from between the lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Polypogon 7b. Awn of glumes 0.7–3 mm long; lemmas without awns; glumes prominently ciliate on the keels [Fig. 251], otherwise glabrous, entire at the apex, the awn terminal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phleum 2b. Inflorescence an open or dense panicle, the branches identifiable 8a. Lemmas indurate and lustrous at maturity, of much firmer texture than the glumes 9a. Lemmas awnless; spikelets with 1 fertile floret and at its base 2 sterile, rudimentary florets represented by pubescent scales . . . . . . . . . . . . . . . . . . . . . Phalaris 9b. Lemmas with awns; spikelets without basal, sterile florets 10a. Lemmas pubescent on only the callus, with 3 persistent awns [Figs. 189, 190]; callus of lemmas slender and acute; plants annual (perennial in 1 species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Aristida

Poac e a e   2 2 1

10b. Lemmas pubescent over the abaxial surface, with a single awn articulated and eventually deciduous from the summit of the lemma [Fig. 254]; callus of lemmas short and oblique; plants perennial 11a. Panicle open to somewhat congested, the branches spreading to ascending, each branch bearing usually 2 or more spikelets; uppermost leaf blades well-developed, 1–35 cm long; basal leaves deciduous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Piptatherum 11b. Panicle very narrow, raceme-like, with appressed branches, each branch bearing usually a single spikelet; uppermost leaf blades absent or with a blade shorter than 1 cm; basal leaves evergreen . . . . . . . . . . . . . . . . . (in part) Oryzopsis 8b. Lemmas soft and dull at maturity, not of conspicuously firmer texture than the glumes 12a. Glumes virtually obsolete, reduced to minute cupules; nodes densely and conspicuously pubescent with spreading, white hairs [Fig. 237] . . . . . . . . . . . . . Leersia 12b. Glumes present and evident; nodes glabrous or pubescent, but not pubescent with spreading, white hairs 13a. Floret stipitate, with 1 stamen; spikelets articulated below the glumes and falling intact leaving the pedicel behind . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinna 13b. Floret sessile, with mostly 3 stamens; spikelets articulated above the glumes, the florets falling and leaving the empty glumes behind (disarticulating below the glumes in Polypogon) 14a. Spikelets 10–15 mm long; plants growing on sandy beaches of the Atlantic coast and Lake Champlain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ammophila 14b. Spikelets 1–9 (–10) mm long; plants growing in a wide variety of habitats, but not regularly of coastal sands 15a. Ligule of hairs; lemmas 1-veined . . . . . . . . . . . . . . . . . (in part) Sporobolus 15b. Ligule a membrane; lemmas 3- or 5-veined (the veins sometimes faint) 16a. Glumes shorter than the lemmas or the lemmas awned from the very apex or both; caryopsis tightly enclosed by the lemma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Muhlenbergia 16b. At least 1 of the glumes equaling the length of the floret; awn of the lemma, if present, originating from below the apex on the keel of the lemma or from between 2 terminal lobes; caryopsis loosely enclosed by the lemma 17a. Glumes obtuse; inflorescence a raceme with usually 6–8 spikelets; delicate annuals 3–10 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mibora 17b. Glumes acute to acuminate; inflorescence a panicle, with more than 8 spikelets; annual or mostly perennial plants (5–) 10–200 cm tall 18a. Lemma glabrous on the callus (pubescent in Agrostis elliottiana); palea nerveless, or, less often, 2-nerved, delicate, up to 65% as long as the lemma; rachilla not prolonged 19a. Spikelets disarticulating above the glumes, borne on pedicels (0.1–) 0.3–9.6 mm long; lemmas (0.8–) 1.2–2.6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agrostis 19b. Spikelets disarticulating below the glumes, borne on stipes 0.1–0.6 mm long; lemmas 0.5–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Polypogon 18b. Lemma evidently, though sometimes shortly, pubescent on the callus [Figs. 202, 203]; palea 2-nerved, membranaceous, ½ to fully as

22 2  MO NOCOTS

long as the lemma; rachilla prolonged beyond the upper floret as a minute bristle 20a. Awn straight, inserted just below the apex and between the terminal teeth of the lemma, 5–12 mm long; pubescence of the callus of the lemma ca. ⅕ as long as the lemma; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apera 20b. Awn often bent, definitely originating from below the apex of the lemma, up to 3 mm long; pubescence of the callus of the lemma ¼ to fully as long as the lemma; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calamagrostis

Group 8 1a. Lemmas 3- or 5-lobed at the apex, with 3 awns; ligules 0.1–0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Bouteloua 1b. Lemmas entire or bilobed at the apex, awnless or with a single awn; ligules 0.5–16 mm long 2a. Inflorescence a spike, with spikelets positioned on alternating sides of the rachis [Figs. 235, 236]; spikelets 6–150 mm long including the awns . . . . . . . . . . (in part) Hordeum 2b. Inflorescence a spike-like panicle, with spirally arranged or verticillate spikelets [Fig. 270]; spikelets 1.6–6 (–8) mm long including the awns 3a. Glumes without awns, the upper with 3–9 veins; spikelets subtended by 1–12 bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Setaria 3b. Glumes with an awn, both with 1 vein; spikelets not subtended by bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Polypogon

Group 9 1a. Spikelets not organized into pairs or triplets, all pedicellate in an open panicle; disarticulation above the glumes, the falling floret leaving the glumes behind . . . . . . . . Milium 1b. Spikelets organized into pairs or triplets, some sessile (all pedicellate in Miscanthus, but that genus with glumes subtended by a ring of long, silky hairs); disarticulation below the glumes or between the rame segments, in either case the spikelets falling intact 2a. Spikelets unisexual, with the carpellate inflorescences or carpellate portion of inflorescence borne below the staminate ones or staminate portions [Fig. 280] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tripsacum 2b. At least the lower spikelet of each pair or triplet bisexual 3a. Reproductive stems decumbent, creeping over the ground or other low vegetation; spikelets solitary at each node, the pediceled spikelet absent, represented only a pedicel up to 3 (–4) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arthraxon 3b. Reproductive stems ascending to erect, not creeping; spikelets solitary, paired, or in trios at each node, when solitary, with a pedicel usually longer than 3 mm 4a. Spikelets in homogamous and homomorphic pairs (i.e., both spikelets of a pair of similar sex and similar morphology, respectively) [Fig. 240] 5a. Spikelets in sessile-pedicellate pairs (i.e., one spikelet of a pair sessile, the other borne on a short pedicel) [Fig. 240]; terminal inflorescence of (1–) 2–4 (–6) branches originating from the summit of the peduncle, without or with a very short rachis less than 4 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microstegium 5b. Spikelets in unequal-pedicellate pairs (i.e., both spikelets of a pair pedicellate, but 1 spikelet on a shorter pedicel than the other); terminal inflorescence usually with more than 15 branches distributed along a rachis 4–15 cm long . . . . . . . . . Miscanthus

Poac e a e   2 23

4b. Spikelets in heterogamous and heteromorphic pairs or trios (i.e., spikelets of a pair or trio not of similar sex nor of similar morphology, respectively) 6a. Fully formed spikelet solitary at each node, the pedicelled spikelet represented by only its pedicel 7a. Sessile spikelet lacking an awn; pedicels and rame internodes glabrous; mat-forming, stoloniferous plants with reproductive stems 10–35 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Eremochloa 7b. Sessile spikelet with an awn; pedicels and rame internodes pubescent; cespitose or rhizomatous plants with reproductive stems 40–300 cm tall 8a. Pedicels and rame internodes pubescent with white hairs [Fig. 186]; glumes glabrous or scaberulous; peduncles subtended by, or even partly included in, a modified, shortened leaf . . . . . . . . . . . . . . (in part) Andropogon 8b. Pedicels and rame internodes pubescent with yellow-brown hairs; lower glume hirsute; peduncles not closely subtended by a modified leaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sorghastrum 6b. Spikelets paired or in trios at each node, with 1 or 2 pedicelled spikelets, these usually smaller than the sessile spikelet 9a. Each inflorescence peduncle supporting a solitary rame (though each plant may have many peduncles) 10a. Inflorescence solitary and terminal; pedicels and rame internodes glabrous; reproductive stems 10–35 cm tall . . . . . . . . . . (in part) Eremochloa 10b. Inflorescences both terminal and axillary; pedicels and rame internodes pubescent; reproductive stems 30–210 cm tall . . . . . . . . . . . . . Schizachyrium 9b. Each inflorescence peduncle supporting 2 or more rames or branches 11a. Inflorescence a panicle with compound branches distributed along an elongate axis, the ultimate branches of the panicle being rames; pedicel of pedicellate spikelet 1–3.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sorghum 11b. Inflorescence composed of simple, digitately arranged rames, lacking an elongate central axis; pedicel of pedicellate spikelet (2.7–) 3–5.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Andropogon

Group 10 1a. Plants monoecious, the carpellate and staminate inflorescences or portions of inflorescences clearly different from one another; glumes absent, highly reduced, or collapsed (i.e., not evident); robust, annual grasses 1–5 (–6) m tall 2a. Each inflorescence with 2 types of flowers, the staminate positioned below the carpellate; carpellate spikelets with elongate awns; aquatic grasses . . . . . . . . . . . . . . Zizania 2b. Each inflorescence with only 1 type of flower, the carpellate inflorescences axillary, the staminate inflorescences terminal; carpellate spikelets lacking awns; cultivated grasses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zea 1b. Plants synoecious or polygamous, the inflorescence with at least some (or entirely of) bisexual flowers; glumes present (though sometimes the lower one absent); short to tall, annual or perennial grasses 0.15–2.5 (–3) m tall 3a. Lemmas without awns [Fig. 269] 4a. Spikelets at maturity with 1 pair of lemmas and paleas indurate and lustrous, of much firmer texture than the glumes 5a. Spikelets each subtended by 1–12 bristles [Fig. 269]; inflorescence a solitary, terminal, spike-like panicle [Figs. 268, 270] . . . . . . . . . . . . . . . . . . . . . . . . (in part) Setaria

22 4  MO NOCOTS

5b. Spikelets not subtended by bristles; inflorescence an open to dense panicle 6a. Upper leaves (or all the leaves) without ligules . . . . . . . . (in part) Echinochloa 6b. Leaves with ligules of hairs, membranes, or ciliate membranes 7a. Panicle branches both unbranched and secund [Figs. 217, 249] 8a. Spikelets lanceoloid to ellipsoid; upper (i.e., fertile) lemma somewhat indurate, firm but flexible, the margin with a hyaline border and flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Digitaria 8b. Spikelets ovoid or obovoid to spherical; upper lemma prominently indurate, rigid to the edges and lacking a hyaline border, the very edges slightly involute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paspalum 7b. Panicle branches rebranched (at least the lower ones) or not secund or usually both [Figs. 216, 246, 247] 9a. Plants with both aerial and subterranean inflorescences, only the subterranean ones producing fruits; lower glumes absent . . . . Amphicarpum 9b. Plants with only aerial inflorescences, at least some spikelets producing fruits; lower glumes present, though sometimes very small 10a. Upper (i.e., fertile) lemma somewhat indurate, firm but flexible, the margin with a hyaline border and flat . . . . . . . . . . . . . . . . . (in part) Digitaria 10b. Upper lemma prominently indurate, rigid to the edges and lacking a hyaline border, the very edges slightly involute 11a. Upper lemma transversely rugose; panicle branches spike-like, secund, with a triquetrous axis . . . . . . . . . . . . . . . . . . . . . . . . . . . Urochloa 11b. Upper lemma not transversely rugose (verrucose in Panicum verrucosum, but the warts not aligned in rows); panicle branches openbranched to highly congested, not secund (+/- secund in Sorengia), without a triquetrous axis 12a. Inflorescence dense, spike-like, concealing at least the apical ½ of the inflorescence rachis; pedicels with a disk-like apex; upper glume saccate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sacciolepis 12b. Inflorescence an open to somewhat congested panicle, the inflorescence rachis visible; pedicels not or barely thickened at apex, not disk-like; upper glume not saccate 13a. Basal leaves lanceolate to ovate, of distinctly different shape and/or length than the stem leaves and forming an overwintering rosette (elongate and similar to stem leaves in 2 species); many species producing branches from the lower and middle stem nodes in summer, these sometimes rebranching; reduced secondary panicles produced bearing cleistogamous flowers and often partly concealed by subtending leaves; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dichanthelium 13b. Basal leaves linear to lanceolate, similar to the stem leaves, not forming an overwintering rosette; many species not producing branches from the lower and middle nodes, or branches produced but not rebranched; secondary panicles similar to the terminal panicles, chasmogamous; plants annual or perennial 14a. Spikelets borne on short pedicels mostly 0.5–1.5 mm long, ± secund [Fig. 271]; leaf sheaths strongly compressed; apex of lemma and palea of upper floret lacking papillae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sorengia

Poac e a e   2 25

14b. Spikelets borne on pedicels (0.5–) 1–20 mm long, not secund; leaf sheaths ± terete or compressed in P. dichotomiflorum; apex of lemma and palea of upper floret with simple or compound papillae . . . . . . . . . . . Panicum 4b. Spikelets with soft to firm lemmas, the lemmas dull and not conspicuously indurate 15a. Spikelet with an upper bisexual, fertile floret and a lower inconspicuous, sterile floret, articulated below the glumes, the entire spikelet falling and leaving the pedicel behind; upper glume and lower lemma usually of ± similar appearance and texture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Digitaria 15b. Spikelet with 1 bisexual, fertile floret, articulated above the glumes, the floret falling leaving the empty glumes behind; upper glume and lemma not of similar appearance and texture 16a. Lemmas 1-veined, unawned; ligule of hairs . . . . . . . . . . . . (in part) Sporobolus 16b. Lemmas 3-veined, awned in most species; ligule a membrane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Muhlenbergia 3b. Lemmas with awns [Figs. 189, 254, 255] 17a. Upper leaves (or all the leaves) without ligules . . . . . . . . . . . . . (in part) Echinochloa 17b. Leaves with ligules of hairs, membranes, or ciliate membranes 18a. Lemmas with 3 awns at the apex; plants annual (or perennial in 2 species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Aristida 18b. Lemmas with a single awn from the apex; plants perennial 19a. Lemmas indurate and lustrous, conspicuously firmer than the glumes, pilose over the abaxial surface (only in the basal ⅓ in Piptochaetium); awn of lemma articulated, persistent or eventually deciduous 20a. Lemma awn 45–70 mm long; spikelets 9–13 mm long excluding the awns; lemmas with 3 distinct zones—villous in basal ⅓, glabrous in the middle, and scabrous near the apex [Fig. 255] . . . . . . . . . . . . . . . . . . . . . . . . . . . . Piptochaetium 20b. Lemma awn 0.5–14 mm long; spikelets 3.5–9 mm long excluding the awns; lemmas sparsely to densely pubescent ± throughout 21a. Panicle open to somewhat congested, the branches spreading to ascending, each branch usually bearing 2 or more spikelets; uppermost leaf blades well-developed, 1–35 cm long; basal leaves deciduous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Piptatherum 21b. Panicle very narrow, raceme-like, with appressed branches, each branch usually bearing a single spikelet; uppermost leaf blades absent or with a blade shorter than 1 cm; basal leaves evergreen . . . . . . . . . (in part) Oryzopsis 19b. Lemmas firm, but not indurate, dull, glabrous or scaberulous, pubescent only on the nerves or callus if at all (minutely pubescent between the nerves in some Muhlenbergia); awn of lemma not articulated, persistent 22a. First glume diminutive or obsolete; rachilla prolonged, located near the abaxial surface of the palea and appearing as a minute bristle; plants with rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachyelytrum 22b. First glume present and evident (except in M. schreberi, which lacks rhizomes); rachilla not prolonged; plants with or without rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Muhlenbergia

22 6  MO NOCOTS

Aegilops 1. Aegilops cylindrica Host E jointed goat-grass. Cylindropyrum cylindricum (Host) A. Löve; Triticum cylindricum (Host) Ces., Pass., & Gib. • MA. Roadsides, fields, railroads.

Agropyron 1a. Spikes 8–23 mm wide; spikelets diverging from the rachis at an angle of 45–90 degrees; glumes not appressed to lemmas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. cristatum 1b. Spikes 5–10 mm wide; spikelets diverging from the rachis at an angle of 25–45 degrees; glumes appressed to lemmas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. desertorum 1. Agropyron cristatum (L.) Gaert. E crested wheat grass. Agropyron pectiniforme Roemer & J.A. Schultes • MA, NH. Fields, roadsides, waste places. 2. Agropyron desertorum (Fisch. ex Link) J.A. Schultes E clustered wheat grass. Agropyron cristatum (L.) Gaert. ssp. desertorum (Fisch. ex Link) A. Löve; A. cristatum (L.) Gaert. var. desertorum (Fisch. ex Link) Dorn • MA. Fields, roadsides, waste places.

Agrostis When determining the presence of an awn borne from the keel of the lemma, refer to the spikelets near the distal portions of branches. Spikelets from near the base of the branches may show shorter awns that are positioned closer to (or at) the tip of the lemma. Reports of Agrostis pallens Trin. from MA (e.g., Kartesz 1999, Sorrie and Somers 1999) were based on a specimen of Agrostis stolonifera. Reference: Harvey (2007). 1a. Palea 2-nerved, 0.6–1.4 mm long, (40–) 50% or more as long as the lemma; plants mostly with rhizomes and/or stolons 2a. Ligules 0.5–2 mm long, usually wider than long; branches of panicle bearing flowers only in the distal half; horizontal stems (i.e., rhizomes or stolons) up to 5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. capillaris 2b. Ligules 2–6 mm long, usually longer than wide; branches of panicle bearing flowers in both distal and proximal halves [Fig. 180]; horizontal stems usually longer than 5 cm 3a. Inflorescence with spreading branches during and after anthesis [Fig. 180], (8–) 10–25 cm tall, usually red-purple; plants with rhizomes and erect stems; leaf blades 3–9 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. gigantea 3b. Inflorescence with ascending to appressed branches after anthesis [Fig. 182], 3–10 (–15) cm tall, usually yellow-brown; plants without rhizomes, but with stolons and basally decumbent stems; leaf blades 2–5 mm wide . . . . . . . . . . . . . . . . A. stolonifera 1b. Palea unnerved and up to 0.5 mm long or absent, less than 33% as long as the lemma; plants cespitose, without prolonged rhizomes or stolons (except A. canina, which often has stolons) 4a. Plants annual; lemmas usually with a delicate, flexuous awn (3–) 6–10 mm long; callus of the lemma densely pubescent with hairs to 0.6 mm long . . . . . . . . . . . . . . . A. elliottiana 4b. Plants perennial; lemmas with or without awns, the awns, if present, straight or geniculate and up to 5 mm long; callus of the lemma glabrous or sparsely to moderately pubescent with hairs to 0.3 mm long 5a. Panicle dense and spike-like, with ascending to appressed lower branches 1–2 (–4) cm long that are partly concealed by the spikelets . . . . . . . . . . . . . . . . A. exarata

P oac e a e   2 27

5b. Panicle open, with ascending to wide-spreading lower branches 2–12 cm long that are readily visible 6a. Lemmas awnless (very rare individuals of A. perennans producing some spikelets with short awns to 2 mm, but these usually not exserted beyond the glumes) 7a. Leaf blades flat, 2–5 mm wide, the basal ones usually withered by anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. perennans 7b. Leaf blades involute, 0.5–2 mm wide, the basal ones persisting all year in a tuft 8a. Anthers globose, 0.2–0.3 (–0.5) mm long; glumes 1–2 mm long; lemmas 0.8–1.2 mm long; spikelets ± evidently clustered near the tips of the panicle branches; pedicels 0.3–2 mm long; reproductive stems often with more than 3 leaf-bearing nodes; plants flowering March through July . . . . . . . . A. hyemalis 8b. Anthers oblong, 0.4–0.8 mm long; glumes 1.8–3.4 mm long; lemmas 1.4–2 mm long; spikelets not appearing clustered; pedicels 0.5–5 mm long; reproductive stems with 3 or fewer leaf-bearing nodes; plants flowering June through November . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) A. scabra 6b. Some or all lemmas with an awn (0.2–) 1.3–5 mm long, the awn produced from near or below the middle of the lemma and commonly exserted from the apex of the glumes [Fig. 181] 9a. Leaf blades involute, 0.2–2 mm wide, the basal ones persisting all year in a tuft; reproductive stems with 3 or fewer leaf-bearing nodes . . . . (in part) A. scabra 9b. Leaf blades flat, 1–3 mm wide, the basal ones usually withered prior to anthesis; reproductive stems with usually more than 3 leaf-bearing nodes 10a. Glumes 1.7–3 mm long; anthers 1–1.5 mm long; caryopses 0.8–1.2 mm long; stolons usually present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. canina 10b. Glumes 2.5–3.8 mm long; anthers 0.5–0.8 mm long; caryopses 1.4–2 mm long; stolons absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. mertensii 1. Agrostis canina L. E dog bentgrass. CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, rarely also found at elevation in boreal and subalpine lawns and clearings. 2. Agrostis capillaris L. E Rhode Island bentgrass. Agrostis tenuis Sibthorp • CT, MA, ME, NH, RI, VT; throughout. Fields, roadsides. 3. Agrostis elliottiana J.A. Schultes E Elliott’s bentgrass. MA, ME. Fields, roadsides. 4. Agrostis exarata Trin. E spike bentgrass. Agrostis exarata Trin. var. monolepis (Torrey) A.S. Hitchc. • VT. Forest clearings, edges of gravel roads. 5. Agrostis gigantea Roth E Fig. 180 redtop bentgrass. Agrostis alba, auct. non L.; A. nigra With.; A. stolonifera L. ssp. gigantea (Roth) Schuebl. & Martens; A. stolonifera L. var. major (Gaudin) Farw. • CT, MA, ME, NH, RI, VT; throughout. Fields, meadows, shorelines, roadsides. 6. Agrostis hyemalis (Walt.) B.S.P. N winter bentgrass. Cornucopiae hyemalis Walt. • CT, MA, ME, NH, RI; mainly on the coastal plain. Fields, roadsides, open woodlands. This uncommon species is over-reported, and many collections thought to be this grass are actually Agrostis scabra (including all of those from VT; collections at NEBC!, VT!).

Fig. 180  Inflorescence of Agrostis gigantea.

22 8  MO NOCOTS

7. Agrostis mertensii Trin. N C Fig. 181 northern bentgrass. Agrostis borealis Hartman; A. borealis Hartman ssp. americana (Hartman) Tzvelev; A. borealis Hartman var. americana (Scribn.) Fern. • ME, NH, VT; northern counties. High-elevation ridges and plateaus, boreal and alpine cliffs. 8. Agrostis perennans (Walt.) Tuckerman N autumn bentgrass. Agrostis altissima (Walt.) Tuckerman; A. perennans (Walt.) Tuckerman var. elata (Pursh) A.S. Hitchc.; Cornucopiae perennans Walt. • CT, MA, ME, NH, RI, VT; throughout. Forests, stream banks, shaded roadsides.

Fig. 181  Spikelet of Agrostis mertensii.

9. Agrostis scabra Willd. N rough bentgrass. Agrostis geminata Trin.; A. hyemalis (Walt.) B.S.P. var. scabra (Willd.) H.A. Blomquist; A. hyemalis (Walt.) B.S.P. var. tenuis (Tuckerman) Gleason; A. scabra Willd. var. geminata (Trin.) Swallen; A. scabra Willd. var. septentrionalis Fern. • CT, MA, ME, NH, RI, VT; throughout. Fields, forest clearings, open shorelines, roadsides, and open balds, tolerant of a wide range of hydrology. A form with shorter spikelets and anthers is sometimes collected from higher elevation peaks (var. tenuis). Another form (var. geminata), with shorter panicle branches and frequently with awned spikelets (the awn arising from the keel of the lemma), is also collected from ridges and summits. This latter form is sometimes confused with A. mertensii. Both are distinctive in their extremes but overlap extensively with typical A. scabra. 10. Agrostis stolonifera L. E Fig. 182 creeping bentgrass. Agrostis alba L. var. palustris (Huds.) Pers.; A. alba L. var. stolonifera (L.) Sm.; A. palustris Huds.; A. stolonifera L. var. compacta Hartman • CT, MA, ME, NH, RI, VT. Hydric or temporarily saturated soils of fields, inland and saline marshes, ditches, and shorelines. This species is often considered to be non-native to North American; however, some northern salt marsh and lakeside populations may be native (Harvey 2007).

Aira Fig. 182  Inflorescence of Agrostis stolonifera.

1a. Panicle narrow, dense, spike-like, with appressed branches, 0.3–0.7 cm wide [Fig. 183]; stems scabrous below the nodes; pedicels mostly shorter than the spikelets . . . . . A. praecox 1b. Panicle open, with ascending branches, 1.5–10 cm wide; stems smooth; pedicels mostly 1–2 times longer than the spikelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. caryophyllea 1. Aira caryophyllea L. var. caryophyllea E common silver-hairgrass. Aspris caryophyllea (L.) Nash • CT, MA, NH, VT. Open, often sandy and disturbed, soils, such as roadsides, fields, and woodland edges. 2. Aira praecox L. E Fig. 183 early silver-hairgrass. Aspris praecox (L.) Nash • CT, MA. Open, often sandy and disturbed, soils, such as roadsides, fields, and woodland edges.

Alopecurus

Fig. 183  Congested inflorescence of Aira praecox.

A genus sometimes confused with Phleum. In addition to unawned glumes and awned lemmas, Alopecurus has appressed-ascending spikelets. Phleum, on the other hand, has awned glumes, unawned lemmas, and spreading to reflexed spikelets (except for the apical ones, which are ascending). Reference: Crins (2007). 1a. Glumes 4–6 mm long, acute to acuminate and firm at the apex [Fig. 185]; anthers 2.2–3.5 mm long 2a. Glumes connate at the base for up to 25% of their length, ciliate on the keel with long hairs [Fig. 185], the hairs on the apical half of the keel 1–1.5 mm long . . . . . . . . A. pratensis 2b. Glumes connate at the base for 35–50% of their length, short-ciliate on the basal half of the keel, scabrous on the apical half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. myosuroides 1b. Glumes 2–3.2 mm long, blunt and scarious at the apex [Fig. 184]; anthers 0.5–2 mm long

Poac e a e   2 2 9

3a. Plants perennial, with decumbent-based stems that usually root at the lower nodes; glumes 2.6–3 mm long; anthers 1.3–2 mm long; inflorescence 4–6 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. geniculatus 3b. Plants annual or short-lived perennial, erect or decumbent at base, usually not rooting from the lower nodes; glumes 2–2.7 mm long; anthers 0.3–1 mm long; inflorescence 3–5 mm thick 4a. Awn inserted on the lemma just below the midpoint, straight, equaling the glumes or exserted from the spikelet up to 0.5 (–1) mm [Fig. 184] . . . . . . . . . . . . . . . . . . A. aequalis 4b. Awn inserted on the lemma halfway between the midpoint and base, geniculate, exserted from the spikelet 1.5–3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. carolinianus 1. Alopecurus aequalis Sobol. N Fig. 184 short-awned meadow-foxtail. Alopecurus geniculatus L. var. aristulatus (Michx.) Torr. • CT, MA, ME, NH, VT. Hydric meadows, ditches, shorelines, wet sand of borrow pits and other disturbed places. 2. Alopecurus carolinianus Walt. E Carolina meadow-foxtail. Alopecurus ramosus Poir. • MA; also reported from CT by Dowhan (1979), but specimens are unknown. Fields, roadsides, areas of cultivation.

Fig. 184  Spikelet of Alopecurus aequalis with glumes that are blunt at the apex.

3. Alopecurus geniculatus L. E marsh meadow-foxtail. CT, MA, ME, NH, RI, VT. Hydric soil or shallow water of meadows, ditches, marshes, and shorelines. 4. Alopecurus myosuroides Huds. E slender meadow-foxtail. Alopecurus agrestis L. • MA, ME. Fields, roadsides, and other open places with human-disturbed soil. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1997) lists no collections and the species is omitted from George (1999). 5. Alopecurus pratensis L. E Fig. 185 field meadow-foxtail. CT, MA, ME, NH, RI, VT. Mesic to hydric fields, roadsides, and ditches.

Ammophila 1. Ammophila breviligulata Fern.

nC

American beach grass.  1a. Ammophila champlainensis Seymour • CT, MA, ME, NH, RI, VT. Coastal dunes and beaches, sandy lake beaches. Ammophila champlainensis has been treated by many past authors as an endemic of Lake Champlain. However, there has been tremendous disagreement over how to treat inland vs. coastal populations of Ammophila breviligulata, because inland populations resembling A. champlainensis also occur around the Great Lakes, in the Saint Lawrence River Valley, and at Lac Saint-Jean. A recent study by Delisle-Oldham et al. (2008) looked at the A. breviligulata complex throughout eastern North America and recommended treating it as one polymorphic species (noting that Lake Champlain populations were the most distinctive group). Their study relied on seven important morphological characters, but phenology and DNA sequence data were not analyzed (note that both of these latter data sets show inland populations of A. champlainensis are distinct from A. breviligulata of the Atlantic coast). Despite the fact that their study shows overlap in all morphological characteristics when populations are examined range-wide, the taxa found in New England are known to be genetically different and flower at different times (as further evidenced by common garden experiments). Therefore, A. champlainensis is recognized here, albeit at a lower rank. 1a. Plants flowering in June through mid-July, restricted to beaches of Lake Champlain, VT; inflorescence (10–) 12–17.3 (–22) cm tall; glumes acute to short-acuminate at apex, the lower one (7–) 9–10.5 (–12) mm long . . . 1a. A. breviligulata ssp. champlainensis (Seymour) Walker, Paris, & Barrington ex Barkworth

Fig. 185  Spikelet of Alopecurus aequalis with glumes that are acuminate at the apex.

230 MO NOCOTS

1b. Plants flowering in late July through September, restricted to Atlantic coast beaches and dunes (except where introduced); inflorescence (14.3–) 21–36.5 cm tall; glumes acuminate at apex, the lower one (7.5–) 11–13 mm long . . . . . . . . . . . . 1b. A. breviligulata ssp. breviligulata Subspecies champlainensis is known from VT and is of regional conservation concern. It occurs on sand beaches of lower (i.e., northern) Lake Champlain. Subspecies breviligulata is known from CT, MA, ME, NH, RI, VT. It is native to coastal dunes and beaches but has been introduced as a sand stabilizer to some inland locations (e.g., VT).

Amphicarpum 1. Amphicarpum amphicarpon (Pursh) Nash

NC

Pursh’s blue maidencane. Amphicarpum purshii Kunth • MA; Nantucket Island. Coastal plain pond shores.

Andropogon Reference (Campbell 2003). 1a. Sessile spikelets 5–11 mm long; pedicellate spikelets usually well-developed and staminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. gerardii 1b. Sessile spikelets (2.6–) 3–5 mm long; pedicellate spikelets usually reduced or absent, often only the pedicel present 2a. Leaves with scabrous sheaths, the blades (13–) 30–109 cm long; ligules 0.6–2.2 mm long; mature peduncles of the rames (4–) 11–35 (–60) mm long . . . . . . . . . . . A. glomeratus 2b. Leaves with smooth sheaths, the blades 11–30 (–52) cm long; ligules 0.2–1 mm long; mature peduncles of the rames (2–) 3–4 (–10) mm long . . . . . . . . . . . . . . . . . . . . . A. virginicus 1. Andropogon gerardii Vitman N big bluestem. Andropogon gerardii Vitman var. chrysocomus (Nash) Fern. • CT, MA, ME, NH, RI, VT. Sandy soils of roadsides, grasslands, and shorelines, also on river shore outcrops. 2. Andropogon glomeratus (Walt.) Britt. var. glomeratus N Fig. 186 bushy bluestem. Andropogon virginicus L. var. corymbosus (Chapman ex Hack.) Fern. & Grisc.; Cinna glomerata Walt. • MA, RI. Pond shores, ditches, seasonally wet depressions, wetland edges. Fig. 186  Inflorescence of Andropogon glomeratus.

3. Andropogon virginicus L. var. virginicus N broomsedge bluestem. CT, MA, RI. Sandy fields, woodland openings, roadsides.

Anthoxanthum See Schouten and Veldkamp (1985) for rationale of an expanded Anthoxanthum (i.e., one that includes Hierochloe). Reference: Allred and Barkworth (2007). 1a. Inflorescence an open to somewhat compact panicle, the lower branches ascending to wide-spreading or even drooping; glumes subequal in length; each spikelet with 1 bisexual floret and 2 unisexual florets that are staminate 2a. Staminate florets with awned lemmas, the awns 0.6–10.5 mm long; glumes 5–8 mm long; panicle moderately compact, 1–8.5 cm tall, with (3–) 10–20 (–35) spikelets; plants subcespitose on short rhizomes up to 2 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . A. monticola 2b. Staminate florets unawned or with short awn-tips to 0.6 (–1) mm long; glumes (2.5–) 3.5–6 mm long; panicle open (less so in subalpine plants), 4–9 (–12.5) cm tall, with 8–100 spikelets; plants with stems produced singly or a few-together on elongate rhizomes

P oac e a e   23 1

3a. Pubescence of lemma of bisexual floret appressed to appressed-ascending, 0.1–0.5 mm long (sometimes longer hairs can be found near the keel), tending to be concentrated toward the keel (i.e., often noticeably sparser near the margins) [Fig. 188] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. nitens 3b. Pubescence of lemma of bisexual floret loosely ascending, 0.5–1 mm long, ± uniformly distributed around the apex of the lemma [Fig. 187] . . . . . . . . . . . . . . . A. hirtum 1b. Inflorescence a densely congested panicle, the lower branches appressed to appressedascending (briefly somewhat open at anthesis); glumes distinctly unequal in length; each spikelet with 1 bisexual floret and 2 sterile florets 4a. Plants perennial, with erect stems; leaves 2–6 mm wide; awn of the lower sterile lemma straight; spikelets 7–9 mm long; glumes scabrous to hirtellous . . . . . . . . . . . . . . A. odoratum 4b. Plants annual, with geniculate, branched stems; leaves mostly narrower than 2 mm; awn of both sterile lemmas geniculate; spikelets 5–7 mm long; glumes glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ovatum 1. Anthoxanthum hirtum (Schrank) Y. Schouten & Veldkamp E Fig. 187 northern sweet grass. Hierochloe hirta (Schrank) Borbás; Savastana hirta Schrank • MA, ME, NH, VT. Roadsides, meadows, shorelines. 2. Anthoxanthum monticola (Bigelow) Y. Schouten & Veldkamp ssp. monticola

Fig. 187  Lemma of bisexual floret of Anthoxanthum hirtum.

NC

alpine sweet grass. Hierochloe alpina (Sw. ex Willd.) Roemer & J.A. Schultes; H. alpina (Sw. ex Willd.) Roemer & J.A. Schultes ssp. orthantha (Sørensen) G. Weim; H. monticola (Bigelow) A. & D. Löve; H. orthantha Sørensen • ME, NH, VT; northern counties. Alpine ridges and plateaus, growing in open turfs and heaths. 3. Anthoxanthum nitens (Weber) Y. Schouten & Veldkamp ssp. nitens N Fig. 188 vanilla sweet grass. Hierochloe odorata (L.) Beauv.; H. odorata (L.) Beauv. var. fragrans (Willd.) Richter; Holcus odoratus L.; Savastana odorata (L.) Scribn. • CT, MA, ME, NH, RI, VT. Brackish and saline marshes, swamps, river shores, rarely ascending to subalpine habitats. 4. Anthoxanthum odoratum L. E large sweet grass. CT, MA, ME, NH, RI, VT; throughout. Fields, roadsides, lawns, forest edges and openings. 5. Anthoxanthum ovatum Lag. var. aristatum Pérez-Lara E small sweet grass. Anthoxanthum aristatum Boiss.; A. odoratum L. var. puelii (Lecoq & Lamotte) Coss. & Durieu; A. puelii Lecoq & Lamotte • MA, ME, NH. Fields, roadsides, disturbed ground. See Pereira et al. (2007) for rationale of including Anthoxanthum aristatum under A. ovatum.

Apera 1a. Leaves 3–5 mm wide; branches of the panicle ascending to spreading at anthesis, bearing flowers in the distal half of the branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. spica‑venti 1b. Leaves 1–3 mm wide; branches of the panicle contracted, erect at anthesis, bearing flowers in both distal and proximal halves of the branches . . . . . . . . . . . . . . . . . . . . . . . . . . . A. interrupta 1. Apera interrupta (L.) Beauv. E dense silky bentgrass. Agrostis interrupta L. • MA, ME. Lawns, roadsides, fields. 2. Apera spica-venti (L.) Beauv. E loose silky bentgrass. Agrostis spica-venti L. • CT, MA, ME, VT. Lawns, roadsides, fields, disturbed sandy places.

Fig. 188  Lemma of bisexual floret of Anthoxanthum nitens.

232  MONOCOTS

Aristida When measuring the length of the lemma, be sure to include the basal callus in the measurement. Reference: Allred (2003). 1a. Lemma awns twisted together at the base forming a spiral column 8–15 mm tall [Fig. 191]; callus of lemma 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. tuberculosa 1b. Lemma awns not twisted together into a column (though they may be coiled at the base independent of one another) [Figs. 189, 190]; callus of lemma 0.3–2 mm long 2a. Lower glume with 3–7 veins; awns subequal, (12–) 35–65 (–70) mm long . . . . A. oligantha 2b. Lower glume with 1 or 2 veins; awns subequal or distinctly unequal, 1–27 (–30) mm long 3a. Central awn at maturity spirally coiled at base with ½ to 3 full turns [Fig. 189] 4a. Central awn 10–15 mm long; lateral awns 5–10 mm long, usually somewhat divergent from near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. basiramea 4b. Central awn 3–8 mm long; lateral awns 1–4 mm long, straight and erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. dichotoma 3b. Central awn not coiled at base [Fig. 190] 5a. Plants annual; lateral awns of lemma usually markedly reduced in length compared with the central awn, up to 66% as long as the central awn . . . . . . . . . . . . A. longespica 5b. Plants perennial; lateral awns of lemma equal or subequal in length to the central awn [Fig. 190] 6a. Lower glume 4–7 mm long, usually less than 75% as long as the upper glume; lemmas 7–13 mm long, with a callus 0.5–1.8 mm long . . . . . . . . . . . . . . . A. purpurea 6b. Lower glume 5–10 mm long, more than 75% as long as the upper glume (and sometimes even exceeding the length of the upper glume); lemmas 4–8 mm long, with a callus 0.4–0.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. purpurascens 1. Aristida basiramea Engelm. ex Vasey n Fig. 189 fork-tipped threeawn. MA, ME, NH, VT. Sandy, often sterile, soils of fields, grasslands, roadsides, and disturbed areas. Though this species is native to New England, it is considered non-native in MA.

Fig. 189  Spikelet of Aristida basiramea.

2. Aristida dichotoma Michx. var. dichotoma N churchmouse threeawn. CT, MA, ME, NH, RI, VT. Sandy, often sterile, soils of fields, grasslands, roadsides, and disturbed areas. 3. Aristida longespica Poir. n red threeawn.  3a. Aristida geniculata Raf.; Aristida intermedia Scribn. & Ball; 3b. Aristida gracilis Ell. • CT, MA, NH, RI, VT. Sandy fields, roadsides, woodland openings, disturbed sandy soils. 1a. Central awn (8–) 12–27 mm long; lateral awns (1–) 6–18 mm long; glumes 4–11 mm long; flowers with 1 or 3 anthers . . . . . . . . . . . . . . . . . . 3a. A. longespica var. geniculata (Raf.) Fern. 1b. Central awn 1–10 (–14) mm long; lateral awns 0–5 (–8) mm long; glumes 2–8 mm long; flowers with 1 anther . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. A. longespica var. longespica Variety geniculata is native and known from CT, MA, NH, RI, VT. Variety longespica is non-native and known from CT, MA. 4. Aristida oligantha Michx. E oldfield threeawn. CT, MA, ME, NH, RI, VT; absent from most of ME. Sandy, often sterile, soils of fields, grasslands, roadsides, and disturbed areas. Though considered native by some authors, this grass is likely non-native in New England.

Fig. 190  Spikelet of Aristida purpurascens.

5. Aristida purpurascens Poir. var. purpurascens N C Fig. 190 arrow-feather threeawn. Aristida purpurascens Poir. var. minor Vasey • CT, MA, RI. Sandy fields, roadsides, woodland openings, grasslands.

Poac e a e   23 3

6. Aristida purpurea Nutt. var. nealleyi (Vasey) Allred E purple threeawn. Aristida glauca (Nees) Walp.; A. nealleyi (Vasey) Vasey • VT. Roadsides, disturbed, sandy soil. 7. Aristida tuberculosa Nutt. N Fig. 191 seaside threeawn. CT, MA, NH. Coastal dunes, dry, sterile, sandy soil near the coast.

Arrhenatherum 1. Arrhenatherum elatius (L.) J. & K. Presl E tall oat grass. 1a. Arrhenatherum elatius (L.) J. & K. Presl var. bulbosum (Willd.) Spenner; A. elatius (L.) J. & K. Presl var. tuberosum Thiel.; 1b. Arrhenatherum elatius (L.) J. & K. Presl var. biaristatum (Peterm.) Peterm.; Avena elatior L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest edges, human-disturbed soil. 1a. Lower internodes of reproductive stems shortened, closely set, and thickened into cormlike sections 5–10 mm in diameter 1a. A. elatius ssp. bulbosum (Willd.) Schübler & Martens 1b. Lower internodes of reproductive stems not shortened or thickened, 1.8–3 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. A. elatius ssp. elatius Subspecies elatius is known from CT, MA, ME, NH, RI, VT. Supbspecies bulbosum has been found only in MA.

Arthraxon 1. Arthraxon hispidus (Thunb.) Makino E small carp grass. Phalaris hispida Thunb. • MA. Hydric soil of roadsides, ditches, and riparian forests.

Avena 1a. Spikelets with predominantly 3 florets; lemmas pubescent with appressed, brown hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. fatua 1b. Spikelets with predominantly 2 florets; lemmas glabrous or scabrous, without brown hairs 2a. Lemmas scabrous near apex, both of each spikelet with an awn; inflorescence secund . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. strigosa 2b. Lemmas glabrous, only the lower of each spikelet with an awn; inflorescence not secund . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sativa 1. Avena fatua L. E wild oat. Avena fatua L. var. glabrata Peterm. • CT, MA, ME, NH, VT. Fields, roadsides, disturbed soil. 2. Avena sativa L. E oat. Avena fatua L. var. sativa (L.) Hausskn. • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil. 3. Avena strigosa Schreb. E lopsided oat. MA. Fields, roadsides, disturbed soil.

Avenula 1. Avenula pubescens (Huds.) Dumort. E downy alpine oat grass. Avena pubescens Huds.; Helictotrichon pubescens (Huds.) Pilger • CT, MA, VT. Fields, roadsides, disturbed ground.

Fig. 191  Spikelet of Aristida tuberculosa.

234  MONOCOTS

Beckmannia 1. Beckmannia syzigachne (Steud.) Fern. E American slough grass. Beckmannia erucaeformis (L.) Host ssp. syzigachne (Steud.) Breitung; Panicum syzigachne Steud. • ME. Wet ditches, low meadows, shorelines, marshes, edges of swamps.

Bouteloua Reference: Wipff (2003a). 1a. Branches of panicle each with 20–130 spikelets [Fig. 193], persistent, the branch axis terminated by progressively reduced spikelets, not prolonged beyond spikelets; disarticulation above the glumes 2a. Plants annual, without rhizomes or stolons; glumes glabrous, sometimes scabrous distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. simplex 2b. Plants perennial, often with rhizomes or stolons; upper glumes often with pustulosebased hairs on the midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. gracilis 1b. Branches of panicle each with (1–) 2–8 (–15) spikelets, deciduous, the branch axis not terminated by reduced florets, the axis prolonged beyond the base of the uppermost spikelet as a bristle 3–6 mm long; disarticulation occurring at base of branch 3a. Inflorescence with (12–) 30–80 branches [Fig. 192]; central awn of lowest lemma not flanked by lateral lobes (i.e., the lemma with 3 total lobes) . . . . . . . . . . . . . . . B. curtipendula 3b. Inflorescence with (3–) 6–12 branches; central awn of lowest lemma arising from between two membranous lobes 0.5–1.5 mm long (i.e., the lemma with 5 total lobes; the lateral lobes sometimes shorter in B. repens ) 4a. Apex of panicle branch axis prominently bifid or trifid; upper glume bilobed, the awn emerging from the sinus; lower glumes 3–4 mm long . . . . . . . . . . . . . . . . . . . . B. rigidiseta 4b. Apex of panicle branch axis entire; upper glume acute at apex, unawned or with a short awn-tip; lower glumes 4–7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. repens Fig. 192  Portion of inflorescence of Bouteloua curtipendula showing branches.

1. Bouteloua curtipendula (Michx.) Torr. var. curtipendula

n C Fig. 192

sideoats grama. Atheropogon curtipendulus (Michx.) Fourn.; Chloris curtipendula Michx. • CT, ME; western portion of CT. Native habitats include dry-mesic to xeric open woodlands and balds, sandy fields, river banks in areas of high-pH bedrock; introduced to ME on wool waste and seeded areas. 2. Bouteloua gracilis (Willd. ex Kunth) Lag. ex Griffiths E Fig. 193 blue grama. Chondrosum gracile Willd. ex Kunth • MA, ME, VT; also reported from CT by Kartesz (1999), but specimens are unknown. Wool waste, disturbed ground. Reports of Bouteloua hirsuta Lag. in New England are based on collections of this species— Parlin 1521 (GH!, NEBC!). 3. Bouteloua repens (Kunth) Scribn. & Merr. E slender grama. Bouteloua filiformis (Fourn.) Griffiths; Dinebra repens Kunth • MA, ME. Waste areas, disturbed ground.

Fig. 193  Branch of Bouteloua gracilis showing spikelet distribution.

4. Bouteloua rigidiseta (Steud.) A.S. Hitchc. E Texas grama. MA. Waste areas, disturbed ground. 5. Bouteloua simplex Lag. E mat grama. Chondrosum prostratum (Lag.) Sweet • ME. Wool waste, disturbed ground.

Poac e a e   23 5

Brachyelytrum Reference: Saarela et al. (2003). 1a. Lemmas hispid with hairs (0.2–) 0.4–0.8 (–0.9) mm long [Fig. 195], the body (0.8–) 1.1–1.5 (–1.8) mm wide, with conspicuous veins, the midvein much more prominent than the lateral veins; anthers 3.3–6 mm long; ligules 2–3.5 mm long . . . . . . . . . . . . . B. erectum 1b. Lemmas glabrous or minutely scabrous with hairs 0.08–0.14 (–0.2) mm long [Fig. 194], the body (0.7–) 0.8–1.2 (–1.4) mm wide, with weak veins of equal prominence; anthers 2–3.5 mm long; ligules 1.8–2.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. aristosum 1. Brachyelytrum aristosum (Michx.) Trel. in Branner & Coville N Fig. 194 northern long-awned wood grass. Brachyelytrum erectum (Schreb.) Beauv. var. septentrionale Babel; B. septentrionale (Babel) G. Tucker; Dilepyrum aristosum Michx. • CT, MA, ME, NH, RI, VT. Mesic forests and forest openings. 2. Brachyelytrum erectum (Schreb.) Beauv. N Fig. 195 southern long-awned wood grass. Brachyelytrum aristosum (Michx.) Trel. in Branner & Coville var. glabratum Vasey; B. erectum (Schreb.) Beauv. var. glabratum (Vasey) T. Koyama & Kawano; Dilepyrum erectum (Schreb.) Farw.; Muhlenbergia erecta Schreb.; M. brachyelytrum Trin. • CT, MA, NH, RI, VT. Mesic forests and forest openings, often in areas of high-pH bedrock in the northern portion of its range.

Fig. 194  Spikelet of Brachyelytrum aristosum.

Brachypodium 1. Brachypodium pinnatum (L.) Beauv. E heath false brome. Bromus pinnatus L. • MA. Disturbed ground, waste areas.

Briza Reports of Briza maxima L. from MA and VT were based on vouchers taken from cultivated specimens. 1a. Ligules 0.5–1 mm long, truncate at apex; leaves mostly confined to the basal portion of the plant; anthers 1.3–2 mm long; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. media 1b. Ligules 4–13 mm long (rarely longer), pointed at apex (infrequently truncate); leaves ± evenly distributed on the stem; anthers 0.4–0.5 mm long; plants annual . . . . . . . . . . B. minor 1. Briza media L. E perennial quaking grass. CT, MA, ME, NH, VT. Roadsides, fields, disturbed soil. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species had questionable naturalization and reported no occurrences (i.e., the author is unaware of any collections). 2. Briza minor L. E little quaking grass. CT. Roadsides, fields, disturbed soil.

Bromus Reference: Pavlick (1995). 1a. Lobes of the lemma apex 0.1–0.5 mm long [Fig. 196]; plants perennial 2a. Lower glume 3- or 5-nerved; upper glume 5- to 9-nerved 3a. Lemmas compressed and strongly keeled; ligules 2–3.5 mm long; first and second glumes 7–9 and 9–11 mm long, respectively . . . . . . . . . . . . . . . . . . . . . . . . . . B. marginatus

Fig. 195  Spikelet of Brachyelytrum erectum.

236  MO NOCOTS

3b. Lemmas convex, neither strongly compressed nor keeled; ligules 0.5–1 mm long; first and second glumes 5–7.5 and 6.5–8.5 mm long, respectively . . . . . . . . . . . . . . . . . B. kalmii 2b. Lower glume 1-nerved; upper glume 3-nerved 4a. Plants with long rhizomes; lemmas awnless or with awns up to 3 mm long [Fig. 198] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. inermis 4b. Plants without rhizomes; lemmas always awned, the awns 3–7 (–8) mm long 5a. Panicle contracted, usually narrower than 2 cm, with erect to ascending branches; anthers 4–6.5 mm long; leaf blades often involute . . . . . . . . . . . . . . . . . . . . . . B. erectus 5b. Panicle open, usually wider than 2 cm, with spreading to drooping branches (rarely ascending); anthers 1–4 (–5) mm long; leaf blades flat 6a. Lemmas conspicuously pubescent along the margins [Fig. 196], glabrous or scabrous over the abaxial surface; anthers 1–2 mm long . . . . . . . . . . . . . . . B. ciliatus 6b. Lemmas sparsely to densely pubescent over the abaxial surface; anthers 2–4 (–5) mm long 7a. Leaf sheaths overlapping and concealing the nodes, the densely pubescent apex prominently outward-flanged and auriculate [Fig. 199]; leaves 9–20 per stem; anthers 2–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. latiglumis 7b. Leaf sheaths not overlapping, at least the upper nodes exposed, the glabrous to pubescent apex neither outwardly flanged nor auriculate; leaves 4–6 per stem; anthers (2–) 2.5–4 (–5) mm long . . . . . . . . . . . . . . . . . B. pubescens 1b. Lobes of the lemma apex 0.6–5 mm long [Figs. 200, 201]; plants annual or biennial 8a. First glume 1 (–3)-veined; second glume 3 (–5)-veined; lemmas narrow from a side view, 1–1.5 mm deep; spikelets oblong to obtriangular in outline, usually wider toward the apex; lemma awns (8–) 10–50 mm long, often longer than the associated lemma 9a. Lemmas 22–28 mm long, with awns 30–50 mm long emerging between 2 apical lobes 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. rigidus 9b. Lemmas 9–20 mm long, with awns 10–30 mm long emerging between 2 apical lobes 0.8–3 mm long [Fig. 201] 10a. Panicle with crowded, ascending to erect branches, 2–10 cm tall; pedicels usually shorter that associated spikelets; ligules pubescent (as well as lacerate at the apex) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. rubens 10b. Panicle with loose, nodding to ascending branches, 5–20 cm tall; pedicels usually longer than associated spikelets excluding the awns; ligules glabrous (though lacerate at the apex) 11a. Lemmas 9–12 mm long; anthers 0.5–1 mm long; awns 10–18 mm long; lower panicle branches bearing 4–8 spikelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. tectorum 11b. Lemmas 14–20 mm long; anthers 1–1.4 mm long; awns 15–30 mm long; panicle branches bearing 1–3 spikelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sterilis 8b. First glume 3- or 5-veined; second glume 5- or 7-veined; lemmas wider from side view, 1.2–2.2 mm deep; spikelets ovate to lanceolate in outline, usually widest below the middle; lemma awns absent or up to 13 mm long, usually as long as or shorter than the associated lemma 12a. Lemmas awnless or with awns up to 0.8 mm long; spikelets ovate in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. briziformis 12b. Lemmas with awns 2–13 mm long; spikelets narrow-ovate to lanceolate in outline

P oac e a e   237

13a. Lemmas 4.5–6.5 mm long, the margins with a sharp bend where the lemma begins to taper toward the apex from its widest point [Fig. 200]; caryopsis longer than the palea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. lepidus 13b. Lemmas 6.5–11.5 mm long, the margins rounded or with a bluntly angled bend where the lemma begins to taper toward the apex from its widest point; caryopsis as long as or shorter than the palea 14a. Lemmas with prominently raised veins and concave regions between the veins; anthers 0.6–1 (–1.5) mm long; pedicels mostly shorter than their spikelets [Fig. 197]; inflorescence compact, erect, 3–10 cm tall . . . . . . . . . . . . . B. hordeaceus 14b. Lemmas with obscure to distinct, but not raised, veins; anthers 1–5 mm long; pedicels as long as or longer than their associated spikelets excluding the awns (usually shorter in B. squarrosus); inflorescence open, ascending to drooping (narrow and erect in B. racemosus), 4–20 cm tall 15a. Margin of the lemma involute at maturity, tightly enclosing the caryopsis and exposing the rachilla 16a. Ligules 2–3 mm long, glabrous; anthers 1–2 mm long; awns straight or flexuous, 3–6 (–9.5) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. secalinus 16b. Ligules 1–1.5 mm long, pubescent; anthers 3–5 mm long; awns straight, 6–11 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) B. arvensis 15b. Margin of the lemma not tightly involute, overlapping the caryopsis and concealing the rachilla 17a. Anthers 3–5 mm long; panicle 15–30 cm tall . . . . . . . (in part) B. arvensis 17b. Anthers 1–3 mm long; panicle 4–22 cm tall 18a. Branches of the panicle erect, straight or nearly so, usually 2 per node; lemmas 6.5–8 mm long; anthers 1.5–3 mm long . . . . . B. racemosus 18b. Branches of the panicle ascending to, more commonly, spreading or even drooping, flexuous or arching, 2–6 per node; lemmas 7–11.5 mm long; anthers 1–2 mm long 19a. Awns terete or narrow and flattened at the base, not or only slightly divergent; apical lobes of lemma up to 1.5 mm long; palea up to 1.5 mm shorter than its associated lemma; branches of the panicle rather stiffly ascending to spreading . . . . . . . . . . . . . . . . B. commutatus 19b. Awns broad and flattened at the base, divergent at maturity; apical lobes of lemma 1.5–3 mm long; palea 1–2.5 mm shorter than its associated lemma; branches of the panicle lax, often arched or drooping 20a. Panicle not secund, with compound lower branches; spikelets with 6–12 florets; lemmas 7–9 mm long, 1.2–2.2 mm deep in side view, the margins with a long, obscure bend where the lemma begins to taper toward the apex from its widest point . . . . . . . . B. japonicus 20b. Panicle ± secund, often with simple branches; spikelets with 8–30 florets; lemmas 8–11 mm long, 2.5–3.5 mm deep in side view, the margins with an obtusely angled bend where the lemma begins to taper toward the apex from its widest point . . . . . . B. squarrosus 1. Bromus arvensis L. E field brome. CT, MA, ME, NH, RI. Fields, roadsides, disturbed soil.

238  MONOCOTS

2. Bromus briziformis Fisch. & C.A. Mey. in Fisch. et al. E rattlesnake brome. CT, MA, VT. Fields, roadsides, disturbed soil, dumps. 3. Bromus ciliatus L. N Fig. 196 fringed brome. Bromopsis ciliata (L.) Holub; Bromus canadensis Michx.; B. ciliatus L. var. intonsus Fern.; B. dudleyi Fern. • CT, MA, ME, NH, RI, VT. Mesic to wet-mesic forests, forest edges and openings, stream banks. 4. Bromus commutatus Schrad. E meadow brome. CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil. 5. Bromus erectus Huds. E Fig. 196  Lemma of Bromus ciliatus.

upright brome. Bromopsis erecta (Huds.) Fourr. • CT, MA, ME, VT. Fields, roadsides, disturbed soil. 6. Bromus hordeaceus L. E Fig. 197 soft brome. 6a. Bromus thominei Hardouin; 6b. Bromus mollis L.; 6c. Bromus mollis L. var. leiostachys Hartman • CT, MA, ME, NH, RI. Fields, roadsides, disturbed soil. 1a. Inflorescence often reduced to 1 or 2 spikelets; reproductive stems slender, 2–16 cm tall; awns sometimes divaricate in fruit . . . . . . . . . . . . . . . . . . . 6a. B. hordeaceus ssp. thominei (Hardouin) Maire in Emberger & Maire 1b. Inflorescence usually not reduced to 1 or 2 spikelets (except in depauperate forms of B. hordeaceus ssp. hordeaceus) [Fig. 197]; reproductive stems medium to stout, (3–) 10–70 cm tall; awns straight and erect 2a. Lemmas 8–11 mm long, usually pubescent, the margins with an obscurely angled bend where the lemma begins to taper toward the apex from its widest point; caryopsis shorter than the palea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6b. B. hordeaceus ssp. hordeaceus

Fig. 197  Inflorescence of Bromus hordeaceus ssp. hordeaceus.

2b. Lemmas 6.5–8 mm long, usually glabrous, the margins with an abrupt bend where the lemma begins to taper toward the apex from its widest point; caryopsis usually as long as the palea . . . . . . . . . . . . . . . . . . . 6c. B. hordeaceus ssp. pseudothominei (P. Sm.) H. Scholz Subspecies thominei is known from MA, RI. Its lemmas are 6.5–7.5 mm long and vary from glabrous to pubescent. Subspecies hordeaceus is known from CT, MA, ME, NH, RI. Subspecies pseudothominei is known from CT, MA. 7. Bromus inermis Leyss. ssp. inermis E Fig. 198 smooth brome. Bromopsis inermis (Leyss.) Holub • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil. Collections of this species with shortly awned lemmas are responsible for the reports of Bromus pumpellianus Scribn. in New England. These collections lack the spikelet and leaf pubescence of the latter species (i.e., they are merely part of the variation displayed by B. inermis). 8. Bromus japonicus Thunb. ex Murr. E Japanese brome. CT, MA, ME, NH, VT. Fields, roadsides, disturbed soil. 9. Bromus kalmii Gray N

Fig. 198  Inflorescence of Bromus inermis.

Kalm’s brome. Bromopsis kalmii (Gray) Holub; Bromus purgans L. • CT, MA, ME, NH, VT. Commonly in dry-mesic soils of outcrops, open forests, and woodlands, less frequently in wet-mesic meadows and riparian forests, usually in soils influenced by high-pH bedrock. 10. Bromus latiglumis (Shear) A.S. Hitchc. N Fig. 199 flanged brome. Bromus altissimus Pursh; B. purgans, auct. non L. • CT, MA, ME, NH, VT. River banks, upper edge of gravel and cobble river shores, riparian forests. 11. Bromus lepidus Holmb. E Fig. 200 slender soft brome. MA; also reported from CT by Dowhan (1979), but specimens are unknown. Fields, roadsides, disturbed soil.

Poac e a e   23 9

12. Bromus marginatus Nees ex Steud. E large mountain brome. Bromus breviaristatus Buckl.; B. carinatus Hook. var. linearis Shear; B. carinatus Hook. var. marginatus (Nees ex Steud.) Barkworth & Anderton; Ceratochloa marginata (Nees ex Steud.) W.A. Weber • CT, MA, ME, NH. Fields, roadsides, disturbed soil. 13. Bromus pubescens Muhl. ex Willd. N hairy wood brome. Bromus purgans, auct. non L. • CT, MA, ME, NH, RI, VT. Forests and woodlands, often associated with rocky slopes and outcrops, usually in soils influenced by circumneutral or basic bedrock. 14. Bromus racemosus L. E bald brome. RI, VT; also reported from CT by Dowhan (1979), for MA by Pavlick (1995), and from ME by Campbell et al. (1995), but specimens are unknown. Fields, roadsides, disturbed soil.

Fig. 199  Summit of leaf sheath of Bromus latiglumis.

15. Bromus rigidus Roth E ripgut brome. Anisantha rigida (Roth) Hyl.; Bromus diandrus Roth ssp. rigidus (Roth) Lainz • MA. Fields, roadsides, waste areas around seaports. This species is closely related to, and

sometimes included in, Bromus diandrus. 16. Bromus rubens L. E foxtail brome. Anisantha rubens (L.) Nevski • MA. Fields, roadsides, disturbed soil. 17. Bromus secalinus L. E rye brome. Bromus secalinus L. var. velutinus (Schrad.) Koch • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil. Specimens with pubescent spikelets occur (the “velutinus” form). 18. Bromus squarrosus L. E corn brome. CT, VT. Fields, roadsides, disturbed soil. 19. Bromus sterilis L. E poverty brome. Anisantha sterilis (L.) Nevski • CT, MA, RI. Fields, roadsides, disturbed soil.

Fig. 200  Lemma of Bromus lepidus showing curvature of margin.

20. Bromus tectorum L. E Fig. 201 cheat brome. Anisantha tectorum (L.) Nevski • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil, cracks in pavement and sidewalks.

Calamagrostis The genera Agrostis, Calamagrostis, and Deyeuxia are closely related and are defined by a combination of characters that are not unique to any genus (i.e., there exist several problematic, intermediate taxa). Based on the criteria used by Phillips and Chen (2003), all our species of Calamagrostis, except C. epigejos, would be transferred to the genus Deyeuxia. However, a phylogeny based on DNA sequence data did not support the monophyly of Deyeuxia. Reference: Marr et al. (2007). 1a. Pubescence of the callus (130–) 150–200 (–250)% as long as the lemma; rachilla not prolonged; glumes exceeding the lemmas by a distance of (1.5–) 2–3 (–4.5) mm; lemmas 2-nerved apical to the insertion of the awn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. epigejos 1b. Pubescence of the callus 20–120 (–150)% as long as the lemma [Figs. 202, 203]; rachilla prolonged behind the palea as a slender, pubescent bristle; glumes exceeding the lemmas by a distance of (0–) 0.1–2.1 mm; lemmas 4-nerved apical to the insertion of the awn 2a. Awn attached well above the middle of the lemma; rachilla pubescent only at apex; caryopsis pubescent at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cinnoides 2b. Awn attached at or below the middle of the lemma [Fig. 203]; rachilla pubescent throughout; caryopsis glabrous

Fig. 201  Lemma of Bromus tectorum.

240 MON OCOTS

3a. Callus hairs scant, up to 1 mm long, 20–30% as long as the lemma [Fig. 203]; awns straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pickeringii 3b. Callus hairs moderate to copious, (1.5–) 1.7–3.5 (–4.5) mm long, 50–120 (–150)% as long as the lemma [Fig. 202]; awns straight or bent 4a. Panicle contracted and dense, (7–) 10–20 (–28) mm wide, the longer branches 1.4–5 (–9.5) cm long; pubescence of callus scant to moderate, concentrated in unequal lateral tufts, 20–75 (–100)% as long as the lemma; lemma awn minutely antrorsely scabrous throughout its length, delicate to stout, usually distinguishable from the callus hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stricta 4b. Panicle loose and somewhat open (less open in C. canadensis var. macouniana), (10–) 20–40 (–80) mm wide, the longer branches 2.7–6 (–12) cm long; pubescence of callus moderate to dense, uniformly distributed, (70–) 90–120 (–150)% as long as the lemma; lemma awn smooth, as least in the basal half, delicate, often difficult to distinguish from the callus hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. canadensis 1. Calamagrostis canadensis (Michx.) Beauv.

Fig. 202  Lemma of Calamagrostis canadensis var. canadensis.

N C Fig. 202

Canada reed grass. 1a. Arundo langsdorfii Link; Calamagrostis canadensis (Michx.) Beauv. var. scabra (J. Presl) A.S. Hitchc.; C. langsdorfii (Link) Trin.; C. nubila Louis-Marie; Deyeuxia langsdorfii (Link) Kunth; D. preslii Kunth; 1b. Arundo canadensis Michx.; Calamagrostis canadensis (Michx.) Beauv. var. robusta (Vasey) Stebbins; Deyeuxia canadensis (Michx.)  Munro ex Hook. f.; 1c. Calamagrostis macouniana (Vasey) Vasey; Deyeuxia macouniana Vasey • CT, MA, ME, NH, RI, VT; throughout. Wet-mesic to hydric soils of marshes, shorelines, open swamps, and low fields, also in mesic soils of open, boreal to alpine slopes and plateaus, and rarely in mesic to dry-mesic soils. 1a. Glumes 4.5–6 mm long, pilose-hirtellous over the abaxial surface, thick and oqaque . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. C. canadensis var. langsdorfii (Link) Inman 1b. Glumes 2.2–4.5 mm long, glabrous or minutely scabrous over the abaxial surface, translucent at least along the margins and at the apex 2a. Glumes 2.8–4.5 mm long, acute to acuminate at the apex, distinctly longer than the lemma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. C. canadensis var. canadensis 2b. Glumes 2.2–2.8 mm long, obtuse to acute at the apex, scarcely or not exceeding the lemma . . . . . . . . . . . . . . . . . . . . . . . 1c. C. canadensis var. macouniana (Vasey) Stebbins Variety langsdorfii is known from NH, VT; also reported from ME by Knowlton (1899), but specimens are unknown. It is restricted to high elevation areas and is of regional conservation concern. Variety canadensis is known from low to high elevation areas of CT, MA, ME, NH, RI, VT. Variety macouniana is known from CT, MA, ME, NH, VT and is also of regional conservation concern. 2. Calamagrostis cinnoides (Muhl.) W. Bart. N Nuttall’s reed grass. Arundo cinnoides Muhl.; Calamagrostis coarcta (Torr.) Torr. ex Eat.; Deyeuxia nuttalliana (Steud.) Vasey • CT, MA, ME, NH, RI. Swamps, peatlands, sandy or peaty hydric openings and rights-of-way. 3. Calamagrostis epigejos (L.) Roth var. georgica (K. Koch) Ledeb. E feathertop reed grass. MA, RI. Fields, roadsides, disturbed soil. 4. Calamagrostis pickeringii Gray

N C Fig. 203

Pickering’s reed grass. Calamagrostis pickeringii Gray var. debilis (Kearney) Fern. & Wieg.; Deyeuxia pickeringii (Gray) Vasey • MA, ME, NH, VT. Bogs, gravel river shores, open alpine and subalpine areas, shores of small, usually mid- to high elevation, ponds, peaty meadows. Fig. 203  Lemma of Calamagrostis pickeringii.

5. Calamagrostis stricta (Timm) Koel.

NC

neglected reed grass.  5a. Calamagrostis fernaldii Louis-Marie; C. inexpansa Gray; C. inexpansa Gray var. novae-angliae Stebbins; C. lacustris (Kearney) Nash; C. stricta (Timm) Koel. var. brevior Vasey; Deyeuxia neglecta (Ehrh.) Kunth var. americana Vasey in Macoun; 

Poac e a e   2 41

5b. Arundo neglecta Ehrh.; Calamagrostis neglecta (Ehrh.) P.G. Gaertn. et al.; C. neglecta P.G. Gaertn. et al. ssp. stricta (Timm) Tzvelev; Deyeuxia neglecta (Ehrh.) Kunth • CT, ME, NH, VT; northern portion of states. Mainly in boreal to alpine settings (CT and MA occurrences excepted), such as streambeds, talus slopes, ridges, rock outcrops, ledges, ice-scoured river shores, lake shores, woodlands, and high-elevation cliffs and plateaus. 1a. Leaf blades 3–6 mm wide, ± flat, scabrous; ligules 3–6 mm long, erose at the apex; glumes 3–6 mm long, thick and opaque; callus hairs 65–100% as long as the lemma; panicles 6–20 cm tall; anthers degenerate and lacking pollen . . . 5a. C. stricta ssp. inexpansa (Gray) C.W. Greene 1b. Leaf blades 2–4 mm wide, involute, scabrous only on the margins and near the apex; ligules 1–3 mm long, entire at the apex; glumes 2–4.5 mm long, translucent at least on the margins and near the apex; callus hairs 50–75% as long as the lemma; panicles 5–12 cm tall; anthers containing pollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5b. C. stricta ssp. stricta Subspecies inexpansa is known from CT, ME, NH, VT. Subspecies stricta is known from ME, NH, VT. Though many named variants exist for ssp. inexpansa, two of these forms that occur in New England may warrant recognition. The “lacustris” variant has lower leaf sheaths minutely pubescent on the collar and relatively stout lemma awns with 1 or 2 twists. The “inexpansa” variant has glabrous leaf sheath collars and slender to stout lemmas with 0–1 twists.

Cenchrus The bur-like fascicles (in our species) of Cenchrus have two cycles of bristles. The outer bristles are positioned closer to the pedicel, are usually retrorsely oriented, and are terete to flattened in cross-section (depending on the species). The inner bristles are fused into a cupule with the apical portions distinct, vary from spreading to erect, and are flattened in cross-section. Reports of C. tribuloides from New England were based on collections of C. longispinus (mostly) and C. spinifex. Reference: Stieber and Wipff (2003). 1a. Bur-like fascicles with 45–75 bristles [Fig. 204], 8.3–11.9 mm long, the outer bristles terete, the inner bristles 0.5–0.9 (–1.4) mm wide at the base; carypsis 1.5–2.6 mm wide . . . . . . C. longispinus 1b. Bur-like fascicles with 8–40 bristles, 5.5–10.2 mm long, the outer bristles absent or present and then usually flattened, the inner bristles 1–3 mm wide; caryopsis 1–2 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. spinifex 1. Cenchrus longispinus (Hack.) Fern. N Fig. 204 long-spined sandbur. Cenchrus carolinianaus Walt.; C. echinatus L. forma longispinus Hack. • CT, MA, ME, NH, RI, VT; mostly southern New England and southern portions of northern states, though disjunct in Chittenden County, VT. Sandy soils of fields, roadsides, woodland edges, and railroads. 2. Cenchrus spinifex Cavanilles E coastal sandbur. Cenchrus incertus M.A. Curtis; C. pauciflorus Benth. • MA, ME, NH. Sandy soils of fields, roadsides, railroads, and disturbed places, wool waste. Reports of this species in VT are based on collections of Cenchrus longispinus.

Chloris Reference: Barkworth (2003a). 1a. Second lemma of spikelet pointed at the apex, not widened distally; plants perennial, 10–15 (–30) dm tall, usually producing stolons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. gayana 1b. Second lemma of spikelet ± truncate at the apex, sometimes widened distally; plants annual or perennial, 1.4–6 (–10) dm tall, not stoloniferous (rarely stoloniferous in C. virgata) 2a. Plants annual; leaf blades 3–6 (–15) mm wide; ligules up to 4 mm long; spikelets with a lower bisexual floret and 1 or 2 sterile, apical florets, the third floret often represented by only a rachilla segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. virgata

Fig. 204  Bur-like fascicle of Cenchrus longispinus.

242  MO NOCOTS

2b. Plants perennial; leaf blades 2–4 mm wide; ligules 0.7–1.3 mm long; spikelets with a lower bisexual floret and an upper sterile floret, without a third floret or its rachilla segment 3a. Upper glumes 1–1.5 mm long; lemma of upper floret unawned or with an excurrent midrib up to 1.5 mm long; panicle branches arranged in several, closely spaced whorls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cucullata 3b. Upper glumes 2.8–3.5 mm long; lemma of upper floret with a prolonged awn 3.2–7 mm long; panicle branches arranged in well-separated whorls with a terminal, solitary, erect branch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. verticillata 1. Chloris cucullata Bisch. E hooded windmill-grass. ME. On wool waste. 2. Chloris gayana Kunth E Rhodes windmill-grass. MA. Fields, roadsides, disturbed soil. The report of this species from Maine by Campbell et al. (1995) is based on a specimen from Hawaii—2 Mar 1930, Lowe s.n. (MAINE!). 3. Chloris verticillata Nutt. E tumble windmill-grass. ct, MA. Fields, roadsides, disturbed soil, lawns. 4. Chloris virgata Sw. E Fig. 205 feather windmill-grass. MA, ME. Wool waste.

Cinna Fig. 205  Inflorescence of Chloris virgata.

1a. Second (i.e., upper) glume 1-veined, 2.6–4.1 mm long, with wide, scarious margins, each scarious margin constituting nearly 25% of the total width of the glume; anthers 0.5–0.8 mm long; branches of the panicle spreading to drooping; lemma awn 0.1–2.5 mm long, commonly longer than 0.5 mm; leaf blades 6–25 cm long; palea with 2 closely spaced nerves (less frequently with only 1 nerve) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. latifolia 1b. Second glume 3-veined, 4.1–6.6 mm long, with narrow, scarious margins; anthers 0.8–1.8 mm long; branches of the panicle ascending to spreading; lemma awn 0.2–1.5 mm long, commonly shorter than 0.5 mm; leaf blades 15–40 cm long; palea with 1 nerve . . . . . . . . . . . . C. arundinacea 1. Cinna arundinacea L. N sweet wood-reed. Cinna arundinacea L. var. inexpansa Fern. & Grisc. • CT, MA, ME, NH, RI, VT. Riparian forests and terraces, stream banks, open marshes, swamps. 2. Cinna latifolia (Trev. ex Goepp.) Griseb. N slender wood-reed. Agrostis latifolia Trev. ex Goepp. • CT, MA, ME, NH, VT. Forested swamps, riparian forests, seepage forests, less commonly in mesic forests.

Corynephorus 1. Corynephorus canescens (L.) Beauv. E gray club-awn grass. Aira canescens L. • MA. Sandy soil of fields, roadsides, and clearings.

Crypsis 1. Crypsis schoenoides (L.) Lam. E swamp pricklegrass. Heleochloa schoenoides (L.) Host ex Roemer • MA, VT. Disturbed soil, waste places.

Poac e a e   2 43

Cynodon 1. Cynodon dactylon (L.) Pers. var. dactylon E Bermuda grass. Cynodon aristiglumis Caro & E.A. Sánchez • CT, MA. Fields, roadsides, waste areas.

Cynosurus 1. Cynosurus cristatus L. E crested dogtail grass. CT, MA, ME, NH, RI, VT. Roadsides, waste places, disturbed soil.

Dactylis 1. Dactylis glomerata L. E orchard grass. CT, MA, ME, NH, RI, VT. Fields, roadsides, cultivated land. This species includes diploid and tetraploid forms that appear to be morphologically distinguishable only on the basis of stomate and pollen size. Thus far, only tetraploid plants are known to have been collected in New England.

Dactyloctenium 1a. Inflorescence subglobose, congested, with branches 4–15 mm long; axis of each branch prolonged beyond the apical-most spikelet for 1–1.5 mm . . . . . . . . . . . . . . . . . . . . . . . D. radulans 1b. Inflorescence not congested, with divergent branches 15–60 mm long; axis of each branch prolonged beyond the apical-most spikelet for 1–6 mm . . . . . . . . . . . . . . . . . . . . . . D. aegyptium 1. Dactyloctenium aegyptium (L.) Willd. E Durban crowfoot grass. Cynosurus aegyptius L. • MA, ME. Waste places, cultivated land, disturbed soil. 2. Dactyloctenium radulans (R. Br.) Beauv. E button grass. Eleusine radulans R. Br. • MA. Wool waste, disturbed soil.

Danthonia Danthonia sericea Nutt. was reported from MA by several recent sources, likely based originally on Hitchcock and Chase (1950). The voucher specimen was misidentified and is Danthonia spicata. 1a. Spikelets (10–) 14–26 (–30) mm long; lemmas 5–10 mm long excluding the awns; apical teeth of lemma (2–) 4–6 (–7) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. californica 1b. Spikelets 7–16 mm long; lemmas 2.5–5 mm long excluding the awns; apical teeth of lemma 0.5–4 mm long 2a. Apical teeth of the lemma 0.5–2 mm long, triangular-acuminate [Fig. 207]; panicle 2–5 cm tall, crowded, the branches ascending to appressed after anthesis; pedicels shorter than or equaling the associated spikelets; leaf blades commonly involute, 0.8–2 (–3) mm wide, usually curling at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. spicata 2b. Apical teeth of the lemma 2–4 mm long, aristate or setaceous [Fig. 206]; panicle 5–10 cm tall, open, the branches often divergent after anthesis; pedicels as long as or longer than the associated spikelets; leaf blades flat, 2–4 mm wide, not curling . . . . . . . . . . . . . . . D. compressa 1. Danthonia californica Boland. E California oatgrass. Danthonia americana Scribn. • MA. Waste areas, roadsides, ditches.

244  MO NOCOTS

2. Danthonia compressa Austin ex Peck N Fig. 206 flattened oatgrass. Danthonia alleni Austin • CT, MA, ME, NH, RI, VT; throughout. Forest edges and openings, edges of shaded trails, old logging road beds, uncommonly in old pastures and fields. 3. Danthonia spicata (L.) Beauv. ex Roemer & J.A. Schultes N Fig. 207 poverty oatgrass. Danthonia spicata (L.) Beauv. ex Roemer & J.A. Schultes var. longipila Scribn. & Merr.; D. spicata (L.) Beauv. ex Roemer & J.A. Schultes var. pinetorum Piper • CT, MA, ME, NH, RI, VT; throughout. Dry-mesic to xeric, often sandy and/or thin soils of woodlands, grasslands, open balds, roadsides, and fields.

Deschampsia Reference: Barkworth (2007a). 1a. Plants annual; leaves few, short, the blades less than 33% as tall as the reproductive stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. danthonioides Fig. 206  Lemma of Danthonia compressa.

1b. Plants perennial; leaves relatively more numerous, longer, 33–50% as tall as the reproductive stem 2a. Panicle very narrow, with appressed branches; both glumes with 3 nerves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. elongata 2b. Panicle open, with ascending to spreading branches; first, and sometimes also the second, glume with 1 nerve 3a. Leaf blades involute, 1–2 mm wide; ligule 1–2.5 mm long; lemmas minutely scabrouspubescent, bearing a conspicuously bent awn that evidently exceeds the lemmas by 1–3 mm [Fig. 208]; palea not bifid at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. flexuosa 3b. Leaf blades flat or conduplicate, 1–5 mm wide; ligule usually 3–12 mm long; lemmas glabrous, bearing a ± straight awn that is shorter than to slightly exceeding the lemmas; palea bifid at the apex 4a. Stems 65–170 cm tall, 2–6 mm thick near the base; lower stem leaves 15–50 cm long, with ligules (2–) 5–10 (–12) mm long; panicles 15–45 cm tall . . . . . . D. cespitosa 4b. Stems 7–75 cm tall, 1–2.5 mm thick near the base; lower stem leaves 1–8 cm long, with ligules (2–) 3–4 (–7) mm long; panicles 2–22 cm tall . . . . . . . . . . . D. anadyrensis

Fig. 207  Lemma of Danthonia spicata.

1. Deschampsia anadyrensis V.N. Vassil. N glaucous hair grass. Deschampsia cespitosa (L.) Beauv. ssp. glauca (Hartman) Hartman; D. cespitosa (L.) Beauv. var. glauca (Hartman) Lindm. f.; D. glauca Hartman • CT, MA, ME, NH, RI, VT; more common in the northern states and becoming rare in southern New England. Rocky and gravelly shorelines, river shore cliffs and outcrops. Morphological review of museum specimens indicates that what has been called Deschampsia glauca (or its various nomenclatural synonyms) in New England is not the same as that from Sweden (where the type of Deschampsia glauca is from; David Murray, personal communication). The name D. anadyrensis may be the appropriate name for our taxon, based on a type from Russia. Though more work is needed to confirm the appropriate epithet for our taxon, it is known that the epithet “glauca” is not correct. 2. Deschampsia cespitosa (L.) Beauv. n tufted hair grass. Aira cespitosa L.; Deschampsia cespitosa (L.) Beauv. ssp. parviflora (Thuill.) Dumort.; D. cespitosa (L.) Beauv. var. parviflora (Thuill.) Coss. & Germ. • CT, MA, ME, NH, RI, VT. Fields, roadsides, and other open, disturbed places, rarely in riparian forests. 3. Deschampsia danthonioides (Trin.) Munro E annual hair grass. Aira danthonioides Trin. • ME. Wool waste, edges of lawns. 4. Deschampsia elongata (Hook.) Munro E slender hair grass. Aira elongata Hook. • ME. Fields, roadsides, disturbed soil. Inclusion of this species in New England is based on Weatherby et al. (1936), but specimens are unknown.

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5. Deschampsia flexuosa (L.) Trin. N Fig. 208 wavy hair grass. Aira flexuosa L.; Avenella flexuosa (L.) Drej.; A. flexuosa (L.) Drej. ssp. montana (L.) A. & D. Löve; Deschampsia flexuosa (L.) Trin. ssp. montana (L.) A. & D. Löve; Lerchenfeldia flexuosa (L.) Schur; L. flexuosa (L.) Schur ssp. montana (L.) Tzvelev • CT, MA, ME, NH, RI, VT. Thin and/or sandy soils of balds, mountains, woodlands, sand plains, forest openings, and ledges, sometimes forming extensive vegetative colonies in evergreen forests on mid-elevation peaks. Plants from high-elevation peaks have been segregated as ssp. montana. These plants have larger spikelets (5–7 mm vs. 4–5 mm in low-elevation plants) and more contracted panicles. They may deserve formal recognition (see Löve and Löve 1966 for discussion).

Desmazeria 1. Desmazeria rigida (L.) Tutin E fern grass. Catapodium rigidum (L.) C.E. Hubbard ex Doney; Poa rigida L.; Scleropoa rigida (L.) Griseb. • MA, RI. Fields, roadsides, ditches, disturbed soil.

Dichanthelium See references under Panicum for evidence for segregation of this genus. Many species of Dichanthelium flower twice during the growing season. In early to mid-summer, the chasmogamous primary panicle is produced at the summit of the main stem. Later, lateral branches and secondary panicles are produced, the secondary panicles are often partially or completely enclosed in leaf sheaths and are cleistogamous. Care needs to be taken when identifying species of Dichanthelium because the character states are sometimes different between the main stems and chasmogamous panicles and the lateral branches and cleistogamous panicles, the latter appearing in the later season. The following key is written for the main stems and chasmogamous panicles. Reference: Freckmann and Lelong (2003a). 1a. Basal and lower stem leaf blades similar in shape, elongate and narrow, (6.3–) 9.6–30 times as long as wide, erect to ascending, crowded [Fig. 213]; reproductive stems with 2–4 leaves 2a. Upper glume and lower lemma pointed at the apex and surpassing the upper (i.e., fertile) lemma by (0.2–) 0.5–1 mm; spikelets 3.2–4.3 mm long; terminal panicle with 7–20 (–25) spikelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. depauperatum 2b. Upper glume and lower lemma rounded to subacute at the apex, not surpassing upper lemma or surpassing it by no more than 0.3 mm; spikelets 2–3.2 mm long; terminal panicle with (12–) 20–70 spikelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. linearifolium 1b. Basal and lower stem leaf blades dissimilar, the crowded basal leaves ovate to lanceolate and spreading, 2.3–7.3 times as long as wide, the stem leaf blades longer and only rarely crowded; reproductive stems with 3–14 leaves 3a. Leaf blades with a white or green-white cartilaginous margin, cordate at base; spikelets 1.4–1.8 mm long, obovoid-spherical, nearly as wide as long . . . . . . . . . . . D. sphaerocarpon 3b. Leaf blades without or with a poorly developed cartilaginous margin, cordate to narrowed at base; spikelets 0.8–5.2 mm long, ellipsoid to obovoid, distinctly longer than wide 4a. Leaf sheaths viscid near the apex, especially on the surface away from the margins, long spreading-pubescent with ± soft hairs except near the base where moderately to densely retorse-pubescent above a prominent, glabrous, viscid band [Fig. 214] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. scoparium 4b. Leaf sheaths not viscid near the apex, variously glabrous or pubescent, without or, less commonly, with a prominent band of retrorse pubescence at the base, lacking a viscid band at the base

Fig. 208  Spikelet of Deschampsia flexuosa.

246 MO NOCOTS

5a. Ligules composed of a membranous base that is ciliate apically; plants from rhizomes (usually from caudices in D. commutatum); leaf blades 5–40 mm wide, truncate to cordate at base (often narrowed to the base in D. xanthophysum) 6a. Leaf blades strictly ascending to erect; panicle usually cylindrical, 1–4 (–5) cm wide, more than 2 times as tall as wide, with strictly ascending to erect branches [Fig. 215]; spikelets rounded at apex . . . . . . . . . . . . . . . . . . . . . . . . . D. xanthophysum 6b. Leaf blades spreading to ascending; panicle usually broader, (3–) 4–13 cm wide, mostly 1–2 times as long as wide, with spreading to ascending branches; spikelets pointed to bluntly pointed at apex 7a. Leaf sheaths pubescent with pustulose-based hairs [Fig. 212]; rhizomes 3–5 mm thick; reproductive stems with (5–) 7–14 leaves 8a. Upper glume and lower lemma obtuse to subacute (note: those of the secondary panicles often sharply acute at the apex), not or hardly surpassing the upper lemma; leaf blades 15–30 mm wide, flat to the tip; spikelets sparsely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. clandestinum 8b. Upper glume and lower lemma acute to long-acute, surpassing the upper lemma; leaf blades 7–15 mm wide, tapering to an involute tip; spikelets glabrous or sparsely pubescent . . . . . . . . . . . . . . . . . . . . . . . . D. scabriusculum 7b. Leaf sheaths glabrous or pubescent with non-pustulose-based hairs; rhizomes up to 2 mm thick or absent; reproductive stems with 4–6 leaves 9a. Stems densely crisp-puberulent; ligules up to 0.3 mm long; spikelets 2.2–3.2 mm long; stem leaf blades 5–10 mm wide; plants from caudices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. commutatum 9b. Stems glabrous or shortly pilose; ligules 0.4–0.9 mm long; spikelets 2.9– 5.2 mm long; stem leaf blades 15–40 mm wide; plants from knotty rhizomes 10a. Spikelets 2.9–3.9 mm long; nodes nearly glabrous to sparsely (rarely moderately) pubescent with retrorsely oriented hairs, the longer hairs up to 0.8 (–1.4) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. latifolium 10b. Spikelets 3.8–5.2 mm long; nodes moderately to densely (rarely sparsely) pubescent with retrorsely oriented hairs, the longer hairs 1.3–3.4 mm long [Fig. 211] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. boscii 5b. Ligules composed entirely of hairs; plants from caudices; leaf blades 2–12 (–15) mm wide, narrowed to rounded at the base (truncate to cordate in D. boreale and some D. dichotomum) 11a. Spikelets 2.7–4.2 mm long; upper glumes often with a purple or orange-tan to orange spot or blotch near the base, strongly veined . . . . . . . . . . . D. oligosanthes 11b. Spikelets 0.8–2.6 mm long; upper glumes usually without an orange or purple area at base, weakly to strongly veined 12a. Ligule a conspicuous band of hairs 1–5 mm tall that protrudes above the summit of the sheath 13a. Spikelets 0.8–1 (–1.1) mm long; stems 0.3–0.8 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. wrightianum 13b. Spikelets 1.3–2.6 mm long; stems thicker than (0.8–) 1 mm 14a. Spikelets 1.3–2.1 mm long; sheaths glabrous or with hairs usually shorter than 2 (–3) mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. acuminatum 14b. Spikelets 2.1–2.6 mm long; sheaths pubescent with hairs 2–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. ovale

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12b. Ligule nearly absent or up to 1 mm long, the hairs hardly protruding above the summit of the sheath 15a. Spikelets pubescent; nodes of stem glabrous; leaf blades truncate to cordate at base; uppermost stem leaf erect or ascending; spikelets (1.9–) 2–2.2 (–2.3) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. boreale 15b. Spikelets glabrous or pubescent; nodes of stem glabrous or the lower nodes slightly to evidently retrorse-pubescent; leaf blades narrowed to the base in common forms (varying to truncate to cordate); uppermost stem leaf usually spreading; spikelets 1.5–2.5 mm long . . . . . . . . . . D. dichotomum 1. Dichanthelium acuminatum (Sw.) Gould & C.A. Clark

N C Fig. 209, 210

hairy rosette-panicgrass.  1a. Dichanthelium columbianum (Scribn.) Freckmann; Panicum acuminatum Sw. var. columbianum (Scribn.) Lelong; P. columbianum Scribn.; P. oricola A.S. Hitchc. & Chase; P.  sabulorum (Lam.) Gould & C.A. Clark var. thinium (A.S. Hitchc. & Chase) C.F. Reed;  1b. Dichanthelium lanuginosum (Ell.) Gould; Panicum acuminatum Sw.; P. auberne Ashe; P. lanuginosum Ell.; 1c. Dichanthelium lanuginosum (Ell.) Gould var. lindheimeri (Nash) Fern.; D. lindheimeri (Nash) Gould; Panicum lanuginosum Ell. var. lindheimeri (Nash) Fern.; P. lanuginosum Ell. var. septentrionale (Fern.) Fern.;  1d. Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var. densiflorum (Rand & Redf.) Gould & C.A. Clark; Panicum nitidum Lam. var. densiflorum Rand & Redf.; P. spretum J.A. Schultes; 1e. Dichanthelium lanuginosum (Ell.) Gould var. fasciculatum (Torr.) Spellenberg; Panicum lanuginosum Ell. var. fasciculatum (Torr.) Fern.; P. lanuginosum Ell. var. tennesseense (Ashe) Gleason; P. subvillosum Ashe; P. tennesseense Ashe; 1f. Panicum lanuginosum Ell. var. implicatum (Scribn.) Fern.; P. acuminatum Sw. var. implicatum (Scribn.) C.F. Reed; P. meridionale Ashe • CT, MA, ME, NH, RI, VT. Frequently in open places with sandy or thin soils, such as shorelines, fields, roadsides, disturbed places, grasslands, balds, ledges, rocky slopes, and woodlands. 1a. Stems and leaf sheaths densely puberulent (the sheaths sometimes less so) with minute, non-pustulose-based hairs, with scattered longer hairs also usually present (i.e., the pubescence of two types) [Fig. 209] . . . . . . . . . . . . . . . . . . . . . . 1a. D. acuminatum ssp. columbianum (Scribn.) Freckmann & Lelong 1b. Stems and leaf sheaths glabrous or pubescent, when pubescent, most or all of the hairs elongate (i.e., exceeding 1 mm) and usually pustulose-based, not of two types (except in the rare ssp. acuminatum and sometimes in ssp. fasciculatum) 2a. Leaf sheaths densely pubescent with long hairs that originate usually from small pustules and also with minute hairs underneath the longer ones (i.e., the pubescence of two types); leaf blades softly pubescent with velvety hairs on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. D. acuminatum ssp. acuminatum 2b. Leaf sheaths glabrous or pubescent, the hairs all long (i.e., of one type), and originating from rather conspicuous pustules or not at all pustulose-based (sometimes with hairs of two types in the common ssp. fasciculatum, but the short hairs usually scarce); leaf blades variously glabrous, puberulent, or pilose on the abaxial surface, but generally not feeling velvety to the touch 3a. Leaf sheaths glabrous or pubescent with non-pustulose-based hairs (though the margins of the sheaths often ciliate near the apex) [Fig. 210] 4a. Terminal panicle 2.5–7 cm tall, less than 2 times as tall as wide, open; leaf blades and sheaths yellow-green, with few or no pustulose-based cilia near the base; spikelets usually obovoid . . . . . 1c. D. acuminatum ssp. lindheimeri (Nash) Freckmann & Lelong 4b. Terminal panicle 4–12 cm tall, 2–4 times as tall as wide, somewhat congested; leaf blades and sheaths green or purple-tinged, often with conspicuous, pustulosebased cilia near the base; spikelets usually ellipsoid . . . . . . . . . . . . . . 1d. D. acuminatum ssp. spretum (J.A. Schultes) Freckmann & Lelong

Fig. 209  Leaf sheath of Dichanthelium acuminatum ssp. columbianum.

Fig. 210  Leaf sheath of Dichanthelium acuminatum ssp. lindheimeri.

248  MO NOCOTS

3b. Leaf sheaths pubescent with pustulose-based hairs 5a. Leaf blades glabrous or pilose on the adaxial surface with hairs shorter than 3 mm, usually 6–12 mm wide, spreading or ascending; spikelets 1.5–2 mm long . . . . . . . . . . . . . . . . . 1e. D. acuminatum ssp. fasciculatum (Torr.) Freckmann & Lelong 5b. Leaf blades pilose on the adaxial surface with hairs 3–6 mm long, usually 2–6 mm wide, erect to ascending; spikelets 1.1–1.5 mm long . . . . . . . . . . . . . . . . . 1f. D. acuminatum ssp. implicatum (Scribn.) Freckmann & Lelong Subspecies columbianum is known from CT, MA, ME, NH, RI, VT. Subspecies acuminatum is known from MA, RI and is of regional conservation concern. Subspecies lindheimeri is known from CT, MA, ME, NH, RI, VT. Subspecies spretum is known from CT, MA, ME, NH, RI. Subspecies fasciculatum is known from CT, MA, ME, NH, RI, VT. Subspecies implicatum is known from CT, MA, ME, NH, RI, VT.

Fig. 211  Node of Dichanthelium boscii.

1‌ × 10. Dichanthelium ×scoparioides (Ashe) Mohlenbrock is a rare rosette panicgrass hybrid in New England known from CT, MA, VT. It resembles D. ovale due to the long, projecting ligule of hairs mostly 2–3 mm long and the spikelets 2.2–2.4 mm long. However, the stems and sheaths are pubescent with short (mostly 1–1.5 mm long) and sparse, spreading-ascending hairs (or these parts ± glabrous; rather than having long, spreading or ascending to appressed hairs in D. ovale). The hybrid is further characterized by leaf blades mostly 6–10 mm wide. It was also reported from RI by Kartesz (1999); however, this record was erroneously based on Collins (1928), who did not report this hybrid from RI. The epithet scopariodes may not properly refer to this hybrid. 2. Dichanthelium boreale (Nash) Freckmann N northern rosette-panicgrass. Panicum boreale Nash • CT, MA, ME, NH, RI, VT. Mesic forests, forest openings, trail edges, shorelines, fields. The names Panicum bicknellii Nash and P. calliphyllum Ashe have been considered as taxonomic synonyms of Dichanthelium boreale by some authors. However, both are likely rare hybrids. The former may be a cross between D. dichotomum ssp. dichotomum and D. linearifolium, and the latter may be a cross between D. boreale and D. xanthophysum. 3. Dichanthelium boscii (Poir.) Gould & C.A. Clark N Fig. 211 Bosc’s rosette-panicgrass. Panicum boscii Poir.; P. boscii Poir. var. molle (Vasey) A.S. Hitchc. & Chase • CT, RI. Woodlands and semi-open rocky slopes, often associated with Quercus and/ or Carya.

Fig. 212  Leaf sheath of Dichanthelium clandestinum.

4. Dichanthelium clandestinum (L.) Gould N Fig. 212 deer-tongue rosette-panicgrass. Panicum clandestinum L. • CT, MA, ME, NH, RI, VT; throughout. Forest openings and edges, fields, open rights-of-way, sandy road shoulders, shorelines. Rare individuals of Dichanthelium clandestinum show sheath morphology similar to that of D. scoparium, especially the orientation and zonation of hairs (see additional diagnostic characters under D. scoparium). 5. Dichanthelium commutatum (J.A. Schultes) Gould ssp. ashei (G. Pearson ex Ashe) Freckmann & Lelong N Ashe’s variable rosette-panicgrass. Panicum ashei Pearson ex Ashe; P. commutatum (J.A. Schultes) Gould var. ashei (Pearson ex Ashe) Fern.; P. umbrosum Le Conte ex Torr. • CT, MA, RI; also reported from NH by Freckmann and Lelong (2003a), but specimens are unknown. Woodlands, openings, sandy fields. Dichanthelium commutatum was reported from ME by Gould and Clark (1978), but the seasonal development of the specimen is in contradiction with the collection date. The specimen is believed to be from much farther south and the result of a label mix-up (Robert Freckmann, personal communication). Reports of Dichanthelium commutatum ssp. commutatum from New England appear to be in error. Those specimens identified as such and seen by Robert Freckmann or me were misidentified.

Fig. 213  Habit of Dichanthelium depauperatum.

6. Dichanthelium depauperatum (Muhl.) Gould N Fig. 213 starved rosette-panicgrass. Panicum depauperatum Muhl.; P. depauperatum Muhl. var. psilophyllum Fern. • CT, MA, ME, NH, RI, VT. Dry-mesic to xeric soils of woodlands, rocky slopes, balds, grasslands, and open, sandy areas.

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7. Dichanthelium dichotomum (L.) Gould

NC

forked rosette-panicgrass.  7a. Panicum barbulatum Michx.; P. dichotomum L.; 7b. Panicum clutei Nash; P. dichotomum L. var. mattamuskeetense (Ashe) Lelong; P. mattamuskeetense Ashe; 7c. Dichanthelium microcarpon (Muhl. ex Ell.) Mohlenbrock; Panicum microcarpon Muhl. ex Ell.; P. nitidum Lam. var. ramulosum Torr. • CT, MA, ME, NH, RI, VT. Mesic to xeric soils of woodlands, rocky forests, balds, and sandy roadsides, often with Quercus and/or Carya, or in wet-mesic to hydric soils of swamps, wetland edges, low fields, stream banks, pond shores, and bog edges. Reports of Dichanthelium dichotomum ssp. lucidum (Ashe) Freckmann & Lelong from New England are based on specimens of D. dichotomum ssp. dichotomum (specimens at CONN!, GH!, MASS!, NEBC!). 1a. Lower nodes glabrous or infrequently sparsely pubescent; leaf blades usually 5–7 mm wide, slightly narrowed or constricted near the base . . . . . . . . 7a. D. dichotomum ssp. dichotomum 1b. Lower nodes pubescent with retrorsely oriented hairs; leaf blades (5–) 7–14 mm wide, not or only scarcely narrowed or constricted near the base 2a. Spikelets 1.8–2.5 mm long, pubescent; leaf sheaths and blades moderately to densely velutinous . . . . . 7b. D. dichotomum ssp. mattamuskeetense (Ashe) Freckmann & Lelong 2b. Spikelets 1.5–1.8 mm long, usually glabrous; leaf sheaths and blade surfaces ± glabrous to sparsely pubescent . . . . . . . . . . . . . . . 7c. D. dichotomum ssp. microcarpon (Muhl. ex Ell.) Freckmann & Lelong Subspecies dichotomum is known from CT, MA, ME, NH, RI, VT. Subspecies mattamuskeetense is known from MA, RI. Both of these subspecies are normally found in dry-mesic to xeric soils, but ssp. mattamuskeetense is occasionally associated with small and/or seasonal wetlands. Subspecies mattamuskeetense is of regional conservation concern. Subspecies microcarpon is known from CT, MA, RI and is usually found in wet-mesic to hydric soils. 8. Dichanthelium latifolium (L.) Harvill N broad-leaved rosette-panicgrass. Panicum latifolium L. • CT, MA, ME, NH, RI, VT. Mesic to drymesic, usually deciduous, forests and woodlands. 9. Dichanthelium linearifolium (Scribn.) Gould N linear-leaved rosette-panicgrass. Dichanthelium depauperatum (Muhl.) Gould var. perlongum (Nash) Boivin; Panicum linearifolium Scribn.; P. perlongum Nash; P. werneri Scribn. • CT, MA, ME, NH, RI, VT. Dry-mesic to xeric soils of woodlands, rocky slopes, balds, grasslands, and open, sandy areas. 10. Dichanthelium oligosanthes (J.A. Schultes) Gould ssp. scribnerianum (Nash) Freckmann & Lelong N few-flowered rosette-panicgrass. Panicum macrocarpon Le Conte ex Torr.; P. oligosanthes J.A. Schultes var. scribnerianum (Nash) Fern. • CT, MA, ME, NH, RI, VT; restricted in ME to extreme south. Sandy openings, grasslands, open woodlands, sandy roadsides. Dichanthelium oligosanthes ssp. oligosanthes was reported from southern New England by Angelo and Boufford (1998); however, the voucher specimens (at MASS! and NEBC!) were misidentified (most were D. oligosanthes ssp. scribnerianum). 11. Dichanthelium ovale (Ell.) Gould & C.A. Clark

NC

stiff-leaved rosette-panicgrass.  11a. Dichanthelium commonsianum (Ashe) Freckmann; Panicum addisonii Nash; P. commonsianum Ashe; P. ovale Ell. var. addisonii (Nash) C.F. Reed; P. ovale Ell. var. pseudopubescens (Nash) Lelong; 11b. Panicum acuminatum Sw. var. villosissimum (Nash) C.F. Reed; P. atlanticum Nash; P. lanuginosum (Ell.) Gould var. villosissimum (Nash) Gould; P. villosissimum Nash • CT, MA, RI. Sandy soils of woodlands, coastal plain pond shores, and disturbed openings. 1a. Lower stem internodes and leaf sheaths pubescent with ascending to erect, nonpustulose-based hairs; principal leaf blades 2–6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . 11a. D. ovale ssp. pseudopubescens (Nash) Freckmann & Lelong

25 0  MONOCOTS

1b. Lower stem internodes and leaf sheaths pubescent with spreading to retrorse, pustulosebased hairs; principal leaf blades 6–10 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11b. D. ovale ssp. villosissimum (Nash) Freckmann & Lelong Subspecies pseudopubescens is known from CT, MA. Subspecies villosissimum is known from CT, MA, RI. Both subspecies are of regional conservation concern. 12. Dichanthelium scabriusculum (Ell.) Gould & C.A. Clark

NC

tall swamp rosette-panicgrass. Panicum aculeatum A.S. Hitchc. & Chase; Panicum scabriusculum Ell. • CT, MA, RI. Dry, sandy roadsides, grasslands, and open rights-of-way, mesic to hydric, rocky or boggy soils adjacent to streams. 13. Dichanthelium scoparium (Lam.) Gould

N C Fig. 214

velvety rosette-panicgrass. Panicum scoparium Lam. • MA, RI; coastal plain. Open, often wet, sandy soils of pond shores, wetland borders, and clearings, rarely also near dunes and along roadsides. This rare species is best identified by its unusual sheaths with a unique combination of characters. It is related to the common Dichanthelium clandestinum but differs in its smaller spikelets (2.2–2.8 mm vs. 2.4–3.6 mm), denser and softer sheath pubescence, ligule 0.5–1 (–2) mm long composed entirely of hairs (rather than a ciliate membrane 0.4–0.9 mm long), and viscid regions on the sheath (among other characters). Fig. 214  Leaf sheath of Dichanthelium scoparium.

14. Dichanthelium sphaerocarpon (Ell.) Gould N round-fruited rosette-panicgrass. Panicum sphaerocarpon Ell. • CT, MA, ME, NH, RI, VT; restricted in ME and NH to the extreme southern portion of states. Woodlands, open rights-of-way, roadsides, fields, usually in sandy soils. Reports of Dichanthelium polyanthes (Schult.) Mohlenb. from New England are largely based on collection of D. sphaerocarpon (specimens at CONN!, NEBC!, YU!). 15. Dichanthelium wrightianum (Scribn.) Freckmann N Wright’s rosette-panicgrass. Panicum acuminatum Sw. var. wrightianum (Scribn.) C.F. Reed; P. minutulum Desv.; P. wrightianum Scribn. • MA, RI. Pond shores and exposed, vegetated floors of temporary pools on the coastal plain. 16. Dichanthelium xanthophysum (Gray) Gould N Fig. 215 pale-leaved rosette-panicgrass. Panicum xanthophysum Gray • CT, MA, ME, NH, VT. Dry-mesic to xeric, well drained soils of woodlands, heaths, and cleared rights-of-way.

Digitaria Reference: Wipff (2003b). Fig. 215  Inflorescence of Dichanthelium xanthophysum.

1a. Inflorescence an open panicle, the primary branches usually branched again [Fig. 216]; spikelets borne on pedicels 10–40 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cognata 1b. Inflorescence a panicle of secund, spike-like racemes, the primary branches usually unbranched [Fig. 217]; spikelets borne on pedicels 0.2–3.8 mm long 2a. Axis of panicle branches triquetrous, but not wing-margined; reproductive stems usually upright, only rarely rooting at the lower nodes . . . . . . . . . . . . . . . . . . . . . . D. filiformis 2b. Axis of panicle branches wing-margined; reproductive stems usually decumbent and rooting from the lower nodes 3a. Lower glume present, 0.2–0.4 mm long; upper glume 33–50 (–60)% as long as the entire spikelet; upper lemmas yellow or gray to brown at maturity; spikelets of the middle portion of the primary panicle branches usually borne in pairs on pedicels that are not adnate to the branch axis; leaf sheaths usually pustulose-pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sanguinalis 3b. Lower glume absent or a veinless, membranous rim; upper glume (50–) 75–100% as long as the entire spikelet; upper lemmas dark purple-brown at maturity; spikelets of the middle portion of the primary panicle branches usually borne in trios on pedicels that

P oac e a e   25 1

are adnate to the branch axis (at least the longer pedicels of each cluster); leaf sheaths glabrous over the surface (rarely pubescent, the upper margins sometimes ciliate) 4a. Spikelets 1.7–2.3 mm long; plants with axillary panicles, these often concealed in the leaf sheaths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. ischaemum 4b. Spikelets 1.2–1.7 mm long; plants without axillary panicles . . . . . . . . D. violascens 1. Digitaria cognata (J.A. Schultes) Pilger N Fig. 216 fall crabgrass. Leptoloma cognatum (J.A. Schultes) Chase • CT, MA, NH, RI, VT. Sandy fields, roadsides, railroads, grasslands, and other open, dry, sandy areas. Thought by Sorrie and Somers (1999) to be introduced; however, this species, like others it resembles (e.g., Eragrostis spectabilis, Panicum capillare), is capable of using human-disturbed habitats and has expanded its range in New England as a result. See Zika (1990) for discussion of this grass in VT. 2. Digitaria filiformis (L.) Koel.

NC

slender crabgrass. 2a. Digitaria laeviglumis Fern.; 2b. Panicum filiforme L. • CT, MA, NH, RI. Dry, open, often sandy, areas, such as fields, roadsides, and disturbed lots, also in peaty depression on granitic ledges. Digitaria filiformis resembles D. ischaemum and D. violescens in several character states, including the highly reduced lower glume, relative length of the upper glume, and color of the upper lemma. In addition to characters stated in the key, D. filiformis differs from the two mentioned species in its pustulose-based hairs on the lower sheaths (vs. glabrous to sparsely pubescent).

Fig. 216  Inflorescence of Digitaria cognata.

1a. Lower lemmas glabrous . . . . . . . . . . . . . . . . 2a. D. filiformis var. laeviglumis (Fern.) Wippf 1b. Lower lemmas pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. D. filiformis var. filiformis Variety laeviglumis is known from (and endemic to) NH. It is of regional conservation concern and occurs in peaty depressions on granitic ledges in Hillborough County. Variety filiformis is known from CT, MA, NH, RI. 3. Digitaria ischaemum (Schreb.) Muhl. E smooth crabgrass. Panicum ischaemum Schreb. • CT, MA, ME, NH, RI, VT. Roadsides, yards, railroads, lawns, sandy, disturbed soil. 4. Digitaria sanguinalis (L.) Scop. E Fig. 217 hairy crabgrass. Panicum sanguinale L. • CT, MA, ME, NH, RI, VT. Roadsides, yards, railroads, lawns, sandy, disturbed soil. 5. Digitaria violascens Link E violet crabgrass. Digitaria ischaemum (Schreb.) Muhl. var. violascens (Link) Radford; Syntherisma chinensis (Nees) A.S. Hitchc. • MA. Fields, roadsides, disturbed places.

Distichlis 1. Distichlis spicata (L.) Greene N saltgrass. Distichlis spicata (L.) Greene var. borealis (J. Presl) Beetle; Uniola spicata L. • CT, MA, ME, NH, RI; coastal counties. Saline marshes and shores.

Echinochloa In the following key, spikelet and scale lengths do not include the awns (when present). Reference: Michael (2003). 1a. Panicle branches 0.7–2 (–4) cm long, few, distant, simple; leaf blades 3–6 (–10) mm wide; spikelets 2–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. colona 1b. Panicle branches (1–) 2–10 cm long, variable in number but often crowded, sometimes the longer branches with secondary branches; leaf blades 5–30 mm wide (sometimes narrower in small forms of E. crus-galli); spikelets 2.5–5 mm long

Fig. 217  Inflorescence of Digitaria sanguinalis.

25 2 MON OCOTS

2a. Lower lemmas usually unawned; upper lemmas wider and longer than the upper glumes at maturity, the margins and apex exposed and visible . . . . . . . . . . . E. frumentacea 2b. Lower lemmas usually awned; upper lemmas hardly, if at all, exceeding the upper glumes in length and width, primarily concealed 3a. Lower sheaths hispid with pustulose-based hairs; lower 1 or 2 nodes evidently pubescent with long, erect-ascending to spreading-ascending hairs . . . . . . . . . E. walteri 3b. Sheaths glabrous; lower nodes glabrous or sparsely pubescent with short hairs 4a. Coriaceous portion of upper lemmas rounded at apex, abruptly transitioning to an early withering, membranous apex, the two regions separated by a line of minute hairs [Fig. 218] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. crus‑galli 4b. Coriaceous portion of upper lemmas acute to acuminate at apex, gradually transitioning from the coriaceous body into the membranous tip, the two regions not offset by a line of minute hairs [Fig. 219] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. muricata 1. Echinochloa colona (L.) Link E awnless barnyard grass. Panicum colonum L. • MA, VT. Cultivated fields, ditches, disturbed soil. 2. Echinochloa crus-galli (L.) Beauv. E Fig. 218 Fig. 218  Upper (i.e., fertile) lemma of Echinochloa crus-galli.

common barnyard grass. Panicum crus-galli L. • CT, MA, ME, NH, RI, VT. Cultivated fields, areas of habitation, ditches, disturbed soil. 3. Echinochloa frumentacea Link E billion-dollar barnyard grass. Echinochloa crus-galli (L.) Beauv. var. frumentacea (Link) W. Wight • CT, MA, ME, NH, RI, VT. Fields, roadsides, seeded areas, disturbed soil. Echinochloa frumentacea resembles E. crus-galli in many respects but on average is a taller plant with wider leaves in New England and has ± unawned lower lemmas. 4. Echinochloa muricata (Beauv.) Fern. N Fig. 219 American barnyard grass.  4a. Echinochloa microstachya (Wieg.) Rydb.; E. pungens (Poir.) Rydb. var. microstachya (Wieg.) Fern. & Grisc.; E. pungens (Poir.) Rydb. var. wiegandii Fassett; 4b. Echinochloa pungens (Poir.) Rydb. • CT, MA, ME, NH, RI, VT. Damp sandy shorelines, fields, roadsides, disturbed soil.

Fig. 219  Upper (i.e., fertile) lemma of Echinochloa muricata.

1a. Spikelets 2.5–3.8 mm long; lower lemmas unawned or with awns to 6 (–10) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a. E. muricata var. microstachya Wieg. 1b. Spikelets 3.5–5 mm long; lower lemmas with awns 6–16 mm long  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. E. muricata var. muricata Variety microstachya is known from CT, MA, ME, NH, RI, VT. Variety muricata is known from CT, MA, ME, NH, RI, VT. 5. Echinochloa walteri (Pursh) Heller N coast barnyard grass. Panicum walteri Pursh • CT, MA, NH, RI. Shorelines, upper borders of saline and brackish marshes, low, wet areas.

Eleusine 1. Eleusine indica (L.) Gaertn. E goosegrass. Cynosurus indicus L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, yards, disturbed soil.

Elymus In the following key, measurements of spike width include the awns. Collections of Elymus should include all of the aerial stem (i.e., collect at the ground level) so that the number of nodes can be accurately assessed. References: Haines (2000d), Barkworth et al. (2007).

Poac e a e   25 3

1a. Spikelets spreading to horizontally divergent at maturity; glumes obsolete or represented by veinless bristles up to 10 mm long (rarely longer in robust spikes) and 0.1–0.2 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. hystrix 1b. Spikelets appressed to spreading-ascending; glumes present, with (1–) 2–7 veins, 6–40 mm long including the awn and 0.2–2.3 mm wide [Figs. 221, 223, 224] 2a. Spikelets mostly solitary at each node of the spike; glumes 6–7 mm long including the awn 3a. Anthers 3–7 mm long (i.e., plants outcrossing); plants rhizomatous; spikelets disarticulating below the glumes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. repens 3b. Anthers 1–2 mm long (i.e., plants self-pollinating); plants cespitose, without rhizomes; spikelets disarticulating above the glumes . . . . . . . . . . . . . . . . . E. trachycaulus 2b. Spikelets 2–4 at each node of the spike, usually paired [Figs. 220, 224]; glumes 10–40 mm long including the awn 4a. Awns arching at maturity, especially in late season and in drying; paleas 7–14 mm long; glumes thin or compressed at the base, indurate basally for up to 1 mm 5a. Leaf blades thin, flat, short-pilose on the adaxial surface, (8–) 15–25 mm wide, numbering mostly 9–15 per reproductive stem; glumes 0.3–0.8 mm wide, with 1–3 (–5) veins; spikes often conspicuously drooping from near the base . . . . . . . . . . E. wiegandii 5b. Leaf blades firm, flat or involute, minutely scabrous on the adaxial surface (rarely pubescent), 3–15 mm wide, numbering mostly 5–9 per reproductive stem; glumes 0.5–1.6 mm wide, with 3–5 veins; spikes nearly erect to arching . . . . . . . E. canadensis 4b. Awns straight or nearly so; paleas 5–10 (–12) mm long; glumes ± terete at the base, indurate basally for 0.5–4 mm 6a. Glumes setaceous, only slightly broadened apically, 0.2–1 mm wide, straight or slightly bowed out at the base [Fig. 222]; spikes arching to nodding 7a. Leaf blades softly villous or pilose on the adaxial surface; internodes of spike (1.5–) 2–3 (–4) mm long; lemmas usually villous, 5.5–9 mm long excluding the awns, 0–1.5 mm longer than the obtuse to emarginate paleas; spikelets with 1 or 2 (rarely 3) florets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. villosus 7b. Leaf blades glabrous or scabrous on the adaxial surface; internodes of spike 3–5 mm long; lemmas usually minutely scabrous, 7–14 mm long excluding the awns, 1–5 mm longer than the acute paleas; spikelets with 2 or 3 (rarely 4) florets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. riparius 6b. Glumes flattened above the base, 0.7–2.3 mm wide, slightly to strongly bowed out at the base and exposing the florets [Fig. 223]; spikes erect [Figs. 221, 224] 8a. Glumes including the awns 27–40 mm long, the glume awns 10–25 mm long; lemmas including the awns 35–50 mm long, the lemma awns (10–) 15–40 mm long; spikes 25–60 mm wide, usually exserted [Fig. 221]; spikelets spreadingascending [Fig. 221] 9a. Spikes with 15–30 nodes separated by internodes 3–5 mm long; auricles 0–2 mm long; glumes and lemmas often pubescent; plants of dry-mesic woodlands and forest openings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. glabriflorus 9b. Spikes with 9–18 nodes separated by internodes 4–7 mm long [Fig. 221]; auricles 2–3 mm long; glumes and lemmas usually glabrous or minutely scabrous; plants of rich, mesic alluvium, less commonly on forested slopes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. macgregorii 8b. Glumes including the awns 7–27 mm long, the glume awns absent or up to 10 (–15) mm long; lemmas including the awns 6–30 mm long, the lemma

25 4 MON OCOTS

awns absent or up to 5–15 (–20) mm long; spikes 7–20 mm wide, exserted or partly included in the upper leaf sheath [Figs. 220, 224]; spikelets appressed to appressed-ascending 10a. Awns absent or up to 3 (­–5) mm long [Fig. 220]; leaf blades often ascending and somewhat involute, the lower frequently narrower and senescing earlier than the upper ones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. curvatus 10b. Awns 3–15 (–20) mm long [Figs. 223, 224]; leaf blades usually laxly spreading, flat or involute, the lower neither noticeably narrower nor earlier senescing than the upper . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. virginicus 1. Elymus canadensis L. N Great Plains wild-rye. Elymus canadensis L. var. glaucifolius (Muhl. ex Willd.) Torr.; E. canadensis L.var. villosus Bates • CT, MA, ME, NH, VT. Riparian forests, river banks, railroads. Some collections from New England show prominently glaucous leaf blades and spikelets and, on average, have stiffer, more involute leaf blades. These plants, referred to as var. glaucifolius, are distinctive in the extremes but appear to intergrade with non-glaucuous plants with laxer leaf blades. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species could be in RI (i.e., the author is unaware of any collections). 2. Elymus curvatus Piper E Fig. 220 awnless wild-rye. Elymus submuticus (Hook.) Smyth & Smyth; E. virginicus L. var. submuticus Hook. • MA, RI. Disturbed soil near coastal beaches and ports. 3. Elymus glabriflorus (Vasey ex Dewey) Scribn. & Ball NC southeastern wild-rye. 3a. Elymus australis Scribn. & Ball; E. virginicus L. var. australis (Scribn. & Ball) A.S. Hitchc.; 3b. Elymus virginicus L. var. glabriflorus (Vasey ex Dewey) Bush • CT, MA, RI. Dry-mesic, deciduous forests, usually occurring on hills ridges, including trap rock, in shallow soils associated with Quercus and/or Carya. Fig. 220  Inflorescence of Elymus curvatus.

1a. Glumes and lemmas pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. E. glabriflorus var. australis (Scribn. & Ball) J.J.N. Campb. 1b. Glumes and lemmas glabrous or minutely scabrous . . . 3b. E. glabriflorus var. glabriflorus Variety australis is known from CT, MA, RI. Variety glabriflorus is known from only CT. Both varieties are of regional conservation concern. 3 × 4. This extremely rare grass hybrid is currently known within New England from a single hill in New Haven County, CT, where it occurs with both parents. It is recognized by the intermediate nature of the glume development, spike width, and spikelet orientation. 4. Elymus hystrix L. N eastern bottle-brush grass.  4a. Hystrix patula Moench var. bigeloviana (Fern.) Deam;  4b. Hystrix hystrix (L.) Millsp.; H. patula Moench • CT, MA, ME, NH, RI, VT. Rich, mesic soils of high-terrace floodplain forests and deciduous forests, often associated with Acer saccharum, Tilia americana, and/or Fraxinus americana; also in dry-mesic woodlands and forests on hills and ridges underlaid by circumneutral to basic bedrock. 1a. Lemmas appressed-puberulent to strigose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a. E. hystrix var. bigeloviana (Fern.) Bowden 1b. Lemmas glabrous or scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. E. hystrix var. hystrix Variety bigeloviana is known from CT, MA, ME, NH, RI, VT. It appears to be most common in mesic soils. Variety hystrix is known from CT, MA, ME, NH, VT. 4 × 7. This very rare wild-rye hybrid is known from VT. It generally resembles Elymus riparius with a large, arching spike. However, the spike internodes are 6–7 mm long (rather than 3–5 mm long), the glumes range in development from nearly absent or merely slender bristles (as in E. hystrix) to fully developed (as in E. riparius), and the spikelets are more loosely spreading than in typical E. riparius.

P oac e a e   25 5

5. Elymus macgregorii R. Brooks & J.J.N. Campb.

N C Fig. 221

early wild-rye. CT, MA, ME, NH, RI, VT. Rich, mesic soils of high-terrace riparian floodplains and deciduous forests, often associated with Acer saccharum, Tilia americana, and/or Fraxinus americana. 6. Elymus repens (L.) Gould E creeping wild-rye. Agropyron repens (L.) Gould; Elytrigia repens (L.) Desv. ex B.D. Jackson; Triticum repens L. • CT, MA, ME, NH, RI, VT; thoughout. Fields, roadsides, areas of cultivation, sometimes colonizing native habitats such as gravel and cobble river shores. 7. Elymus riparius Wieg. N Fig. 222 eastern riverbank wild-rye. Elymus canadensis L. var. riparius (Wieg.) Boivin • CT, MA, ME, NH, RI, VT. Riparian forests, river banks. 7 × 11. This very rare wild-rye hybrid is known from VT. It is largely sterile (as shown by its abortive pollen) and on the surface appears to be a less robust, narrower-leaved form of Elymus wiegandii (i.e., it often has the conspicuously drooping spikes of that species). However, the hybrid plants show spikelets with mostly 2 or 3 florets, indurate and usually unstriate glume bases, and paleas mostly 10–10.5 mm long (with nodding or arching spikes, spikelets with mostly 2 florets, indurate and unstriate glume bases, and paleas 6–9 mm long in E. riparius vs. conspicuously drooping spikes, spikelets with mostly 4–6 florets, slightly indurate and somewhat striate glume bases, and paleas 9–14 mm long in E. wiegandii).

Fig. 221  Inflorescence of Elymus macgregorii.

8. Elymus trachycaulus (Link) Gould ex Shinners N slender wild-rye.  1a. Agropyron caninum (L.) Beauv. var. pubescens Scribn. & J.G. Sm.; A. trachycaulum (Link) Malte var. glaucum (Pease & Moore) Malte; 1b. Agropyron trachycaulum (Link) Malte ex H.F. Lewis; A. trachycaulum (Link) Malte ex H.F. Lewis var. majus (Vasey) Fern.; A. trachycaulum (Link) Malte ex H.F. Lewis var. novae-angliae (Scribn.) Fern.; Triticum trachycaulum L. • CT, MA, ME, NH, RI, VT. Gravelly and ledgy river shores, cliffs and talus fields, open areas near the coast, peatlands, pond shores, roadsides, fields, swamp edges. 1a. Lemma awns 5–17 (–24) mm long, straight or curving outward from the axis of the spike; glume awns 0–10 mm long . . . . . . . . 8a. E. trachycaulus ssp. glaucus (Pease & Moore) Cody 1b. Lemma awns wanting or represented by a short awn-tip (rarely as long as 5 mm and straight); glume awns absent or represented by a short awn-tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8b. E. trachycaulus ssp. trachycaulus Subspecies glaucus is known from CT, MA, ME, NH, VT. Subspecies trachycaulus is known from CT, MA, ME, NH, RI, VT. Elymus subsecundus (synonym: E. trachycaulus var. unilateralis (Cassidy) Beetle) was reported from New England based on specimens with evident lemmas awns. These plants are, in fact, intermediate between E. trachycaulus and E. subsecundus and have been referred to by the epithet “glaucus ” by Fernald (1950b). Jozwik (1966) suggested that such plants with intermediate awn length and sometimes divergent awns are often hybrids between E. trachycaulus and E. subsecundus. If this hypothesis is confirmed, ssp. glaucus should be recognized at the specific level to ensure its name does not imply an incorrect evolutionary relationship. 9. Elymus villosus Muhl. ex Willd.

NC

downy wild-rye. 9a. Elymus canadensis L. var. villosus (Muhl. ex Willd.) Shinners; E. striatus Willd. var. villosus (Muhl. ex Willd.) Gray; 9b. Elymus arkansanus Scrib. & Ball • CT, MA, RI, VT. High-terrace floodplain forests, river banks, rich, rocky forests, open soils near the coast. 1a. Glumes and lemmas villous to hirsute . . . . . . . . . . . . . . . . . . . . . . 9a. E. villosus var. villosus 1b. Glumes and lemmas glabrous or scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9b. E. villosus var. arkansanus (Scribn. & Ball) J.J.N. Campb. Variety villosus is known from CT, MA, RI, VT. Variety arkansanus is known from CT, MA, VT and is much rarer than variety villosus. Both varieties are of conservation concern.

Fig. 222  Spikelet of Elymus riparius.

25 6  MONOCOTS

10. Elymus virginicus L. N Fig. 223, 224 common eastern wild-rye.  10a. Elymus hirsutiglumis Scribn.; E. virginicus L. var. hirsutiglumis (Scribn.) A.S. Hitchc.;  10b. Elymus striatus Willd.; 10c. Elymus halophilus Bickn.; 10d. Elymus jejunus (Ramaley) Rydb. • CT, MA, ME, NH, RI, VT. Riparian forests, river banks, rocky forests, cliff bases, coastal beaches, upper margin of saline marshes. 1a. Lemmas pubescent [Fig. 223]; spikes glaucous; ligules and auricles usually absent; plants flowering July through August . . . . . 10a. E. virginicus var. intermedius (Vasey ex Gray) Bush 1b. Lemmas glabrous or minutely scabrous; spikes green or glaucous; ligules and auricles often present; plants flowering prior to August 2a. Spikes often partly included in the upper leaf sheath [Fig. 224]; glumes 1–2.3 mm wide, strongly indurate and bowed out in the basal (1.5–) 2–4 mm; plants usually green to yellowbrown in life; nodes often covered by overlapping leaf sheaths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10b. E. virginicus var. virginicus Fig. 223  Spikelet of Elymus virginicus var. intermedius.

2b. Spikes usually exserted from the upper leaf sheath; glumes 0.7–1.5 (–1.8) mm wide, moderately indurate and bowed out in the basal 1–2 mm; plants usually glaucous, sometimes with red-brown coloration, in life; nodes usually exposed 3a. Reproductive stems mostly 3–8 dm tall, with 4–6 nodes; leaf blades 2–9 mm wide, becoming involute; spikes 3.5–11 cm tall, strongly glaucous; glumes indurate in the basal 1–2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10c. E. virginicus var. halophilus (Bickn.) Wieg. 3b. Reproductive stems mostly 7–10 dm tall, with 6–8 nodes; leaf blades 3–15 mm wide, flat; spikes 4–20 cm tall, pale green to glaucous; glumes indurate only in the basal 1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10d. E. virginicus var. jejunus (Ramaley) Bush Variety intermedius is known from CT, MA, ME, NH, RI, VT. Variety virginicus is known from CT, MA, ME, NH, RI, VT. Variety halophilus is known from CT, MA, ME, NH, RI. Variety jejunus is known from CT, MA, ME, NH. Varieties intermedius and jejunus are less common than the other varieties. 11. Elymus wiegandii Fern. N Wiegand’s wild-rye. Elymus canadensis L. var. wiegandii (Fern.) Bowden • CT, MA, ME, NH, VT. Riparian forests, usually on rich soils of high terraces.

Eragrostis Eragrostis hirsuta (Michx.) Nees was reported from MA by Hitchcock and Chase (1950), but specimens are unknown. Reference: Peterson (2003a). Fig. 224  Inflorescence of Elymus virginicus var. virginicus.

1a. Plants extensively creeping by stolons, rooting at the nodes, forming mats; inflorescence 1–3.5 (–6) cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. hypnoides 1b. Plants without stolons, the stems erect or ascending, sometimes decumbent at the base, not rooting at the nodes; inflorescence (3–) 4–40 (–60) cm tall 2a. Leaf blades with elevated glands or glandular pits along the margin [Fig. 225] 3a. Lemmas 1.5–2 mm long, lacking glandular pits along the keel (rarely with 1 or 2 obscure glands); sheaths pubescent on the margin and often near the apex; spikelets 4–7 (–11) mm long, with 7–12 (–20) florets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. minor 3b. Lemmas 2–2.8 mm long, with 1–3 glandular pits along the keel; sheaths glabrous along the margin, sometimes pubescent near the apex; spikelets 6–20 mm long, with 10–40 florets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. cilianensis 2b. Leaf blades without glands along the margin 4a. Plants perennial, with basal innovations, sometimes with short, knobby rhizomes

Poac e a e   257

5a. Plants with short, knobby rhizomes; inflorescence usually more than 40% of the total plant height; anthers 0.3–0.5 mm long; paleas falling with the lemmas during spikelet disarticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. spectabilis 5b. Plants without rhizomes; inflorescence usually less than 40% of the total plant height; anthers 0.5–1.2 mm long; paleas usually not falling with the lemmas and persistent on the rachilla during spikelet disarticulation 6a. Lemmas 1.6–2.2 mm long, usually rounded on the abaxial surface and hardly keeled, with inconspicuous lateral veins; upper glume 1.3–2 mm long; caryopsis red-brown, opaque; ligules 0.2–0.4 mm long . . . . . . . . . . . . . . . . . . . . . E. intermedia 6b. Lemmas 1.8–3 mm long, keeled on the abaxial surface, with conspicuous lateral veins; upper glume 2–3 mm long; caryopsis light brown, transluscent; ligules 0.6–1.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. curvula 4b. Plants annual, from fibrous roots, without basal innovations or rhizomes 7a. Panicle large and diffuse, (10–) 15–45 (–55) cm tall and usually more than 50% of the total plant height; pedicels (4–) 5–25 mm long; spikelets with 2–5 (–7) florets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. capillaris 7b. Panicle often smaller, 4–20 (–28) cm tall (up to 40 cm in E. mexicana) and usually less than 50% of the total plant height; pedicels 1–7 (–10) mm long; spikelets with 5–22 florets (with only 3–6 florets in E. frankii) 8a. Spikelets with 3–6 florets, narrow-ovate to ovate in broad outline at maturity; plants often with glandular pits on the sheaths, below the nodes of the stem, on the rachis of the inflorescence, and/or on the panicle branches . . . . . . . . E. frankii 8b. Spikelets with 5–22 florets, narrow-lanceolate to broad-lanceolate in broad outline at maturity; plants lacking glandular pits (rarely with a few scattered glandular pits on the stems in E. pilosa) 9a. Caryopsis with a shallow to deep, longitudinal groove running the entire length; leaf blades (2–) 3–7 (–9) mm wide . . . . . . . . . . . . . . . . . . . . . . . E. mexicana 9b. Caryopsis without a longitudinal groove; leaf blades 1–3 (–4.5) mm wide 10a. Lower glume 0.3–0.6 (–0.8) mm long, less than ½ as long as the lowest lemma [Fig. 227]; branches of the panicle usually whorled at the 2 lowest nodes; paleas mostly deciduous with the lemmas and not persistent on the rachilla; lemmas with inconspicuous, lateral veins . . . . . . . . . . . . . . . . E. pilosa 10b. Lower glume 0.5–1.1 mm long, ½ or more as long as the lowest lemma [Fig. 226]; branches of the panicle usually solitary or paired at the lowest 2 nodes; paleas persistent on the rachilla after the lemmas have fallen; lemmas with conspicuous, lateral veins . . . . . . . . . . . . . . . . . . . . . E. pectinacea 1. Eragrostis capillaris (L.) Nees

NC

lace lovegrass. CT, MA, ME, NH, ri, vT. Open balds and rocky ridges, sandy roadsides, river banks, and waste lots, railroads and railroad yards. Some stations for this species are very likely the result of unintentional introduction. However, given that disturbed, sandy areas mimic this species natural habitat, it is difficult to know for sure which stations are native and which are introduced. 2. Eragrostis cilianensis (All.) Lut. ex Janchen E Fig. 225 stinking lovegrass. Eragrostis major Host; E. megastachya (Koel.) Link; Poa cilianensis All. • CT, MA, ME, NH, RI, VT. Roadsides, areas of cultivation, railroads, disturbed soil. 3. Eragrostis curvula (Schrad.) Nees E weeping lovegrass. MA, RI. Waste areas, disturbed soil. 4. Eragrostis frankii C.A. Mey. ex Steud. n

Fig. 225  Portion of leaf blade of Eragrostis cilianensis showing elevated glands along margin.

25 8  MONOCOTS

sandbar lovegrass. Eragrostis frankii C.A. Mey. ex Steud. var. brevipes Fassett • CT, MA, ME, NH, VT. Sandy riverbanks, sand and silt accretion bars, roadsides, disturbed soil. Some occurrences of this plant are likely the result of unintentional human introduction. The occurrence in Penobscot County, ME, is likely introduced. 5. Eragrostis hypnoides (Lam.) B.S.P. N teel lovegrass. Poa hypnoides Lam. • CT, MA, ME, NH, VT; very rare in northern portion of range. Sand, silt, and mud shorelines. 6. Eragrostis intermedia Hitchc. E plains lovegrass. MA, ME. Dry, disturbed soil. 7. Eragrostis mexicana (Hornem.) Link ssp. virescens (J. Presl) S.D. Koch & Sánchez E Mexican lovegrass. Eragrostis orcuttiana Vasey; E. virescens J. Presl • MA, ME. Wool waste, disturbed soil. 8. Eragrostis minor Host E little lovegrass. Eragrostis eragrostis (L.) Beauv.; E. poaeoides Beauv. ex Roemer & J.A. Schultes • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, disturbed soil. Fig. 226  Spikelet of Eragrostis pectinacea with persistent paleas in apical half.

9. Eragrostis pectinacea (Michx.) Nees ex Steud. var. pectinacea N Fig. 226 tufted lovegrass. Eragrostis diffusa Buckl.; Poa pectinacea Michx. • CT, MA, ME, NH, RI, VT. Roadsides, fields, railroads, sand and silt bars, disturbed soil. 10. Eragrostis pilosa (L.) Beauv. var. pilosa E Fig. 227 India lovegrass. Eragrostis multicaulis Steud.; Poa pilosa L. • CT, MA, ME, NH, RI, VT. Roadsides, sidewalks, yards, railroads, disturbed sandy and gravelly areas. 11. Eragrostis spectabilis (Pursh) Steud. N purple lovegrass. Eragrostis spectabilis var. sparsihirsuta Farw.; Poa spectabilis Pursh • CT, MA, ME, NH, RI, VT. Dry-mesic to xeric soil of roadsides, fields, railroads, and coastal sands.

Eremochloa Fig. 227  Spikelet of Eragrostis pilosa.

1. Eremochloa ophiuroides (Munro) Hack. E centipede grass. Ischaemum ophiuroides Munro • MA. Roadsides, fields, disturbed soil.

Festuca Lemma awns tend to be longest on the distal florets of a given spikelet. Leaf sheaths readily split in age. Therefore, when assessing closure of the sheath, only new leaves should be examined. Reference: Darbyshire and Pavlick (2007). 1a. Leaf blades flat, (3–) 4–10 mm wide; apex of ovary pubescent; panicles with eventually wide-spreading or even somewhat deflexed branches with triangular cross-section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. subverticillata 1b. Leaf blades involute or rarely flat, 0.2–3 mm wide; apex of ovary glabrous; panicle branches erect to ascending (rarely spreading-ascending), without triangular cross-section 2a. Most of the flowers proliferous, replaced by vegetative bulbils [Fig. 229]; anthers rarely present, usually abortive . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. prolifera 2b. Flowers fertile, not proliferous; anthers present 3a. Basal sheaths red-brown, usually minutely pubescent in the apical half, shredding into persistent fibers [Fig. 230]; sheaths of new leaves closed ¾ or more of their length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. rubra 3b. Basal sheaths white-brown to light brown, usually glabrous, not shredding [Fig. 228]; sheaths of new leaves not closed or closed up to or slightly more than ½ their length

Poac e a e   25 9

4a. Lemmas awnless or with a tiny mucro up to 0.4 mm long; leaf blades of vegetative shoots with 1 rib . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. filiformis 4b. Lemmas with awns 0.5–2.5 mm long; leaf blades of vegetative shoots with 1–7 ribs (the ribs often obscure in F. ovina) 5a. Anthers (0.5–) 0.7–1.1 (–1.3) mm long; spikelets with 2–4 (–6) florets; reproductive stems 8–35 cm tall; native grass of alpine habitats . . . . . . . . . . . . . . . . F. brachyphylla 5b. Anthers (1.8–) 2–3.4 mm long; spikelets with 3–7 (–8) florets; reproductive stems (10–) 20–75 cm tall; European species introduced as turf grasses and soil stabilizers 6a. Anthers (1.8–) 2–2.5 mm long; leaf blades 0.3–0.7 (–1.2) mm wide, with 1–3 obscure ribs; lemmas 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. ovina 6b. Anthers 2.3–3.4 mm long; leaf blades (0.5–) 0.8–1.2 mm wide, with 5–7 ribs; lemmas 3.8–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. trachyphylla 1. Festuca brachyphylla J.A. Schultes & J.A. Schultes f. ssp. brachyphylla

NC

alpine fescue. Festuca ovina L. var. alpina (Gaudin) W.D.J. Koch; F. ovina L. var. brachyphylla (J.A. Schultes & J.A. Schultes f.) Piper ex A.S. Hitchc.; F. ovina L. ssp. brevifolia (S. Wats.) Hack., pro parte • VT; also reported from NH by Löve and Löve (1966), but specimens are unknown. Alpine cliffs and ravines. Löve and Löve (1966) reported this species from near the summit of Mount Washington, Coos County, NH. Based on their report, it is unsure if they observed reproductive stems or relied solely on chromosome counts for identification. The voucher specimen was not located at the stated repository and has yet to be discovered.

Fig. 228  Pale, non-shredding basal sheaths of Festuca ovina.

2. Festuca filiformis Pourret E fine-leaved sheep fescue. Festuca capillata Lam.; F. ovina L. var. capillata (Lam.) Alef.; F. tenuifolia Sibthorp • CT, MA, ME, NH, RI, VT. Sandy soils of roadsides, fields, grasslands, and open woodlands. 3. Festuca ovina L. E Fig. 228 sheep fescue. CT, MA, ME, NH, RI, VT. Lawns, fields, roadsides, seeded areas. Festuca ovina was used introduced as a turf grass, but it not widely used anymore. 4. Festuca prolifera (Piper) Fern.

N C Fig. 229

proliferous fescue. Festuca rubra L. var. prolifera (Piper) Piper • ME, NH. Alpine cliffs, seeps, and ravines. 5. Festuca rubra L. n Fig. 230 red fescue. 5a. Festuca diffusa Dumort.; F. heteromalla Pourret; F. multiflora Hoffmann; F. rubra L. var. multiflora (Hoffmann) Aschers. & Graebn.; 5b. Festuca rubra L. ssp. densiuscula Hack. ex Piper; F. rubra L. ssp. juncea (Hack.) Soó; F. rubra L. var. juncea (Hack.) Richter;  5c. Festuca duriuscula L.; F. ovina L. var. rubra (L.) Sm.;  5d. Festuca nigrescens Lam.; F. rubra L. var. commutata Gaudin • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, disturbed soil, coastal beaches and headlands.

Fig. 229  Inflorescence of Festuca prolifera showing proliferous spikelets.

1a. Leaf blades of vegetative shoots 0.8–3 mm wide, flat or loosely folded; anthers 3.5–4.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5a. F. rubra ssp. fallax (Thuill.) Nyman 1b. Leaf blades of vegetative shoots 0.3–1 mm wide, tightly folded (rarely up to 2 mm wide and flat in F. rubra ssp. rubra); anthers 1.8–3.5 mm long 2a. Leaf blades stiff, wiry, and strongly glaucous; plants of Atlantic coast beaches, headlands, and near coast habitats . . . . . . . . . . . . . . 5b. F. rubra ssp. pruinosa (Hack.) Piper 2b. Leaf blades relatively soft, not wiry, green or slightly glaucous; plants primarily of human-disturbed and cultivated areas 3a. Anthers 2.4–3.5 mm long; plants rhizomatous, the stems loosely cespitose; upper glumes 4–6.4 mm long; lemmas (4–) 6–7.5 (–8) mm long . . . . . . . . 5c. F. rubra ssp. rubra

Fig. 230  Red-brown, shredding basal sheaths of Festuca rubra.

26 0  MO NOCOTS

3b. Anthers 1.8–2.2 (–3) mm long; plants without rhizomes, the stems densely cespitose; upper glumes 3.5–5 mm long; lemmas 4.5–6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5d. F. rubra ssp. commutata Gaudin Subspecies fallax is known from MA, ME, NH, VT. Subspecies pruinosa is known from MA, ME, NH. Subspecies rubra is known from CT, MA, ME, NH, RI, VT. Subspecies commutata is known from CT, MA, ME, NH, RI, VT. Festuca rubra ssp. pruinosa represents plants of the F. rubra complex that are native to North America, whereas the other subspecies are non-native. This taxon is found on coastal beaches and headlands (as opposed to more human-modified habitats). It is a cespitose species with upper glume and lemma measurements comparable to subspecies commutata, except the foliage is different and the anthers are 2.3–3.2 mm long (see key above). Subspecies pruinosa has moderate sized anthers, similar to ssp. rubra; however, it commonly has smaller floral parts compared with ssp. rubra (lower glumes 2.2–3.2 (–4.5) mm long and lemmas 4.5–6 (–6.5) mm long in ssp. pruinosa vs. lower glumes 3–4.5 mm long and lemmas (4–) 6–7.5 (–8) mm long in ssp. rubra). 6. Festuca subverticillata (Pers.) Alexeev N nodding fescue. Festuca obtusa Biehler • CT, MA, ME, NH, RI, VT. Rich, mesic forests, often associated with colluvial deposits or cicumneutral to basic bedrock. 7. Festuca trachyphylla (Hack.) Krajina E hard fescue. Festuca ovina L. var. duriuscula, auct. non (L.) W.D.J. Koch • CT, MA, ME, NH, RI, VT. Fields, lawns, seeded soils.

Gastridium 1. Gastridium phleoides (Nees & Meyen) C.E. Hubbard E nit grass. Gastridium ventricosum, auct. non (Gouan) Schinz & Thellung • MA, ME. Wool waste, rubbish heaps, disturbed soil.

Glyceria The genera Glyceria, Puccinellia, and Torreyochloa are often confused because of superficial similarities in morphology and occurrence in hydric habitats. Among these three genera, only Glyceria has closed leaf sheaths and 1-veined upper glumes (the other two genera have open leaf sheaths and 3-veined upper glumes). Only Puccinellia is tolerant of saline conditions and has inconspicuous veins on the lemmas (the other two genera are restricted to freshwater habitats and generally have conspicuous veins on the lemmas). Reference: Barkworth and Anderton (2007). 1a. Spikelets 10–40 mm long, with 7–16 florets [Fig. 231]; internodes of the rachilla 1–4 mm long; stamens 3 per floret; sheaths conspicuously compressed 2a. Spikelets 20–40 mm long; lemmas 7–8.5 mm long, acute at the apex [Fig. 231]; paleas exceeding lemmas by 1.5–3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. acutiflora 2b. Spikelets 10–20 mm long; lemmas 3.1–7 mm long, truncate to obtuse at the apex; paleas equaling or exceeding lemmas by no more than 1 mm 3a. Lemmas glabrous between the veins (infrequently minutely scabrous), 3.1–3.9 mm long, the veins terminating ca. 0.2 mm from the lemma apex; anthers 0.6–1 mm long; internodes of rachilla 1–1.3 mm long; mid-stem leaf blades minutely, but densely, papillose on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. borealis 3b. Lemmas scabrous between the veins, 3.7–7 mm long, the veins extending to within 0.1 mm of lemma apex; anthers 1–3 mm long; internodes of rachilla 1.1–2.5 mm long; mid-stem leaf blades smooth or scabridulous (sometimes minutely papillose in G. septentrionalis)

P oac e a e   2 6 1

4a. Lemmas 3.7–5.3 mm long; internodes of rachilla 1.1–1.8 mm long; anthers 1–1.7 mm long; adaxial surface of the mid-stem leaf blades sometimes minutely papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. septentrionalis 4b. Lemmas 5.4–7 mm long; internodes of rachilla 1.9–2.5 mm long; anthers 2–3 mm long; adaxial surface of the mid-stem leaf blades smooth . . . . . . . . . . . . . . . G. fluitans 1b. Spikelets 2–8 mm long, with 3–10 florets [Fig. 232]; internodes of the rachilla shorter than 1 mm; stamens 2 or 3 per floret; sheaths not conspicuously compressed 5a. Panicle dense, with strictly ascending to erect branches; ligules 0.5–0.9 mm long 6a. Inflorescence arching to nodding, narrow-cylindric, with 1–3 branches at each node, mostly 15–25 cm long; internodes of the inflorescence 2–7 cm long; spikelets 3.5–5 mm long; new stem leaves numbering 3–6 per stem . . . . . . . . . . . . . . . . . . . . . . . . G. melicaria 6b. Inflorescence erect, ovoid to obloid [Fig. 233], with 3–8 branches at each node, mostly 5–15 cm long; internodes of the inflorescence up to 2.5 cm long, commonly less than 2 cm; spikelets 4–7 mm long; new stem leaves numbering 7–9 per stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. obtusa 5b. Panicle open, the branches ascending to drooping; ligules 1–6 mm long 7a. Spikelets (2.5–) 3–5 mm wide; veins of the lemma visible but not raised; palea less than 2 times as long as wide 8a. Lemmas (2.4–) 2.9–4 mm long, acute at the apex, projecting beyond the paleas at least 0.5 mm; spikelets 5–8 mm long, with 4–10 florets; upper glume acute at the apex [Fig. 232] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. canadensis 8b. Lemmas 1.8–2.5 (–2.9) mm long, obtuse at the apex, not, or scarcely, exceeding the associated paleas; spikelets 4–5 mm long, with 2–5 florets; upper glumes rounded at the apex (rarely varying to acute) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. laxa 7b. Spikelets 1.5–2.5 mm wide; veins of the lemma conspicuous and raised; palea 3–5 times as long as wide 9a. Spikelets 2–4.5 mm long; glumes obtuse, the first and second 0.5–1 mm and 0.8–1.3 mm long, respectively; lemmas firm, caducous; stamens 2 per floret; new cauline leaves 5–10 per stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. striata 9b. Spikelets 4–6.5 mm long; glumes acute, the first and second 1.2–1.9 mm and 1.5–2.4 mm long, respectively; lemmas membranaceous, deciduous; stamens mostly 3 per floret; new cauline leaves 3–6 per stem 10a. Leaves with blades 4.5–12 (–15) mm wide and usually smooth sheaths; second glume 1.5–2.7 mm long; apex of lemma ± flat; anthers 0.5–1.2 mm long . . . G. grandis

Fig. 231  Spikelet of Glyceria acutiflora.

10b. Leaves with blades (6–) 8–18 (–20) mm wide and minutely scabrous sheaths; second glume (2.5–) 3–4 mm long; apex of lemma cucullate; anthers (1–) 1.2–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. maxima 1. Glyceria acutiflora Torr. N Fig. 231 sharp-scaled manna grass. Panicularia acutiflora (Torr.) Kuntze • CT, MA, ME, NH, RI, VT; rare in and generally restricted to the southern portion of the northern states. Shallow water of pools, lakes, and streams. 2. Glyceria borealis (Nash) Batchelder N northern manna grass. Panicularia borealis Nash • CT, MA, ME, NH, RI, VT. Shallow water of lakes and streams, saturated, often muddy, soils of pools and beaver flowages. 3. Glyceria canadensis (Michx.) Trin. N Fig. 232 rattlesnake manna grass. Panicularia canadensis (Michx.) Kuntze • CT, MA, ME, NH, RI, VT; throughout. Graminoid marshes, open swamps and stream courses, shorelines. 4. Glyceria fluitans (L.) R. Br. E

Fig. 232  Spikelet of Glyceria canadensis.

262  MONOCOTS

water manna grass. Festuca fluitans L.; Panicularia fluitans (L.) Kuntze • MA. Shallow water of pools, lakes, and streams. 5. Glyceria grandis S. Wats. var. grandis N American manna grass. Glyceria maxima (Hartman) Holmb. ssp. grandis (S. Wats.) Hultén; G. maxima (Hartman) Holmb. var. americana (Torr.) Boivin; Panicularia grandis (S. Wats.) Nash • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Shallow water of pools, lakes, and streams, ditches, wetland edges. 6. Glyceria laxa (Scribn.) Scribn. N flaccid manna grass. Panicularia laxa Scribn. • CT, MA, ME, NH, VT. Graminoid marshes, open swamps and stream courses, shorelines. Glyceria laxa is often treated as a hybrid of G. canadensis and G. striata. Though this grass may have originated through the specified cross, it often produces normal anthers and well-formed fruits. Therefore, it appears best to treat this plant as an orthospecies. 7. Glyceria maxima (Hartman) Holmb. E reed manna grass. Glyceria spectabilis Mert. & Koch; Molinia maxima Hartman • CT, MA. Ditches and other wet, disturbed places, muddy stream shores. 8. Glyceria melicaria (Michx.) F.T. Hubbard N northeastern manna grass. Panicularia melicaria (Michx.) A.S. Hitchc. • CT, MA, ME, NH, RI, VT. Graminoid marshes, open swamps, shorelines. 9. Glyceria obtusa (Muhl.) Trin. N Fig. 233 Atlantic manna grass. Panicularia obtusa (Muhl.) Kuntze • CT, MA, ME, NH, RI; primarily coastal counties. Open swamps, stream courses, shorelines. 10. Glyceria septentrionalis A.S. Hitchc. var. septentrionalis N floating manna grass. Panicularia septentrionalis (A.S. Hitchc.) Bickn. • CT, MA, ME, NH, RI. Shallow water of lakes and streams, saturated, often muddy, soils of pools and beaver flowages. Fig. 233  Inflorescence of Glyceria obtusa.

11. Glyceria striata (Lam.) A.S. Hitchc. N fowl manna grass. Glyceria striata (Lam.) A.S. Hitchc. var. stricta (Scribn.) Fern.; Panicularia nervata (Willd.) Kuntze • CT, MA, ME, NH, RI, VT; throughout. Graminoid marshes, open swamps, wetland borders, shorelines.

Graphephorum 1. Graphephorum melicoides (Michx.) Desv.

NC

graphephorum. Aira melicoides Michx.; Trisetum melicoides (Michx.) Vasey ex Scribn.; T. melicoides (Michx.) Vasey ex Scribn. var. majus (Gray) A.S. Hitchc. • ME, NH, VT. Ice-scoured river shores, river shore ledges, inland cliffs, usually in regions of high-pH bedrock or till.

Holcus 1a. Lower internodes of stem softly villous; awn of upper lemma 1–2 mm long, hooked, scarcely, if at all, exserted from the glumes [Fig. 234]; plants without rhizomes . . . . H. lanatus 1b. Lower internodes of stem glabrous; awn of upper lemma 3–5 mm long, bent but not hooked, well exserted beyond the glumes; plants with slender rhizomes . . . . . . . . . . . H. mollis 1. Holcus lanatus L. E Fig. 234 common velvet grass. Nothoholcus lanatus (L.) Nash • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil. Fig. 234  Spikelet of Holcus lanatus.

2. Holcus mollis L. ssp. mollis E creeping velvet grass. MA, VT. Fields, roadsides, waste areas.

P oac e a e   2 63

Hordeum Hordeum marinum Huds. ssp. gussoneanum (Parl.) Thellung was reported from MA by Hitchcock and Chase (1950), but specimens are unknown. Reference: von Bothmer et al. (2007). 1a. Anthers 2–2.5 mm long; leaf blades 5–15 mm wide; rachis continuous and not disarticulating . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. vulgare 1b. Anthers 0.5–1.8 mm long; leaf blades 1.5–5 (–7) mm wide; rachis disarticulating 2a. Plants perennial; awns very slender 3a. Spikes arching or nodding [Fig. 235]; glumes 25–150 mm long including the awns; glume awns strongly bent outward and wide-ascending to spreading at maturity; fertile lemmas with an awn 10–60 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. jubatum 3b. Spikes erect [Fig. 236]; glumes 7–20 mm long including the awns; glume awns not or scarcely bent outward, erect to ascending; fertile lemmas with an awn 5–10 (–20) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. brachyantherum 2b. Plants annual; awns stout 4a. Glumes of the central spikelet of each triad with ciliate margins; auricles present at the summit of most leaf sheaths, well-developed, up to 8 mm long; body of fertile lemmas (6–) 7–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. murinum 4b. Glumes of the central spikelet of each triad with scabrous margins; auricles absent or poorly developed and very short; body of fertile lemmas 5–7 mm long . . . . . . . H. pusillum 1. Hordeum brachyantherum Nevski ssp. brachyantherum E meadow barley. Critesion brachyantherum (Nevski) Barkworth & D.R. Dewey; Hordeum boreale Scrib. & J.G. Sm.; H. nodosum L., pro parte; H. nodosum L. var. boreale (Scrib. & J.G. Sm.) A.S. Hitchc. • MA, NH; also reported from ME by Campbell et al. (1995), but specimens are unknown. Railroads, disturbed soil.

Fig. 235  Inflorescence of Hordeum jubatum.

2. Hordeum jubatum L. ssp. jubatum N Fig. 235 fox-tail barley. Critesion jubatum (L.) Nevski • CT, MA, ME, NH, RI, VT. Coastal beaches, upper edge of saline marshes, roadsides, railroads. 3. Hordeum murinum L. ssp. leporinum (Link) Arcang. E Fig. 236 mouse barley. Critesion murinum (L.) A. Löve var. leporinum (Link) A. Löve; Hordeum leporinum Link • CT, MA, ME. Roadsides, fields, disturbed soil. 4. Hordeum pusillum Nutt. E little barley. Critesion pusillum (Nutt.) A. Löve • CT, MA, ME, NH. Fields, pastures, roadsides, borders of marshes. 5. Hordeum vulgare L. E common barley. Hordeum distichon L.; H. hexastichon L.; H. vulgare L. var. trifurcatum (Schlecht.) Alef. • CT, MA, ME, NH, RI, VT. Roadsides, fields, disturbed soil. Two forms of this cultivated grass can be found—those with all 3 spikelets of a triad sessile and fertile, called 6-row barley, and those with only the central spikelet of a triad fertile, the lateral spikelets sterile and pedicelled, called 2-row barley.

Koeleria 1. Koeleria macrantha (Ledeb.) J.A. Schultes E prairie Koeler’s grass. Koeleria cristata, auct. non Pers.; K. pyramidata, auct. non (Lam.) Beauv. • ME, VT. Fields, areas of cultivation, disturbed soil.

Fig. 236  Inflorescence of Hordeum murinum ssp. leporinum.

26 4  MO NOCOTS

Leersia 1a. Branches of the panicle 2 or more at some of the nodes; florets with 3 stamens; lemmas 4–7.5 mm long; principal leaf blades 15–30 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . L. oryzoides 1b. Branches of the panicle 1 per node; florets with 2 stamens; lemmas 2.9–4.1 mm long; principal leaf blades 5–20 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. virginica 1. Leersia oryzoides (L.) Sw. N Fig. 237 rice cut grass. Homalocenchrus oryzoides (L.) Pollich; Phalaris oryzoides L. • CT, MA, ME, NH, RI, VT. Shorelines, low areas in fields, open swamps, ditches, pools, beaver flowages. 2. Leersia virginica Willd. N Fig. 237  Node of Leersia oryzoides.

white cut grass. Homalocenchrus virginicus (Willd.) Britt.; Leersia virginica Willd. var. ovata (Poir.) Fern. • CT, MA, ME, NH, RI, VT. Riparian forests, river banks, silt and mud bars, often in more shade than Leersia oryzoides.

Leptochloa Reference: Snow (2003). 1a. Leaf sheaths sparsely to densely pubescent with pustulose-based hairs; ligule 0.6–3.2 mm long, truncate at apex; spikelets 2–4 mm long, with 2–5 (–6) florets . . . . . . . . . . . . . . L. panicea 1b. Leaf sheaths glabrous or scabrous, but not pustulose-pubescent; ligule 2–8 mm long, attenuate at apex and becoming lacerate at maturity; spikelets 5–12 mm long, with 6–20 florets [Fig. 238] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. fusca 1. Leptochloa fusca (L.) Kunth n Fig. 238 bearded sprangletop.  1a. Diplachne acuminata Nash; D. fascicularis (Lam.) Beauv.; D. maritima Bickn.; Leptochloa fascicularis (Lam.) Gray; L. fascicularis (Lam.) Gray var. maritima (Bickn.) Gleason; 1b. Diplachne uninervia (J. Presl) Parodi; Leptochloa uninervia (J. Presl) A.S. Hitchc. & Chase • CT, MA, ME, NH, RI, VT. Saline marshes, coastal beaches, railroads, disturbed ground.

Fig. 238  Spikelet of Leptochloa fusca ssp. fascicularis.

1a. Uppermost leaf usually partly enclosing the inflorescence in the sheath, the blade usually exceeding the inflorescence; mature lemmas gray-white, usually with a dark spot in the basal portion; reproductive stems prostrate to erect . . . . 1a. L. fusca ssp. fascicularis (Lam.) N. Snow 1b. Uppermost leaf usually not partly enclosing the inflorescence (i.e., the inflorescence completely exserted), the blade usually not exceeding the inflorescence; mature lemmas light brown, dark-green, or gray, usually lacking a dark, basal spot; reproductive stems erect to ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. L. fusca ssp. uninervia (J. Presl) N. Snow Subspecies fascicularis is native and known from CT, MA, NH, RI, VT. Subspecies uninervia is non-native and known from MA. Reports of ssp. uninervia from ME (e.g., Campbell et al. 1995) are based on collections taken from cultivated plants— Sep 1896, Harvey & Harvey s.n. (MAINE!). Coastal plants with long awns have been segregated under the epithet “maritima”. Given that awn length and salinity tolerance are known to vary within this species, these plants are best referred to as ecological variants. 2. Leptochloa panicea (Retz.) Ohwi ssp. mucronata (Michx.) Nowack E needle sprangletop. Leptochloa filiformis (Lam.) Beauv. var. attenuata (Nutt.) Steyermark & Kucera; L. mucronata (Michx.) Kunth • MA. Fields, roadsides, gardens.

Leymus 1. Leymus mollis (Trin.) Hara var. mollis N American lyme grass. Elymus arenarius L. var. villosus E. Mey.; E. capitatus Scrib.; E. mollis Trin.; Leymus arenarius (L.) Hochst. ssp. mollis (Trin.) Tzvelev • MA, ME, NH. Atlantic coast beaches.

P oac e a e   2 6 5

Lolium Reference: Terrell (2007). 1a. Glume (5.5–) 7–28 mm long, the body 0.75–1.5 times as tall as the column of florets; plants annual, with 2- to 10-flowered spikelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. temulentum 1b. Glume 3.5–15 mm long, the body 0.25–0.75 times as tall as the column of florets [Fig. 239]; plants annual or perennial, with (2–) 5- to 22-flowered spikelets (when annual, with 10 or more florets per spikelet) 2a. Rachis of the inflorescence rough on the surface opposite the spikelet; spikelets with (10–) 11–22 florets; apex of the glume of upper spikelets not reaching the tip of the lowest lemma on the same side; lemmas of the upper florets with a conspicuous awn to 10 mm (rarely absent); leaf blades rolled in young shoots, (2–) 3–8 (–13) mm wide at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. multiflorum 2b. Rachis of the inflorescence smooth on the surface opposite the spikelet, rough only on the angles; spikelets with mostly (2–) 5–10 florets [Fig. 239]; apex of the glume of upper spikelets more or less extending to the tip of the lowest lemma on the same side; lemmas with relatively short awns (awns sometimes absent or rarely prolonged to 8 mm); leaf blades folded in young shoots, (1–) 2–4 (–6) mm wide at maturity . . . . . . . . . . . . . L. perenne 1. Lolium multiflorum Lam. E Italian rye grass. Lolium multiflorum Lam. ssp. italicum (A. Braun) Volk. ex Schinz & R. Keller; L. perenne L. var. aristatum Willd.; L. perenne L. ssp. multiflorum (Lam.) Husnot; L. perenne L. var. multiflorum (Lam.) Parnell • CT, MA, ME, NH, VT. Fields, roadsides, seeded areas. 2. Lolium perenne L. E Fig. 239 perennial rye grass. Lolium multiflorum Lam. var. ramosum Guss. ex Arcang.; L. perenne L. var. cristatum Pers. ex B.D. Jackson • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, seeded areas. 3. Lolium temulentum L. ssp. temulentum E poison darnel. Lolium arvense With.; L. temulentum L. var. arvense (With.) Lilja • CT, MA, ME, VT. Areas of cultivation, disturbed soil.

Mibora

Fig. 239  Inflorescence of Lolium perenne.

1. Mibora minima (L.) Desv. E early sand grass. Agrostis minima L. • MA. Waste heaps, around nurseries.

Microstegium 1. Microstegium vimineum (Trin.) A. Camus E Fig. 240 Japanese stiltgrass. Andropogon vimineum Trin.; Eulalia viminea (Trin.) Kuntze • CT, MA, RI. Riparian forests and terraces, fields, roadsides.

Milium 1. Milium effusum L. ssp. cisatlanticum (Fern.) A. Haines N millet grass. Milium effusum L. var. cisatlanticum Fern. • CT, MA, ME, NH, VT; primarily in the western counties in southern New England. Mesic, deciduous forests, often at mid-elevations and frequently associated with circumneutral or basic bedrock.

Miscanthus Reference: Barkworth (2003b).

Fig. 240  Homogamous and homomorphic spikelet pair of Microstegium vimineum.

26 6  MO NOCOTS

1a. Awn of lemma elongate, 6–12 mm long, geniculate at base, conspicuously surpassing glumes; callus of lemma pubescent with hairs 3.5–12 mm long, up to twice as long as the associated spikelet; ligules 1–2 mm long; plants cespitose, with short rhizomes, forming large clumps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. sinensis 1b. Awn of lemma absent or short, not exceeding the glumes of intact spikelets; callus of lemma pubescent with hairs ca. 8–24 mm long, 2–4 times as long as the associated spikelet; ligules 0.5–1 mm long; plants long-rhizomatous, forming extensive colonies . . . . . . . . . M. sacchariflorus 1. Miscanthus sacchariflorus (Maxim.) Franch. E Amur silvergrass. CT, MA, ME. Fields, roadsides, disturbed soil. 2. Miscanthus sinensis Anderss. E Chinese silvergrass. Miscanthus sinensis Anderss. var. gracillimus A.S. Hitchc. • CT, MA, RI. Fields, roadsides, disturbed soil.

Molinia 1. Molinia caerulea (L.) Moench E purple moorgrass. Aira caerulea L. • CT, MA, ME, RI, VT. Fields, roadsides, and other open areas.

Muhlenbergia See Peterson et al. (2010) for rationale for an expanded Muhlenbergia (i.e., one that includes Lycurus and other genera). Reference: Peterson (2003b). 1a. Panicle dense and spike-like, the branches virtually absent; lower glume with 2 (–3) awns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. alopecuroides 1b. Panicle open and diffuse to slender and contracted, but the branches discernable; lower glume with a single awn 2a. Panicle open and diffuse, (2.5–) 3–30 (–41) cm wide; spikelets on long, thin pedicels (0.2–) 3–40 (–50) mm long 3a. Lemmas without an awn; leaf blades 0.8–2 mm wide; pedicels (0.2–) 3–7 mm long; callus of the lemma glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. uniflora 3b. Lemmas usually with an awn 2–13 (–18) mm long; leaf blades 2–4 mm wide; pedicels 10–40 (–50) mm long; callus of lemma shortly pubescent . . . . . . . . . . . . . . . . M. capillaris 2b. Panicle slender and contracted, 0.1–3 cm wide; spikelets subsessile or on short pedicels to 5 mm long 4a. Callus of the lemma glabrous; leaves 0.5–2 mm wide, involute; plants cespitose or mat-forming . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. richardsonis 4b. Callus of the lemma pubescent; leaves 1–7 mm wide, normally flat; plants with stems distributed along rhizomes or the stems sprawling, neither cespitose nor mat-forming 5a. Glumes (3–) 4–8 mm long including the awns, 1.3–2 times as long as the lemmas 6a. Internodes of the stem pubescent and dull, terete in cross-section, not keeled; anthers 0.8–1.5 mm long; ligules 0.2–0.6 mm long; mature caryopsis 1.0–1.6 mm long; margin of the lemma pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . M. glomerata 6b. Internodes of the stem glabrous, except sometimes immediately below the node, lustrous, elliptic in cross-section and keeled; anthers 0.4–0.8 mm long; ligules 0.6–1.5 (–1.7) mm long; mature caryopsis (1.2–) 1.4–2.3 mm long; margin of the lemma glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. racemosa 5b. Glumes 0.1–3.7 (–4) mm long including the awns, shorter than to equaling (up 1.2 times as long as) the lemmas

P oac e a e   2 67

7a. Glumes minute and veinless, the larger (i.e., upper) only 0.1–0.3 mm long; plants with sprawling stems that root at the lower nodes [Fig. 243], without rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. schreberi 7b. Glumes 1–3.7 (–4) mm long, 1- or 3-veined; plants not rooting at the nodes, with rhizomes [Fig. 244] 8a. Internodes of the stem glabrous and lustrous [Fig. 241] 9a. Glumes 1–2.5 mm long, broad and overlapping at base, narrowing above to a slender, acuminate apex (i.e., the margins sigmoid curved); plants usually few-branched above the base, axillary panicles, if present, generally well exserted from the leaf sheaths; panicles 2–5 (–8) mm wide; ligules 0.3–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. sobolifera 9b. Glumes 2–4 mm long, narrow and only slightly overlapping at base and gradually tapering throughout the length to a slender apex (i.e., the margins ± straight); plants usually much branched above the base with numerous axillary panicles that are partly included in the leaf sheaths; panicles (3–) 4–15 (–20) mm wide; ligules 0.7–1.7 mm long . . . . . . M. frondosa 8b. Internodes of the stem pubescent and dull [Fig. 242] 10a. Glumes definitely shorter than the lemmas, broad and overlapping at base, narrowing above to a slender, acuminate apex (i.e., the margins sigmoid curved) [Fig. 245]; anthers 1.1–2.2 mm long; caryopsis 2–2.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. tenuiflora 10b. Glumes nearly as long as or slightly longer than the lemmas, narrow and only slightly overlapping at base and gradually tapering throughout the length to a slender apex (i.e., the margins ±straight); anthers 0.3–0.8 mm long; caryopsis 1.1–2 mm long 11a. Ligule 1–2.5 mm long; panicle loose, with pedicels 0.8–3.5 mm long; spikelets usually without anthocyanic pigments; anthers 0.4–0.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. sylvatica 11b. Ligule 0.4–1 mm long; panicle dense, with pedicels up to 2 mm long; spikelets often tinged with anthocyanin; anthers 0.3–0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. mexicana 1. Muhlenbergia alopecuroides (Griseb.) P.M. Peterson E bristly wolfstail. Lycurus alopecuroides Griseb.; L. setosus (Nutt.) C.G. Reeder; Pleopogon setosus Nutt. • ME. Wool waste. This species has long been reported as Muhlenbergia phleoides (Kunth) P.M. Peterson (synonym: Lycurus phleoides Kunth). However, introduced plants in New England show upper leaf blades terminating in an elongate, fragile awn tip and relatively long ligules (mostly longer than 3 mm). Muhlenbergia phleoides has upper blades lacking an awn tip or with a very short one and ligules mostly shorter than 3 mm. 2. Muhlenbergia capillaris (Lam.) Trin.

NC

hair-awned muhly. Stipa capillaris Lam. • CT, MA. Rocky forests and woodlands, ridges, ledges, often occurring on trap rock. 3. Muhlenbergia frondosa (Poir.) Fern. N Fig. 241 wire-stemmed muhly. Agrostis frondosa Poir. • CT, MA, ME, NH, RI, VT. Forests and forest edges, along streams, sometimes on more disturbed soil and along lawn edges. 3 ‌ × 8. Muhlenbergia ×curtisetosa Scribn. is a rare hybrid grass in New England known from VT. It combines some features of each parent, sometimes being rhizomatous and often rooting from the lower 1 or 2 nodes (but not as weak and sprawling as M. schreberi, which always lacks rhizomes). Its lower and upper glumes are 0.4–1.5 mm and 0.8–1.9 mm long, respectively (those of M. frondosa are subequal and 2–4 mm

Fig. 241  Glabrous and lustrous apical internode of Muhlenbergia frondosa.

26 8  MO NOCOTS

long, and those of M. schreberi are distinctly unequal—the lower rudimentary or lacking, the upper 0.1–0.3 mm long). 4. Muhlenbergia glomerata (Willd.) Trin. N spike muhly. Muhlenbergia glomerata (Willd.) Trin. var. cinnoides (Link) F.J. Herm. • CT, MA, ME, NH, RI, VT; throughout. Swamps, fens, wet meadows, infrequently on upland soils. 5. Muhlenbergia mexicana (L.) Trin. N Fig. 242 Mexican muhly. Agrostis mexicana L.; Muhlenbergia mexicana (L.) Trin. var. filiformis (Torr.) Scribn. • CT, MA, ME, NH, RI, VT. Forests, swamps and wetland borders, trail edges, roadsides. 6. Muhlenbergia racemosa (Michx.) B.S.P. E Fig. 242  Dull and minutely pubescent apical internode of Muhlenbergia mexicana.

marsh muhly. Agrostis racemosa Michx. • ME, NH. Dry, often disturbed, soil of railroads, roadsides, and other open areas. Reports of this species from VT are based on a specimen of Muhlenbergia glomerata. 7. Muhlenbergia richardsonis (Trin.) Rydb.

NC

matted muhly. Muhlenbergia squarrosa (Trin.) Rydb. • ME. Ice-scoured river shores, river shore ledges and cobble pavement, in regions of high-pH bedrock or till. 8. Muhlenbergia schreberi J.F. Gmel. N Fig. 243 nimblewill muhly. Muhlenbergia schreberi J.F. Gmel. var. palustris (Scribn.) Scribn. • CT, MA, ME, NH, RI, VT. Mesic to wet-mesic, often disturbed, soils of lawns, trail edges, roadsides, gardens, and stream courses. 9. Muhlenbergia sobolifera (Muhl. ex Willd.) Trin. N Fig. 244 rock muhly. Agrostis sobolifera Muhl. ex Willd. • CT, MA, ME, NH, RI, VT. Mesic to dry-mesic forests and woodlands, often on rocky slopes or in the vicinity of cliffs. 10. Muhlenbergia sylvatica Torr. ex Gray N Fig. 243  Sprawling and rooting stems of Muhlenbergia schreberi.

woodland muhly. Agrostis sylvatica Torr. ex Gray; Muhlenbergia sylvatica Torr. ex Gray var. robusta Fern. • CT, MA, ME, NH, RI, VT. Forests, stream courses, shaded ledges. 11. Muhlenbergia tenuiflora (Willd.) B.S.P. N Fig. 245 slender muhly. Agrostis tenuiflora Willd. • CT, MA, NH, VT; primarily southern New England, extending northward only in western VT; also reported from RI by George (1992), but specimens are unknown. Deciduous woodlands and forests, usually those dominated by Quercus and/or Carya, shaded ledges. 12. Muhlenbergia uniflora (Muhl.) Fern. N bog muhly. Agrostis serotina Torr.; Sporobolus serotinus Gray; S. uniflorus (Muhl.) Scribn. & Merr. • CT, MA, ME, NH, RI, VT. Sandy and/or peaty soils of pond shores, boggy depressions, abandoned borrow pits, and fen-like meadows.

Nardus 1. Nardus stricta L. E doormat grass. MA, NH, VT. Fields, roadsides, river banks, ski trails. Fig. 244  Rhizomes of Muhlenbergia sobolifera.

Oryzopsis 1. Oryzopsis asperifolia Michx. N white-grained rice grass. CT, MA, ME, NH, RI, VT. Mainly on coarse, xeric to dry-mesic soils of woodlands, forests, clearings, and open rights-of-way.

Poac e a e   2 6 9

Panicum Recent phylogenetic research showed convincingly that a broadly defined Panicum is artificial (Aliscioni et al. 2003, Zuloaga et al. 2006). Not only are Dichanthelium and Sorengia wellsupported segregates, but also P. verrucosum will need to be segregated into a separate genus. References: Darbyshire and Cayouette (1995), Freckmann and Lelong (2003b). 1a. Upper glumes and lower lemmas distinctly verrucose with hemispherical protruberences, faintly veined [Fig. 248] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. verrucosum 1b. Upper glumes and lower lemmas not verrucose, with distinct veins 2a. Plants perennial, from rhizomes 3a. Panicle open, with ascending to spreading branches; plants sometimes pubescent on the leaf sheaths and/or blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. virgatum 3b. Panicle contracted, with strictly ascending to appressed branches [Fig. 246]; plants glabrous throughout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. amarum 2b. Plants annual, from fibrous roots 4a. Lower glume truncate to subacute at the apex, 20–33% as long as the upper glume; sheaths ± compressed, glabrous; plants often somewhat spongy near base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. dichotomiflorum

Fig. 245  Spikelet of Muhlenbergia tenuifolia.

4b. Lower glume acute to acuminate at the apex, 33–75% as long as the upper glume; sheaths ± terete, pubescent; plants not spongy near base 5a. Spikelets 4–6 mm long; upper (i.e., fertile) florets 2–2.5 mm wide; panicles dense, often arching or nodding at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. miliaceum 5b. Spikelets 1.4–4 mm long; upper florets 0.4–0.9 mm wide; panicles open, upright 6a. Panicles with ascending to spreading-ascending branches, more than 2 times as tall as wide, 1–6 cm wide, with glabrous axillary pulvini; anthers 1.2–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. flexile 6b. Panicles with spreading-ascending to spreading branches, less than 2 times as tall as wide, 4–24 cm wide (narrower in depauperate individuals), with pubescent or glabrous axillary pulvini; anthers 0.7–1.2 mm long 7a. Panicle usually more than 50% of the total plant height [Fig. 247], often breaking at the base of the peduncle and wind-dispersed; spikelets (1.9–) 2.3– 4 mm long; upper floret usually light brown-yellow to light brown (rarely dark brown), 0.45–0.60 times as long as the entire spikelet . . . . . . . . . . . P. capillare 7b. Panicle usually less than 50% of the total plant height, not breaking at the base of the peduncle; spikelets 1.4–2.4 mm long; upper floret dark brown at maturity (light brown in P. philadelphicum ssp. gattingeri), 0.54–0.79 times as long as the entire spikelet 8a. Spikelets 1.8–2.4 at maturity; anthers 0.9–1.2 mm long; upper florets 1.20–1.55 × 0.60–0.85 mm at maturity; uppermost leaf (i.e., the one immediately below the inflorescence) with a blade length to sheath length ratio of 0.8–2.6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. philadelphicum 8b. Spikelets 1.4–1.8 (–2) mm long at maturity; anthers 0.7–0.9 mm long; upper florets 0.75–1.3 × 0.42–0.65 mm at maturity; uppermost leaf with a blade length to sheath length ratio of 2–3 . . . . . . . . . . . . . . . . P. tuckermannii 1. Panicum amarum Ell.

n C Fig. 246

bitter panicgrass. 1a. Panicum amarulum A.S. Hitchc. & Chase; P. amarum Ell. var. amarulum (A.S. Hitchc. & Chase) P.G. Palmer • CT, MA, RI. Atlantic coast beaches, dunes, sandy roadsides. Fig. 246  Inflorescence of Panicum amarum.

270  MONOCOTS

1a. Lower glume with 3–5 veins, the midvein smooth apically; spikelets 4–5.9 mm long; panicles with 2 or more primary branches per node; rhizomes short and/or ascending; reproductive stems often cespitose, mostly 100–250 cm tall . . . . . . . . . . . . . . . . . . 1a. P. amarum ssp. amarulum (A.S. Hitchc. & Chase) Freckmann & Lelong 1b. Lower glume with 7–9 veins, the midvein minutely scabrous apically; spikelets 4.7–7.7 mm long; panicles with 1 or 2 primary branches per node; rhizomes horizontally elongate; reproductive stems usually solitary, mostly 20–150 cm tall . . . . . 1b. P. amarum ssp. amarum Subspecies amarulum is known from CT, Ma, ri and is considered non-native (original populations are believed to have been intentionally introduced). Subspecies amarum is known from CT, RI. It is both native and of conservation concern. 2. Panicum capillare L. ssp. capillare N Fig. 247 witch panicgrass. Panicum capillare L. var. agreste Gattinger • CT, MA, ME, NH, RI, VT. Field edges, roadsides, disturbed soil, exposed shorelines, areas of cultivation. 3. Panicum dichotomiflorum Michx. N Fig. 247  Habit and inflorescences of Panicum capillare.

fall panicgrass. 3a. Panicum dichotomiflorum Michx. var. geniculatum (Wood) Fern.; 3b. Panicum dichotomiflorum ssp. puritanorum (Svens.) Freckmann & Lelong • CT, MA, ME, NH, RI, VT. Exposed shorelines, wet sand, roadsides, disturbed soil, areas of cultivation. 1a. Spikelets 2.3–3.8 mm long, narrow-ovoid, acuminate at the apex, thick and subcoriaceous; pedicels usually shorter than 3 mm . . . . . . . . . . 3a. P. dichotomiflorum var. dichotomiflorum 1b. Spikelets 1.8–2.2 mm long, ellipsoid to narrow-ovoid, acute at the apex; upper glumes and lower lemmas thin-membranaceous; pedicels commonly longer than 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. P. dichotomiflorum var. puritanorum Svens. Variety dichotomiflorum is known from CT, MA, ME, NH, RI, VT. Variety puritanorum is known from MA, NH, RI; also reported from CT by Dowhan (1979), but specimens are unknown. It is more ecologically restricted, typically found on wet soil of exposed shorelines (as opposed to frequently found in human-disturbed areas in the typical subspecies). 4. Panicum flexile (Gattinger) Scribn.

NC

wiry panicgrass. Panicum capillare L. var. flexile Gattinger • CT, VT. Ledges, shorelines, fen-like meadows, usually in soils influenced by high-pH bedrock. 5. Panicum miliaceum L. ssp. miliaceum E Proso millet. CT, MA, ME, NH, RI, VT. Roadsides, yards, waste places, disturbed soil. 6. Panicum philadelphicum Bernh. ex Trin.

NC

Philadelphia panicgrass.  6a. Panicum capillare L. var. campestre Gattinger; P. gattingeri Nash; P. philadelphicum Bernh. ex Trin. ssp. gattingeri (Nash) Freckmann & Lelong • CT, MA, ME, NH, RI, VT. River shores, sand/silt bars, low areas in fields, roadsides, ditches, open woodlands. Darbyshire and Cayouette (1995) showed that the two taxa treated below as varieties are morphologically similar and are closer to each other than any other members of the Panicum capillare complex. 1a. Leaf blades usually 6–12 mm wide; uppermost leaf (i.e., the one immediately below the inflorescence) with a blade length to sheath length ratio of 1.4–2.6, the blade usually more than 50% as tall as the panicle; pedicels spreading, provided with scabrules that slightly, if at all, increase in length toward the apex of the pedicel; apex of the upper glume and lower lemma straight; spikelets usually disarticulating below the glumes, leaving behind an empty pedicel . . . . . . . . . . . . . . . . . . . . . 6a. P. philadelphicum var. campestre (Gatt.) A. Haines 1b. Leaf blades usually 2–6 mm wide; uppermost leaf with a blade length to sheath length ratio of 0.8–1.6, the blade usually less than 50% as tall as the panicle; pedicels appressed, provided with scabrules that noticeably increase in length toward the apex of the pedicel; apex of the upper glume and lower lemma curving over the upper floret; spikelets usually disarticulating on the rachilla first, the upper floret dropping prior to the more tardily disarticulating glumes and sterile lemma . . . . . . 6b. P. philadelphicum var. philadelphicum

Poac e a e   27 1

Variety campestre is known from CT, MA and is of regional conservation concern. Variety philadelphicum is known from CT, MA, ME, NH, RI, VT. 7. Panicum tuckermannii Fern. N Tuckermann’s panicgrass. Panicum philadelphicum Bernh. ex Trin. var. tuckermanii (Fern.) Steyermark & Schmoll • CT, MA, ME, NH, RI, VT. Shorelines, especially frequent on receding shorelines, roadsides, ditches, low areas in fields. Though sometimes considered to be a dwarf state of Panicum philadelphicum, study by Darbyshire and Cayouette (1995) demonstrated that P. tuckermannii is distinct (in fact, more distinct than var. campestre, making it appropriate to treat this taxon as a species). In addition to characters used in the identification key, P. tuckermannii and P. philadelphicum often differ in their spikelet disarticulartion. The former has spikelets that consistently disarticulates below the glumes, leaving behind an empty pedicel. The latter has spikelets that often disarticulate on the rachilla first, the fertile floret dropping prior to the more tardily disarticulating glumes and sterile lemma. 8. Panicum verrucosum Muhl. N Fig. 248 warty panicgrass. CT, MA, RI. Sandy and/or peaty pond shores, low areas, and wetland edges. 9. Panicum virgatum L. N switch panicgrass. Panicum virgatum L. var. cubense Griseb.; P. virgatum L. var. spissum Linder • CT, MA, ME, NH, RI, VT. Fields, roadsides, pond shores, railroads, upper edge of saline to brackish marshes. Fig. 248  Spikelet of Panicum verrucosum.

Pappophorum 1. Pappophorum vaginatum Buckl. E whiplash pappus grass. ME. Wool waste.

Pascopyrum 1. Pascopyrum smithii (Rydb.) A. Löve E western wheat grass. Agropyron smithii Rydb.; Elytrigia smithii (Rydb.) Nevski • MA, NH. Railroad rights-of-way, roadsides, disturbed soil.

Paspalum Reference: Allen and Hall (2003). 1a. Spikelets 1.4–2.5 mm long, paired at each node; ligules 0.2–0.5 mm long. . . . . P. setaceum 1b. Spikelets 2.3–3.3 mm long, solitary at each node [Fig. 249]; ligules 1.5–3.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. laeve 1. Paspalum laeve Michx.

N C Fig. 249

field beadgrass. Paspalum circulare Nash; P. laeve Michx. var. ciculare (Nash) Stone; P. longipilum Nash • CT, MA. Mesic to wet-mesic fields, shorelines, meadows, river banks. 2. Paspalum setaceum Michx. N slender beadgrass. 2a. Paspalum psammophilum Nash; 2b. Paspalum ciliatifolium Michx. var. muhlenbergii (Nash) Fern.; P. muhlenbergii Nash; P. pubescens Muhl. • CT, MA, NH, RI, VT. Sandy fields, roadsides, and forest edges. 1a. Plants prostrate to wide-spreading; leaf blades pubescent with short, soft hairs shorter than 1.5 mm . . . . . . . . . . . . . . . . . . . . . . . . 2a. P. setaceum var. psammophilum (Nash) D. Banks 1b. Plants erect to spreading; leaf blades pubescent, at least in part, with stiffer hairs longer than 1.5 mm (shorter hairs may also be present)

Fig. 249  Branch of inflorescence of Paspalum laeve.

272  MONOCOTS

2a. Spikelets 1.8–2.5 × 1.5–2 mm; lower lemmas usually with a midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. P. setaceum var. muhlenbergii (Nash) D. Banks 2b. Spikelets 1.4–1.9 × 1.1–1.6 mm; lower lemmas usually without a midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2c. P. setaceum var. setaceum Variety psammophilum is known from CT, MA, RI and occurs primarily on the coastal plain. Variety muhlenbergii is known from CT, MA, NH, RI, VT. Variety setaceum is known from CT, MA, RI.

Phalaris 1a. Plants perennial, with creeping rhizomes; keel of glumes not winged; panicle mostly 7–25 cm tall, some branching visible; glumes 4–6.5 mm long; sterile lemmas 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. arundinacea 1b. Plants annual, without rhizomes; keel of glumes winged [Fig. 250]; panicle 1.5–4 cm tall, very dense; glumes 7–10 mm long; sterile lemmas 2.5–4.5 mm long . . . . . . . . . . . P. canariensis 1. Phalaris arundinacea L. N reed canary grass. Phalaris arundinacea L. var. picta L.; Phalaroides arundinacea (L.) Raeusch.; P. arundinacea (L.) Raeusch. var. picta (L.) Tzvelev • CT, MA, ME, NH, RI, VT; throughout. Shorelines, ditches, low fields, graminoid marshes. Based on the behavior of Phalaris arundinacea in New England (invasive in areas with a history of human modification), there exists a possibility that the plants seen on the landscape represent a mixture of native and introduced genotypes. A cultivated form with white-striped leaf blades (picta) is occasionally found naturalized in New England. 2. Phalaris canariensis L. E Fig. 250 common canary grass. CT, MA, ME, NH, RI, VT. Roadsides, lawns, railroads, disturbed sites. Fig. 250  Spikelet of Phalaris canariensis.

Phleum 1a. Plants annual; reproductive stems 5–30 cm tall 2a. Inflorescence 1–3 cm tall; glumes acuminate at apex . . . . . . . . . . . . . . . . . . . P. arenarium 2b. Inflorescence 3–8 cm tall; glumes subacute at apex . . . . . . . . . . . . . . . . . . . . P. subulatum 1b. Plants perennial; reproductive stems 20–100 cm tall 3a. Longer glume awns of a spike 1.5–3 mm long [Fig. 251]; stems not bulbous-thickened at the base, smooth near the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. alpinum 3b. Glume awns 0.7–1.5 mm long; stems bulbous-thickened at the base, scabrous near the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pratense 1. Phleum alpinum L.

Fig. 251  Spikelet of Phleum alpinum.

N C Fig. 251

mountain Timothy. Phleum alpinum L. var. commutatum (Gaudin) Griseb.; P. commutatum Gaudin; P. commutatum Gaudin var. americanum (Fourn.) Hultén • ME, NH. Open, mesic sites in alpine areas, such as snowbank lawns, stream shores, and bases of headwalls, also northern, ice-scoured river shores in high-pH bedrock and/or till regions. 2. Phleum arenarium L. E sand Timothy. MA. Ballast dumps, waste areas. 3. Phleum pratense L. E common Timothy. Phleum nodosum L.; P. pratense L. var. nodosum (L.) Huds. • CT, MA, ME, NH, RI, VT; nearly throughout. Fields, roadsides, and other open, human-disturbed or human-maintained sites. 4. Phleum subulatum (Savi) Aschers. & Graebn. E Italian Timothy. Phleum bellardii Willd. • MA. Ballast dumps, waste areas.

P oac e a e   27 3

Phragmites 1a. Ligules 0.4–0.9 (–1.1) mm long; lower glumes 2.5–5 mm long; upper glumes 4.5–7.5 mm long; lemmas 7.5–12 mm long; middle and upper internodes of stem dull, ridged, tan during the growing season [Fig. 253]; leaf sheaths of middle and upper stem persistent on plant, removed with difficulty in the fall; rhizomes usually thicker than 15 mm, often compressed; leaf blades dark green or dark gray-green (yellow-green in some coastal populations); clones with densely set stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. australis 1b. Ligules 1–1.7 mm long; lower glumes 3–6.5 mm long; upper glumes 5.5–11 mm long; lemmas 8–13.5 mm long; middle and upper internodes of stem smooth and highly lustrous, red-brown to dark red-brown during the growing season [Fig. 252]; leaf sheaths of middle and upper stem sometimes deciduous, easily removed in the fall; rhizomes usually thinner than 15 mm, terete; leaf blades yellow-green; clones with sparsely set stems . . . . . . P. americanus 1. Phragmites americanus (Saltonstall, P.M. Peterson, & Soreng) A. Haines

N C Fig. 252

American reed. Phragmites australis (Cav.) Trin. ex Steud. ssp. americanus Saltonstall, P.M. Peterson, & Soreng • CT, MA, ME, NH, RI, VT. Tidal river shores, fens, lake shores.

Fig. 252  Smooth, lustrous, and dark internode of Phragmites americanus.

2. Phragmites australis (Cav.) Trin. ex Steud. var. australis n Fig. 253 common reed. Phragmites communis Trin. • CT, MA, ME, NH, RI, VT. Fresh to brackish marshes, shores, ditches, fens.

Phyllostachys 1. Phyllostachys dulcis McClure E sweet-shoot bamboo. MA. Fields, yards.

Piptatherum Reference: Barkworth (2007b). 1a. Lemmas dark brown to nearly black; leaves with minute ligules or none, the uppermost blades flat, 8–15 mm wide, longer than (10–) 12 cm . . . . . . . . . . . . . . . . . . . . . . . . . P. racemosum

Fig. 253  Ridged, dull, and pale internode of Phragmites australis.

1b. Lemmas pale green, gray, or pale brown; leaves with ligules, those of the upper leaves 1.5–3 mm long, the uppermost blades involute, 1–2 mm wide, shorter than 10 (–12) cm 2a. Awn of lemma 6–11 mm long, often bent or twisted in the basal half [Fig. 254]; glumes glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. canadense 2b. Awn of lemma 0.5–2 mm long (sometimes absent), straight or slightly bent; glumes minutely scabrous in the apical portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pungens 1. Piptatherum canadense (Poir.) Dorn

N C Fig. 254

Canada mountain-rice grass. Oryzopsis canadensis (Poir.) Torr.; Stipa canadensis Poir. • ME, NH. Dry-mesic forests, woodlands, fields, and heath barrens, rock outcrops and balds, talus slopes. 2. Piptatherum pungens (Torr. ex Spreng.) Dorn N short-awned mountain-rice grass. Milium pungens Torr. ex Spreng.; Oryzopsis pungens (Torr. ex Spreng.) A.S. Hitchc. • CT, MA, ME, NH, RI, VT. Deciduous or mixed evergreen-deciduous woodlands and barrens, dry-mesic to xeric openings. 3. Piptatherum racemosum (Sm.) Eat. N black-seeded mountain-rice grass. Milium racemosum Sm.; Oryzopsis racemosa (Sm.) Ricker • CT, MA, ME, NH, RI, VT. Mesic to dry-mesic, often rocky or ledgy, forests, usually in regions of circumneutral or basic bedrock.

Fig. 254  Spikelet of Piptatherum canadense.

274 MO NOCOTS

Piptochaetium 1. Piptochaetium avenaceum (L.) Parodi N Fig. 255 black-seeded spear grass. Stipa avenacea L. • CT, MA, RI. Sandy and/or rocky woodlands and openings.

Poa Ligule measurements for the key should be performed on the upper one or two leaves of the stem. Assessment of pubescence (of any kind) within the spikelet should occur on the lower florets. Reference: Soreng (2007). 1a. Plants annual, with soft stem bases 2a. Callus of the lemma without arachnoid hairs; reproductive stems mostly decumbent to ascending, sometimes even rooting at the lower nodes [Fig. 257]; anthers (0.5–) 0.8–1 mm long; all 5 veins of the lemma prominent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. annua Fig. 255  Lemma of Piptochaetium avenaceum.

2b. Callus of the lemma with arachnoid hairs; reproductive stems strongly ascending to erect; anthers 0.1–0.2 mm long; only 3 veins of the lemma prominent, the intermediate 2 veins inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. chapmaniana 1b. Plants perennial, with firm stem bases and persistent, dried leaves from the previous season 3a. Many of the florets converted to bulbils with overlapping scales, some of the scales prolonged into narrow, elongate tips 5–15 mm long [Fig. 258]; new shoots bulbous at the base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bulbosa 3b. None of the florets converted to bulbils, lacking narrow, elongate tips; new shoots without bulbous bases 4a. Plants with elongate, slender rhizomes 5a. Stems strongly compressed, with 3–7 leaves; panicle branches mostly 2 per node [Fig. 259]; only 3 veins of the lemma prominent, the intermediate 2 veins inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. compressa 5b. Stems terete to slightly compressed, with 2–4 leaves; panicle branches 2–5 per node [Fig. 262]; all 5 veins of the lemma prominent . . . . . . . . . . . . . . . . . . . . P. pratensis 4b. Plants without rhizomes (though sometimes with stolons) 6a. Marginal veins of the lemma pubescent, at least toward the base [Fig. 261] 7a. Ligules 1 mm or shorter 8a. Glumes narrow-lanceolate, long-acuminate at the apex [Fig. 260], the upper one 4.6–6.2 times as long as wide in sideview and shorter than to subequal in length to the adjacent lemma; uppermost nodes on stem positioned at ½ to ¾ stem height; ligules 0.2–0.7 (–1) mm long; plants often forming extensive colonies in the understory of forested communities . . . . . . . . . . . . . P. nemoralis 8b. Glumes lanceolate to broad-lanceolate, acute to short-acuminate at the apex, the upper one 3.1–5 times as long as wide in side view and shorter than adjacent lemma; uppermost nodes on stem positioned at ⅓ to ⅗ stem height; ligules 0.5–1.5 mm long; plants not forming colonies, the individuals more scattered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) P. interior 7b. Ligules longer than 1 mm

Fig. 256  Lemma of Poa alsodes.

9a. Panicle (9–) 13–30 cm tall, with spreading to spreading-ascending branches that often arch or nod in the distal portion; internodes of the rachilla glabrous; plants primarily of temperate and low elevation mesic to hydric soils, sometimes weakly stoloniferous and sometimes with stems branched above the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. palustris

Poac e a e   27 5

9b. Panicle 1–10 (–15) cm tall, with ascending to erect, relatively straight branches (some branches often ±spreading in P. interior); internodes of rachilla minutely pubescent (view at 30× magnification) or glabrous in P. laxa and sometimes P. interior; plants primarily of boreal, subalpine, and alpine cliffs, talus, and plateaus, without stolons and with simple stems above the base 10a. Anthers 0.7–0.9 mm long; ligules 2.5–3.5 mm long; branches of panicle smooth or sparsely scabrous on the angles, produced 1–3 at each node, usually paired . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. laxa 10b. Anthers 1–2.2 (–2.5) mm long; ligules 0.5–2 (–2.5) mm long; branches of panicle moderately to densely scabrous on the angles (rarely smooth for much of length), produced 2–5 at each node 11a. Lemmas usually at least sparsely pubescent on the lateral veins (often some within a panicle glabrous); uppermost stem node usually positioned at ¹⁄10 to ⅓ stem height; reproductive stems with 0 (–1) exposed nodes, the nodes usually concealed within the leaf sheaths; ligules (1–) 1.2–2 (–2.5) mm long; mature, intact spikelets (1.7–) 2–3.9 mm wide in side-view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. glauca 11b. Lemmas glabrous on the lateral veins; uppermost stem node usually positioned at ⅓ to ⅗ stem height; reproductive stems with (0–) 1 or 2 (–3) exposed nodes; ligules 0.5–1.5 mm long; mature, intact spikelets 1.2–2 (–2.6) mm wide in side view . . . . . . . . . . . . . . . . . . . . (in part) P. interior

Fig. 257  Habit of Poa annua.

6b. Marginal veins of the lemma glabrous [Fig. 256] 12a. Glumes and lemmas ± rounded on the abaxial surface, scarcely keeled; lemmas minutely pubescent with stiff or soft hairs between the veins, at least in the basal ½ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. secunda 12b. Glumes and lemmas keeled; lemmas usually glabrous between the veins 13a. Branches of the panicle mostly 2 per node; lemmas glabrous on the keel [Fig. 263] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. saltuensis 13b. Branches of the panicle 4–8 per node; lemmas pubescent or scabrous on the keel [Fig. 256] 14a. Leaf sheaths usually scabrous; ligule 2.5–7 mm long; all 5 veins of the lemma prominent; plants usually weakly stoloniferous . . . . . . . . . . . P. trivialis

Fig. 258  Inflorescence of Poa bulbosa.

14b. Leaf sheaths glabrous; ligule 0.7–3 mm long; only 3 veins of the lemma prominent, the intermediate 2 veins inconspicuous; plants cespitose, without stolons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. alsodes 1. Poa alsodes Gray N Fig. 256 grove blue grass. CT, MA, ME, NH, VT. Mesic, deciduous, upland and riparian forests. 2. Poa annua L. E Fig. 257 annual blue grass. CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, areas of cultivation. 3. Poa bulbosa L. ssp. vivipara (Koel.) Arcang. E Fig. 258 bulbous blue grass. Poa bulbosa L. var. vivipara Koel. • CT, MA, ME, NH, VT. Fields, lawns, disturbed soil. 4. Poa chapmaniana Scribn. E Chapman’s blue grass. MA. Fields, lawns, areas of cultivation, disturbed soil. 5. Poa compressa L. E Fig. 259 flat-stemmed blue grass. CT, MA, ME, NH, RI, VT. Roadsides, disturbed soil, occasionally found in ± pristine communities on outcrops and cliffs.

Fig. 259  Inflorescence of Poa compressa.

276  MONOCOTS

6. Poa glauca Vahl ssp. glauca

NC

glaucous blue grass. Poa glaucantha Gaudin • ME, NH, VT; mainly in northern portions of the states. River shore outcrops and banks, cliffs, talus, alpine and subalpine plateaus and gullies, lake headlands, usually in areas of high-pH bedrock and/or till. Poa glauca is often stated as having a glabrous lemma callus. This is generally incorrect as most collections show at least a few arachnoid hairs on the callus. Poa glauca has been confused with P. interior, P. laxa, and P. nemoralis in New England, and can be difficult to separate from these species, especially in herbarium collections. Most collections thought to be this species in VT were in fact P. interior. As its name indicates, P. glauca has gray-green to conspicuously glaucous-green foliage. 7. Poa interior Rydb.

NC

interior blue grass. Poa nemoralis L. ssp. interior (Rydb.) W.A. Weber; P. nemoralis L. var. interior (Rydb.) Butters & Abbe • VT; northern portion of state. Talus, cliffs, rocky woodlands, limestone headlands, subalpine gullies. Most collections determined as Poa glauca or as P. nemoralis from pristine locations on high-pH substrate in VT were in fact P. interior. Therefore, P. glauca is rarer than previously believed. 8. Poa laxa Haenke ssp. fernaldiana (Nannf.) N. Hylander

NC

wavy blue grass. ME, NH, VT. Alpine plateaus, ravines, and summits. Fig. 260  Spikelet of Poa nemoralis.

9. Poa nemoralis L. E Fig. 260 wood blue grass. CT, MA, ME, NH, RI, VT. Riparian forests, mesic to dry-mesic upland forests, roadsides, areas of habitation, rarely on rivershore ledges. Herbarium collections are sometimes confused with Poa glauca (see discussion under that species). 10. Poa palustris L. N Fig. 261 fowl blue grass. Poa triflora Gilib. • CT, MA, ME, NH, RI, VT; throughout. Open swamps, shorelines, ditches, wet meadows, graminoid marshes. 11. Poa pratensis L.

n C Fig. 262

Kentucky blue grass. 11a. Poa alpigena (Fries ex Blytt) Lindm. f.; P. pratensis L. var. alpigena Fries ex Blytt; 11b. Poa agassizensis Boivin & D. Löve; 11c. Poa angustifolia L.; P. pratensis L. var. angustifolia (L.) Gaudin; 11d. Poa pratensis L. var. domestica Laestad. • CT, MA, ME, NH, RI, VT. Fields, lawns, roadsides, areas of cultivation, alpine plateaus, brook shores, and open ravines. 1a. Panicle branches smooth or with a few scabrules [Fig. 262]; intermediate nerves of the lemma pubescent or glabrous Fig. 261  Lemma of Poa palustris.

2a. Leaf blades glabrous on the adaxial surface (rarely sparsely pubescent), flat or folded, relatively soft; intermediate veins of lemma usually pubescent; plants of open, alpine plateaus . . . . . . . . . . . . . . . . . . . . . . . 11a. P. pratensis ssp. alpigena (Fries ex Blytt) Hiitonen 2b. Leaf blades sparsely pubescent on the adaxial surface, folded or involute, firm; intermediate veins of lemma glabrous; plants of lower elevation communities . . . . . . . . . . . . . . . . 11b. P. pratensis ssp. agassizensis (Boivin & D. Löve) Taylor & MacBryde 1b. Panicle branches scabrous; intermediate nerves of the lemma glabrous 3a. Basal leaf blades 0.5–1 (–1.5) mm wide, involute to folded (note: senesced blades can be flat), relatively firm, sparsely pubescent on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11c. P. pratensis ssp. angustifolia (L.) Arcang. 3b. Basal leaf blades 1.5–4 mm wide, flat or folded, relatively lax, usually glabrous on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11d. P. pratensis ssp. pratensis

Fig. 262  Panicle branches of Poa pratensis ssp. alpigena.

Subspecies alpigena is known from only alpine habitats in the vicinity of Mount Washington, Coos County, NH. It is native and of regional conservation concern. The report of this species in VT by Kartesz (1999) was erroneous. Subspecies agassizensis is known from only Aroostook County, ME. It is native and of regional conservation concern. Subspecies angustifolia is known MA, ME, NH, VT. It is non-native. Subspecies pratensis is known from CT, MA, ME, NH, RI, VT. It is also non-native.

P oac e a e   27 7

12. Poa saltuensis Fern. & Wieg.

N C Fig. 263

weak spear grass. 12a. Poa saltuensis Fern. & Wieg. var. microlepis Fern.; 12b. Poa debilis Torr. • CT, MA, ME, NH, RI, VT. Dry-mesic to hydric forests, woodlands swamps, meadows, cliff bases, rock outcrops; ssp. languida is usually found on relatively drier and higher pH substrate than ssp. saltuensis, but many exceptions exist. 1a. Anthers 0.9–1.5 mm long; lemmas acute to acuminate at the apex, the keel and lateral margins of lemma forming an apical angle of 10–47 degrees, pliable at the apex, with a prominent scarious tip 0.25–0.5 mm long; upper ligules mostly 0.6–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12a. P. saltuensis ssp. saltuensis 1b. Anthers 0.6–0.9 (–1) mm long; lemmas broad-acute to shortly truncate at apex [Fig. 263], the keel and lateral margins of lemma forming an apical angle of 42–82 degrees, firm at the apex, the scarious tip absent or up to 0.25 mm long; upper ligules mostly 2–4.6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12b. P. saltuensis ssp. languida (A.S. Hitchc.) A. Haines Subspecies saltuensis is known from CT, MA, ME, NH, RI, VT. Subspecies languida is known from CT, MA, RI, VT and is of regional conservation concern. 13. Poa secunda J. Presl ssp. juncifolia (Scribn.) Soreng E curly blue grass. Poa juncifolia Scribn.; Poa nevadensis Vasey ex Scribn.; Poa nevadensis Vasey ex Scribn. var. juncifolia (Scribn.) Beetle • ME; southern portion of state. Wool waste. 14. Poa trivialis L. ssp. trivialis E rough-sheathed blue grass. CT, MA, ME, NH, RI, VT. Low, mesic to hydric ground, including seeps and springs, riparian forests, and stream shores.

Polypogon 1a. Glumes tipped by an awn 4–10 mm long; lemmas tipped by an awn 0.5–1 (–4) mm long; ligules 2.5–16 mm long; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. monspeliensis 1b. Glumes unawned; lemmas unawned; ligules up to 5 mm long; plants perennial . . . . P. viridis 1. Polypogon monspeliensis (L.) Desf. E annual rabbit’s-foot grass. Alopecurus monspeliensis L. • CT, MA, ME, NH. Waste areas, disturbed soil, wool waste. 2. Polypogon viridis (Gouan) Breistr. E beardless rabbit’s-foot grass. Agrostis semiverticillata (Forsk.) C. Christens.; A. verticillata Vill.; A. viridis Gouan; Polypogon semiverticillatus (Forsk.) Hyl. • CT. Ballast dumps, ditches, riparian areas.

Pseudosasa 1. Pseudosasa japonica (Sieb. & Zucc. ex Steud.) Makino ex Nakai E Japanese arrow bamboo. Arundinaria japonica Sieb. & Zucc. ex Steud. • CT. Forest edges, disturbed soil, areas of cultivation.

Puccinellia Lemma measurements should be made on the lower ones of a given spikelet. The orientation of panicle branches should be assessed when the plant is in fruit. Puccinellia nutkaensis (J. Presl) Fern. & Weatherby was reported from ME (under the name P. laurentiana Fern. & Weatherby) by Angelo and Boufford (1998); however, the specimen was in fact P. pumila—Somes Harbor, Rand s.n. (MASS!). References: Hughes and Halliday (1980), Davis and Consaul (2007). 1a. Anthers 1.5–2.6 mm long; first glume 2–3.5 mm long; second glume 2.8–4 mm long; spikelets with 5–11 florets; plants often stoloniferous . . . . . . . . . . . . . . . . . . . . . . . . . . P. maritima

Fig. 263  Spikelet of Poa saltuensis ssp. languida.

278  MONOCOTS

1b. Anthers 0.3–1.5 (–2) mm long; first glume 0.7–2.7 mm long; second glume 1.2–3 mm long; spikelets with 3–9 florets; plants not stoloniferous, or infrequently so in P. pumila (and then usually the result of buried stems) 2a. Inflorescence contracted, even the lower branches ascending; spikelets borne in the apical as well as the basal portion of branches; lemma firm at the tip, with an excurrent midrib; anthers 0.6–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. fasciculata 2b. Inflorescence often open, at least the lower branches spreading or divergent; spikelets borne mostly in the apical portion of branches; lemma hyaline and weak at the apex, the midrib evanescent, usually not reaching the tip; anthers 0.7–1.5 mm long 3a. Branches of the panicle usually smooth or only slightly scabrous; distal margin of the lemma entire or with a few scattered cilia or scabrules [Fig. 265]; caryopsis 1.3–2.1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pumila 3b. Branches of the panicle prominently scabrous; distal margin of the lemma prominently scabrous-ciliate [Fig. 264]; caryopsis 1–1.6 mm long 4a. Lemmas broad-obtuse to truncate at the apex [Fig. 264], the lowest of the spikelet 1.5–2.5 mm long; inflorescence open, the lower branches spreading or reflexed; anthers 0.4–0.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. distans 4b. Lemmas acute to narrow-obtuse at the apex, the lowest of the spikelets (2–) 2.2– 3 (–3.5) mm long; inflorescence more upright, the branches commonly ascending; anthers 0.6–1.5 (–2) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. nuttalliana 1. Puccinellia distans (Jacq.) Parl. E Fig. 264

Fig. 264  Lemma of Puccinellia distans.

spreading alkali grass. Poa distans Jacq.; P. distans (Jacq.) Parl. var. tenuis (Uechtr.) Fern. • CT, MA, ME, VT; also reported from RI by Magee and Ahles (1999), but specimens are unknown. Roadsides and other salted places, ditches, railroads, waste areas. Two infraspecific taxa have been recognized in New England primarily on differences in leaf blade morphology (Fernald and Weatherby 1916). Variety distans has broader blades that are ± flat and var. tenuis has narrower blades that are folded or involute. Review of herbarium specimens shows a continuum of leaf morphologies. In light of this, Puccinellia distans is here treated broadly without formal recogniztion of infraspecific taxa. 2. Puccinellia fasciculata (Torr.) Bickn. E saltmarsh alkali grass. CT, MA, ME, RI. Atlantic coast shores, saline to brackish marshes. 3. Puccinellia maritima (Huds.) Parl. E seaside alkali grass. Poa maritima Huds.; Puccinellia americana Sörenson • CT, MA, ME, NH, RI. Puccinellia maritima is sometimes considered native to northeastern North America. Under such an interpretation, some authors have separated the northeastern plants from European plants as P. americana. However, it does not appear to be possible to separate many collections in the absence of geographic data. Further, this species is believed to be largely or wholly introduced by Davis and Consaul (2007). Given that its distribution is limited to areas of the northeastern coastline with a long history of settlement and that it is known to have been introduced to some states on ballast dumps, it is here considered non-native as well. 4. Puccinellia nuttalliana (J.A. Schultes) A.S. Hitchc. E Nuttall’s alkali grass. Puccinellia airoides (Nutt.) S. Wats. & Coult. • ME, VT. Wet-mesic, often basic, soil. 5. Puccinellia pumila (Vasey) A.S. Hitchc. N Fig. 265 tundra alkali grass. Glyceria paupercula (Holmb.); P. paupercula Holmb.; P. paupercula (Holmb.) Fern. & Weatherby var. longiglumis Fern. & Weatherby • CT, MA, ME, NH, RI. Saline marshes and beaches.

Fig. 265  Lemma of Puccinellia pumila.

P oac e a e   27 9

Sacciolepis 1. Sacciolepis striata (L.) Nash E American cup-scale. Holcus striatus L.; Panicum aquaticum Bosc ex Spreng. • MA. Ditches, shorelines.

Schedonorus Schedonorus has traditionally been included in Festuca. However, many morphological features unite this genus with Lolium rather than Festuca (e.g., auricle morphology, ovary indumentum, style attachment, leaf blade morphology). Reference: Darbyshire (2007). 1a. Lemma awns 10–18 mm long; leaf blades (4–) 6–18 mm wide . . . . . . . . . . . . . . . S. giganteus 1b. Lemma awns absent or up to 2 (–4) mm long; leaf blades 2–10 (–12) mm wide 2a. Auricles of leaves eciliate; basal sheaths brown to red-brown, soon shredding into many fibers; internodes of the rachilla smooth or nearly so; upper glumes 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pratensis 2b. Auricles of leaves ciliate (often sparsely so) [Fig. 266]; basal sheaths white-brown to light brown, persistent, not soon shredding into fibers; internodes of the rachilla antrorsely scabrous; upper glumes 4.5–7 (–9) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . S. arundinaceus 1. Schedonorus arundinaceus (Schreb.) Dumort. E Fig. 266 tall rye grass. Festuca arundinacea Schreb.; F. elatior L. var. arundinacea (Schreb.) C.F.H. Wimmer; Lolium arundinaceum (Schreb.) Darbyshire • CT, MA, ME, VT. Fields, roadsides, lawns. 2. Schedonorus giganteus (L.) Holub E giant rye grass. Bromus giganteus L.; Festuca gigantea (L.) Vill.; Lolium giganteum (L.) Darbyshire • CT. Waste areas, disturbed soil. 3. Schedonorus pratensis (Huds.) Beauv. E meadow rye grass. Festuca elatior L., pro parte; F. pratensis Huds.; Lolium pratense (Huds.) Darbyshire • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns.

Fig. 266  Ciliate auricles at summit of leaf sheath of Schedonorus arundinaceus.

Schizachne 1. Schizachne purpurascens (Torr.) Swallen N false melic grass. Melica purpurascens (Torr.) A.S. Hitchc.; Trisetum purpurascens Torr. • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic, deciduous or mixed evergreen-deciduous forests.

Schizachyrium Reference: Wipff (2003c). 1a. Reproductive stems decumbent and rooting at nodes in contact with soil; apex of leaf sheath with a prominent and elongate constriction proximal to the leaf blade [Fig. 267] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. littorale 1b. Reproductive stems upright, not rooting from lower nodes; apex of leaf sheath with an inconspicuous and short constriction proximal to the leaf blade . . . . . . . . . . . . . . S. scoparium 1. Schizachyrium littorale (Nash) Bickn.

N C Fig. 267

dune bluestem. Andropogon littoralis Nash; A. scoparius Michx. var. littoralis (Nash) A.S. Hitchc.; Schizachyrium scoparium (Michx.) Nash var. littorale (Nash) Gould • CT; also reported from MA and ME by Wipff (2003c), but specimens are unknown. Dunes, upper margin of beaches.

Fig. 267  Constriction at apex of leaf sheath of Schizachyrium littorale.

28 0  MO NOCOTS

2. Schizachyrium scoparium (Michx.) Nash var. scoparium N little bluestem. Andropogon scoparius Michx.; A. scoparius Michx. var. ducis Fern. & Grisc.; A. scoparius Michx. var. frequens F.T. Hubbard; A. scoparius Michx. var. neomexicanus (Nash) A.S. Hitchc.; A. scoparius Michx. var. septentrionalis Fern. & Grisc. • CT, MA, ME, NH, RI, VT; throughout. Grasslands, dry fields, roadsides, rock outcrops, open woodlands.

Secale 1. Secale cereale L. E cultivated rye. Triticum cereale (L.) Salisb. • CT, MA, ME, NH, RI, VT. Roadsides, seeded banks, areas of cultivation.

Setaria Reference: Rominger (2003). 1a. Plants perennial from short, knotty rhizomes; axis of inflorescence scabrous-hispid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. parviflora 1b. Plants annual from fibrous roots; axis of the inflorescence villous and/or hispid or scabrous-hispid in S. verticillata 2a. Each spikelet subtended by 4–12 bristles [Fig. 269]; spikelets 3–3.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pumila 2b. Each spikelet subtended by 1–3 bristles (up to 6 in S. faberi); spikelets 1.6–3 mm long 3a. Axis of the panicle retrorsely scabrous-hispid; branches of the panicle verticillate; inflorescence 5–12 mm thick; leaf blades usually spreading; bristles of the inflorescence retrorsely scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. verticillata 3b. Axis of the panicle villous to hispid, but not scabrous; branches of the panicle not verticillate; inflorescence 7–35 mm thick; leaf blades usually ascending (at least the basal portion of the blade); bristles of the inflorescence antrorsely scabrous 4a. Fertile lemma smooth (rarely very finely transerve rugose), lustrous; disarticulation above the glumes, the caryopsis falling free from the more persistent glumes; mature caryopsis yellow to red or brown to black . . . . . . . . . . . . . . . . . S. italica 4b. Fertile lemma evidently transversely rugulose, dull; disarticulation below the glumes, the entire spikelet, including the caryopsis, falling intact; mature caryopsis green 5a. Leaves pubescent with soft hairs and scabrous on the adaxial surface; spikelets 2.5–3 mm long; inflorescence nodding from near the base [Fig. 268], 20–30 mm wide; second glume 65–75% as long as the spikelet . . . . . . . . . . . . . . . . . . . S. faberi 5b. Leaves scabrous on the adaxial surface; spikelets 1.8–2.2 mm long; inflorescence erect or nodding from near the apex [Fig. 270], 10–23 mm wide; second glume nearly as long as the spikelet . . . . . . . . . . . . . . . . . . . . . . . . . . S. viridis 1. Setaria faberi Herrm. E Fig. 268 Chinese foxtail. CT, MA, ME, NH, RI, VT. Roadsides, fields, disturbed soil. 2. Setaria italica (L.) Beauv. E Fig. 268  Inflorescence of Setaria faberi.

millet foxtail. Panicum verticillatum L. • CT, MA, ME, NH, RI, VT. Roadsides, railroads, fields, areas of cultivation. 3. Setaria parviflora (Poir.) Kerguélen N knotroot foxtail. Panicum geniculatum Lam.; Setaria geniculata (Lam.) Beauv. • CT, MA, RI. Borders of saline marshes, disturbed soil.

Poac e a e   2 8 1

4. Setaria pumila (Poir.) Roemer & J.A. Schultes ssp. pumila E Fig. 269 yellow foxtail. Panicum pumilum Poir.; Setaria glauca, auct. non (L.) Beauv. • CT, MA, ME, NH, RI, VT. Roadsides, fields, disturbed soil, areas of habitation. 5. Setaria verticillata (L.) Beauv. E hooked foxtail. Panicum verticillatum L. • CT, MA, ME, RI, VT; also reported from NH by Fernald (1950b), but specimens are unknown. Roadsides, disturbed soil. 6. Setaria viridis (L.) Beauv. var. viridis E Fig. 270 green foxtail. Panicum viride L.; Setaria viridis (L.) Beauv. var. breviseta (Döll) A.S. Hitchc.; S. viridis (L.) Beauv. var. weinmannii (Roemer & J.A. Schultes) Brand • CT, MA, ME, NH, RI, VT. Roadsides, disturbed soil, cultivated fields, railroads.

Sorengia 1. Sorengia longifolia (Torr.) Zuloaga & Morrone N C Fig. 271 long-leaved redtop-panicgrass. 1a. Panicum anceps Michx. var. pubescens Vasey; P. longifolium Torr.; P. longifolium Torr. var. pubescens (Vasey) Fern.; P. rigidulum Bosc ex Nees ssp. pubescens (Vasey) Freckmann & Lelong; P. rigidulum Bosc ex Nees var. pubescens (Vasey) Lelong;  1b. Panicum agrostoides Muhl. var. elongatum Scribn.; P. elongatum Pursh; P. rigidulum Bosc ex Nees ssp. elongatum (Scribn.) Freckmann & Lelong; P. rigidulum Bosc ex Nees var. elongatum (Scribn.) Lelong; P. stipitatum Nash; 1c. Panicum agrostoides Muhl.; P. condensum Nash; P. rigidulum Bosc ex Nees • CT, MA, ME, NH, RI, VT. Sandy and/or peaty pond shores, meadows, edges of marshes, lacustrine and riparian flood plains.

Fig. 269  Spikelet and subtending bristles of Setaria pumila.

1a. Ligules 0.5–3 mm long, conspicuously fimbriate-ciliate along the apex; leaf blades usually 2–7 mm wide, often folded or involute, usually pilose on the adaxial surface (at least near the base) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. S. longifolia ssp. longifolia 1b. Ligules 0.3–1 mm long, merely erose along the apex; leaf blades usually 5–12 mm wide, usually flat, usually glabrous (but often scabrous) 2a. Spikelets 2.4–3 mm long, usually narrower than 0.6 mm, purple, the upper floret borne on a stipe up to 0.4 mm long . 1b. S. longifolia ssp. elongata (Scribn.) Zuloaga & Morrone 2b. Spikelets 1.6–2.5 mm long, usually wider than 0.6 mm, green or purple-tinged, the upper floret sessile or scarcely stipitate . . . . . . . . . . . . . . . . . . . . . . . 1c. S. longifolia ssp. rigidula (Bosc ex Nees) Zuloaga & Morrone Subspecies longifolia is known from CT, MA, NH, RI. Subspecies elongata is known from CT, RI. Both of these subspecies are considered to be of regional conservation concern. Subspecies rigidula is known from CT, MA, ME, NH, RI, VT. It is the most common form of this species found in New England. Note: as this manuscript went to print, the genus Sorengia was shown to be illegitimate. Coleataenia is the correct genus for this grass with the following names—C. longifolia (Torr.) Soreng ssp. elongata (Scribn.) Soreng, C. longifolia (Torr.) Soreng ssp. longifolia, and C. longifolia (Torr.) Soreng ssp. rigidula (Bosc ex Nees) Soreng.

Fig. 270  Inflorescence of Setaria viridis.

Sorghastrum 1. Sorghastrum nutans (L.) Nash N Indian grass. Andropogon nutans L. • CT, MA, ME, NH, RI, VT. Fields, woodland openings, rocky shorelines, roadsides.

Sorghum 1a. Plants annual or short-lived perennial, lacking rhizomes; leaf blades (5–) 20–100 mm wide; mature spikelets not or only tardily disarticulating . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. bicolor 1b. Plants perennial, with rhizomes; leaf blades 8–20 (–40) mm wide; mature spikelets disarticulating . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. halepense

Fig. 271  Panicle branches with subsecund spikelets of Sorengia longifolia.

28 2  MO NOCOTS

1. Sorghum bicolor (L.) Moench E sorghum.  1a. Holcus bicolor L.; H. sorghum L.; Sorghum vulgare Pers.; S. vulgare Pers. var. technicum (Koern.) Jáv.; 1b. Andropogon drummondii Nees ex Steud.; Sorghum bicolor (L.) Moench var. drummondii (Nees ex Steud.) Mohlenbrock; S. bicolor (L.) Moench var. sudanense (Piper) A.S. Hitchc.; S. drummondii (Nees ex Steud.) Millsp. & Chase; S. sudanense (Piper) Stapf; S. vulgare Pers. var. drummondii (Nees ex Steud.) Hack. ex Chiov. • CT, MA, RI, VT; also reported from ME by Campbell et al. (1995), but specimens are unknown. Fields, disturbed soil, areas of cultivation. 1a. Rames remaining intact at maturity; sessile spikelets 3–9 mm long, ellipsoid to obloid; caryopses exposed at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. S. bicolor ssp. bicolor 1b. Rames tardily disarticulating at maturity; sessile spikelets 5–6 (–7) mm long, lanceoloid to ellipsoid; caryopses not exposed at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. S. bicolor ssp. drummondii (Nees ex Steud.) de Wet & Harlan Subspecies bicolor is known from CT, MA, RI, VT. It is a cultivated strain that is derived from S. bicolor ssp. arundinaceum (Desv.) de Wet & Harlan. Subspecies drummondii is known from MA. It represents hybrid derivatives of S. bicolor ssp. arundinaceum and ssp. bicolor. 2. Sorghum halepense (L.) Pers. E Johnson grass. Holcus halepensis L. • CT, MA, VT; also reported from NH by Magee and Ahles (1999) and from RI by Seymour (1982), but specimens are unknown. Fields, disturbed soil, areas of cultivation.

Spartina Reference: Barkworth (2003c). 1a. Reproductive stems 1–4 (–6) mm thick near base; leaf blades 0.5–4 mm wide, involute when fresh; ligules 0.3–1 mm long; panicles with 2–9 (–15) branches 1–7 cm long . . . . . . . . . S. patens 1b. Reproductive stems (2.5–) 5–20 mm thick near base; leaf blades 5–20 mm wide, flat when fresh (sometimes involute in S. cynosuroides); ligules 1–3 mm long; panicles with (3–) 5–67 branches (1.5–) 5–15 cm long 2a. Leaf blades smooth or slightly scabrous on the margins and near the apex; rhizomes ± white when fresh, with non-imbricate or only slightly overlapping scales; axis of panicle branches prolonged and often exceeding uppermost spikelet . . . . . . . . . . . . . S. alterniflora 2b. Leaf blades strongly scabrous on the margins and near the apex; rhizomes light brown to purple-brown when fresh, with closely imbricate scales; axis of panicle branches not prolonged or shortly prolonged and then not exceeding the uppermost spikelet 3a. Upper glumes with awns 3–8 mm long [Fig. 273], the lateral veins usually glabrous; lower glume ¾ to fully as long as adjacent lemma [Fig. 273] . . . . . . . . . . . . . . S. pectinata 3b. Upper glumes unawned or with awns up to 2 mm long [Fig. 272], the lateral veins usually hispid; lower glume ½ to ⅔ as long as adjacent lemma [Fig. 272] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. cynosuroides 1. Spartina alterniflora Loisel. N smooth cordgrass. Spartina alterniflora Loisel. var. glabra (Muhl. ex Bigelow) Fern. • CT, MA, ME, NH, RI. Saline marshes, Atlantic coast beaches and strands. 2. Spartina cynosuroides (L.) Roth

N C Fig. 272

big cordgrass. Spartina cynosuroides (L.) Roth var. polystachya (Michx.) Beal ex Fern.; S. polystachya Michx. • CT, MA, RI. Saline and brackish marshes. 3. Spartina patens (Ait.) Muhl. N Fig. 272  Spikelet of Spartina cynosuroides.

saltmeadow cordgrass. Spartina patens (Ait.) Muhl. var. monogyna (M.A. Curtis) Fern. • CT, MA, ME, NH, RI. Saline marshes, Atlantic coast beaches and strands.

P oac e a e   2 83

‌ × 4. Spartina ×caespitosa A.A. Eat. is a rare hybrid that grows in disturbed saline 3 and brackish marshes. It is highly variable due to its polyphyletic origin. It usually demonstrates the few and short panicle branches, short ligules, and involute leaf blades of S. patens; however, the rhizomes (when present) are usually purple-brown and have closely imbricate scales and the spikelets are 10–17 mm long (vs. white rhizomes with non-imbricate scales and spikeletes 7–12 mm in S. patens). This nothospecies often has a cespitose habit, unlike any other Spartina in New England. It is known from CT, MA, ME, NH, RI. 4. Spartina pectinata Link N Fig. 273 prairie cordgrass. Spartina pectinata (Ait.) Muhl. var. suttiei (Farw.) Fern. • CT, MA, ME, NH, RI, VT. River and lake shores, roadsides, fresh water marshes, upper edge of saline and brackish marshes, low areas in fields.

Sphenopholis

Fig. 273  Spikelet of Spartina pectinata.

Reference: Daniel (2007). 1a. Spikelets 5–9.5 mm long; awn of second lemma 3.5–7 mm long . . . . . . . . . S. pensylvanica 1b. Spikelets 1.5–4.2 mm long; second lemma lacking an awn or rarely with a short point 2a. Second lemma prominently scabrous [Fig. 275]; anthers 1.2–1.6 mm long . . . . . S. nitida 2b. Second lemma smooth to minutely scabrous [Fig. 274]; anthers 0.3–0.7 mm long 3a. Upper glume acute at the apex [Fig. 274]; spikelets 3–4.2 mm long; lowest rachilla segment 0.8–1 mm long; inflorescence looser and somewhat open; anthers 0.3–0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. intermedia 3b. Upper glume broad-rounded to truncate at the apex; spikelets 2.5–3 mm long; lowest rachilla segment 0.5–0.7 mm long; inflorescence dense; anthers 0.5–0.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. obtusata 1. Sphenopholis intermedia (Rydb.) Rydb. N Fig. 274 slender wedgescale. Eatonia intermedia Rydb.; Sphenopholis obtusata (Michx.) Scribn. var. major (Torr.) K.S. Erdman • CT, MA, ME, NH, RI, VT. River shores, forest openings, marsh edges, slopes.

Fig. 274  Spikelet of Sphenopholis intermedia.

2. Sphenopholis nitida (Biehler) Scribn. N Fig. 275 shiny wedgescale. Eatonia dudleyi Vasey; E. nitida Nash • CT, MA, RI, VT. Dry-mesic to mesic forests and woodlands, often on hillsides and rocky slopes, sometimes associated with cliff bases and outcrops. 3. Sphenopholis obtusata (Michx.) Scribn.

NC

prairie wedgescale. Eatonia obtusata Gray; Sphenopholis obtusata (Michx.) Scribn. var. lobata (Trin.) Scribn. • CT, MA, ME, NH, RI, VT. Upper edges and drier portions of saline and brackish marshes, woodlands, ledges, river and lake shores, roadsides. 4. Sphenopholis pensylvanica (L.) A.S. Hitchc.

NC

swamp wedgescale. Eatonia pensylvanica Gray; Trisetum pensylvanicum (L.) Beauv. ex Roemer & J.A. Schultes • CT, MA, RI. Open swamps, graminoid marshes, wet meadows, mossy seeps and banks along streams.

Sporobolus Sporobolus indicus (L.) R. Br. was reported for ME by Peterson et al. (2003), but specimens are unknown. Sporobolus nealleyi Vasey was reported for ME by Hitchcock and Chase (1950), but specimens are unknown. Reference: Peterson et al. (2003). 1a. Plants annual, from fibrous roots; panicles terminal and axillary, 10–50 × 2–5 mm

Fig. 275  Spikelet of Sphenopholis nitida.

28 4  MO NOCOTS

2a. Lemmas sparsely pubescent; spikelets 2.3–6 mm long; first glume (2.2–) 2.8–4.7 mm long; caryopsis (1.1–) 1.8–2.7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. vaginiflorus 2b. Lemmas glabrous; spikelets 1.6–3 mm long; first glume 1.5–2.4 mm long; caryopsis 1.2–1.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. neglectus 1b. Plants perennial, often with hard, knotty stem bases or rhizomes; panicles terminal or both terminal and axillary, (30–) 50–450 (–500) × 4–120 (–140) mm 3a. Spikelets 1.5–3.2 mm long; anthers 0.3–1 mm long; apex of leaf sheath with a conspicuous tuft of hairs 4a. Panicle dense and contracted throughout flowering and fruiting, 2–8 (–10) mm wide [Fig. 277]; branches of panicle bearing spikelets throughout their length; anthers 0.3–0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. contractus 4b. Panicle initially contracted, ultimately becoming open, 20–120 (–140) mm wide; branches of panicle bearing spikelets only in the apical 75–88% of the branch; anthers 0.5–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. cryptandrus 3b. Spikelets 3–9 (–10) mm long; anthers (1.5–) 1.7–3.2 mm long; apex of leaf sheath glabrous or pubescent, but without a conspicuous tuft of hairs 5a. Panicle (6–) 10–110 mm wide, with appressed-ascending to spreading branches; first glume tapering to a subulate tip from the broader base; glumes equaling or exceeding the lemmas; plants lacking rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. heterolepis 5b. Panicle 4–16 mm wide, with appressed branches [Fig. 276]; first glume lanceolate, gradually tapering to an acuminate tip; glumes shorter than the lemmas; plants sometimes with rhizomes 6a. Lemmas pubescent or scabrous, chartaceous, opaque; caryopsis (1.5–) 2.4–3.5 mm long, the pericarp not becoming gelatinous and slipping off the seed when wet; lower sheaths often sparsely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . S. clandestinus Fig. 276  Inflorescence of Sporobolus compositus var. compositus.

6b. Lemmas glabrous and smooth, membranceous to chartaceous, hyaline; caryopsis 1–2 mm long, the pericarp gelatinizing and slipping off the seed when wet; sheaths glabrous except near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. compositus 1. Sporobolus clandestinus (Biehler) A.S. Hitchc.

NC

hidden dropseed. Sporobolus asper (Beauv.) Kunth var. clandestinus (Biehler) Shinners; S. compositus (Poir.) Mer. var. clandestinus (Biehler) Wipff & S.D. Jones • CT; also reported from MA by Peterson et al. (2003), but specimens are unknown. On ledge and thin soils of ridges and rocky slopes, usually on trap rock, but also occurring on limestone. 2. Sporobolus compositus (Poir.) Mer.

N C Fig. 276

rough dropseed. 2a. Sporobolus asper (Beauv.) Kunth; 2b. Sporobolus asper (Beauv.) Kunth var. pilosus (Vasey) A.S. Hitchc.; S. drummondii (Trin.) Vasey • CT, MA, ME, RI, VT. River shores, banks, and outcrops, lakeshore headlands, sandy fields and coastal beaches, dry openings and barrens. 1a. Reproductive stems 2–5 mm thick; upper sheaths 2.6–6 mm wide; inflorescences with 30–90 spikelets per cm² when pressed . . . . . . . . . . . . . . . . 2a. S. compositus var. compositus 1b. Reproductive stems 1–2 (–2.5) mm thick; upper sheaths 0.8–2.5 mm wide; inflorescences with 16–36 spikelets per cm² when pressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. S. compositus var. drummondii (Trin.) Kartesz & Gandhi Variety compositus is known from CT, MA, RI, VT. Variety drummondii is known from ME and is disjunct from other populations of this taxon. Both varieties are of regional conservation concern.

Poac e a e   2 8 5

3. Sporobolus contractus A.S. Hitchc. E Fig. 277 narrow-spiked dropseed. Sporobolus cryptandrus (Torr.) Gray var. strictus Scribn. • ME; also reported from CT by Dowhan (1979), but specimens are unknown. Dry-mesic to mesic, sandy soils. 4. Sporobolus cryptandrus (Torr.) Gray n sand dropseed. Agrostis cryptandra Torr. • CT, MA, ME, NH, VT. Sandy soils of roadsides, railroads, and fields. Some records of this grass represent introductions (e.g., railroad yards in VT). This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species could be in RI (i.e., the author is unaware of any collections). 5. Sporobolus heterolepis (Gray) Gray

NC

prairie dropseed. Vilfa heterolepis Gray • CT, MA. Thin soils and exposed ledges of woodlands ridges, often on trap rock. 6. Sporobolus neglectus Nash

NC

small dropseed. Sporobolus vaginiflorus (Torr. ex Gray) Wood var. neglectus (Nash) Scribn. • CT, MA, ME, NH, VT. Ledges, river shore outcrops, dry sandy soil of roadsides and fields, often in regions of high-pH bedrock and/or till, more recently naturalizing along heavily salted roadsides. 7. Sporobolus vaginiflorus (Torr. ex Gray) Wood var. vaginiflorus N poverty dropseed. Sporobolus vaginiflorus (Torr. ex Gray) Wood var. inaequalis Fern. • CT, MA, ME, NH, RI, VT. Dry-mesic to xeric, often sterile, soils of fields, roadsides, and disturbed places.

Fig. 277  Inflorescence of Sporobolus contractus.

Taeniatherum 1. Taeniatherum caput-medusae (L.) Nevski ssp. caput-medusae E Medusa-head. Elymus caput-medusae L. • CT. Disturbed soil, waste places.

Thinopyrum 1. Thinopyrum pycnanthum (Godr.) Barkworth E Fig. 278 tick quack grass. Agropyron pycnanthum (Godr.) Godr. & Gren.; Elymus pungens (Pers.) Melderis subsp. pycnanthus (Godr.) O. Bolòs & Vigo; E. pycnanthus (Godr.) Melderis; Elytrigia pycnantha (Godr.) A. Löve; Triticum pycnanthum Godr. • MA, ME, RI. Saline marshes, Atlantic coast beaches. This species was erroneously reported from NH by Kartesz (1999) based on Hodgdon et al. (1979), who merely stated that the species was to be expected in NH.

Torreyochloa 1. Torreyochloa pallida (Torr.) Church N pale false manna grass.  1a. Glyceria fernaldii (A.S. Hitchc.) St. John; Puccinellia fernaldii (A.S. Hitchc.) E.G. Voss; Torreyochloa fernaldii (A.S. Hitchc.) Church; 1b. Glyceria pallida (Torr.) Trin.; Panicularia pallida (Torr.) Kuntze; Puccinellia pallida (Torr.) Kuntze • CT, MA, ME, NH, RI, VT. Shallow water of lakes and pools, swamps, slow-moving streams. 1a. Spikelets 3–5 mm long; lemmas 2–2.8 mm long; larger leaf blades 1–3.5 mm wide, bearing ligules 2–4 mm long; anthers nearly globose, 0.2–0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . 1a. T. pallida var. fernaldii (A.S. Hitchc.) Dore ex Koyama & Kawano 1b. Spikelets 5–7 mm long; lemmas 2.5–3.5 mm long; larger leaf blades mostly 4–10 mm wide, bearing ligules 5–8 mm long; anthers definitely longer than wide, 0.6–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. T. pallida var. pallida Variety fernaldii is known from CT, MA, ME, NH, VT; also reported from RI by Seymour (1982), but specimens are unknown. Variety pallida is known from CT, MA, ME, NH, RI, VT.

Fig. 278  Spikelet of Thinopyrum pungens.

28 6  MO NOCOTS

Tragus 1a. Upper glume 7-veined, 3.8–6.6 mm long, with (5–) 6 or 7 longitudinal rows of spine-like projections; panicle branches 2.1–4.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. racemosus 1b. Upper glume 5-veined, 1.8–4.3 mm long, with 5 longitudinal rows of spine-like projections; panicle branches (0.5–) 0.7–2.7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. berteronianus 1. Tragus berteronianus J.A. Schultes E Fig. 279 spike burgrass. Nazia aliena, auct. non (Spreng.) Scribn. • MA, ME. Wool waste, ballast. 2. Tragus racemosus (L.) All. E stalked burgrass. Cenchrus racemosus L.; Nazia racemosa (L.) Kuntze • ME. Wool waste, ballast. Fig. 279  Spikelet of Tragus berteronianus.

Tridens 1. Tridens flavus (L.) A.S. Hitchc. var. flavus n purpletop tridens. Triodia flava (L.) Smyth • CT, MA, NH, RI, VT. Dry-mesic to xeric fields, roadsides, open woodlands. Some populations of this grass represent introductions (e.g., railroad yards in VT).

Triplasis 1. Triplasis purpurea (Walt.) Chapman var. purpurea N purple sandgrass. Triplasis intermedia Nash • CT, MA, ME, NH, RI. Upper sand beaches and dunes along the Atlantic coast, less frequently inland in sandy soils.

Tripsacum 1. Tripsacum dactyloides (L.) L.

N C Fig. 280

eastern gamagrass. Coix dactyloides L. • CT, MA, RI. Upper border of saline marshes, salt meadows, river shores and banks, dry fields near the coast.

Triraphis Fig. 280  Inflorescence of Tripsacum dactyloides.

1. Triraphis mollis R. Br. E purpleheads. MA. Wool waste.

Trisetum Reference: Rumely (2007). 1a. Inflorescence 3–10 cm tall, dense and spike-like; anthers 0.6–1.4 mm long; leaf blades 1–3 (–4) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. spicatum 1b. Inflorescence 8–15 cm tall, congested, but not at all spike-like; anthers 1.5–2.5 mm long; leaf blades 2–6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. flavescens 1. Trisetum flavescens (L.) Beauv. E Fig. 281 yellow false oat. Avena flavescens L. • MA, VT. Fields, roadsides. 2. Trisetum spicatum (L.) Richter N

Fig. 281  Spikelet of Trisetum flavescens.

narrow false oat. Aira spicata L.; Trisetum spicatum (L.) Richter var. molle (Kunth) Beal; T. spicatum (L.) Richter var. psilosiglume Fern.; T. triflorum (Bigelow) A. & D. Löve • CT, MA, ME, NH, VT; also reported from RI by Kartesz (1999), but specimens are unknown. River shore outcrops, ledges, often in regions of high-pH bedrock and ascending to subalpine situations.

P oac e a e   2 87

Triticum Triticum spelta L. was reported from VT by Atwood et al. (1973); however, the voucher specimen was collected from a cultivated plant—6 Aug 1912, Dutton s.n. (VT!). 1a. Stems hollow below spikes; glumes usually with a well-developed keel only in the apical half, terminating in a short tooth or long awn as long as 40 mm; spikes always simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. aestivum 1b. Stems solid for ca. 1 cm below spikes; glumes with a well-developed keel in the basal and apical halves, terminating in a short tooth as long as 3 mm; spikes sometimes branched near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. turgidum 1. Triticum aestivum L. E bread wheat. Triticum vulgare Vill. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste places, areas of cultivation. 2. Triticum turgidum L. E rivet wheat. CT. Waste areas.

Urochloa 1. Urochloa texana (Buckley) R.D. Webster E Texas liverseed grass. Brachiaria texana (Buckl.) S.T. Blake; Panicum texanum Buckley • MA. Fields, disturbed soil.

Vahlodea 1. Vahlodea atropurpurea (Wahlenb.) Fries ex Hartman

N C Fig. 282

arctic hair grass. Aira atropurpurea Wahlenb.; Deschampsia atropurpurea (Wahlenb.) Scheele • ME, NH, VT. Usually wet, often mossy, places in subalpine and alpine areas, such as ledges, gullies, and brook borders, less frequently also found on plateaus ridges.

Ventenata 1. Ventenata dubia (Leers) Durieu E North African grass. Avena dubia Leers; Ventenata avenacea Koel. • ME. Fields, roadsides, disturbed soil.

Vulpia Reference: Lonard (2007). 1a. Lower glume 0.5–2.5 mm long, 25–33 (–50)% as long as the upper glume; awn of lowest lemma 7.5–22 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. myuros 1b. Lower glume 1.7–5 mm long, mostly 66–75% as long as the upper glume [Fig. 283]; awn of lowest lemma 0.3–12 mm long 2a. Spikelets with 5–13 close florets [Fig. 283]; rachilla internodes mostly 0.5–0.7 mm long; awn of lowest lemma 0.3–6 (–9) mm long [Fig. 283] . . . . . . . . . . . . . . . . . . . . . . . . . V. octoflora 2b. Spikelets with 4–7 loose florets; rachilla internodes ca. 1 mm long; awn of lowest lemma (3–) 9–12 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. bromoides 1. Vulpia bromoides (L.) S.F. Gray E brome six-weeks grass. Bromus dertonensis All.; Festuca bromoides L.; F. dertonensis (All.) Aschers. & Graebn.; Vulpia dertonensis (All.) Gola • MA; also reported from ME by Kartesz (1999), but specimens are unknown. Fields, roadsides, disturbed soil.

Fig. 282  Spikelet of Vahlodea atropurpurea.

28 8  MO NOCOTS

2. Vulpia myuros (L.) K. C. Gmel. E rat-tail six-weeks grass. Festuca megalura Nutt.; F. myuros L.; Vulpia megalura (Nutt.) Rydb. • CT, MA, ME, RI; also reported from NH by Magee and Ahles (1999), but specimens are unknown.

Dry, often sandy, fields, roadsides, and disturbed areas. 3. Vulpia octoflora (Walt.) Rydb. N Fig. 283 eight-flowered six-weeks grass. 3a. Festuca octoflora Walt. var. glauca (Nutt.) Fern.; F. octoflora Walt. var. tenella (Willd.) Fern.; F. tenella Willd.; F. tenella Willd. var. glauca Nutt.; Vulpia octoflora (Walt.) Rydb. var. glauca (Nutt.) Fern.; 3b. Festuca octoflora Walt. • CT, MA, ME, NH, RI, VT. Sandy roadsides, edges of parking lots, and fields, open ledges. 1a. Spikelets 4–5.5 mm long; awn of lowest lemma 2.5–6 (–9) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. V. octoflora var. tenella (Willd.) Fern. 1b. Spikelets 5.5–10 mm long; awn of lowest lemma 0.3–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. V. octoflora var. octoflora Variety tenella is known from CT, MA, ME, NH, RI, VT. Variety octoflora is known only from CT. Some populations of Vulpia octoflora may be introductions. However, this is hard to assess given the species’ affinity to sites with a history of human disturbance. Fig. 283  Spikelet of Vulpia octoflora var. octoflora.

Zea 1. Zea mays L. ssp. mays E corn. CT, MA, ME, NH, VT. Fields, roadsides, areas of cultivation.

Zizania Zizania palustris var. interior (Fassett) Dore is mapped for western VT by Terrell et al. (1997); however, this species is not listed for VT in the written distribution of states and provinces (i.e., the manuscript is internally inconsistent). Therefore, this taxon is not here attributed to VT. Vegetative characters (e.g., leaf blade width, ligule length) are highly variable in New England and show tremendous overlap. Identifications based on such characters should be considered tentative. Reference: Terrell et al. (1997).

Fig. 284  Spikelet of Zizania aquatica.

1a. Fertile carpellate lemmas coriaceous to indurate, lustrous, glabrous or with scabrules confined to the veins and a dense patch at the apex [Fig. 285]; abortive carpellate spikelets 0.6–2.6 mm wide; carpellate portion of inflorescence 1–8 (–15) cm wide, the branches bearing mature carpellate spikelets appressed (rarely some ascending); leaf blades 3–15 (–20) mm wide; ligules 3–8 (–16) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. palustris 1b. Fertile carpellate lemmas chartaceous, flexible, dull or sublustrous, with scattered scabrules on and between the veins, the scabrules not or only slightly concentrated near the apex [Fig. 284]; abortive carpellate spikelets 0.4–1 mm wide; carpellate portion of inflorescence (5–) 10–50 cm wide, the branches bearing mature carpellate spikelets spreading-ascending to ascending (rarely appressed when bearing mainly abortive spikeletes); leaf blades (3–) 10–55 (–72) mm wide; ligules (5–) 10–15 (–30) mm long . . . . . . . . . . Z. aquatica 1. Zizania aquatica L. var. aquatica N Fig. 284 southern wild rice. Ceratochaete aquatica (L.) Lunell; Zizania aquatica L. var. subbrevis Boivin • CT, MA, ME, RI, VT. Fresh to brackish-tidal river shores, shallow water of lakes and inland rivers. Some populations of this species have been intentionally introduced as food for waterfowl. 2. Zizania palustris L. var. palustris N Fig. 285 northern wild rice. Melinum palustre (L.) Link; Zizania aquatica L. var. angustifolia A.S. Hitchc. • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving water of lakes and rivers.

Fig. 285  Spikelet of Zizania palustris.

P o nt e d e r i ac e a e   2 8 9

Pontederiaceae 1a. Inflorescence densely and many flowered, commonly with more than 50 flowers; perianth blue-purple (rarely white), the upper middle lobe with 2 contiguous, yellow spots; fruit achene-like, with a single seed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pontederia 1b. Inflorescence open to dense, with 2–15 flowers; perianth white, yellow, or blue, with or without spots, but never with 2 yellow spots; fruit a capsule, with 7–30 seeds 2a. Petioles at least slightly inflated; perianth blue, the lobes 16–37 mm long; flowers with 6 stamens; plants typically free-floating . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eichhornia 2b. Petioles absent or not inflated; perianth white or yellow, the lobes 3–11 mm long; flowers with 3 stamens; plants rooted in substrate . . . . . . . . . . . . . . . . . . . . . . . . Heteranthera

Eichhornia 1. Eichhornia crassipes (Mart.) Solms-Laubach E common water-hyacinth. Eichhornia speciosa Kunth; Piaropus crassipes (Mart.) Raf. • CT. Slow-moving streams, ponds.

Heteranthera Reference: Horn (2002). 1a. Only linear sessile leaves produced; flowers yellow; filaments glabrous . . . . . . . . . H. dubia 1b. Both linear to oblanceolate sessile leaves and petiolate leaves with reniform blades produced on each plant; flowers largely white; filaments pubescent with multicellular hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. reniformis 1. Heteranthera dubia (Jacq.) MacM. N grass-leaved mud-plantain. Commelina dubia Jacq.; Heteranthera graminea (Michx.) Vahl; Zosterella dubia (Jacq.) Small • CT, MA, ME, NH, VT. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers. This vegetatively plastic species has been considered to belong to a different genus (Zosterella) based on convincing morphological characters. However, phylogenetic work has shown it to be nested within Heteranthera (see Horn 2002 for discussion and additional references). 2. Heteranthera reniformis Ruiz & Pavón

NC

kidney-leaved mud-plantain. CT; southwestern portion of state. Mud flats of tidal river shores.

Pontederia 1. Pontederia cordata L. N Fig. 286 pickerelweed. Narukila cordata (L.) Nieuwl.; Pontederia cordata L. var. angustifolia (Pursh) Torr. & Ell.; P. cordata L. var. lanceolata (Nutt.) Griseb.; P. lanceolata Nutt.; Unisema cordata (L.) Farw. • CT, MA, ME, NH, RI, VT; nearly throughout. Shallow, still or slow-moving water of lakes, rivers, and backwater sloughs, typically in waters with mud or muck substrate.

Fig. 286  Flower of Pontederia cordata.

29 0  MO NOCOTS

Potamogetonaceae The leaf sheaths in this family are usually separate from the blade and appear as stipules. For circumscription of the family, see Lindquist et al. (2006). References: Hellquist and Crow (1980), Wiegleb and Kaplan (1998). 1a. Leaves opposite; flowers unisexual; staminate flowers with 1 stamen; stigma funnelform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zannichellia 1b. Leaves alternate; flowers bisexual, with 2 or 4 stamens; stigma slender to capitate 2a. Leaf blades narrow-linear, 0.2–1.5 mm wide, all submersed, cross-septate throughout; stipules connate to the blade for a distance of 4–30 mm, commonly more than 10 mm; spikes submersed or sometimes floating, with well-separated lower whorls of flowers, on thin, flexuous peduncles; drupes not compressed, with an indistinctly coiled embryo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stuckenia 2b. Leaf blades filiform to orbicular, 0.1–75 mm wide, all submersed or some floating, cross-septate only in the central lacunar bands (if present); stipules distinct from the blade or connate for a distance of less than 10 mm (long-connate in P. robbinsii); spikes usually with contiguous whorls of flowers, on thick, stiff peduncles, usually emersed; drupes compressed, with a distinctly coiled embryo (indistinctly coiled in P. robbinsii) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Potamogeton

Potamogeton Species of Potamogeton produce one or both of two types of leaves—submersed and floating. All species of Potamogeton produce submersed leaves, which are very thin, translucent, and generally occur below the surface of the water (unless stranded). Floating leaves are thick, opaque, and occur on the surface of the water during normal seasons. Sometimes submersed leaves appear absent due to their decay in the later season or mechanical damage (e.g., wave action). Oil glands, referred to as nodal glands, occur in many species (most common in those with sessile, submersed leaves). They appear as a pair of green- to gold-colored circular bumps at the nodes. They are sometimes difficult to detect and should be sought on several nodes to assess their presence/absence. Hybrid plants are infrequently collected but are most common among the broad-leaved species and in riverine communities. They are typically sterile and do not produce fruits (though they do flower). Potamogeton alpinus × P. epihydrus was reported from ME by Angelo and Boufford (2000), but the specimens (at MASS!) have qualified (i.e., tentative) determinations and do not have morphologies that suggest the involvement of P. epihydrus (e.g., the lacunar bands of the leaf blade are too few). Potamogeton ×‌nericius Hagstr. was reported from ME by Angelo and Boufford (2000), but the only collection observed (at MASS!) has a qualified (i.e., tentative) determination and appears equivocal. Potamogeton illinoensis × P. perfoliatus was reported from MA and P. ×‌prussicus Hagstr. was reported from VT by Angelo and Boufford (2000), but specimens are unknown. 1a. Submersed leaf blades narrower than 4 mm wide 2a. Submersed leaf blades with the stipule basally fused to the blade, only the tip of the stipule distinct from the blade [Fig. 299]; drupe with an embryo coiled more than 1 full spiral [Fig. 289] 3a. Submersed leaf blades 0.1–0.4 mm wide, the apex acute to long-tapering; fused portion of the stipule usually shorter than the distinct portion; apex of floating leaves acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bicupulatus 3b. Submersed leaf blades 0.5–2 mm wide, the apex obtuse to acute; fused portion of the stipule usually equal to or longer than the distinct portion; apex of floating leaves obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. spirillus

Pota m o g e to n ac e a e   2 9 1

2b. Submersed leaf blades with stipules that are distinct from the blade for their entire length [Fig. 287]; drupe with an embryo coiled 1 full spiral or less [Fig. 294] 4a. Floating leaves, which are thicker and greatly expanded in width compared with the submersed leaves, produced and generally present on some of the plants in each colony [Fig. 296] 5a. Floating leaf blades 6–15 mm long, 5- to 9-veined; submersed leaf blades 0.1–1 mm wide, thin, transparent, lacking lacunar cells; spikes (3–) 6–8 mm long; drupes 1.5–2.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. vaseyi 5b. Floating leaf blades 15–120 mm long, 7- to 37-veined; submersed leaf blades 0.25–10 mm wide, either with abundant lacunar cells or with thick and nearly opaque texture; spikes 10–50 mm long; drupes (2–) 2.5–4.5 mm long 6a. Submersed leaf blades (1–) 2–10 mm wide, thin, transparent, with prominent bands of lacunar cells 1–2 mm wide on each side of the midrib [Fig. 291]; drupes with abaxial and lateral keels . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) P. epihydrus 6b. Submersed leaf blades 0.25–2 mm wide, thick and nearly opaque, petiole-like, lacking lacunar cells; drupes with only an abaxial keel 7a. Submersed leaf blades 0.8–2 mm wide; floating leaf blades 5–10 × 2.5–6 cm, usually subcordate to rounded at the base [Fig. 296]; petioles pale at the apex; stipules 4–10 cm long; drupes 3.5–5 mm long, deeply wrinkled on each side, with an inconspicuous abaxial keel . . . . . . . . . . . . . . . . . . P. natans 7b. Submersed leaf blades 0.25–1 mm wide; floating leaf blades 2.5–6 × 1–3 cm, cuneate to rounded at the base; petioles usually lacking a pale band at the apex; stipules 2.5–4 cm long; drupes 2–3.5 mm long, not wrinkled on the sides, with a prominent abaxial keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. oakesianus 4b. Floating leaves not produced, all the leaves submerged [Figs. 290, 292] 8a. Rhizome elongate; peduncles (3–) 5–25 cm long [Fig. 290]; leaf blades 0.1–0.5 mm wide, 1-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. confervoides 8b. Rhizome scarcely developed (elongate in P. foliosus); peduncles 0.5–6 cm long [Fig. 292]; leaf blades 0.2–5 mm wide, 1- to 35-veined 9a. Stems conspicuously compressed; leaf blades 15- to 25 (–35)-veined; spikes with 7–11 whorls of flowers; drupes 4–4.5 mm long . . . . . (in part) P. zosteriformis 9b. Stems terete or slightly compressed; leaf blades 1- to 13-veined; spikes with 1–5 whorls of flowers; drupes 1.5–3.5 mm long 10a. Nodal glands absent (very rarely present); spikes 1.5–7 mm long, with 1 or 2 whorls of flowers [Fig. 292]; drupes with a conspicuous abaxial keel 11a. Leaf blades acute at apex; drupes 1.4–2.3 (–2.7) mm long, with a single, wing-like and undulate dorsal keel to 0.4 mm tall; peduncles 3–11 (–37) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. foliosus 11b. Leaf blades bristle-tipped (rarely only apiculate or blunt) [Fig. 295]; drupes 2.3–4 mm long, with 3 keels, the dorsal keel ridge-like, to 0.2 mm tall; peduncles 6–14 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. hillii 10b. Nodal glands present on at least some of the nodes [Fig. 288]; spikes 2–18 mm long, with 1–5 whorls of flowers; drupes without an abaxial keel (sometimes with a low keel in P. obtusifolius) 12a. Stipules fibrous, white (brown in P. ogdenii) 13a. Leaf blades obtuse to rounded at the apex, sometimes with an apiculus; winter buds with inner leaves oriented at a right angle to the outer leaves, the outer leaves corrugated at the base, the inner leaves modified into a fan-shaped structure [Fig. 293] . . . . . . . . . . . . . . . . . P. friesii

29 2  MO NOCOTS

13b. Leaf blades obtuse to attenuate at the apex; winter buds flattened, the inner and outer leaves oriented in the same plane, the outer leaves not corrugated at the base (rarely corrugated in P. strictifolius), the inner leaves undifferentiated or modified into a fusiform structure [Fig. 300] 14a. Leaf blades with 3–5 (–7) veins, 0.6–2 mm wide, without lacunae; stipules usually white, the margins connate; drupes 1.9–2.1 mm long; inner leaves of winter bud undifferentiated . . . . . . . . . . . . P. strictifolius 14b. Leaf blades with 3–9 (–13) veins, 1.2–2.9 mm wide, with 0–3 rows of lacunae on each side of the midrib; stipules usually brown, convolute; drupes 2.5–3 mm long; inner leaves of winter bud commonly rolled into a hard, fusiform structure . . . . . . . . . . . . . . . . . . . . . . P. ogdenii 12b. Stipules delicate, not fibrous, green, brown, or white 15a. Leaf blades 1–3.5 mm wide, obtuse to rounded at the apex, sometimes with an apiculus, commonly suffused with red; drupe 2.5– 3.5 mm long, with a beak 0.6–0.7 mm long, the embryo coiled 1 full spiral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. obtusifolius 15b. Leaf blades 0.2–2.5 mm wide, acute to obtuse at the apex, green; drupe 1.5–2.2 mm long, with a beak 0.1–0.6 mm long, the embryo coiled less than 1 full spiral 16a. Leaf blades 0.2–0.7 mm wide, acute at the apex, 1-veined, with a single row of lacunae on each side of the midvein; peduncles 10–35 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  P. gemmiparus 16b. Leaf blades 0.2–2.5 mm wide, acute to obtuse (rarely apiculate) at the apex, 3- or 5-nerved, with 0–5 rows of lacunae on each side of the midvein; peduncles 5–62 mm long 17a. Leaf blades with up to 2 rows of lacunae on each side of the midvein, acute at the apex (rarely apiculate); stipules with united margins, forming a tube surrounding the stem; spikes mostly terminal, 1–3 per plant, with peduncles 10–62 mm long, with 2–4 separate whorls of flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pusillus 17b. Leaf blades with up to 5 rows of lacunae on each side of the midvein, acute to obtuse at the apex; stipules convolute, but without united margins; spikes terminal and/or axillary, often more than 3 per plant, with peduncles 5–30 (–45) mm long, with 1–3 crowded whorls of flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. berchtoldii 1b. Submersed leaf blades 4 mm wide or wider (those of the axillary shoots may be narrower in some species) 18a. Stipules basally fused to the leaf blade, only the tip of the stipule distinct from the blade; leaf blades usually minutely serrulate, especially in the apical portion, with a pair of auricles at the base; well-formed inflorescences branched [Fig. 298] . . . . . . . . . . P. robbinsii 18b. Stipules of submersed leaves distinct from the leaf blade their entire length; leaf blades entire or toothed in P. crispus, without auricles; inflorescences unbranched 19a. Leaf margins conspicuously serrulate; beak of drupe 2–3 mm long; turions commonly formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. crispus 19b. Leaf margins entire; beak of drupe 0.3–0.9 mm; turions absent or rarely formed 20a. Submersed leaf blades sessile and conspicuously cordate-clasping at the base [Fig. 297]; floating leaves not produced 21a. Leaf blades strictly entire, (60–) 80–280 mm long, the apex cucullate and usually splitting when pressed; rhizome spotted with red-brown; stipules persistent; drupes 4–5.7 mm long, with a prominent, abaxial keel . . . . . . . . . . . . . . P. praelongus

Pota m o g e to n ac e a e   2 9 3

21b. Leaf blades provided with minute, caducous, 1-celled spicules along the margin, 9–130 mm long, the apex flat and not splitting when pressed; rhizome unspotted; stipules disintegrating and remaining as fibers or completely absent; drupes 1.6–4.2 mm long, with an inconspicuous keel or the keel absent 22a. Leaf blades ovate to suborbicular, 9–76 (–97) mm long, delicately 7- to 15-veined; stipules disintegrating and absent on the lower part of the stem; drupes 1.6–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. perfoliatus 22b. Leaf blades narrow-ovate to narrow-lanceolate, (16–) 30–130 mm long, coarsely 13- to 21-veined [Fig. 297]; stipules disintegrating into persistent fibers; drupes 2.2–4.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. richardsonii 20b. Submersed leaf blades sessile or petiolate, not cordate-clasping; floating leaves produced and generally present on some of the plants in each colony or not produced in P. zosteriformis 23a. Stems red-brown- or black-spotted [Fig. 287]; floating leaf blades rounded to cordate at base; submersed leaf blades usually crisped and arcuate [Fig. 287]; drupes 5–6.7 mm long 24a. Submersed leaf blades narrow-lanceolate to lanceolate, with 7–19 veins; floating leaf blades with 15–19 veins; stems black-spotted . . . . . . . . . . P. pulcher 24b. Submersed leaf blades ovate to broad-oblanceolate, with 19–49 veins; floating leaf blades with 27–49 veins; stems red-brown-spotted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. amplifolius 23b. Stems and petioles unspotted; floating leaf blades cuneate to rounded at the base or absent altogether in P. zosteriformis; submersed leaf blades usually flat and not arcuate (sometimes crisped and/or arcuate in P. illinoensis); drupes 1.9–4.3 mm long 25a. Stems conspicuously compressed; floating leaves not produced; submersed leaf blades 15- to 25 (–35)-veined . . . . . . . . (in part) P. zosteriformis 25b. Stems terete to somewhat compressed; floating leaves produced, though not always present; submersed leaf blades 3- to 19 (–21)-veined 26a. Stipules (1–) 3–9 cm long; submersed leaf blades with 7–19 (–21) veins, sessile or with petioles up to 13 cm long 27a. Submersed leaves with petioles (2–) 5–13 cm long, the larger leaves with blades 10–35 mm wide and acute at the apex; drupes red to redbrown, the lateral keels with blunt to sharp teeth . . . . . . . . . . . . P. nodosus 27b. Submersed leaves sessile or with petioles up to 4 cm long, the larger leaves with blades 20–45 mm wide and mucronate at apex; drupes graygreen to olive-green, the lateral keels +⁄– without teeth . . . . . . P. illinoensis 26b. Stipules 1–3 cm long; submersed leaf blades with 3–13 veins, sessile 28a. Submersed leaves distichous, the blades ± linear, with prominent lacunar bands 1–2 mm wide on each side of the midrib; each lacunar band composed of 9–18 rows of cells . . . . . . . . . . . . . . . . . . (in part) P. epihydrus 28b. Submersed leaves ± spirally arranged, the blades linear to narrowobovate, with inconspicuous lacunar bands up to 0.5 mm wide on each side of the midrib; each lacunar band composed of 0–4 rows of cells 29a. Submersed leaf blades 4.5–18 (–25) cm long, strictly entire, redtinged, especially in drying; stems unbranched; drupes (2.5–) 3–3.5 mm long, plump and turgid, the exocarp hard, the embryo coiled 1 full spiral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. alpinus

29 4  MO NOCOTS

29b. Submersed leaf blades (1.5–) 3.1–9.1 (–10) cm long, provided with minute, caducous, 1-celled spicules along the margin, green; stems sparingly to many-times branched; drupes 1.9–2.3 mm long, laterally compressed, the exocarp soft, the embryo coiled less than 1 full spiral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. gramineus 1. Potamogeton alpinus Balbis N reddish pondweed. Potamogeton alpinus Balbis var. tenuifolius (Raf.) Ogden • MA, ME, NH, VT; most common in the northern portions of the states. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers. 1 × 19. Potamogeton ×‌prussicus Hagstr. is a very rare pondweed hybrid known from VT. It is intermediate in morphology between the parental species. 2. Potamogeton amplifolius Tuckerman N Fig. 287 big-leaved pondweed. CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving, slightly acidic to basic water of lakes and rivers, often in relatively deep water relative to some other congenerics. 2 × 11. This very rare pondweed hybrid is known from CT. It has sessile submersed leaves that are weakly arched and have 9–11 veins. Further, they are narrower than generally observed in Potamogeton amplifolius. The stems are unspotted. Fig. 287  Submersed leaves and stipules of Potamogeton amplifolius.

‌ × 13. Potamogeton ×scoliophyllus Hagstr. is a rare pondweed hybrid known from CT, 2 MA, VT. It generally resembles P. amplifolius, but the submersed leaves are weakly arched, have mostly 13–15 veins (rather than 19–49), and are relatively elongate and narrow for that species. The floating leaf blades are broad-cuneate at the base (rather than rounded to cordate), and the stems are unspotted (rather than often spotted with red-brown). 2 × 20. This rare pondweed hybrid known from ME, VT. It has leaf blades that are weakly arched, sessile to somewhat clasping, and with mostly 15–21 veins. The tips of the leaf blades split in pressing (as in Potamogeton praelongus). The stems have dark blotches. 2 × 21. This rare pondweed hybrid is known from NH. It has smaller (though still somewhat arched) submersed blades than P. amplifolius (mostly 31–57 mm long, rather than 50–125 mm long), and there is some black spotting on the stems. 3. Potamogeton berchtoldii Fieber N Fig. 288 Berchtold’s pondweed. Potamogeton pusillus L. var. mucronatus (Fieber) Graebn.; P. pusillus L. ssp. tenuissimus (Mert. & Koch) Haynes & Hellquist; P. pusillus L. var. tenuissimus Mert. & Koch • CT, MA, ME, NH, RI, VT; throughout. Shallow, still or slow-moving, acidic to basic water of lakes and rivers. ‌ × 19. Potamogeton ×mysticus Morong is a rare pondweed hybrid known from MA, 3 ME, VT. It generally resembles P. gramineus, but the submersed blades vary from sessile to weakly clasping. 4. Potamogeton bicupulatus Fern. N Fig. 289

Fig. 288  Nodal glands of Potamogeton berchtoldii.

Fig. 289  Achene of Potamogeton bicupulatus showing coiling of embryo.

snail-seed pondweed. Potamogeton capillaceus Poir., pro parte; P. diversifolius Raf. var. trichophyllus Morong • CT, MA, ME, NH, RI, VT; mainly on the coastal plain but disjunct to Essex County, VT. Shallow, still or slow-moving, acidic to circumneutral water of lakes and rivers. ‌4 × 7. Potamogeton ×aemulans Z. Kaplan, Hellquist, & Fehrer is a rare pondweed hybrid responsible for reports of Potamogeton diversifolius Raf. in New England. It has submersed leaves 0.2–1.8 mm wide that are 3 (–5)-veined, with or without a narrow band of lacunae bordering the midrib. The stipules are 3–34 mm long, mostly distinct from the leaf but some connate for a short distance (up to 2.8 mm). The floating leaves, which are always present on flowering plants, are 18–35 × 5–11 mm and have 9–15 veins. The hybrid shows dimorphic inflorescences (within a population), with some short ones (2–3 mm) in the axils of submerged leaves and longer ones (3–6 mm) in emersed spikes. This hybrid is known from CT, MA, and is found primarily in slow-moving, circumneutral water of rivers.

Pota m o g e to nac e a e   2 9 5

5. Potamogeton confervoides Reichenb. N Fig. 290 alga-like pondweed. CT, MA, ME, NH, RI, VT. Shallow acidic to, less commonly, circumneutral water of lakes and ponds, frequently occurring in north-temperate to boreal areas. 6. Potamogeton crispus L. E curly pondweed. CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers, including fresh and brackish systems. 7. Potamogeton epihydrus Raf. N Fig. 291 ribbon-leaved pondweed. Potamogeton epihydrus Raf. var. nuttallii (Cham. & Schlecht.) Fern.; P. epihydrus Raf. var. ramosus (Peck) House • CT, MA, ME, NH, RI, VT; throughout. Shallow, still or slow-moving, acidic to basic water of lakes and rivers. 7 ‌ × 19. Potamogeton ×versicolor Z. Kaplan, Hellquist, & Fehrer is a rare pondweed known from ME. It ± resembles Potamogeton alpinus, but the submersed blades have prominent lacunar bands, and the narrow-lanceolate blades sometimes have weakly clasping bases. Floating leaves are sometimes produced on mature plants and have short petioles 4–22 mm long (compared with 20–125 mm in P. epihydrus). This hybrid closely resembles P. ×‌nitens (P. gramineus × P. perfoliatus) but has submersed leaves 4–11 mm wide and floating leaves 22–32 mm long with apically winged petioles 4–22 mm long (vs. submersed leaves 5–23 mm and floating leaves 27–65 mm with apically unwinged or barely winged petioles 12–40 mm long).

Fig. 290  Spikes borne on elongate peduncles of Potamogeton confervoides.

8. Potamogeton foliosus Raf. ssp. foliosus N Fig. 292 leafy pondweed. Potamogeton foliosus Raf. var. genuinus Fern.; P. foliosus Raf. var. macellus Fern. • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving, circuneutral to basic water of lakes and rivers. 9. Potamogeton friesii Rupr.

N C Fig. 293

flat-stalked pondweed. CT, MA, ME, VT. Shallow, circumneutral to basic water of lakes and ponds. In addition to characters used in the key, Potamogeton friesii is separated from P. strictifolius, which it most closely resembles, by its compressed peduncles (they are terete in P. strictifolius). 10. Potamogeton gemmiparus (J.W. Robbins) J.W. Robbins ex Morong

NC

budding pondweed. Potamogeton berchtoldii Fieber ssp. gemmiparus (J.W. Robbins) K. Les & N.P. Tippery; P. pusillus L. ssp. gemmiparus (J.W. Robbins) Haynes & Hellquist; P. pusillus L. var. gemmiparus J.W. Robbins • CT, MA, ME, NH, RI; also reported from VT by Haynes and Hellquist (2000c), but specimens are unknown. Shallow, still or slow-moving, circumneutral water of lakes and rivers. Potamogeton gemmiparus is a near-endemic of New England; it also is known from nearby Quebec.

Fig. 291  Portion of leaf blade showing prominent lacunar bands of Potamogeton epihydrus.

11. Potamogeton gramineus L. N Fig. 294 grassy pondweed. Potamogeton gramineus L. var. maximus Morong; P. gramineus L. var. myriophyllus J.W. Robbins; P. gramineus L. var. typicus Ogden • CT, MA, ME, NH, RI, VT; throughout. Shallow, still or slow-moving, acidic to basic water of lakes and rivers. ‌ × 13. Potamogeton ×spathuliformis (J.W. Robbins) Morong is a rare pondweed 11 hybrid known from CT, MA, VT. It has submersed leaves that are subsessile to petiolate, the petioles as long as 19 mm. The submersed leaves have 5–11 veins (usually 7) and have stipules 11–35 mm long. ‌11 × 14. Potamogeton ×sparganiifolius Laestad. ex Fries is a rare pondweed hybrid known from NH; also reported from MA by Angelo and Boufford (2000), but specimens are unknown. It ± resembles P. oakesianus, but the submersed leaves have expanded, narrow, green blades near the apex (instead of all being phyllodial and lacking blades). ‌ × 15. Potamogeton ×argutulus Hagstr. is a rare pondweed hybrid known from MA, 11 ME. It has submersed leaves on petioles mostly 8–65 mm long (rarely subsessile). The submersed leaves have 7–11 veins and have stipules 13–42 mm long.

Fig. 292  Habit of Potamogeton foliosus showing short spikes.

29 6  MO NOCOTS

‌11 × 16. Potamogeton ×mirabilis Z. Kaplan, Hellquist, & Fehrer is a very rare pondweed hybrid known from NH. It is a multibranched plant that has floating leaves on mature plants like Potamogeton oakesianus, but the submersed leaves are green, narrow (mostly 0.3–1.6 mm wide), some of them with expanded green blades near the apex. This hybrid is unusual in that some leaf blades show stipules connate to the leaf blades for a distance of up to 1.2 mm (neither parent shows this character state). The spikes are 22–26 mm long.

Fig. 293  Winter bud of Potamogeton friesii.

‌11 × 19. Potamogeton ×nitens G. Weber is an uncommon pondweed hybrid known from CT, MA, ME, NH, VT. It ± resembles P. gramineus, but some of the leaf blades weakly clasp the stem. It is further characterized by submersed leaves 5–23 mm wide with 7–17 veins, floating leaves sometimes produced on mature plants that are 27–65 mm long and with 11–17 veins, and stipules 5–28 mm long. It is very similar to P. ×‌versicolor (7 × 19; see that nothospecies for discussion). 12. Potamogeton hillii Morong

N C Fig. 295

Hill’s pondweed. Potamogeton porteri Fern. • CT, MA, VT. Shallow, still or slow-moving, circumneutral to basic water of lakes and streams. 13. Potamogeton illinoensis Morong N Fig. 294  Achene of Potamogeton gramineus showing coiling of embryo.

Illinois pondweed. Potamogeton angustifolius Bercht. & K. Presl; P. heterophyllus Schreb. • CT, MA, VT; also reported from NH by Haynes and Hellquist 2000, but specimens are

unknown. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers. Potamogeton illinoensis can be difficult to separate from P. gramineus. The former has submersed leaves with an acute and apiculate apex and with 7–15 veins and the drupes are 2.5–3.6 mm long. The latter has submersed leaves with a short-acuminate to acuminate apex and with 3–9 veins and the drupes are 1.9–2.3 mm long. ‌ × 15. Potamogeton ×faxonii Morong is a rare and difficult to detect pondweed 13 hybrid known from VT; also reported from MA by Angelo and Boufford (2000), but the specimens (at MASS!) either are misidentified (a specimen is actually P. ×‌spathuliformis) or have qualified (i.e., tentative) determinations. This hybrid is recognized by its leaves that are sessile or borne on petioles up to 35 (–55) mm long, the principal ones mostly 16–20 mm wide, and are scarcely or not mucronate at the apex. 13 × 23. This very rare pondweed hybrid is known from CT. It has sessile, broadlanceolate to narrow-ovate leaf blades that vary from not clasping to weakly clasping, have mostly 18–21 veins, the principal ones mostly 48–77 mm long. The lower stipules disintegrate into fibers.

Fig. 295  Leaf apex of Potamogeton hillii.

14. Potamogeton natans L. N Fig. 296 floating pondweed. CT, MA, ME, NH, RI, VT; throughout. Shallow, still or slow-moving, acidic to basic water of lakes and rivers, sometimes stranded on mud along tidal streams. Potamogeton natans, like many other species of pondweed, is capable of tremendous morphological variation. In still water its leaf blades are broad and subcordate at the base. In flowing or tidal water, the leaf blades become narrower and the bases tapered. 15. Potamogeton nodosus Poir. N long-leaved pondweed. Potamogeton americanus Cham. & Schlecht.; P. fluitans Roth • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers, often in circumneutral riverine systems in eastern New England and basic lacustrine or riverine systems in western New England. 16. Potamogeton oakesianus J.W. Robbins N Oakes’ pondweed. CT, MA, ME, NH, RI, VT. Shallow, acidic to basic water of lakes and ponds, less frequently in streams. 17. Potamogeton obtusifolius Mert. & Koch N

Fig. 296  Floating leaf blade and petiole of Potamogeton natans.

blunt-leaved pondweed. CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers.

Pota m o g e to n ac e a e   2 97

18. Potamogeton ogdenii Hellquist & Hilton

NC

Ogden’s pondweed. CT, MA, VT. Shallow, basic water of lakes, less frequently in slow-moving streams. Potamogeton ogdenii is believed to have originated from a cross between P. hillii and P. zosteriformis with later backcrossing to one or both parents. 19. Potamogeton perfoliatus L. N clasping-leaved pondweed. Potamogeton amplexicaulis Kar.; P. bupleuroides Fern.; P. perfoliatus L. var. bupleuroides (Fern.) Farw. • CT, MA, ME, NH, RI, VT; throughout. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers, sometimes periodically stranded on mud along tidal rivers. ‌ × 23. Potamogeton ×absconditus Z. Kaplan, Fehrer, & Hellquist is a very rare 19 pondweed hybrid known from CT, ME, VT. It is very difficult to distinguish from its parental taxa due to the close morphological similarity. It is most similar to P. perfoliatus with regard to habit and leaf blade shape, but the stipules, even though translucent and delicate, disintegrate into ± white fibers like P. richardsonii (but the fibers can be very fine and difficult to detect). 20. Potamogeton praelongus Wulfen N white-stemmed pondweed. Potamogeton praelongus Wulfen var. angustifolius Graebn. • CT, MA, ME, NH, VT. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers. 20 × 23. This rare pondweed hybrid is known from CT; also reported from MA by Angelo and Boufford (2000), but the specimen (at MASS!) has a qualified (i.e., tentative) determination and is equivocal. It is intermediate in morphology between its parental species.

Fig. 297  Leaf blade of Potamogeton richardsonii.

21. Potamogeton pulcher Tuckerman N spotted pondweed. CT, MA, ME, NH, RI; rare in northern New England. Shallow, still or slowmoving, acidic to circumneutral water of lakes and rivers, including streams and pools within acid swamps. 22. Potamogeton pusillus L. N small pondweed. P. lateralis Morong, pro parte; P. pusillus L. var. minor (Biv.) Fern. & Schub. • CT, MA, ME, RI, VT; also reported from NH by Haynes and Hellquist (2000c), but specimens are unknown. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers. See Potamogeton vasey for comments regarding P. lateralis. 23. Potamogeton richardsonii (Benn.) Rydb. N Fig. 297 Richardson’s pondweed. Potamogeton perfoliatus L. var. richardsonii Benn. • CT, MA, ME, nh, vT. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers. 24. Potamogeton robbinsii Oakes N Fig. 298 Robbins’ pondweed. CT, MA, ME, NH, RI, VT; nearly throughout. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers. 25. Potamogeton spirillus Tuckerman N Fig. 299 northern snail-seed pondweed. CT, MA, ME, NH, RI, VT; throughout. Shallow, still or slow-moving, acidic to basic water of lakes and rivers. 26. Potamogeton strictifolius Benn.

Fig. 298  Habit of Potamogeton robbinsii showing branched inflorescences.

N C Fig. 300

straight-leaved pondweed. Potamogeton strictifolius Benn. var. rutiloides Fern. • CT, MA, ME, VT. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers. ‌26 × 28. Potamogeton ×haynesii Hellquist & Crow is a rare pondweed hybrid known from VT. It closely resembles P. strictifolius and, like that species, has light-colored, fibrous stipules and usually lacks lacunar bands on leaf blades. The hybrid also has similar winter buds to those of P. strictifolius. The hybrid shows leaf blades with 7–21 veins and rarely produces fruit. Potamogeton strictifolius has leaf blades with 3–5 (–7) veins and routinely produces fruits.

Fig. 299  Basally connate stipule of Potamogeton spirillus.

29 8  MO NOCOTS

27. Potamogeton vaseyi J.W. Robbins

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Vasey’s pondweed. Potamogeton lateralis Morong, pro parte • CT, MA, ME, NH, VT. Shallow, still or slow-moving, circumneutral water of lakes and rivers. Potamogeton lateralis was believed to be an extremely rare taxon that occurred in New England. However, the original material (including the lectotype) was a mixture of P. pusillus and P. vaseyi. Potamogeton vaseyi senesces early, and the stems are typically gone by mid-August. It is very difficult to separate from P. pusillus when floating leaves are absent; however, P. vaseyi lacks nodal glands, often has narrower submersed blades, lacks lacunar bands on submersed leaves, and has turions on short, divergent, axillary branches (P. pusillus has nodal glands on at least some nodes, often has wider leaf blades (see identification key), sometimes has 1 or 2 more rows of lacunae on submersed leaves, and has turions on elongate, ascending, terminal branches). 28. Potamogeton zosteriformis Fern. N flat-stem pondweed. Potamogeton zosterifolius Schumacher ssp. zosteriformis (Fern.) Hultén • CT, MA, ME, NH, VT. Shallow, still or slow-moving, slightly acidic to basic water of lakes and rivers.

Fig. 300  Winter bud of Potamogeton strictifolius.

Stuckenia 1a. Stipular sheaths open with overlapping edges; leaf blades commonly acute at the apex, with an apiculus on young plants; stigma borne on a short style, the style persistent on the achene and forming a tiny beak on mature drupes; drupes 3–4.5 mm long . . . . . . S. pectinata 1b. Stipular sheaths closed; leaf blades commonly blunt to obtuse at the apex, sometimes retuse, rarely apiculate; stigma broad and sessile, not forming a beak on mature drupes; drupes 2–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. filiformis 1. Stuckenia filiformis (Pers.) Boerner

NC

thread-leaved false pondweed. Coleogeton filiformis (Pers.) D. Les & R. Haynes ssp. alpinus (Blytt) D. Les & R. Haynes; C. filiformis (Pers.) D. Les & R. Haynes ssp. occidentalis (J.W. Robbins) D.H. Les & Haynes; Potamogeton filiformis Pers. var. alpinus (Blytt) Aschers. & Graebn.; P. filiformis Pers. var. borealis (Raf.) St. John; P. filiformis Pers. var. macounii Morong; P. filiformis Pers. var. occidentalis (J.W. Robbins) Morong; Stuckenia filiformis (Pers.) Börner ssp. alpina (Blytt) Haynes, D.H. Les, & M. Kral; S. filiformis (Pers.) Börner ssp. occidentalis (J.W. Robbins) Haynes, D.H. Les, & M. Kral • ME, NH, VT; northern portion of states. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers. Two infraspecific taxa have traditionally been recognized in New England—a robust plant with broader leaves and sterile fruits most commonly found in moving water (occidentalis) and a smaller plant with narrower leaves and fertile fruits from still and slow-moving water (alpina). Various studies, including transplant experiments, show that much of the variation observed in Stuckenia filiformis is under environmental control (Kaplan 2008), including leaf width, stem length, and fruit production. Given these facts and other complications (e.g., the epithet alpina is based on a European type but is applied only to plants from North America and Asia by Haynes and Hellquist 2000c), S. filiformis is recognized here in the broad sense without infraspecific taxa. Note that the more robust, moving-water forms are responsible for reports of Stuckenia vaginata (Turcz.) Holub from New England (e.g., Magee and Ahles 1999). 2. Stuckenia pectinata (L.) Boerner N Sago false pondweed. Coleogeton pectinatus (L.) D.H. Les & Haynes; Potamogeton pectinatus L. • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving, circumneutral to basic water of lakes and rivers. Identification of species of Stuckenia in North America have relied too heavily on leaf blade apex shape (Kaplan 2008). Leaf sheath morphology is the most important feature for distinguishing vegetative plants.

Zannichellia 1. Zannichellia palustris L. N Fig. 301 Fig. 301  Fruit of Zannichellia palustris.

horned-pondweed. Zannichellia palustris L. var. major (Hartman) Koch • CT, MA, ME, NH, RI, VT. Primarily in fresh to brackish-tidal waters, uncommonly inland in basic waters (e.g., Lake Champlain, VT).

Ru p p i ac e a e   2 9 9

Ruppiaceae Ruppia 1. Ruppia maritima L. N beaked ditch-grass. Ruppia maritima L. var. longipes Hagstr.; R. maritima L. var. obliqua (Schur) Aschers. & Graebn.; R. maritima L. var. rostrata Agardh; R. maritima L. var. subcapitata Fern. & Wieg. • CT, MA, ME, NH, RI; also reported from VT by Magee and Ahles (1999), but specimens are unknown. Saline to brackish waters and pools.

Ruscaceae 1a. Leaves all basal (though not apparent due to sheathing nature of petioles); inflorescence a secund raceme of white flowers with drooping pedicels subtended by scarious bracts 4–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Convallaria 1b. Leaves borne on a stem [Fig. 302]; inflorescence not combining above features 2a. Tepals evidently connate and forming a tube, 10–22 mm long; inflorescence of axillary flowers or 1- to 10 (–15)-flowered racemes; berry dark blue to black; rhizome thick and knotty, mostly 10–30 mm thick [Fig. 303] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polygonatum 2b. Tepals distinct, 1.5–6 mm long; inflorescence a terminal raceme or panicle; berry initially, or ultimately becoming, red; rhizome mainly more slender, 1–14 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maianthemum

Convallaria 1. Convallaria majalis L. E European lily-of-the-valley. CT, MA, ME, NH, RI, VT. Fields, abandonded homesteads, roadsides, edges of lawns, forest openings.

Maianthemum Reference: LaFrankie (2002). 1a. Flowers with 4 tepals; leaf blades cordate at the base . . . . . . . . . . . . . . . . . . . . M. canadense 1b. Flowers with 6 tepals; leaf blades narrowed near the base [Fig. 302] 2a. Inflorescence a panicle; tepals 0.5–1 mm long, shorter than the stamens; rhizome 8–14 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. racemosum 2b. Inflorescence a raceme [Fig. 302]; tepals 2.5–6 mm long, longer than the stamens; rhizome 1–4.5 mm thick 3a. Leaves 7–11 per stem, the blades sessile and somewhat clasping at the base [Fig. 302], pubescent on the abaxial surface; peduncle of the raceme short, often shorter than 2 cm [Fig. 302] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. stellatum 3b. Leaves 2–4 per stem, the blades narrowed to a subpetiolar base, glabrous; peduncle of raceme 2–6 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. trifolium

30 0  MO NOCOTS

1. Maianthemum canadense Desf. N Canada-mayflower. Maianthemum canadense Desf. var. pubescens Gates & Ehlers; Unifolium canadense (Desf.) Greene • CT, MA, ME, NH, RI, VT; throughout. Forests, woodlands, clearings. 2. Maianthemum racemosum (L.) Link ssp. racemosum N feathery false Solomon’s-seal. Convallaria racemosa L.; Smilacina racemosa (L.) Desf.; S. racemosa (L.) Desf. var. lanceolata Boivin; Vagnera racemosa (L.) Morong; Unifolium racemosum (L.) Britt. • CT, MA, ME, NH, RI, VT; throughout. Forests, usually in mesic soils with deciduous overstories, including upland and riparian types. 3. Maianthemum stellatum (L.) Link N Fig. 302 star-like false Solomon’s-seal. Convallaria stellata L.; Smilacina stellata (L.) Desf.; Vagnera stellata (L.) Morong • CT, MA, ME, NH, RI, VT. Sandy soils of woodlands, dry fields, dunes and near coastal areas, also forest openings, swamps, and fens. 4. Maianthemum trifolium (L.) Sloboda N three-leaved false Solomon’s-seal. Convallaria trifolia L.; Smilacina trifolia (L.) Desf.; Vagnera trifolia (L.) Morong • CT, MA, ME, NH, RI, VT. Fens, bogs, laggs, often with at least a partial overstory. Fig. 302  Upper leaves and inflorescence of Maianthemum stellatum.

Polygonatum Reference: Utech (2002). 1a. Leaves glabrous on both surfaces, with 7–19 prominent veins; axillary peduncles with 2–10 (–15) flowers; distinct portion of tepals (i.e., the lobes) 4–6.5 mm long . . . . . . . P. biflorum 1b. Leaves pubescent on the veins of the abaxial surface, with 3–9 prominent veins; axillary peduncles with 1–3 (–5) flowers; distinct portion of tepals 2–3 mm long 2a. Tepals green-yellow; filaments papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pubescens 2b. Tepals white with green at apex; filaments smooth, glabrous or pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. latifolium 1. Polygonatum biflorum (Walt.) Ell. N King Solomon’s-seal. Convallaria biflora Walt., pro parte; Polygonatum biflorum (Walt.) Ell. var. commutatum (J.A. & J.H. Schultes) Morong; P. commutatum (J.A. & J.H. Schultes) A. Dietr. • CT, MA, ME, NH, VT. Rich, dry-mesic to mesic, upland forests and woodlands, riparian forests, embankments, fields, roadsides. Though native, this species is also commonly planted in New England (generally it is robust forms that are cultivated). Polygonatum biflorum is sometimes treated as being comprised of two species—a diploid (P. biflorum) and a tetraploid (P. commutatum). Though the tetraploid is usually larger (rhizomes 15–30 mm thick, stems 5–13 mm thick at the lowest leaves, peduncles 2- to 10-flowered, tepals 17–20 mm long) compared with the diploid (rhizomes 6–15 mm thick, stems 1.5–5 mm thick at the lowest leaves, peduncles 2- or 3 (–5)-flowered, tepals 10–17 mm long), some diploids have been collected that are fully as large as any tetraploid. This has led some authors to treat the entire complex as a single, variable species (which is followed here until more detailed studies are performed). That stated, local ecological differences have been observed between diploids and tetraploids, lending support to the hypothesis that plants with different ploidy levels may represent real taxa that deserve recognition at some level. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species is not naturalized in RI. 2. Polygonatum latifolium (P. Mill.) Desf. E broad-leaved Solomon’s-seal. Convallaria latifolia P. Mill.; Polygonatum hirtum (Bosc ex Poir.) Pursh • MA, NH, VT. Roadsides, forest fragments. 3. Polygonatum pubescens (Willd.) Pursh N Fig. 303

Fig. 303  Rhizome of Polygonatum pubescens.

hairy Solomon’s-seal. Convallaria pubescens Willd. • CT, MA, ME, NH, RI, VT. Mesic, deciduous forests, often found in rich soils and on rocky slopes.

S c he uc hz e r i ac e a e   3 0 1

Scheuchzeriaceae Scheuchzeria 1. Scheuchzeria palustris L. N Fig. 304 pod-grass. Scheuchzeria americana (Fern.) G.N. Jones; S. palustris L. ssp. americana (Fern.) Hultén; S. palustris L. var. americana Fern. • CT, MA, ME, NH, RI, VT. Bogs, fens.

Smilacaceae

Fig. 304  Leaf apex of Scheuchzeria palustris showing apical pore.

Smilax Smilax pulverulenta Michx. was reported from RI by Angelo and Boufford (2000). The voucher was misidentified and is S. herbacea—20 Jun 1882, Leland s.n. (NEBC!). Reference: Holmes (2002). 1a. Plants vines (i.e., stems herbaceous), without prickles [Fig. 305]; ovules 2 (rarely 1) per locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. herbacea 1b. Plants lianas (i.e., stems woody), with prickles, especially toward the base; ovule 1 per locule 2a. Leaf blades glaucous on the abaxial surface; anthers longer than filaments . . . . S. glauca 2b. Leaf blades not glaucous; anthers equal to or shorter than filaments 3a. Prickles thin, bristly, nearly black at maturity; peduncle 1.5–6.5 cm long; berry not glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hispida 3b. Prickles stout, green with red to dark brown tips at maturity; peduncle 0.5–1.5 cm long; berry glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rotundifolia 1. Smilax glauca Walt. N glaucous-leaved greenbrier. Smilax glauca Walt. var. leurophylla Blake • CT, MA, RI. Dry-mesic to wet-mesic forests, forest edges, sandy openings, roadsides. Smilax glauca is well marked in New England; it is the only woody Smilax with abaxially glaucous leaves. However, caution is needed because this character can be altered by heated drying. 2. Smilax herbacea L. N Fig. 305 carrion-flower. Coprosmanthus herbaceus (L.) Kunth; Nemexia herbacea (L.) Small • CT, MA, ME, NH, RI, VT. River banks, riparian forests, swamp edges, meadows. 3. Smilax hispida Raf.

NC

bristly greenbrier. Smilax hispida Muhl. ex Torr.; S. tamnoides L.; S. tamnoides L. var. hispida (Muhl. ex Torr.) Fern. • CT; southwestern portion of state. Mesic soils of forests, forest edges, riparian areas. 4. Smilax rotundifolia L. N common greenbrier. Smilax rotundifolia L. var. crenulata Small & Heller; S. rotundifolia L. var. quadrangularis (Muhl. ex Willd.) Wood • CT, MA, ME, NH, RI. Dry-mesic to mesic forests, woodlands, fields, and pond borders.

Fig. 305  Leaves and stipular tendrils of Smilax herbacea.

302 MO NOCOTS

Tofieldiaceae Triantha 1. Triantha glutinosa (Michx.) Baker N sticky false asphodel. Narthecium glutinosum Michx.; Tofieldia glutinosa (Michx.) Pers.; T. racemosa (Walt.) Small var. glutinosa (Michx.) Ahles • ME, NH, VT. Fens, meadows, rivershore seeps in regions of high-pH bedrock or till.

Typhaceae 1a. Spikes spherical, most or all of carpellate ones subtended by prominent, leafy bracts [Figs. 306, 307]; fruit an achene with a persistent stigma beak; leaves flaccid to firm, parallelveined and checkered with rectangular areoles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sparganium 1b. Spikes cylindric, the carpellate ones without leafy bracts; fruit a wind-dispersed follicle; leaves firm, parallel-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Typha

Sparganium Angelo and Boufford (2000) reported Sparganium americanum × S. fluctuans for MA. Though this collection may be accurately determined, it is not included here until further study is taken given that these two species below to different subgenera and Cook and Nichols (1986, 1987) did not mention the occurrence of this hybrid. References: Cook and Nicholls (1986, 1987). 1a. All or most flowers and fruits with 2 stigmas; all or most ovaries with 2 locules; body of achenes truncate or abruptly tapering to the beak, with 3–7 angles (i.e., not terete in cross-section), not constricted near center, 4–8 mm wide . . . . . . . . . . . . . . . . . . . S. eurycarpum 1b. Flowers and fruits with 1 stigma; ovary with 1 locule; body of achenes tapering to beak, terete or weakly angled in cross-section, often slightly constricted near center, up to 4 mm wide 2a. Flowering stems with 1 staminate spike; fruiting spikes 5–12 mm in diameter; beak of achene 0.5–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. natans 2b. Flowering stems with 2–20 staminate spikes; fruiting spikes 10–35 mm in diameter; beak of achene 1.5–7 mm long 3a. Beak of mature fruit flattened and strongly curved; tepals adnate to fruit stipe for ca. ½ their length; anthers (0.5–) 0.6–0.7 (–0.9) mm long; stigmas (0.4–) 0.7–0.8 (–0.9) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. fluctuans 3b. Beak of mature fruit ± terete, straight or only slightly curved (sometimes strongly curved in S. androcladum); tepals free from fruit stipe; anthers 0.8–1.5 mm long; stigmas (0.6–) 0.8–4 mm long 4a. Both the sessile carpellate spikes of the main axis and the peduncle bases of the lateral branches borne directly in the axil of leaves or bracts [Fig. 306]; tepals ± opaque, with a dark brown pad of tissue near apex, the apical margins emarginate to entire 5a. Fruiting spikes 25–35 mm in diameter; achenes usually dull in basal half and lustrous in apical half, with a beak (4–) 4.5–7 mm long; anthers 1–1.6 mm long; lateral branches of inflorescence, when present, bearing only staminate spikes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. androcladum

Ty ph ac e a e   3 03

5b. Fruiting spikes 15–25 mm in diameter; achenes usually dull throughout, with a beak (1.5–) 3–4 (–5) mm long; anthers 0.8–1.2 mm long; lateral branches of inflorescence, when present, bearing 1–3 carpellate spikes in addition to staminate spikes [Fig. 306] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. americanum 4b. Some of the sessile carpellate spikes of the main axis and/or the peduncle bases of the lateral branches borne above the axil of leaves or bracts [Fig. 307]; tepals ± translucent, without a pad of dark tissue at the apex, the apical margins distinctly erose 6a. Leaves usually erect and emergent, though sometimes flaccid and floating, the distal portion with a prominent abaxial keel or triangular in cross-section; beak of fruit 2–4.5 (–6) mm long; at least some of the staminate spikes separated by short internodes [Fig. 307] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. emersum 6b. Leaves flaccid and floating (or submerged), the distal portion flat, without an abaxial keel; beak of fruit 1.5–2 (–2.2) mm long; staminate spikes contiguous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. angustifolium 1. Sparganium americanum Nutt. N Fig. 306 American bur-reed. CT, MA, ME, NH, RI, VT; nearly throughout. Shorelines and shallow, slightly acidic to basic, still or slow-moving water. This species sometimes has flaccid leaves at the deep-water edge of a population. 2. Sparganium androcladum (Engelm.) Morong N branched bur-reed. Sparganium lucidum Fern. & Eames • CT, MA, ME, NH, RI, VT. Shorelines and shallow, circumneutral, still or slow-moving water. 3. Sparganium angustifolium Michx. N narrow-leaved bur-reed. Sparganium emersum Rehmann var. angustifolium (Michx.) Taylor & MacBryde; S. multipedunculatum (Morong) Rydb. • CT, MA, ME, NH, VT; rare or absent from some southern New England states. Shallow, circumneutral to basic, still or slow-moving water. This species rarely somewhat emergent in shallow water.

Fig. 306  Inflorescences of Sparganium americanum showing axillary carpellate spikes.

3 × 4b. This very rare bur-reed hybrid is known from NH, VT. It is intermediate in both vegetative and floral characters, and the F₁ individuals are known to be highly fertile. It has anthers 0.9–1.2 mm long and stigmas 0.8–1.4 (–1.7) mm, whereas Sparganium angustifolium has anthers (0.8–) 0.9–1 (–1.2) mm long and stigmas (0.6–) 0.8–1 (–1.2) mm long, and S. emersum has anthers 1–1.5 (–2) mm long and stigmas (1–) 1.5–2 (–2.5) mm long. 4. Sparganium emersum Rehmann N Fig. 307 simple-stemmed bur-reed.  4a. Sparganium acaule (Beeby ex Macoun) Rydb.; S. chlorocarpum Rydb. var. acaule (Beeby ex Macoun) Fern.; S. emersum ssp. acaule (Beeby ex Macoun) C.D.K. Cook & M.S. Nicholls; S. simplex Huds. var. acaule Beeby ex Macoun; 4b. Sparganium chlorocarpum Rydb. • CT, MA, ME, NH, VT. Shorelines and shallow, circumneutral to basic, still or slow-moving water. 1a. Carpellate spikes approximate to crowded, sessile (the lowest one sometimes remote and shortly pedunculate); bract subtending uppermost carpellate spike usually exceeding the uppermost staminate spike; achene body 3–4 mm long, equal to or shorter than the beak; inflorescence with 2–5 staminate spikes; anthers 0.8–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a. S. emersum var. acaule (Beeby ex Macoun) A. Haines 1b. Carpellate spikes usually remote, the lowest often pedunculate [Fig. 307]; bract subtending uppermost carpellate spike usually exceeded by the uppermost staminate spike; achene body 3.5–5.5 mm long, longer than the beak; inflorescence with 4–9 staminate spikes; anthers 1–1.5 (–2) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. S. emersum var. emersum Variety acaule is known CT, MA, ME, NH, VT. Subspecies variety is known from CT, MA, ME, NH, VT. 5. Sparganium eurycarpum Engelm. ex Gray N great bur-reed. Sparganium eurycarpum Engelm. ex Gray var. greenei (Morong) Graebn. • CT, MA, ME, NH, RI, VT. Shorelines and shallow, circumneutral to basic, still or slow-moving water.

Fig. 307  Inflorescences of Sparganium emersum var. emersum showing supra-axillary carpellate spikes.

304 MON OCOTS

6. Sparganium fluctuans (Morong) B.L. Robins. N floating bur-reed. CT, MA, ME, NH, VT. Shallow, circumneutral, still or slow-moving water. The leaves of Sparganium fluctuans are flaccid and unkeeled, as in S. angustifolium. In addition to the differences noted in the key, these two species can also be separated by the inflorescence architecture. Sparganium fluctuans frequently has at least one inflorescence branch bearing 2 or more spikes (vs. all spikes solitary, even if borne on a peduncle), and the staminate spikes are usually not contiguous (vs. contiguous). 7. Sparganium natans L. NC arctic bur-reed. Sparganium minimum (Hartman) Wallr. • CT, MA, ME, NH, VT. Shorelines, pools in peatlands, and shallow, acidic to basic, still or slow-moving water.

Typha Leaf blades shrink considerably in drying, and users of this key are warned to follow measurements carefully in the following key. Also, carpellate spike measurements are for fruiting individuals, and measurements may be narrower on individuals collected prior to maturation. Leaf mucilage glands can be difficult to detect in fresh material, so it is best to dry specimens before judging their conspicuousness. In order to confidently identify plants (especially when hybrids may be present), examination of flowers and associated micromorphological structures is crucial, where 20× or higher magnification is useful. The carpellate bracteoles, found in Typha angustifolia and its hybrids, appear as clavate-shaped structures with a truncate and thickened apex (as opposed to the carpellate flowers, which are more fusiform and taper at each end). The staminate scales are very narrow and appear to be stamen filaments on quick examination (but they do not support an anther). Hybrid plants are generally sterile and produce few or no seeds and have abortive pollen. Reference: Smith (2000). 1a. Carpellate spikes (20–) 24–36 mm thick in fruit; leaves glaucous when fresh, the widest ones 10–23 (–29) mm wide when fresh, 5–20 mm wide when dry; compound pedicels on denuded inflorescence axis bristle-like, 1.5–3.5 mm tall [Fig. 309]; carpellate bracteoles absent; stigmas persistent on mature fruit, ovate to broad-lanceolate in outline, 0.1–0.2 (–0.25) mm wide at the widest point; carpellate spike and staminate spike contiguous, rarely separated by a gap as much as 4 cm; staminate scales colorless to stramineous; mucilage glands at transition from blade to sheath usually colorless, obscure . . . . . . . . . . . . . . . . . . . . . . . T. latifolia 1b. Carpellate spikes 13–22 mm thick in fruit; leaves not glaucous, the widest ones 4–12 mm wide when fresh, 3–8 (–10) mm wide when dry; compound pedicels on denuded axis peg-like, 0.5–0.7 mm tall [Fig. 308]; carpellate bracteoles present; stigmas sometimes deciduous in fruit, narrow-lanceolate in outline, 0.04–0.08 (–0.1) mm wide at the widest point; carpellate spike and staminate spike separated by a gap of 1–8 cm; staminate scales stramineous to medium brown; mucilage glands at transition from blade to sheath brown, visible in mid- to late season . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. angustifolia Fig. 308  Compound pedicels on denuded spike axis of Typha angustifolia.

1. Typha angustifolia L. E Fig. 308 narrow-leaved cat-tail. Typha angustifolia L. var. elongata (Dudley) Wieg. • CT, MA, ME, NH, RI, VT; most abundant on the coastal plain and along heavily traveled throughways. Saline and brackish marshes, ditches near heavily salted highways. ‌1 × 2. Typha ×glauca Godr. is an often robust, cat-tail hybrid that is more common in wetlands with a history of human disturbance. It is known from CT, MA, ME, NH, RI, VT and is characterized by carpellate spikes 19–25 mm thick in fruit, usually separated from the staminate spike by a short distance, compound pedicels of the denuded axis 0.6–2 mm tall, carpellate bracteoles present (but inconspicuous and evident only at magnification), and brown staminate scales. Additionally, the stigmas are 0.06–0.11 mm wide at the widest point and usually more resemble T. angustifolia (i.e., they are relatively narrow).

Fig. 309  Compound pedicels on denuded spike axis of Typha latifolia.

2. Typha latifolia L. N Fig. 309 broad-leaved cat-tail. CT, MA, ME, NH, RI, VT; nearly throughout. Saturated soils and shallow water of lake shores, river shores, marshes, ditches, impoundments, and fens. Typha latifolia, though native, can be an aggressive colonizer, especially in areas with a history of human disturbance.

Xy r i dac e a e   3 0 5

Xyridaceae Bracts and sepals continue to grow after anthesis. The measurements used in the following key are for those bracts and sepals that subtend mature fruits. Reference: Kral (2000b).

Xyris 1a. Keel of lateral sepals firm, ciliate except at the red-fimbriate tip; leaves twisted; base of plant abruptly bulbous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. torta 1b. Keel of lateral sepals weak, scarious, entire to lacerate; leaves plane; base of plant not abruptly bulbous 2a. Sheaths minutely papillose; leaves 0.8–2 (–3) mm wide; floral scales 3–4 (–4.5) mm long, erose or minutely fimbriate along the margins, lacking a well-defined, green central region; lateral sepals red at apex, the keel entire or lacerate near the apex . . . . . . . . . . X. montana 2b. Sheaths smooth; leaves 2–15 mm wide; floral scales (4–) 5–8 mm long, entire, with a well-defined, green central area [Fig. 310]; lateral sepals not red at apex, the keel lacerate much of its length 3a. Tips of lateral sepals exserted beyond floral scales; seeds (0.6–) 0.7 (–0.8) mm long; spikes 10–20 (–25) mm tall, the apex usually rounded . . . . . . . . . . . . . . . . . . . . X. smalliana 3b. Tips of lateral sepals concealed by floral scales [Fig. 310]; seeds 0.5 mm long; spikes 5–15 mm tall, the apex usually acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. difformis 1. Xyris difformis Chapman var. difformis N Fig. 310 bog yellow-eyed-grass. CT, MA, ME, NH, RI, VT. Sandy and peaty lake and pond shores, bogs, acidic fens. 2. Xyris montana Ries N northern yellow-eyed-grass. CT, MA, ME, NH, RI, VT. Peaty pond and lake shores, bogs, acidic fens, boggy rights-of-way. 3. Xyris smalliana Nash N Small’s yellow-eyed-grass. Xyris caroliniana Walt. var. olneyi Wood; X. congdonii Small; X. smalliana Nash var. olneyi (Wood) Gleason • CT, MA, ME, RI; primarily along the coastal plain. Peaty and sandy shorelines and shallow water of ponds and lakes, lakeside floating fens. 4. Xyris torta Sm. N slender yellow-eyed-grass. Xyris bulbosa Kunth • CT, MA, NH, RI. Peaty and sandy pond and lake shores, boggy openings.

Zosteraceae Zostera 1. Zostera marina L. N eel-grass. Zostera marina L. var. stenophylla Aschers. & Graebn. • CT, MA, ME, NH, RI; coastal areas. Shallow saline water of the Atlantic Ocean, mostly in sheltered bays and inlets.

Fig. 310  Spike and expanded flower of Xyris difformis.

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3 07

Tricolpates Acanthaceae Justicia 1. Justicia americana (L.) Vahl

NC

American water-willow. Dianthera americana L. • VT. River and lake shores, backwater sloughs, swamp margins.

Actinidiaceae Actinidia 1. Actinidia arguta (Sieb. & Zucc.) Planch. ex Miq. E taravine. MA, ME. Forest margins, abandonded homesteads, roadsides.

Adoxaceae 1a. Leaf blades pinnately divided into 3–11 leaflets; drupe with 3–5 pyrenes; terminal bud absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sambucus 1b. Leaf blades simple, lobed or unlobed [Figs. 314, 315]; drupe with a single seed; terminal bud present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Viburnum

Sambucus Reference: Bolli (1994). 1a. Inflorescence with a main axis that extends beyond the lowermost branches [Fig. 311]; pith of branches orange-brown; winter buds large, ovoid; drupes red (rarely yellow) at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. racemosa 1b. Inflorescence branched from near the base and lacking a prominent main axis; pith of branches white; winter buds small, conical; drupes becoming purple-black to black at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nigra 1. Sambucus nigra L. n black elderberry. 1a. Sambucus canadensis L. • CT, MA, ME, NH, RI, VT. Swamps, wetland margins, stream banks, mesic openings; also areas of habitation and roadsides for ssp. nigra.

30 8   tricolpate s

1a. Leaf with 5–11 leaflets, usually 7; petals white in life; carpels with (3–) 4 (–5) stigmas; branchlets with sparse lenticels; drupe bright red prior to turning purple-black, 4–5 mm wide; plants up to 2.5 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. S. nigra ssp. canadensis (L.) R. Bolli 1b. Leaf with 3–7 leaflets, usually 5; petals yellow-white in life; carpels with 3 (–4) stigmas; branchlets with abundant lenticels; drupe dingy purple prior to turning black, 6–8 mm wide; plants 3–10 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. S. nigra ssp. nigra Subspecies canadensis is native and known from CT, MA, ME, NH, RI, VT. Subspecies nigra is non-native and known from CT. 2. Sambucus racemosa L. N Fig. 311 red elderberry. Sambucus pubens Michx.; S. racemosa L. ssp. pubens (Michx.) House; S. racemosa L. var. pubens (Michx.) Koehne • CT, MA, ME, NH, RI, VT. Forests, edges, forest openings, wooded slopes, ridges, edges of talus.

Viburnum Fig. 311  Inflorescence of Sambucus racemosus.

Species with 3-lobed leaf blades often will produce unlobed blades at the apex of shoot growth. Therefore, these leaves should not be used for identification purposes. Reference: McAtee (1956). 1a. Leaf blades 3-lobed (sometimes the most apical pair of the shoot with simple blades) [Fig. 314] 2a. Bud scales 3 or more, generally at least sparsely pubescent; branchlets pubescent with stellate hairs; petioles lacking glands; drupe dark blue to dark purple . . . . . . V. acerifolium 2b. Bud scales 2 (sometimes appearing as 1), glabrous; branchlets glabrous or stipitateglandular, not stellate-pubescent; petioles with raised glands; drupe red 3a. Marginal flowers of inflorescence not enlarged, fertile, similar in morphology to other flowers in the inflorescence; leaves without stipules, the blades usually toothed in the sinuses; petiole glands usually at very summit near edge of leaf blade; winter buds dark purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. edule 3b. Marginal flowers of inflorescence enlarged and neutral; leaves with slender stipules, the blades usually lacking teeth in the sinuses; petiole glands usually near apex of petiole but separated a short distance from the blade; winter buds red . . . . . . V. opulus 1b. Leaf blades not lobed [Figs. 313, 315] 4a. Winter buds naked [Fig. 312]; branchlets, winter buds, leaves, and inflorescence axes stellate-pubescent; drupes red (eventually turning very dark) 5a. Marginal flowers of inflorescence not enlarged, fertile, similar in morphology to other flowers in the inflorescence; leaf blades 5–12 cm long; pubescence gray; stems erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. lantana 5b. Marginal flowers of inflorescence enlarged and sterile; leaf blades 10–18 cm long; pubescence yellow-brown to red-brown; stems straggling or loosely ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. lantanoides 4b. Winter buds perulate; stellate hairs absent or present and then generally restricted to the leaves (abundant in V. plicatum, also on the winter buds in V. sieboldii); drupes dark blue to dark purple (red in V. dilatatum, at first pink in V. sieboldii) 6a. Leaf blades with prominent, subpalmate or pinnate, simple to twice-forked (infrequently thrice-forked) veins that extend to the tip of each tooth; winter buds with 2 or 4 visible scales 7a. Inflorescence a panicle, with opposite branches; leaf blades elliptic-obovate, cuneate at the base, with a strong smell of green pepper when bruised . . . . V. sieboldii

A d oxac e a e   3 0 9

7b. Inflorescence an umbel-like cyme, with whorled lower branches; leaf blades narrow-ovate or ovate to oblong-ovate or suborbicular, rounded to cordate at the base (rarely broad-cuneate), without a strong smell when bruised 8a. Winter buds with 2 visible, valvate outer scales 9a. Winter buds scurfy-pubescent; flowers all fertile, ca. 5 mm in diameter; leaf blades abaxially appressed-pubescent; drupe red at maturity . . . . V. dilatatum 9b. Winter buds stellate-pubescent; usually all the flowers of cyme enlarged, 15–30 mm in diameter, and sterile; leaf blades abaxially glabrate to sparsely appressed-pubescent; drupe at first red, turning blue-black (but this rarely produced in New England populations) . . . . . . . . . . . . . . . . . . . . . . . . . V. plicatum 8b. Winter buds with 4 visible scales, the outer imbricate, the inner valvate 10a. Leaves stipulate, with petioles 2–15 mm long [Fig. 315]; stylopodium pubescent; stone of drupe compressed . . . . . . . . . . . . . . . . . . V. rafinesquianum 10b. Leaves exstipulate, with petioles 8–25 mm long; stylopodium glabrous; stone of drupe plump . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. dentatum 6b. Leaf blades with many-forked, less conspicuous, pinnate veins that anastomose near the margin and do not extend to the tip of each tooth; winter buds with 2 visible, valvate scales 11a. Cymes elevated on peduncles 5–50 mm long [Fig. 313]; leaf blades entire to crenulate (rarely crenulate-serrulate) [Fig. 313]; outer bud scales ribbed with vein primordia and upon bud expansion developing into foliage leaves . . . . . . . . V. nudum 11b. Cymes sessile or rarely slightly elevated on peduncles up to 5 mm long; leaf blades serrulate; outer bud scales not ribbed, upon bud expansion either soon detaching or slightly expanding but not long persistent 12a. Some leaf blades acuminate at apex; petioles 10–20 (–35) mm long, with an undulate- or denticulate-margined wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. lentago 12b. Leaf blades acute to obtuse at apex; petioles 5–12 mm long, without or with an obscure wing margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. prunifolium 1. Viburnum acerifolium L. N maple-leaved viburnum. Viburnum acerifolium L. var. glabrescens Rehd.; V. acerifolium L. var. ovatum (Rehd.) McAtee • CT, MA, ME, NH, RI, VT. Deciduous forests and woodlands. Rare collections of this shrub show ± unlobed leaf blades (the blades with merely 3 tooth-like lobes) rather than the typical 3-lobed blade outline found in most populations. These plants were referred to as var. ovatum. 2. Viburnum dentatum L. N smooth arrowwood. 2a. Viburnum recognitum Fern.; 2b. Viburnum scabrellum Torr. & Gray ex Chapman var. venosum (Britt.) McAtee; V. venosum Britt • CT, MA, ME, NH, RI, VT. Swamps, stream banks, mesic to xyric sandy plains and woodland borders. The names of the races within this complex have been thoroughly confused and are called differently by various authors (i.e., the names have been misapplied). Therefore, one needs to examine the characters used to identify the different taxa in order to sort out the synonymy in various manuals. 1a. Abaxial leaf blade, petioles, and peduncles glabrous (at most with some simple hairs in the axils of the leaf blade veins); hypanthium eglandular . . . . . . 2a. V. dentatum var. lucidum Ait. 1b. Abaxial leaf blade (at least on the veins), petioles, and peduncles stellate-pubescent; hypanthium usually with sessile and/or stipitate-glands (rarely eglandular) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. V. dentatum var. venosum (Britt.) Gleason Variety lucidum is known from CT, MA, ME, NH, RI, VT. Variety venosum is known from MA, RI. Viburnum dentatum var. dentatum has been reported from CT, MA, and ME by McAtee (1956) and others. All specimens seen by me were either var. lucidum or var. venosum. McAtee

310  tricolpate s

noted that some plants of var. lucidum from well beyond the main range of var. dentatum were “setulose” on the adaxial surface. These plants bear only simple hairs on the leaves but, due to a letter left by McAtee in the collections at the Harvard University Herbaria, became the basis for V. dentatum var. deamii (Rehd.) Fern. in MA and ME by Seymour (1982). Seymour also followed Fernald (1950b) in treating var. venosum as synonymous with var. dentatum. It is currently unknown whether or not true V. dentatum occurs in New England. It is clear that most of the reports to this fact are based on confused synonymies and the voucher specimens are either misidentified or determined only to the level of species (and actually represent a variety other than the typical one). 3. Viburnum dilatatum Thunb. E linden arrowwood. CT, MA. Areas of habitation, forests. 4. Viburnum edule (Michx.) Raf. N squashberry. ME, NH, VT. Shorelines, wetland edges, and forests in boreal and subalpine regions. 5. Viburnum lantana L. E wayfaring tree. Viburnum lantana L. var. sphaerocarpum Gray ex Fern. • CT, MA, ME, VT. Abandoned lots, fields, roadsides. Fig. 312  Naked winter bud of Viburnum lantanoides.

6. Viburnum lantanoides Michx. N Fig. 312 hobblebush. Viburnum alnifolium Marsh.; V. grandifolium Ait. • CT, MA, ME, NH, RI, VT. Temperate to boreal, mesic to wet-mesic forests. 7. Viburnum lentago L. N nannyberry. CT, MA, ME, NH, RI, VT. Margins of fields, swamp edges, riparian forests. 8. Viburnum nudum L.

N C Fig. 313

withe-rod.  8a. Viburnum cassinoides L.; 8b. Viburnum cassinoides L. var. angustifolium (Torr. & Gray) Shinners; V. nudum L. var. angustifolium Torr. & Gray • CT, MA, ME, NH, RI, VT. Swamps, wetland margins, wet-mesic forests.

Fig. 313  Inflorescence and leaf blade of Viburnum nudum var. cassinoides.

1a. Leaf blades crenulate to nearly entire, cuneate to rounded at the base, usually bluntly short-acuminate at the apex [Fig. 313], dull on the adaxial surface, scurfy-ciliate when young with some hairs persisting on the blade at maturity, abaxially with prominent, strongly curving veins; winter buds brown to yellow-brown; peduncle of cyme 5–25 mm long at anthesis; stone of drupe elliptic or elliptic-oblong in outline, without longitudinal grooves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8a. V. nudum var. cassinoides (L.) Torr. & Gray 1b. Leaf blades entire to merely undulate, cuneate at the base, obtuse to acute at the apex, somewhat lustrous on the adaxial surface, not scurfy-ciliate, the blade glabrous at maturity, abaxially with less prominent and less curving veins; winter buds red-brown; peduncle of cyme 20–50 mm long at anthesis; stone of drupe orbicular-ovate in outline, with longitudinal grooves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8b. V. nudum var. nudum Variety cassinoides is known from CT, MA, ME, NH, RI, VT. Variety nudum is known from CT and is of conservation concern. 9. Viburnum opulus L. n Fig. 314 highbush-cranberry.  9a. Viburnum opulus L. var. americanum Ait.; V. trilobum Marsh.;  9b. Viburnum opulus L. var. roseum L. • CT, MA, ME, NH, RI, VT. Swamps, wetland margins, shorelines; also introduced to field edges, roadsides, and abandoned lots. 1a. Petiole glands stalked, taller than wide, truncate to convex at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9a. V. opulus ssp. trilobum (Marsh.) Clausen

Fig. 314  Leaf blade and petiole with concave glands of Viburnum opulus ssp. opulus.

1b. Petiole glands sessile, wider than tall, concave at the apex [Fig. 314] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9b. V. opulus ssp. opulus Subspecies trilobum is native and known from CT, MA, ME, NH, RI, VT. Subspecies opulus is non-native and known from CT, MA, ME, NH, VT. Subspecies opulus was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this taxon was not naturalized in RI.

A d oxac e a e   3 1 1

10. Viburnum plicatum Thunb. E Japanese snowball. Viburnum plicatum Thunb. var. tomentosum (Thunb.) Miq.; V. tomentosum Thunb. • MA. Abandoned lots, roadsides, fields. 11. Viburnum prunifolium L.

NC

smooth blackhaw. Viburnum bushii Ashe • CT; southwestern portion of state. Rocky forests and thickets. 12. Viburnum rafinesquianum J.A. Schultes var. rafinesquianum N Fig. 315 downy arrowwood. CT, MA, NH, VT. Woodlands and dry-mesic, deciduous forests, often in regions of moderate to high-pH bedrock. 13. Viburnum sieboldii Miq. E Siebold’s arrowwood. CT, MA. Areas of habitation, abandoned lots.

Fig. 315  Leaf of Viburnum rafinesquianum.

Aizoaceae 1a. Calyx lobes white or pink to purple adaxially, with small, horn-like, subterminal appendages; leaves opposite, the blades 1–2.5 (–3) × 0.1–0.5 cm, linear to spatulate or ovate, and cuneate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sesuvium 1b. Calyx lobes yellow adaxially, without subapical appendages; leaves alternate, the blades (0.4–) 1–10 × 2.5–8 cm, ovate-rhombic to triangular, and broad-cuneate to truncate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetragonia

Sesuvium 1. Sesuvium maritimum (Walt.) Britt.

NC

annual sea-purslane. Pharnaceum maritimum Walt.; Sesuvium pentandrum Ell. • RI. Coastal beaches, dunes, brackish marshes.

Tetragonia 1. Tetragonia tetragonioides (Pallas) Kuntze E New Zealand-spinach. Demidovia tetragonioides (Pallas) Kuntze; Tetragonia expansa Murr. • CT, MA. Waste areas, roadsides, abandoned lots.

Altingiaceae Liquidambar 1. Liquidambar styraciflua L.

NC

sweet-gum. CT, MA. Swamps and swamp margins. The MA record is based on observations by Tuckerman (and perhaps others). The plants were reported to be growing outside of cultivation, but whether they represent native or introduced populations is unknown. This occurrence was discussed by Knowlton (1916).

31 2   tricolpate s

Amaranthaceae Reference: Clemants (1992). 1a. Leaves opposite 2a. Stems appearing jointed; leaves connate into a sheath that surrounds the stem, with scarcely projecting distinct tips [Fig. 340]; flowers in groups of 3, sunken into the fleshy stem [Figs. 339, 340] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Salicornia 2b. Stems not appearing jointed; leaves neither connate nor scale-like; flowers not arranged in groups of 3, not sunken into the stem 3a. Flowers unisexual; stems and inflorescence glabrous or farinose, but not pubescent; leaves glabrous or farinose, but not pubescent; filaments distinct; fruits concealed by a pair of accrescent bracteoles [Fig. 323]; leaf blades entire or, more commonly, toothed or with a pair of lobes near the base [Fig. 325] . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Atriplex 3b. Flowers bisexual; stems sparsely to densely villous or villous-tomentose; leaves pubescent on one or both surfaces (at least on the abaxial midrib); filaments connate (at least at the base); fruits not concealed by a pair of accrescent bracteoles (though often concealed by the persistent perianth); leaf blades entire 4a. Spikes 20–28 mm in diameter, white, yellow, red, or purple, showy; style terminated by 2 subulate or filiform stigmas; plants annual, from fibrous roots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gomphrena 4b. Spikes 6–8 mm in diameter, mainly white or gray to white-green or gray-green, not especially showy; styles terminated by a single capitate stigma or the stigma minutely 2-lobed; plants annual with a taproot or perennial 5a. Filaments monadelphous, the connate tube suggesting a gamopetalous corolla; leaf blades linear to narrow-lanceolate; rachis of inflorescence moderately to densely villous-tomentose; calyx segments ± monomorphic; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Froelichia 5b. Filaments connate only at the very base; leaf blades obovate to orbicular; rachis of inflorescence glabrous or sparsely pubescent; calyx segments polymorphic—the outer 2 longest and spine-tipped, the next inner shorter and narrower, the inner 2 shortest and with hairs along the abaxial keel; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alternanthera 1b. Leaves alternate or all basal 6a. Sepals usually dry and scarious; filaments distinct or, more commonly, connate near the base; ovary with 3 stigmas (often only 2 in A. palmeri) [Fig. 318] 7a. Flowers unisexual; utricle 1-seeded; inflorescence not fasciated . . . . . . . Amaranthus 7b. Flowers bisexual; utricle 6- to 10-seeded; inflorescence fasciated . . . . . . . . . . Celosia 6b. Sepals membranaceous to herbaceous (hyaline in Axyris); filaments distinct; ovary with 2 (–5) stigmas 8a. Leaves stellate-pubescent on the abaxial surface; sepals hyaline; fruit obovate in outline, with a bilobed wing at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Axyris 8b. Leaves glabrous, farinose, or pubescent, but not stellate-pubescent; sepals usually pigmented; fruit orbicular to obovate or oblong in outline, without a bilobed wing at the apex 9a. Plants monoecious or dioecious (i.e., flowers unisexual); fruit or seed concealed by a pair of accrescent bracteoles [Fig. 323]

A m a r a n t h ac e a e   3 13

10a. Stigmas 2; herbage glabrous or sparsely to densely farinose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Atriplex 10b. Stigmas 4 or 5; herbage glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spinacea 9b. Plants synoecious (i.e., with bisexual flowers only) or sometimes polygamous (i.e., with bisexual and unisexual flowers); fruit visible or concealed by a persistent calyx, the bracteoles not accrescent 11a. Leaf blades linear to narrow-lanceolate or oblanceolate in outline, entire, sessile or subsessile (with obscure petioles up to 3 mm long in Kochia) [Fig. 338] 12a. Leaves tipped with spines (especially those from the upper portion of the stem) 13a. Flowers with usually 2 styles and 5 stamens; fruits globose to ovoid, with a horizontally oriented seed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Salsola 13b. Flowers with usually 5 styles and 3 stamens; fruits compressed, with a vertically oriented seed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polycnemum 12b. Leaves not tipped with spines, at most pointed 14a. Stems, leaves, and/or perianth villous or short-pilose with eglandular hairs 15a. Fruiting sepals with a prominent, transverse wing on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kochia 15b. Fruiting sepals without a wing, with either conical tubercles or hooked spines on the abaxial surface [Fig. 327] . . . . . . . . . . . . . . . . . . Bassia 14b. Plants glabrous or farinose or stipitate-glandular in Dysphania 16a. Leaves terete to planoconvex in cross-section; flowers borne in groups of 1–7, usually 3, in the axils of the leaves . . . . . . . . . . . . . . . Suaeda 16b. Leaves flat, at least in the basal portion; flowers borne variously, but not consistently in groups of 3 17a. Plants glabrous or farinose, but not glandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Chenopodium 17b. Plants with stipitate glands or subsessile glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Dysphania 11b. Leaf blades oblong or lanceolate to ovate or triangular, entire, toothed, or lobed, at least the lower ones petioled (caution: the lower leaves in some Chenopodium and often all the leaves in Cycloloma deciduous by fruiting) [Fig. 329] 18a. Fruiting perianth with a continuous, transverse wing; inflorescence a diffusely branched panicle with solitary flowers at each node . . . . . . Cycloloma 18b. Fruiting perianth lacking a horizontally oriented wing; inflorescence usually either a panicle with spike-like branches or of axillary clusters of flowers, the flowers occurring singly or in glomerules at each node 19a. Ovary semi-inferior; roots moderately to strongly swollen, dark red, yellow, or white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Beta 19b. Ovary superior; roots not swollen, usually pale or brown 20a. Calyx composed of 1 sepal, not enclosing the fruit at maturity; androecium composed of 1 stamen . . . . . . . . . . . . . . . . . . . . . . . . . Monolepis 20b. Calyx composed of 3–5 sepals, enclosing and often somewhat concealing the fruit at maturity; androecium composed of 1–5 stamens, commonly 5

31 4   tricolpate s

21a. Plants glabrous or farinose, but not glandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Chenopodium 21b. Plants with stipitate glands or subsessile glands  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Dysphania

Alternanthera 1. Alternanthera pungens Kunth E khaki joyweed. Achyranthes repens L.; Alternanthera repens (L.) Link • MA. Wool waste.

Amaranthus Determining the type of flowers (i.e., staminate or carpellate) on a plant or branch is very important and requires careful study because some species may show very few staminate flowers among the carpellate ones. Carpellate plants are sometimes necessary for identification of dioecious species. Hybridization has been documented in various parts of the world within Amaranthus, with A. hybridus, A. retroflexus, and A. tuberculatus most frequently involved in cross-species mating. References: Costea et al. (2001a, 2001b), Mosyakin and Robertson (2003). 1a. Plants bearing 2 rigid, sharp-pointed spines (2–) 5–11 (–25) mm long at most nodes; terminal spike with staminate flowers in apical portion and carpellate flowers near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. spinosus 1b. Plants without nodal spines; terminal spikes either entirely unisexual or with scattered flowers of each sex 2a. Plants dioecious; carpellate flowers lacking sepals or with 1 or 2 vestigial ones (or with 5 sepals in A. palmeri); fruit an indehiscent utricle or an irregularly rupturing utricle (or a pyxis in A. palmeri) 3a. Carpellate flowers with 5 spatulate sepals 2–4 mm long and often only 2 style branches; bracteoles of staminate flowers (2.5–) 3–4 mm long; fruit circumscissily dehiscent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. palmeri 3b. Carpellate flowers lacking sepals or with 1 or 2 vestigial ones shorter than 1 mm and 3 style branches; bracteoles of staminate flowers 0.6–1.5 mm long; fruit indehiscent or irregularly dehiscent (dehiscent in rare forms of A. tuberculatus) 4a. Utricle 2.2–3.5 (–4) mm long, fleshy, not dehiscent; seeds 1.9–3 mm long; bracteoles of staminate flowers 0.6–1 (–1.3) mm long, the midrib weakly excurrent; leaf blades linear to lanceolate; plants primarily of saline and brackish wetlands and shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. cannabinus 4b. Utricle 1.5–2 mm long, membranaceous, sometimes dehiscent, though irregularly in most forms; seeds 0.8–1 mm long; bracteoles of staminate flowers 1–1.5 mm long, the midrib evidently excurrent; leaf blades lanceolate to ovate [Fig. 321]; plants of fresh water wetlands and shores, gardens, dumps, and open rights-of-way . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. tuberculatus 2b. Plants monoecious; flowers with a calyx of 3–5 sepals (rarely only 2), the carpellate sepals persistent in fruit; fruit a pyxis or less commonly indehiscent 5a. Inflorescence composed entirely of small, axillary glomerules 2–11 mm in diameter [Fig. 316]; leaf blades 0.5–4.2 (–7) cm long 6a. Fruit warty-rugose, evidently exceeding the sepals . . . . . . . . . . . . A. macrocarpus 6b. Fruit neither warty nor rugose, not or only somewhat exceeding the sepals

A m a r a n th ac e a e   3 15

7a. Carpellate sepals 2.5–3 (–4) mm long; fruit indehiscent, fleshy, 2.5–4 (–5) mm long; seed oval in outline, 1.7–2.2 (–2.5) mm long; leaf blades fleshy, obovate to orbicular [Fig. 319] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pumilus 7b. Carpellate sepals 0.8–2.7 mm long; fruit circumscissilely dehiscent (indehiscent in A. blitum), dry, 0.9–2.2 (–2.5) mm long; seeds orbicular in outline, 0.7–1.6 (–1.8) mm long; leaf blades herbaceous, elliptic or oblong to obovate, rhombic-ovate, or spatulate [Fig. 316] 8a. Leaf blades shallowly to deeply retuse at the apex [Fig. 316]; fruit indehiscent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. blitum 8b. Leaf blades rounded to obtuse at the apex (rarely with an inconspicuous apical notch); fruit circumscissilely dehiscent 9a. Bracteoles of carpellate flowers 1–2.5 mm long, subequal in length with the sepals; carpellate flowers with 4 or 5 sepals 1.2–2.7 mm long; seeds 1.3–1.6 (–1.8) mm long; plants prostrate . . . . . . . . . . . . . . . . . . . . . . . A. blitoides 9b. Bracteoles of carpellate flowers 2–2.6 mm long, mostly 2–3 times as long as the sepals; carpellate flowers with 3 sepals 0.8–1.2 mm long; seeds 0.7–1 (–1.2) mm long; plants upright, bushy branched . . . . . . . . . . . . . . A. albus 5b. Inflorescence composed of a terminal spike or panicle 20–300 mm tall, as well as often axillary spikes or glomerules [Figs. 317, 320]; leaf blades 1–30 cm long, the larger usually longer than 4 cm (shorter in A. deflexus) 10a. Carpellate flowers with 2 or 3 sepals; fruit indehiscent (i.e., a utricle) 11a. Fruits rugose when dry; terminal panicle mostly 10–20 cm tall; stems erect, 2–10 dm tall; fruit 1–1.6 mm long; seed orbicular in outline . . . . . . . . . . . . . . A. viridis 11b. Fruits smooth to slightly wrinkled; terminal panicle mostly 2–5 cm tall; stems prostrate to erect, 1–4 (–5) dm tall; fruit 2.3–2.6 mm long; seed obovate in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. deflexus 10b. Carpellate flowers with (3–) 5 sepals; fruit circumscissilely dehiscent (i.e., a pyxis) 12a. Seeds dark brown to black; bracteoles exceeding the stigma branches in intact infructescences (except in A. dubius); inflorescence not showy, dull green (sometimes tinged with red in A. hybridus) 13a. Bracteoles shorter than 2 mm; styles spreading; seeds 0.8–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. dubius 13b. Bracteoles 2.5–6 (–8) mm long; styles erect to ascending; seeds 1–1.4 mm long 14a. Carpellate sepals rounded to truncate to emarginate at the apex, (2–) 2.5–3.5 (–4) mm long, outwardly curved; terminal inflorescence usually with many short, crowded, thick branches [Fig. 320] . . . . . . . . . A. retroflexus 14b. Carpellate sepals acute to subacute at the apex, 1.2–3 mm long, erect; terminal inflorescence unbranched or with a few, widely spaced, long branches or with many, crowded, slender branches [Fig. 317] 15a. Bracteoles 4.5–6 (–8) mm long, 2–3 (–4) times as long as the sepals; sepals numbering 3–5, with yellow midveins, evidently unequal, 1 sepal 2.2–3 mm long, the others 1.2–1.6 mm long; apex of fruit gradually narrowed; inflorescence with few, erect, rigid, widely spaced branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. powellii 15b. Bracteoles 2.5–4 (–6) mm long, 1.2–2 times as long as the sepals; sepals numbering 5, with green midveins, subequal or 1 longer, 1.5–2.5 (–3) mm long; fruit abruptly narrowed to apex; inflorescence with few to many, slender, lax or flexuous branches [Fig. 317] . . . . . A. hybridus

316 tricolpates

12b. Seeds white-yellow to red-brown (often dark brown in ornamental forms); bracteoles equaling or exceeded by stigma branches in intact infructescences; inflorescence often large and showy, usually red to purple (less frequently yellow or white) 16a. Terminal inflorescence with stiff, thick branches 9–14 mm wide; bracteoles subequal in length to the stigma branches in intact infructescences [Fig. 318]; stigma branches thick at the base; fruit ± truncate at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. hypochondriacus 16b. Terminal inflorescence with lax, narrower branches 4–10 (–12) mm wide; bracteoles shorter than the stigma branches in intact infructescences; stigma branches slender at the base; fruit gradually or abruptly tapering to apex 17a. Sepals narrow-oblong, acute at the apex, not overlapping with one another; stigma branches erect from the base; bracteoles 2–3 mm long, equal to or slightly longer than the sepals . . . . . . . . . . . . . . . . . . . . A. cruentus Fig. 316  Inflorescence and leaves of Amaranthus blitum.

17b. Sepals obovate to spatulate, obtuse to rounded at the apex, overlapping with one another; stigma branches spreading from the base; bracteoles 3–4 mm long, 1.2–1.5 times as long as the sepals . . . . . . . . . . . . . . A. caudatus 1. Amaranthus albus L. E tumbleweed amaranth. Amaranthus albus L. var. pubescens (Uline & Bray) Fern.; A. pubescens (Uline & Bray) Rydb. • CT, MA, ME, NH, RI, VT. Gardens, waste areas, pastures. 2. Amaranthus blitoides S. Wats. E prostrate amaranth. Amaranthus graecizans, auct. non L. • CT, MA, ME, NH, RI, VT. Waste areas, yards, gardens, railroad rights-of way. 3. Amaranthus blitum L. ssp. emarginatus (Moq. ex Uline & Bray) Carretero var. emarginatus E Fig. 316 livid amaranth. Amaranthus ascendens Loisel. var. polygonoides (Moq.) Thellung; A. blitum L. ssp. polygonoides (Zollinger ex Moq.) Carretero; A. lividus L. var. polygonoides (Moq.) Thellung ex Druce; Euxolus viridis L. var. polygonoides Moq. • MA, RI; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Sandy, disturbed areas, dumps. 4. Amaranthus cannabinus (L.) Sauer N

Fig. 317  Inflorescence of Amaranthus hybridus.

salt marsh water-hemp. Acnida cannabina L. • CT, MA, ME, NH, RI. Tidal river shores, brackish marshes, rarely inland wetlands. 5. Amaranthus caudatus L. E purple amaranth. Amaranthus edulis Spegazzini • CT, MA, VT; also reported from ME by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, waste areas. 6. Amaranthus cruentus L. E blood amaranth. Amaranthus hybridus L. var. cruentus (L.) Moq.; A. paniculatus L.; A. sanguineus L. • CT, MA, VT; also reported from ME by Campbell et al. (1995) and from NH by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, waste areas. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species was not naturalized in RI. 7. Amaranthus deflexus L. E large-fruited amaranth. MA. Wool waste.

Fig. 318  Flower of Amaranthus hypochondriacus showing relative style length.

8. Amaranthus dubius Mart. ex Thellung E spleen amaranth. MA. Waste areas, landings. 9. Amaranthus hybridus L. ssp. hybridus N Fig. 317 green amaranth. Amaranthus chlorostachys, auct. non Willd.; A. hybridus L. ssp. patulus (Bertol.) Carretero; A. patulus Bertol. • CT, MA, ME, NH, RI, VT. Fields, gardens, roadsides, waste areas, disturbed sandy soils.

A m a r a n t h ac e a e   3 17

10. Amaranthus hypochondriacus L. E Fig. 318 prince’s-feather amaranth. Amaranthus chlorostachys Willd. var. erythrostachys (Moq.) Aellen; A. flavus L.; A. hybridus L. ssp. hypochondriacus (L.) Thellung; A. leucocarpus S. Wats. • MA. Gardens, cultivated fields. 11. Amaranthus macrocarpus Benth. var. macrocarpus E dwarf amaranth. Euxolus macrocarpus (Benth.) F. Muell. • MA. Wool waste. 12. Amaranthus palmeri S. Wats. E Palmer’s amaranth. MA. Wool waste, disturbed areas. 13. Amaranthus powellii S. Wats. ssp. powellii E Powell’s amaranth. Amaranthus chlorostachys Willd. var. pseudo-retroflexus Thellung; A. retroflexus L. var. powellii (S. Wats.) Boivin • CT, MA, RI, VT; also reported from ME and NH by Mosyakin and Robertson (2003), but specimens are unknown. Fields, roadsides, railroads, disturbed soil, gardens.

N C Fig. 319

14. Amaranthus pumilus Raf.

Fig. 319  Habit of Amaranthus pumilus.

sea side amaranth. CT, MA, RI. Sandy sea beaches, frequently near the wrack line. 15. Amaranthus retroflexus L. N Fig. 320 red-rooted amaranth. Amaranthus retroflexus L. var. salicifolius I.M. Johnston • CT, MA, ME, NH, RI, VT. Gardens, fields, roadsides, waste areas, disturbed sandy soils, sandy shorelines. 16. Amaranthus spinosus L. E spiny amaranth. CT, MA, RI, VT; also reported from ME by Kartesz (1999), but specimens are unknown. Roadsides, railroads, waste areas. 17. Amaranthus tuberculatus (Moq.) Sauer

n C Fig. 321

rough-fruited water-hemp. Acnida altissima (Riddell) Moq. ex Standl.; A. tamariscina (Nutt.) Wood var. prostrata Uline & Bray; A. tuberculata Moq.; Amaranthus altissimus Riddell; A. ambigens Standl.; A. rudis Sauer; A. tuberculatus (Moq.) Sauer var. prostratus (Uline & Bray) Seymour; A. tuberculatus (Moq.) Sauer var. rudis (Sauer) Costea & Tardif; A. tuberculatus (Moq.) Sauer var. subnudus S. Wats. • CT, MA, ME, NH, VT. Fresh water wetlands and shores, gardens, dumps, and open rights-of-way. Introduced to much of New England but some populations appearing native in VT (and these of conservation concern). Two species within this complex have frequently been recognized—Amaranthus tuberculatus and A. rudis. The former is eastern and characterized by carpellate flowers with 1 or 2 sepals and indehiscent fruits, while the latter is western and characterized by carpellate flowers that lack sepals (or have vestigial ones) and dehiscent fruits (among other characters). Midwestern populations have long been known to problematic (they intergrade) and have been considered to be hybrids. However, Pratt and Clark (2001) showed that a geographic cline (east to west) exists for the morphology (i.e., the phenotypic expression cannot be described simply as two species with hybridization where they overlap). Costea and Tardif (2003a) proposed a solution that treats these two species as varieties. However, changing the rank does not alter the fact that two forms cannot be confidently identified over a significant proportion of their range. Therefore, A. tuberculatus is treated here as a single, variable entity (as suggested by Pratt and Clark).

Fig. 320  Inflorescence of Amaranthus retroflexus.

18. Amaranthus viridis L. E slender amaranth. Amaranthus gracilis Desf.; Euxolus viridis L. • MA. Roadsides, railroads, waste areas, dumps.

Atriplex The lower leaves are important for identification. Unfortunately, they are often shed by fruiting. Most species of Atriplex produce two types of fruits—brown fruits and black fruits. The brown fruits (ranging in color from brown to red-brown or yellow-brown) are larger, compressed, possess a radical, and have a dull or barely glossy, somewhat soft outer surface. The black fruits are smaller, biconvex, barely produce a radical, and have a glossy, hard outer surface. Reference: Taschereau (1972).

Fig. 321  Inflorescence of Amaranthus tuberculatus.

318 tricolpates

1a. Principal leaf blades coarsely dentate and with Kranz-type venation (i.e., with a fine reticulum of dark green veins, seen at 10× magnification after scraping surface of leaf blade) 2a. Flowers borne both in small axillary glomerules and in terminal spikes 20–50 (–110) mm long; brown fruits 1.5–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. tatarica 2b. Flowers borne in small axillary glomerules and in terminal spikes up to 20 mm long; brown fruits 2–2.5 mm long 3a. Principal leaf blades sparsely dentate and with a pair of larger teeth near the base or ± entire except for a pair of lobe-like teeth near the base; fruiting bracteoles tuberculate and echinate on the faces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sibirica 3b. Principal leaf blades coarsely dentate (rarely some ± entire); fruiting bracteoles tuberculate to ± smooth on the faces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. rosea 1b. Principal leaf blades ± entire or with a single pair of tooth-like lobes near the base of the blade (rarely the blades sparsely and irregularly dentate, particularly near the apex) and with normal dicotyledenous venation (i.e., many vein branches free and not rejoining to form a reticulum) 4a. Fruiting bracteoles obovate-orbicular to obtriangular, with 3–5 (–7) apical teeth [Fig. 323]; lower leaf blades oval or broad-oblong to broad-obovate, rounded to acute at the apex; fruits with the radicle apical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. cristata 4b. Fruiting bracteoles triangular or ovate to rhombic-triangular or rhombic, without apical teeth (though the lateral margins of the bracteoles may be toothed); lower leaf blades linear to triangular-hastate, acute to acuminate at the apex; fruits with the radicle basal or lateral 5a. Inflorescence with leafy bracts nearly to tip [Fig. 324]; black fruits rare or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. glabriuscula 5b. Inflorescence with leafy bracts only near base [Fig. 322]; black fruits common, sometimes even more abundant on a given plant than the larger brown fruits 6a. Carpellate flowers dimorphic—some with no perianth and ovate-orbicular to suborbicular bracteoles and others with a 5-merous perianth and no bracteoles; fruits trimorphic­­—vertically oriented brown fruits, vertically oriented black fruits enclosed within bracteoles, and horizontally oriented black fruits partly enclosed in a 5-merous perianth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. hortensis 6b. Carpellate flowers monomorphic, all lacking perianth but with a pair of accrescent bracteoles; fruits dimorphic—vertically oriented brown fruits and vertically oriented black fruits 7a. Lower leaf blades linear to lanceolate or narrow-oblong, lacking a basal pair of outward-pointing, tooth-like lobes 8a. Brown fruits orbicular, as long as wide, basally rounded; tip of radicle ascending to outwardly pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. littoralis 8b. Brown fruits elliptic-orbicular, wider than long, basally flattened; tip of radicle inwardly curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) A. subspicata 7b. Lower leaf blades (and often the middle ones) lanceolate or oblong to triangular-hastate, with a basal pair of outward-pointing, tooth-like lobes (caution: the lower leaves are frequently deciduous by maturation of the fruits) 9a. Brown fruits elliptic-orbicular, wider than long, basally flattened [Fig. 326]; tip of radicle inwardly curved . . . . . . . . . . . . . . . . . . . . . . . . . (in part) A. subspicata 9b. Brown fruits orbicular, as long as wide, basally rounded; tip of radicle ascending to outwardly pointed 10a. Lower leaf blades triangular-hastate with outwardly pointed basal lobes, the base truncate to subcordate [Fig. 325]; brown fruits 1.5–2.5 mm long; black fruits 1–1.5 mm wide; bracteoles with a spongy inner layer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. prostrata

Am a r a n th ac e a e   3 19

10b. Lower leaf blades broad-lanceolate to ovate or triangular with outwardly or forwardly pointed lobes, the base cuneate to broad-cuneate or rounded; brown fruits 2.5–3.5 (–4) mm long; black fruits 1.2–2.9 mm long; bracteoles lacking a spongy inner layer 11a. Obligate halophyte of coastal saltmarshes; foliage (especially the margins) and stems often conspicuously tinged with red; black fruits 2–2.9 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. acadiensis 11b. Ruderal weed, not of saltmarshes; foliage and stems green or rarely with weak red-tinging; black fruits 1.2–2 mm long . . . . . . . . . . . . . A. patula 1. Atriplex acadiensis Tascher. N Fig. 322 maritime orache. Atriplex glabriuscula Edmonston var. acadiensis (Tascher.) Welsh • CT, MA, ME, NH, RI. Saline marshes, sea beaches and strands. Long referred to as Atriplex patula, but that species a ruderal weed of inland sites.

Fig. 322  Inflorescence of Atriplex acadiensis.

2. Atriplex cristata Homb. & Bonpl. ex Willd. N Fig. 323 seabeach orache. Atriplex arenaria Nutt.; A. cristata Homb. & Bonpl. ex Willd. var. arenaria (Nutt.) Kuntze; A. pentandra (Jacq.) Standl.; A. pentandra (Jacq.) Standl. ssp. arenaria (Nutt.) Hall & Clements; Obione arenaria (Nutt.) Moq. • CT, MA, RI. Saline marshes, sea beaches and strands. This species was long known by the name Atriplex arenaria, but that name published later than (therefore, without priority over) A. cristata. 3. Atriplex glabriuscula Edmondston N Fig. 324 bracted orache. Atriplex patula L. var. oblanceolata (Victorin & Rouss.) Boivin • CT, MA, ME, NH, RI. Saline marshes, sea beaches and strands. Atriplex glabriuscula and A. prostrata are morphologically similar in some respects (e.g., usual leaf blade outline), frequently grow together, and possess a similar chromosome number. Therefore, it is no surprise that these two species are sometimes confused, especially when A. glabriuscula shows variation in the proportion of the inflorescence that possesses leafy bracts. In addition to the characters used in the key, they can be separated by the size of the brown fruits and length of the carpellate bracteoles. In A. glabiuscula, they are 2.5–3.9 mm wide and 5–13 mm long, respectively. In A. prostrata, they are 1.5–2.5 mm wide and 3–5 mm long, respectively.

Fig. 323  Carpellate bracteoles of Atriplex cristata.

4. Atriplex hortensis L. E garden orache. Atriplex hortensis L. var. atrosanguinea hort.; A. nitens Schkuhr • CT, MA. Gardens, waste areas. 5. Atriplex littoralis L. E linear-leaved orache. Atriplex patula L. var. littoralis (L.) Gray • MA, ME, NH; also reported from CT and RI by Seymour (1982), but specimens are unknown. Saline marshes, sea beaches and strands. All specimens of this species seen by me from ct are either misidentified or equivocal (specimens at CONN! and NEBC!).

Fig. 324  Inflorescence of Atriplex glabriuscula.

6. Atriplex patula L. E spearscale orache. CT, MA, ME, NH, VT. Fields, roadsides, waste areas, ballast. Atriplex patula has long been considered a coastal halophyte. However, it is actually a non-native weed of human-disturbed habitats. Previous reports of A. patula from Atlantic coast shores marshes have been based on A. acadiensis. 7. Atriplex prostrata Bouchér ex DC. n Fig. 325 hastate-leaved orache. Atriplex patula L. var. hastata, auct. non (L.) Gray; A. patula L. var. triangularis (Willd.) Rauschert; A. triangularis Willd. • CT, MA, ME, NH, RI, VT. Saline marshes, sea beaches and strands, rarely inland along waste areas and railroads (e.g., VT). Coastal populations are believed to be native, whereas inland populations in North America are likely non-native and may have originated from Europe. See comments under Atriplex glabriuscula for difficult specimens.

Fig. 325  Lower leaf blade of Atriplex prostrata.

32 0   tricolpate s

8. Atriplex rosea L. E red orache. MA; also reported from RI by Magee and Ahles (1999), but specimens are unknown. Roadsides, ballast, railroads. 9. Atriplex sibirica L. E Siberian orache. Obione muricata Gaertn.; O. sibirica (L.) Fisch. • MA. Wool waste. 10. Atriplex subspicata (Nutt.) Rydb.

Fig. 326  Achene of Atriplex subspicata.

N C Fig. 326

saline orache. Atriplex patula L. var. subspicata (Nutt.) S. Wats; Chenopodium subspicatum Nutt. • CT, MA, ME, NH; coastal regions; also reported from RI by Welsh (2003), but specimens are unknown. Saline marshes, sea beaches and strands, infrequently inland in disturbed soil. The leaf blades of Atriplex subspicata vary in that they may possess forward- or outwardpointing lobes or the leaf blades can be entire (which is more typical in New England). Formerly more abundant in eastern MA, but development in the coastal region has likely had an impact on populations of this species. 11. Atriplex tatarica L. E Tatarian orache. CT, ma; also reported from NH by Welsh (2003), but specimens are unknown. Waste areas and ballast near the coast.

Axyris 1. Axyris amaranthoides L. E Russian-pigweed. MA, ME. Waste areas, barn yards, roadsides.

Bassia 1a. Perianth segments each with a curved spine on the abaxial surface [Fig. 327]; leaf blades flat, linear to lanceolate or narrow-elliptic; axis of inflorescence straight . . . . . B. hyssopifolia 1b. Three (rarely only 2) of the perianth segments with a conical (i.e., non-hooked) projection on the abaxial side; leaf blades semicircular in cross-section, linear to filiform; axis of inflorescence flexuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. hirsuta 1. Bassia hirsuta (L.) Aschers. E hairy smotherweed. Chenopodium hirsutum L.; Kochia hirsuta (L.) Nolte • CT, MA, NH, RI; coastal counties. Saline marshes and sea beaches, usually in areas with a history of human habitation and/or disturbance. 2. Bassia hyssopifolia (Pallas) Kuntze E Fig. 327 five-horned smotherweed. Kochia hyssopifolia (Pallas) Roth; Salsola hyssopifolia Pallas; Suaeda hyssopifolia (Pallas) Pallas • MA, ME, NH; coastal counties. Waste areas, disturbed sea beaches.

Fig. 327  Flower of Bassia hyssopifolia.

Beta 1. Beta vulgaris L. E beet. Beta hortensis Mill. • CT, MA, ME, NH, VT; also reported from RI by Shultz (2003), but specimens are unknown. Fields, roadsides, dumps. Beta vulgaris is sometimes divided into two taxa—ssp. vulgaris (the cultivated type) and ssp. maritima (L.) Arcang. Shultz (2003) noted the work of various taxonomists who had documented a morphological continuum between B. vulgaris ssp. vulgaris and B. vulgaris ssp. maritima. Therefore, infraspecific taxa are not recognized here.

A m a r a n th ac e a e   3 2 1

Celosia 1. Celosia cristata L. E crested cock’s-comb. Celosia argentea L. var. cristata (L.) Kuntze • CT; also reported from RI by Robertson (2003), but specimens are unknown. Fields, roadsides, waste areas, dumps. Celosia cristata is a cultivated species that likely has as a wild progenitor C. argentea. These two species are sometimes combined (under the name C. argentea). However, they differ morphologically and cytologically. The report from VT was based on a collection by Blanchard of a cultivated plant (specimen at HNH).

Chenopodium Size, orientation, surface patterns, and nature of the pericarp (e.g., adherent, loose) are all very important characters of the fruits. Most species of goosefoots have an adherent pericarp that covers the lustrous, often black, seed in fruit. Sometimes the shiny surface of the seed is visible in places where the pericarp has been abraded due to pressing and storage. This is very different from the utricle-type fruit where the pericarp loosens, splits, and small flakes or sheets detach from the seed. Though many species can be confidently identified while in flower, some groups of taxa require mature fruits for determination. Reference: Wahl (1954). 1a. Fruits arranged vertically or both vertically and horizontally in the calyx; calyx with 3–5 sepals [Fig. 333] 2a. Plants perennial from a thick, woody root system; leaf blades entire except for the basal pair of outwardly pointing lobes; achenes 1.5–2 mm wide; styles persistent on the fruit, 0.8–1.5 mm long; calyx with 4 or 5 sepals . . . . . . . . . . . . . . . . . . . C. bonus‑henricus 2b. Plants annual from a taproot; leaf blades dentate or serrate (sometimes ± entire in C. rubrum var. humile); achenes 0.5–1 mm wide; styles inconspicuous in fruit, shorter than 0.8 mm; calyx with 3 sepals 3a. Principal leaf blades lanceolate to elliptic or oblong, conspicuously farinose on the abaxial surface; glomerules of flowers 1.8–2.5 mm wide; most fruits horizontally oriented, some vertically oriented [Fig. 333] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. glaucum 3b. Principal leaf blades triangular to rhombic-ovate (rarely oblong), green or sparsely farinose on the abaxial surface; glomerules of flowers 2–12 mm wide; most or all fruits vertically oriented 4a. Glomerules of flowers 2–5 mm wide; sepals at maturity herbaceous (somewhat fleshy in var. humile); horizontal seeds frequently present; vertical seeds with a basal radicle; plants coastal halophytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rubrum 4b. Glomerules of flowers (3–) 5–12 mm wide; sepals at maturity fleshy; fruits all vertically oriented; horizontal seeds absent; vertical seeds with an apical radicle; plants inland weeds of forest openings and disturbed areas 5a. Glomerules without subtending bracteal leaves (except sometimes the lower ones of each branch); seeds with convex sides, not grooved around periphery; flowers maturing from apex of inflorescence toward base . . . . . . . . . . C. capitatum 5b. Glomerules subtended by bracts; seeds with a flattened groove around the periphery; flowers maturing from base of inflorescence toward apex . . . . C. foliosum 1b. Fruits arranged horizontally in the calyx; calyx with 5 sepals [Figs. 328, 334, 335] 6a. Sepals flat or with an inconspicuous keel on the abaxial surface, at maturity conforming to the outline of the fruit and the outline of the calyx appearing ± circular (except where individual sepals spread or reflex away from fruit) [Fig. 334]; stems, leaf blades, and sepals usually glabrous and green (plants sometimes sparsely farinose on some surfaces, especially when young)

32 2   tricolpate s

7a. Leaf blades entire, or rarely with a single tooth on each margin; stems quadrangular; sepals apiculate at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. polyspermum 7b. At least the lower leaf blades with prominent teeth; stems terete to angled, but not quadrangular; sepals retuse or obtuse to acute at the apex, without an apiculus 8a. Pericarp evidently separating from mature seed [Fig. 332]; leaf blades lanceolate to oblong-ovate, entire or with short, ascending teeth . . . . . . . . . . . C. standleyanum 8b. Pericarp firmly attached to weakly separating from mature seed; leaf blades ovate or rhombic-ovate to triangular, sinuate-dentate to coarsely dentate 9a. Flowers developing asynchronously (i.e., flowers and fruits will be present in the same glomerule until late season when only fruits are present); achenes 1.5–2.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. simplex 9b. Flowers developing synchronously (i.e., all the flowers in a glomerule of similar stage); achenes 1–1.5 mm wide 10a. Achenes 1–1.2 mm wide, with rounded margins; inflorescence typically much longer and standing well above the principal foliage leaves . . . . . . . . . C. urbicum 10b. Achenes 1.2–1.5 mm wide, with acutely keeled margins; inflorescence typically shorter than to shortly exceeding the principal foliage leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. murale 6b. Sepals keeled on the abaxial surface, not conforming to the fruit, the calyx appearing pentagonal or star-shaped [Fig. 328]; stems, leaf blades, and sepals moderately to densely farinose (foliage sometimes ± glabrous in forms of C. album) 11a. Pericarp of mature fruits brown, minutely echinate (view at 20×), eventually separating and flaking away, leaving the lustrous, black seed [Fig. 332] 12a. Leaf blades predominantly narrow-ovate, the larger usually wider than 10 mm and with some form of dentition or lobing; native plants of high-pH outcrops and woodlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. foggii 12b. Leaf blades linear to lanceolate or oblong, (2–) 4–10 mm wide, entire to weakly lobed; naturalized plants of disturbed, often saline, soils . . . . . . . . . . . . . C. pratericola

Fig. 328  Fruiting calyx of Chenopodium album.

11b. Pericarp of mature fruits pale to dark, smooth, irregularly roughened, or cellularreticulate, remaining firmly attached to the seed [Figs. 330, 335] 13a. Achenes with acutely keeled margins; leaf blades irregularly dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. murale 13b. Achenes with rounded margins; leaf blades entire to serrate or infrequently dentate 14a. Principal leaf blades ± linear, 1–3 mm wide, with a single vein, entire; achenes 0.8–1 (–1.1) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. leptophyllum 14b. Leaf blades narrow-lanceolate to rhombic, ovate, or triangular, 5–75 (–90) mm wide, pinnately veined, entire to, more commonly, toothed or lobed; achenes 0.9–2 mm wide 15a. Achenes (1–) 1.3–2 mm wide, with a conspicuously cellular-reticulate pericarp [Fig. 330]; a minute style base (up to 0.2 mm long) sometimes present below the stigma branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. berlandieri 15b. Achenes 0.9–1.5 (–1.6) mm wide, with a smooth or obscurely marked pericarp; style base absent, the 2 stigma branches sessile at the summit of the ovary

Fig. 329  Inflorescence of Chenopodium berlandieri var. bushianum.

16a. Sepals of many flowers spreading to reflexed at maturity, not covering the oval achene [Fig. 335]; lower leaf blades usually serrate with small, low teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. strictum

Am a r a n t h ac e a e   3 23

16b. Sepals curved over the orbicular achene and concealing it (rarely some flowers with exposed fruits) [Fig. 328]; lower leaf blades variable, usually prominently serrate or with tooth-like lobes . . . . . . . . . . . . . . . . . . . . . C. album 1. Chenopodium album L. E Fig. 328 white goosefoot. Chenopodium album L. var. lanceolatum (Muhl. ex Willd.) Coss. & Germ.; C. lanceolatum Muhl. ex Willd.; C. missouriense Aellen; C. opulifolium Schrad. ex Kock. & Ziz; C. viride L. • CT, MA, ME, NH, RI, VT; throughout. Roadsides, gardens, cultivated fields, waste areas, occasionally in saline communities. Chenopodium album typically occurs in one of two intergrading forms in New England. One form (C. album sensu stricto) is a farinose plant with yellow-green leaf blades, those from the lower and middle portion of the plant ± ovate and dentate or nearly 3-lobed. Another form (C. lanceolatum) is a sparsely farinose plant with dark green leaf blades, those from the lower and middle portion of the plant ± lanceolate and with few or no teeth. However, numerous intergrading individuals make it difficult to separate some collections. Further, it is still not completely understood how much of a role that habitat plays into the morphology expressed by a given plant. Therefore, a broad interpretation of the species boundaries of C. album is taken here. 2. Chenopodium berlandieri Moq.

Fig. 330  Achene of Chenopodium berlandieri.

n C Fig. 329, 330, 331

pit-seeded goosefoot.  2a. Chenopodium berlandieri Moq. ssp. zschackei (J. Murr) Zobel;  2b. Chenopodium bushianum Aellen; 2c. Chenopodium macrocalycium Aellen • CT, MA, ME, NH, RI, VT. Saline marshes, sea beaches and strands, roadsides, field edges, ledges, and disturbed areas. 1a. Achenes 1–1.4 mm wide, with a prominent ± yellow area around the style base . . . . . . . . . . . . . . . . . . . . . . 2a. C. berlandieri var. zschackei (J. Murr) J. Murr in Urban & Graebn. 1b. Achenes 1.3–2 mm wide, lacking a prominent yellow area around the style base 2a. Achenes 1.7–2 mm wide; branches of inflorescence elongate, often arching to drooping [Fig. 329]; lower leaf blades 6–10 (–15) cm long; plants mostly 70–150 cm tall, mostly of disturbed soil, particularly in regions of cultivation, sometimes near the coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. C. berlandieri var. bushianum (Aellen) Cronq.

Fig. 331  Inflorescence of Chenopodium berlandieri var. macrocalycium.

2b. Achenes 1.3–1.7 (–1.9) mm wide; branches of inflorescence erect [Fig. 331]; lower leaf blades 3–4 (–6) cm long; plants rarely taller than 50 cm, of coastal beaches and marshes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2c. C. berlandieri var. macrocalycium (Aellen) Cronq. Variety zschakei is non-native and known from MA. Variety bushianum is native, is of conservation concern, and known from CT, MA, ME, NH, RI, VT. Variety macrocalycium is native and known from CT, MA, ME, RI. 3. Chenopodium bonus-henricus L. E perennial goosefoot. CT, MA, ME, RI. Fields, waste areas, disturbed soil. 4. Chenopodium capitatum (L.) Aschers. var. capitatum N strawberry-blite. Blitum capitatum L. • CT, MA, ME, NH, VT. Abandoned lots, fields, clearings. 5. Chenopodium foggii H.A. Wahl

Fig. 332  Utricle of Chenopodium foggii showing detaching pericarp.

N C Fig. 332

Fogg’s goosefoot. CT, MA, ME, NH, VT. Woodlands, ledges, and talus in regions of high-pH bedrock. 6. Chenopodium foliosum (Moench) Aschers. E leafy goosefoot. Blitum virgatum L.; Chenopodium virgatum (L.) Ambrosi • MA, ME. Fields, waste areas, disturbed soil. 7. Chenopodium glaucum L. ssp. glaucum E Fig. 333 oak-leaved goosefoot. Chenopodium glaucum L. ssp. euglaucum Aellen • CT, MA, ME, NH, RI, VT. Roadsides, waste areas, gardens, disturbed soil. 8. Chenopodium leptophyllum (Moq.) Nutt. ex S. Wats. E narrow-leaved goosefoot. Chenopodium album L. var. leptophyllum Moq. • ME. Railroads, waste areas.

Fig. 333  Calyx and vertically oriented fruit of Chenopodium glaucum.

32 4   tricolpate s

9. Chenopodium murale L. E nettle-leaved goosefoot. CT, MA, ME, RI, VT. Roadsides, waste areas, gardens, disturbed soil. 10. Chenopodium polyspermum L. E many-seeded goosefoot. 10b. Chenopodium polyspermum L. var. obtusifolium Gaud. • MA, ME. Roadsides, waste areas, gardens, disturbed soil. 1a. Leaf blades acute at the apex; inflorescence relatively congested, consisting of small cymes or spike-like cymes from the axils of leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10a. C. polyspermum var. acutifolium (Sm.) Gaud. 1b. Leaf blades obtuse at the apex; inflorescence relatively open, consisting of lax, diffuse cymes mostly from the axils of branches . . . . . . . . 10b. C. polyspermum var. polyspermum Variety acutifolium is known from MA, ME; also reported from VT by Hultén and Fries (1986), but specimens are unknown. Variety polyspermum is known from MA; also reported from ME by Kartesz (1999), but specimens are unknown. 11. Chenopodium pratericola Rydb. E desert goosefoot. Chenopodium desiccatum A. Nels. var. leptophylloides (J. Murr) H.A. Wahl; C. pratericola Rydb. var. leptophylloides (J. Murr) Aellen • CT, MA, ME, NH, RI, VT. Roadsides, railroads, waste areas, disturbed soil, sometimes in areas of high salinity. Reports of Chenopodium desiccatum A. Nelson from New England (e.g., Seymour 1982, as C. pratericola var. oblongifolium (S. Wats.) Wahl) are based on small and/or narrow-leaved specimens of C. pratericola. 12. Chenopodium rubrum L. n red goosefoot.  12a. Chenopodium humile Hook. • CT, MA, ME, NH. Saline marshes; ditches and other disturbed sites in var. humile. 1a. Mature achenes 0.8–1 mm in diameter; sepals somewhat fleshy; inflorescence consisting of axillary glomerules; plants spreading-ascending, 3–18 (–25) cm tall; lower leaves subentire or nearly so, the blade 0.7–3 cm long . . . . . . . . . . . 12a. C. rubrum var. humile (Hook.) S. Wats. 1b. Mature achenes 0.5–0.6 mm in diameter; sepals herbaceous; inflorescence consisting, at least in part, of leafy-bracted spikes; plants erect, 30–60 (–80) cm tall; blades of lower leaves toothed, up to 15 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12b. C. rubrum var. rubrum Variety humile is non-native and known from MA, ME. Variety rubrum is native and known from CT, MA, ME, NH; also reported from RI by George (1992), but specimens are unknown. 13. Chenopodium simplex (Torr.) Raf. N Fig. 334

Fig. 334  Fruiting calyx and horizontally oriented fruit of Chenopodium simplex.

giant-seeded goosefoot. Chenopodium gigantospermum Aellen; C. hybridum L. var. gigantospermum (Aellen) Rouleau; C. hybridum L. var. simplex Torr. • CT, MA, ME, NH, RI, VT. Woodlands, ledges, and talus, often in areas of high-pH bedrock, roadsides, fields, waste areas, gardens, dumps. Chenopodium simplex appears to have two very different ecological forms in New England. One of these occurs in natural communities and the other occurs in anthropogenic communities. The former is primarily known from northern and western New England, whereas the latter is primarily known from southern New England. Though the ecological differences suggest two taxa may be involved, preliminary review of herbarium specimens has failed to find any substantive morphological differences. 14. Chenopodium standleyanum Aellen E Standley’s goosefoot. Chenopodium boscianum, auct. non Moq.; C. gigantospermum Aellen var. standleyanum (Aellen) Aellen; C. hybridum L. var. standleyanum (Aellen) Fern. • CT, MA, VT. Roadsides, railroads, waste areas. Chenopodium standleyanum has been over-reported in New England based on misapplication of the name C. boscianum. Many older collections labeled as C. boscianum are actually records of the native C. foggii (both C. standleyanum and C. foggii share the separating pericarp on mature fruits). Though native to North America, C. standleyanum is only known from human-disturbed habitats in New England and is very likely introduced to the region.

Am a r a n t h ac e a e   3 25

15. Chenopodium strictum Roth ssp. glaucophyllum (Aellen) Aellen & Just. E Fig. 335 oval-seeded goosefoot. Chenopodium strictum Roth var. glaucophyllum (Aellen) H.A. Wahl • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads. This taxon is considered native to North America by many authors; however, it is known almost exclusively from humandisturbed habitats in New England and was considered adventive in the northern part of its range (which includes New England) by Wahl (1954). 16. Chenopodium urbicum L. E city goosefoot. CT, MA, ME, NH, RI, VT. Roadsides, waste areas, gardens, disturbed soil. Chenopodium urbicum can be confused with C. murale. In addition to characters stated in the key, C. urbicum has lustrous achenes with rounded margins and anthers 0.2–0.3 mm long (vs. dull achenes with acutely keeled margins and anthers 0.4–0.5 mm long in C. murale).

Cycloloma 1. Cycloloma atriplicifolium (Spreng.) Coult. E winged-pigweed. Cyclolepis platyphylla (Michx.) Moq.; Cycloloma platyphylla (Michx.) Moq.; Kochia atriplicifolia Spreng.; Salsola platyphylla Michx. • CT, MA. Roadsides, fields, disturbed sandy areas.

Dysphania Formerly included in Chenopodium, the genus Dysphania is considered to be generically distinct on the basis of indument. References: Wahl (1954), Clemants and Mosyakin (2003). 1a. Calyx connate except at the very apex, prominently reticulate-veiny, completely enclosing the mature fruit [Fig. 337]; leaves not aromatic, the blades lacking glands . . . . . . . D. multifida 1b. Calyx distinct or connate only at base, not reticulate-veiny, enclosing the fruit or not at maturity; leaves aromatic, the blades with either resin glands or glandular hairs (very rarely lacking both) 2a. Fruits mostly vertically oriented in the calyx, oval in outline, 0.5–0.7 mm wide 3a. Sepals with a fimbriate keel on the abaxial (i.e., outer) surface; glomerules of flowers 4–6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cristata 3b. Sepals lacking a keel on the abaxial surface; glomerules of flowers 1.2–2.8 (–3.5) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. pumilio 2b. Fruits mostly horizontally oriented in the calyx, suborbicular or oval in outline, 0.6–1 mm wide 4a. Leaf blades 1.3–4.5 cm long, the lower typically pinnately lobed; pericarp of fruit often with translucent white strips or blotches, lacking glands, firmly adherent to seed; gynoecium with 2 stigmas; flowers mostly solitary, many with pedicels 5a. Sepals acute to acuminate at the apex, glandular-pubescent, lacking tubercles; cymes densely arranged on raceme-like branches . . . . . . . . . . . . . . . . . . . . . . . D. botrys 5b. Sepals acute to obtuse, with sessile glands, with a single tubercle on the abaxial surface; cymes diffusely arranged on raceme- to panicle-like branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. graveolenst 4b. Larger leaf blades (2–) 4.5–8 (–12) cm long, serrate to laciniately toothed; pericarp of fruits lacking translucent white marks, glandular, not adherent to seed and separating at maturity; gynoecium with 3 stigmas; flowers in small glomerules, sessile 6a. Glomerules of flowers subtended by obvious bracts; lower leaf blades typically serrate; plants flowering in summer through fall . . . . . . . . . . . . . . . . . . . D. ambrosoides

Fig. 335  Fruiting calyx of Chenopodium strictum.

32 6   tricolpate s

6b. Glomerules of flowers lacking bracts throughout or with very small bracts shorter than the glomerules [Fig. 336]; lower leaf blades typically laciniately serrate; plants flowering in late summer through fall . . . . . . . . . . . . . . . . . . . . . . . . . . . D. anthelmintica 1. Dysphania ambrosioides (L.) Mosyakin & Clemants E Mexican-tea. Ambrina ambrosioides (L.) Spach; Chenopodium ambrosioides L.; Teloxys ambrosioides (L.) W.A. Weber • CT, MA, ME, NH, RI, VT. Roadsides, waste areas, disturbed soil, sometimes introduced in bird seed. 2. Dysphania anthelmintica (L.) Mosyakin & Clemants E Fig. 336 wormseed. Chenopodium ambrosioides L. var. anthelminticum (L.) Gray; C. anthelminticum L. • MA. Roadsides, waste areas, disturbed soil. 3. Dysphania botrys (L.) Mosyakin & Clemants E Jerusalem-oak. Botrydium botrys (L.) Small; Chenopodium botrys L.; Teloxys botrys (L.) W.A. Weber • CT, MA, ME, NH, RI, VT. Roadsides, waste areas, gardens, disturbed soil. Fig. 336  Inflorescence of Dysphania anthelmintica.

4. Dysphania cristata (F. Muell.) Mosyakin & Clemants E crested glandular-goosefoot. Blitum cristatum F. Muell.; Chenopodium cristatum (F. Muell.) F. Muell. • MA. Wool waste. 5. Dysphania graveolens (Willd.) Mosyakin & Clemants E fetid glandular-goosefoot. Chenopodium graveolens Willd.; Teloxys graveolens (Willd.) W.A. Weber • MA, ME. Roadsides, waste areas, gardens, disturbed soil. 6. Dysphania multifida (L.) Mosyakin & Clemants E Fig. 337 cut-leaved glandular-goosefoot. Chenopodium multifidum L.; Roubieva multifida (L.) Moq.; Teloxys multifida (L.) W.A. Weber • ma. Roadsides, waste areas, gardens, disturbed soil. 7. Dysphania pumilio (R. Br.) Mosyakin & Clemants E

Fig. 337  Fruiting calyx of Dysphania multifida.

clammy glandular-goosefoot. Chenopodium carinatum, auct. non R. Br.; C. pumilio R. Br.; Teloxys pumilio (R. Br.) W.A. Weber • CT, MA, RI. Roadsides, waste areas, gardens, disturbed soil. This species has often been referred to as Chenopodium carinatum R. Br. in regional herbaria. However, that name properly belongs to a separate species that has not been collected in New England now called Dysphania carinata (R. Br.) Mosyakin & Clemants.

Froelichia The report of Froelichia arizonica Thornb. ex Standl. from MA by Bean et al. (1958) was based on specimens of Froelichia gracilis. 1. Froelichia gracilis (Hook.) Moq. E slender cotton-weed. Oplotheca gracilis Moq. • CT, MA, NH, RI, VT. Roadsides, fields, waste areas, railroads.

Gomphrena 1. Gomphrena globosa L. E common globe-amaranth. MA. Waste areas, dumps.

Kochia 1. Kochia scoparia (L.) Schrad. E Fig. 338

Fig. 338  Inflorescence of Kochia scoparia.

summer-cypress. Bassia scoparia (L.) A.J. Scott; Chenopodium scoparium L.; Kochia scoparia (L.) Schrad. var. pubescens Fenzl; K. scoparia (L.) Schrad. var. subvillosa Moq. • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Roadsides, fields, waste areas, railroads, ballast.

A m a r a n th ac e a e   3 27

Monolepis 1. Monolepis nuttalliana (J.A. Schultes) Greene E Nuttall’s poverty-weed. MA, ME. Roadsides, waste areas.

Polycnemum 1. Polycnemum majus L. E giant needle-leaf. NH. Roadsides, railroads, sandy, disturbed soil.

Salicornia In recent years, it has become customary to separate the annual species of Salicornia (Salicornia s.s.) from the perennial species (Sarcocornia), a separation that is also supported by details of the flowers and their position. However, Sarcocornia has been shown to be composed of two separate clades with Salicornia derived from within it (i.e., the genus Sarcocornia is not monophyletic; Kadereit et al. 2007). Therefore, an inclusive Salicornia is recognized here that includes annual and perennial species. Measurements in the key are based on fresh material. References: Wolff and Jefferies (1987), Ball (2003a, 2003b). 1a. Plants perennial,l with woody rhizomes; all 3 flowers of each cluster inserted at the same level [Fig. 339] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. ambigua 1b. Plants annual, without rhizomes; the middle flower of each cluster conspicuously elevated above the 2 lateral flowers [Fig. 340] 2a. Leaf and scale apex acute to acuminate, with a prominent mucro; inflorescence 4.5–6.2 mm thick, notably wider than the stem; flowers all concealed by bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. bigelovii 2b. Leaf and scale apex rounded to acute [Fig. 340], without a mucro; inflorescence 2.9–5 mm thick, usually of similar thickness to that of the stem; central flower exceeding the bract and visible 3a. Inflorescences cylindric to long-tapering, the terminal with (5–) 7–23 (–25) fertile segments; scarious margin of leaves 0.3–0.4 mm wide; flowers with exserted stamens; fertile segments cylindric [Fig. 340] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. depressa 3b. Inflorescences swollen and rounded near apex, the terminal with (3–) 5–10 (–14) fertile segments; scarious margin of leaves narrower than 0.3 mm; flowers not exserting stamens, or sometimes, but then after dehiscence; fertile segments widened in the apical portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. maritima 1. Salicornia ambigua Michx. N Fig. 339

Fig. 339  Flowers and nodes of Salicornia ambigua.

perennial glasswort. Salicornia virginica, auct. non L. • CT, MA, NH, RI. Saline marshes and coastal strands. This species has long been called Salicornia virginica L. in New England herbaria and literature. The type specimen of S. virginica is clearly annual and cannot be used for this perennial species (Ball 2003b). This species has also, more recently, been known by the name S. perennis P. Mill., but that species is restricted to the Old World. 2. Salicornia bigelovii Torr. N dwarf glasswort. Salicornia mucronata Bigelow • CT, MA, ME, NH, RI. Saline marshes, usually in areas of sparse and/or low vegetation. 3. Salicornia depressa Standl. N Fig. 340 common glasswort. CT, MA, ME, NH, RI. Saline marshes, usually in areas of sparse and/or low vegetation. This species is usually referred as Salicornia europaea L., a species restricted to the Old World. The name S. virginica L. may actually be the earliest name for this plant, but the specimens are vegetative and cannot be determined with confidence (Ball 2003a).

Fig. 340  Flowers and nodes of Salicornia depressa.

32 8   tricolpate s

4. Salicornia maritima Wolff & Jefferies

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sea glasswort. Salicornia prostrata, sensu Standley (1916) • ME. Saline marshes, usually in areas of sparse and/or low vegetation. Reports of this species in New England are based on Standley (1916) and his interpretation of the name Salicornia prostrata. Its actual presence in the region requires confirmation.

Salsola Reference: Mosyakin (2003). 1a. Leaf blades linear, fleshy in life, 1–2 mm wide in drying, gradually narrowed to a firm, spine tip 0.5–1.5 mm long; plants of Atlantic coast shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. kali 1b. Leaf blades narrow-linear to filiform, not or only scarcely fleshy in life, narrower than 1 mm wide drying, abruptly narrowed to a relatively soft, pointed tip; plants of human-disturbed or eroded soils 2a. Inflorescence usually interrupted in the basal half, at maturity with reflexed bracts that are not imbricate and ± abruptly narrowed to a spinulose or mucronulate apex; sepals usually with a transverse, membranous wings; fruiting calyx 4–10 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tragus 2b. Inflorescence relatively dense, with contiguous spikes, at maturity with appressed bracts that are imbricate and gradually narrowed to a spinulose apex; sepals without a transverse wing or with a narrow, erose wing; fruiting calyx 2–5 mm in diameter . . . . . . . . . . . . S. collina 1. Salsola collina Pallas E slender saltwort. MA, NH, VT. Roadsides, cultivated fields, railroads, waste areas. 2. Salsola kali L. E saltwort.  2b. Salsola kali L. var. caroliniana (Walt.) Nutt.; S. kali L. var. pontica Pallas; S. pontica (Pallas) A. Degen • CT, MA, ME, NH, RI. Atlantic coast beaches, saline marshes, sandy areas near the coast, rarely inland in disturbed areas. 1a. Sepals with a distinct midvein prolonged into a rigid, spine tip; bracteoles not swollen, distinct; stems usually papillose to hispid (rarely glabrous) . . . . . . . . 2a. Salsola kali ssp. kali 1b. Sepals with an obscure midvein and a weak apex; bracteoles swollen, connate at the base; stems usually glabrous (infrequently papillose to hispid) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. Salsola kali ssp. pontica (Pallas) Mosyakin Subspecies kali is known from CT, MA, ME, NH, RI. Subspecies pontica is known from CT, MA, NH, RI, VT. 3. Salsola tragus L. E prickly saltwort. Salsola kali L. var. angustifolia Fenzl; S. kali L. var. tenuifolia Tausch ex Moq.; S. kali L. ssp. tragus (L.) Aellen • CT, MA, ME, NH, RI, VT. Roadsides, cultivated fields, railroads, waste areas.

Spinacea 1. Spinacea oleracea L. E spinach. ct, mA, ME, NH, RI. Waste areas, gardens, dumps.

Am a r a n t h ac e a e   3 2 9

Suaeda Reference: Bassett and Crompton (1978). 1a. Sepals rounded on the abaxial surface, ± equal in size [Fig. 342]; utricle 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. maritima 1b. Sepals keeled on the abaxial surface or 1 or 2 of them larger and corniculate-appendaged; utricle 1–1.5 mm long 2a. Sepals unequal at maturity, 1–3 of them larger and corniculate-appendaged; anthers 0.3–0.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. calceoliformis 2b. Sepals ± equal in size, keeled at maturity [Fig. 341]; anthers 0.2–0.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. linearis 1. Suaeda calceoliformis (Hook.) Moq.

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American sea-blite. Dondia americana (Pers.) Britt. in Britt. & Brown; D. calceoliformis (Hook.) Rydb.; D. depressa, auct. non (Pursh) Britt.; Suaeda americana (Pers.) Fern.; S. maritima (L.) Dumort. var. americana (Pers.) Boivin • CT, MA, ME, NH, RI. Saline marshes, coastal beaches.

Fig. 341  Fruiting calyx of Suaeda linearis showing keels on sepals.

2. Suaeda linearis (Ell.) Moq. N Fig. 341 annual sea-blite. Dondia linearis (Ell.) Heller • CT, MA, ME, NH, RI. Saline marshes, coastal beaches. 3. Suaeda maritima (L.) Dumort.

n C Fig. 342

herbaceous sea-blite.  3a. Chenopodium maritimum L.; Dondia maritima (L.) Druce;  3b. Suaeda richii Fern. • CT, MA, ME, NH, RI. Saline marshes, coastal beaches. 1a. Utricle 1.5–2 mm long; plants prostrate to erect; leaf blades 10–30 (–50) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. S. maritima ssp. maritima 1b. Utricle 1–1.5 mm long; plants prostrate and forming mats up to 50 cm in diameter; leaf blades 3–15 mm long . . . . . . . . . . . 3b. S. maritima ssp. richii (Fern.) Bassett & C.W. Crompton Suaeda maritima ssp. maritima is non-native and known from CT, MA, ME, NH, RI. Subspecies richii is native, of conservation concern, and known from MA, ME, NH.

Anacardiaceae 1a. Leaves simple, entire; fruiting panicle with long-plumose, sterile pedicels . . . . . . . . Cotinus 1b. Leaves pinnately compound, the leaflets entire or, more commonly, toothed; fruiting panicle without long-plumose sterile pedicels (though the fertile pedicels may be short-pubescent) 2a. Drupes red, conspicuously pubescent; inflorescence terminal or borne laterally on the previous season’s branchlets, the flowers and fruits crowded; leaves with 7–31 leaflets or only 3 leaflets in R. aromatica; petals often pubescent on the adaxial surface . . . . . . . . Rhus 2b. Drupes white to yellow, glabrous or sparsely pubescent; inflorescence axillary, relatively more open; leaves with 3–13 leaflets; petals glabrous . . . . . . . . . . . . . . . . . . . . Toxicodendron

Cotinus 1. Cotinus coggygria Scop. E European smoketree. Rhus cotinus L. • CT, MA, NH, VT. Roadsides, waste areas.

Fig. 342  Fruiting calyx of Suaeda maritima showing unkeeled sepals.

33 0   tricolpate s

Rhus 1a. Leaves with 3 leaflets, the terminal leaflet elliptic to rhombic-ovate and coarsely dentate in the apical half [Fig. 343]; inflorescence of spike-like, lateral clusters from the previous season’s branchlet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. aromatica 1b. Leaves with 7–31 leaflets, the terminal leaflet lanceolate to oblong or narrow-ovate and either ± entire or serrulate throughout; inflorescence a terminal panicle 2a. Leaflets entire or with a few low, undulate teeth near the apical margin; rachis of leaf winged in distal portion; bud less than half encircled by the leaf scar . . . . . . . R. copallinum 2b. Leaflets serrulate; rachis of leaf not winged; bud nearly encircled by the leaf scar 3a. Branchlets and petioles densely hirsute; drupes pubescent with hairs 1–2 mm long that are ± pointed at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. hirta 3b. Branchlets and petioles glabrous, the branchlets sometimes glaucous; drupes pubescent with hairs ca. 0.2 mm long that are rounded at the apex . . . . . . . . . . R. glabra 1. Rhus aromatica Ait. var. aromatica n Fig. 343 fragrant sumac. Rhus canadensis Marsh.; Schmaltzia crenata (P. Mill.) Greene; S. aromatica Desv.; Toxicodendron crenatum P. Mill. • CT, MA, VT; also reported from NH by Kartesz (1999), but specimens are unknown. Woodlands, ledges, and balds. This species occurs (or occurred) as a native in CT and VT and is considered introduced to MA. 2. Rhus copallinum L. var. latifolia Engl. N Fig. 343  Leaf of Rhus aromatica.

winged sumac. CT, MA, ME, NH, RI, VT. Woodlands, sandy fields and openings, roadsides. 3. Rhus glabra L. N smooth sumac. Rhus borealis Greene • CT, MA, ME, NH, RI, VT. Roadsides, forest edges, dry fields. 3 ‌ × 4. Rhus ×pulvinata Greene is a rare sumac hybrid known from MA, ME, NH. It is recognized by its short-pubescent branchlets (the pubescence much shorter than that on the branchlets of R. hirta). The branches in this hybrid are glabrous (in R. hirta, the branches do not become glabrous until the 3rd year or later). 4. Rhus hirta (L.) Sudworth N staghorn sumac. Datisca hirta L.; Rhus typhina L. • CT, MA, ME, NH, RI, VT. Roadsides, forest edges, dry fields.

Toxicodendron Reference: Gillis (1971). 1a. Leaves with 7–13 entire leaflets; inflorescence 7.5–20 cm long; stamens exserted from corolla; buds sessile, covered by scales; plants erect shrubs 2–7 m tall . . . . . . . . . . . . . T. vernix 1b. Leaves with 3 entire to toothed leaflets; inflorescence 1.5–8 (–10) cm long; stamens included within corolla; buds usually stalked, naked; plants straggling to erect shrubs up to 1 m tall or lianas 2a. Stems straggling to erect, without aerial roots; leaves tending to be clustered near the tip of the stem, often slightly folded; inflorescence simple or few-branched, usually with fewer than 25 flowers; drupes glabrous, sessile or subsessile . . . . . . . . . . . . . . . . T. rydbergii 2b. Stems climbing or trailing-straggling by aerial roots; leaves alternately scattered throughout the stem, usually flat; inflorescence normally branched, usually with more than 25 flowers; drupes sparsely pubescent, scabrous, or papillose (usually glabrous in ssp. negundo), pedicellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. radicans

An ac a r d i ac e a e   3 3 1

1. Toxicodendron radicans (L.) Kuntze

NC

poison-ivy.  1a. Toxicodendron aboriginum Greene; T. negundo Greene; T. radicans (L.) Kuntze var. negundo (Greene) Reveal; 1b. Rhus radicans L.; R. toxicodendron L. var. radicans (L.) Torr.; Toxicodendron vulgare P. Mill. • CT, MA, ME, NH, RI, VT; extending into the northern states primarily along the coastal plain and the Lake Champlain valley. Roadsides, forests, coastal areas. 1a. Leaflets softly pubescent across abaxial surface; petioles pubescent; drupes usually glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. T. radicans ssp. negundo (Greene) Gillis 1b. Leaflets glabrous on the abaxial surface except along main veins and tufts of hairs in the axils of the veins; petioles glabrous; drupes pubescent, scabrous, or papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. T. radicans ssp. radicans Subspecies radicans is known from CT, MA, ME, NH, RI, VT. Subspecies negundo is known from VT and is of conservation concern. 2. Toxicodendron rydbergii (Small ex Rydb.) Greene N western poison-ivy. Rhus radicans L. var. rydbergii (Small ex Rydb.) Rehd.; Toxicodendron radicans (L.) Kuntze var. rydbergii (Small ex Rydb.) Erskine • CT, MA, ME, NH, RI, VT; becoming rare in southern New England. Roadsides, forests, cliff bases, talus. 3. Toxicodendron vernix (L.) Kuntze N poison-sumac. Rhus vernix L. • CT, MA, ME, NH, RI, VT. Forested and shrub swamps, wetland margins.

Apiaceae The concept of an expanded Apiaceae (i.e., one that includes the Araliaceae) has always been challenged. However, recognition of a monophyletic Araliaceae s.s. forces inclusion of some “Apioid” genera within the Araliaceae, and also requires recognition of the Myodocarpaceae, an Australasian group that combines morphological characters of both the Apiaceae and Araliaceae. In light of such evidence, one inclusive family is recognized here—Apiaceae. Schizocarp compression is an important morphological character in this family. How compression is referenced varies by author. Those species that have schizocarps compressed parallel to the commissure are sometimes referred to as “dorsally compressed.” Those species that have schizocarps compressed perpendicular to the commissure are sometimes referred to as “laterally compressed.” 1a. Gynoecium with 2–5 carpels, maturing as a drupe or a berry with 2–5 pyrenes or seeds, respectively; plants woody or herbaceous 2a. Plants lianas, with aerial roots; leaves evergreen . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hedera 2b. Plants trees, shrubs, or herbs, without aerial roots; leaves deciduous 3a. Leaf blades 2 or more times pinnately divided; gynoecium with usually 5 carpels, therefore, the fruit with 5 pyrenes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aralia 3b. Leaf blades palmately lobed or once palmately divided; gynoecium with 2 or 3 carpels, therefore, the fruit with 2 or 3 pyrenes or seeds (up to 5 carpels and seeds in Eleutherococcus) 4a. Plants herbaceous, unarmed, with a single whorl of leaves near the summit of the stem; petals imbricate in bud; styles distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . Panax 4b. Plants woody, armed, with many alternate leaves; petals valvate in bud; styles basally connate

33 2   tricolpate s

5a. Leaf blades palmately lobed; petiole 8–50 cm long . . . . . . . . . . . . . . . Kalopanax 5b. Leaf blades palmately divided; petiole up to 12 cm long . . . . . Eleutherococcus 1b. Gynoecium with 2 carpels, maturing as schizocarp that separates into 2 mericarps; plants herbaceous 6a. Inflorescence composed of dense, head-like clusters of ± sessile flowers [Fig. 349], each flower subtended by a bractlet; most leaf blades spinulose-toothed . . . . . . . Eryngium 6b. Inflorescence dense to open, with pedicellate flowers (sessile in some Hydrocotyle) [Fig. 347], the bractlets, when present, forming a whorl at the base of the umbellet; leaf blades toothed or not, but the teeth not spinulose 7a. Bractlets of umbel connate basally, white to pink, usually equal to or longer than the umbel they subtend; schizocarps covered with bladdery, triangular scales . . . . Astrantia 7b. Bractlets of umbel (when present) distinct, green, and/or shorter than umbel, commonly with all three traits; schizocarps without scales (though sometimes with hairs, bristles, or prickles) 8a. Umbels simple [Fig. 351]; stems prostrate, creeping over the substrate 9a. Leaves with orbicular to ovate-orbicular blades, the blades crenate to inconspicuously lobed, not hollow, without septa [Fig. 351] . . . . . . . . . Hydrocotyle 9b. Leaves phyllodial, linear, entire, hollow, each with 4–6 transverse septa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lilaeopsis 8b. Umbels compound [Figs. 347, 348, 358]; stems ascending to erect 10a. Stem leaves simple and entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bupleurum 10b. Stem leaves divided (simple and toothed in the tidal form of Sium suave) 11a. Leaf blades palmately divided into 3–7 broad segments; plants polygamous, with staminate flowers in separate umbellets or intermixed with the bisexual flowers; ovary and fruit with uncinate bristles [Figs. 355, 356] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sanicula 11b. Leaf blades compound to decompound; plants mostly synoecious (polygamous in Taenidia); ovary and fruit glabrous or provided with hairs, bristles, or prickles, but these not hooked at the tip (except in Torilis) 12a. Ovary and fruit pubescent, bristly, or prickly 13a. Ultimate segments of leaf blades narrow-linear to filiform, up to 1 mm wide, all entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cuminum 13b. Ultimate segments of leaf blades linear to ovate or obovate, wider than 1 mm, often toothed 14a. Bractlets at base of umbellets bilobed at apex; mericarp with a quadrangular body 6–15 mm long, tipped by an elongate, flattened beak 20–70 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scandix 14b. Bractlets simple or pinnately dissected or absent; mericarps 2–20 mm long, without a beak or with a short, inconspicuous beak 1–2 mm long 15a. Bracts of inflorescence pinnately dissected [Fig. 348]; central flower of inflorescence usually purple . . . . . . . . . . . . . . . . . . . . Daucus 15b. Bracts of inflorescence simple or absent; central flower of inflorescence white, colored similar to the other flowers

Ap i ac e a e   3 3 3

16a. Inflorescence with 15–150 primary branches; upper leaf sheaths dilated, wider than 10 mm; schizocarps strongly compressed parallel with the commissure 17a. Principal leaf blades once divided into 3 (rarely as many as 7) segments [Fig. 350], the segments again lobed and 5–30 (–60) cm long; schizocarp 7–12 mm long, with promiment oil tubes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Heracleum 17b. Principal leaf blades 2 or 3 times pinnately compound, the ultimate segments 2–4 (–7) cm long; schizocarp 4–7 mm long, with inconspicuous oil tubes. . . . . . . . . . (in part) Angelica 16b. Inflorescence with 2–15 (–20) primary branches; upper leaf sheaths not conspicuously dilated and narrower than 10 mm; schizocarps weakly compressed perpendicular to the commissure 18a. Ultimate segments of leaf blades narrower than 10 mm; ovary and fruit with stiff, thin, ascending to spreading prickles [Fig. 358] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Torilis 18b. Ultimate segments of leaf blades wider than 10 mm; ovary and fruit with hairs or appressed-ascending bristles [Figs. 352, 353] 19a. Principal leaf blades twice pinnately compound; schizocarps 10–22 mm long, narrow-clavate in outline [Figs. 352, 353] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Osmorhiza 19b. Principal leaf blades simple to once pinnately compound; schizocarps 3–5 mm long, ovate in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Pimpinella 12b. Ovary and fruit glabrous 20a. Principal stem leaf blades with ultimate segments wider than 10 mm, frequently wider than 15 mm, the segments clearly recognizable and often of uniform shape, not further dissected into narrow lobes 21a. Upper leaf sheaths dilated, wider than 10 mm [Fig. 350]; marginal flowers of umbel enlarged, zygomorphic, the outer petals often bifid; schizocarp 7–12 mm long, with conspicuous oil tubes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Heracleum 21b. Upper leaf sheaths not dilated, narrower than 10 mm; marginal flowers of umbel not or only slightly enlarged, the outer petals entire or bifid, similar to the inner petals; schizocarp 2–8 mm long, without conspicuous oil tubes (except in Pastinaca) 22a. Schizocarp 1.5–2 mm long; umbels mostly sessile or shortly pedunculate from the axils of leaves (some terminal ones with evident peduncles also present) . . . . . . . . . . . . . . . . . . . . . . . . . . Apium 22b. Schizocarp 2–10 mm long; umbels, whether lateral or terminal, mostly with evident peduncles 23a. Principal leaf blades with 3 leaflets, the leaflets sometimes with conspicuous lobes but not again divided (the basal leaves with simple blades in Thaspium) 24a. Flowers with white petals and minute sepals; primary branches of umbel distinctly unequal in length [Fig. 347]; basal leaf blades compound; schizocarp 5–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptotaenia

33 4   tricolpate s

24b. Flowers with purple petals and small, but evident, sepals; primary branches of umbel ± equal in lengh; basal leaf blades simple; schizocarp 3.5–5 mm long . . . . . . . . . . . . . . . . Thaspium 23b. Principal leaf blades with more than 3 leaflets 25a. Principal leaf blades (or the emersed ones in Sium) once pinnately compound 26a. Basal leaf blades simple, cordate at base; central flower of each umbellet sessile, the others pedicelled; stylopodium wanting . . . . . . . . . . . . . . . . . . . . . . (in part) Zizia 26b. Basal leaf blades compound (simple in a tidal form of Sium suave); all the flowers pedicellate; stylopodium present 27a. Petals yellow; schizocarp conspicuously compressed parallel to the commissure [Fig. 354], with conspicuous lateral wings much wider than the abaxial ribs . . . . . . . . . . . . . . . . . . . . (in part) Pastinaca 27b. Petals white; schizocarp weakly compressed perpendicular to the commissure [Fig. 357], the lateral ribs not evidently wider than the abaxial ribs 28a. Leaflets of emersed leaves linear to narrowovate, with veins not consistently directed to teeth or sinuses; bractlets at base of umbellets present; native plants of wetlands, when submersed with bi- or tripinnately dissected lower leaf blades . . . . . . . . . . Sium 28b. Leaflets ovate to suborbicular, the veins generally directed to the tips of teeth or lobes; bractlets at base of umbellets usually absent; introduced plants of fields and disturbed ground, not producing highly dissected blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Pimpinella 25b. Principal leaf blades two or three times pinnately compound 29a. Leaflets entire; umbellets near margin of inflorescence bearing bisexual flowers around the periphery and staminate flowers toward the middle, the umbellets near the center of the inflorescence entirely staminate . . . . . . . . . . . . . . . Taenidia 29b. Leaflets toothed; umbellets not varying as to sex of flowers 30a. Central flower of each umbellet sessile, the others pedicelled; stylopodium wanting . . . . . . . . . . . . (in part) Zizia 30b. All the flowers pedicelled; stylopodium present 31a. Central rachis of leaf prominently foliaceous-winged below the terminal triad of leaf segments . . . . . . Falcaria 31b. Central rachis of leaf not wing-margined 32a. Veins of the leaf blade segments directed to the sinuses [Fig. 346]; stems hollow and cross-septate near the base; sepals well-developed, triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cicuta 32b. Veins of the leaf blade segments mostly directed to the tips of the teeth or lobes; stems solid or hollow at base, but without cross septa; sepals minute or obsolete (present in Ligusticum)

Ap i ac e a e   3 3 5

33a. Ultimate segments of leaf blades usually entire in the basal ⅓ to ½; bractlets at base of umbellet mostly numbering 6–10 34a. Petals yellow to green-yellow; sepals obsolete; schizocarp conspicuously compressed parallel to the commissure; introduced plants of fields and disturbed ground . . . . . . . . Levisticum 34b. Petals white; sepals well-developed, triangular; schizocarp weakly compressed perpendicular to the commissure; native plants of Atlantic coast beaches . . . . . . . . . Ligusticum 33b. Ultimate segments of leaf blades ± uniformly serrate or serrulate; bractlets at base of umbellet absent or few (present in Angelica lucida) 35a. Upper leaf sheaths diliated, wider than 10 mm; umbels with 18–50 primary branches; native plants . . . . . . . . . . . . . . . (in part) Angelica 35b. Upper leaf sheaths not dilated, narrower than 10 mm; umbels with 12–25 primary branches; introduced plants, sometimes invasive 36a. Petals yellow; schizocarp 5–7 mm long, conspicuously compressed parallel to the commissure [Fig. 354], with deciduous or inconspicuous styles . . . . (in part) Pastinaca 36b. Petals white; schizocarp 3–4 mm long, compressed perpendicular to the commissure, with persistent, reflexed styles extending back to near midlength of fruit [Fig. 344] . . . . . . . . . . . . . . . . . . . . Aegopodium 20b. Principal stem leaf blades with ultimate segments narrower than 10 mm (frequently narrower than 6 mm), the segments less organized and the limits of each not always obvious 37a. Ultimate segments of leaf blades narrow-linear to filiform, up to 1 mm wide, all entire (the lower leaves in Coriandrum with broader segments); mature schizocarps spicy or pleasantly aromatic (except Bifora and Ptilimnium) 38a. Bracts at base of primary umbel often pinnately divided into filiform segments; sepals present, ± equal in size; native plants of saline marshes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ptilimnium 38b. Bracts at base of primary umbel absent or present and simple; sepals none or present in Coriandrum and then distinctly unequal in size; spice plants escaped from cultivation (Bifora foliage unpleasantly aromatic) 39a. Petals yellow; umbel with (10–) 15–40 primary branches; styles short and stout in anthesis, up to 0.5 mm long excluding stylopodium; bractlets at base of umbellets absent 40a. Principal leaves with petiolar sheaths 1–2.5 (–3) cm long; schizocarp compressed parallel with commissure, with prominent lateral wings; plants annual . . . . . . . Anethum

33 6   tricolpate s

40b. Principal leaves with petiolar sheaths 3–9 (–10) cm long; schizocarp subterete or slightly compressed perpendicular to the commissure, the lateral wings no more prominent than those of the abaxial surface; plants short-lived perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Foeniculum 39b. Petals white to pink; umbel with 3–14 (–16) primary branches; styles slender, 0.5–2 mm long excluding stylopodium; bractlets (at least 1) usually present at base of umbellets 41a. Schizocarp nearly biglobose, each mericarp subglobose, faintly rugulose-patterned; umbel with 3–8 primary branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bifora 41b. Schizocarp not biglobose, each mericarp hemispherical or narrower, ridged or ribbed, but not rugulose; umbel commonly with 8 or more primary branches or less in Coriandrum, but that species with the lower leaf blades merely lobed with relatively broad segments 42a. Umbel with 3–7 (–8) primary branches; schizocarp subglobose, tardily separating; marginal flowers of umbel with sepals and conspicuously enlarged petals 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coriandrum 42b. Umbel with (6–) 8–14 (–16) primary branches; schizocarp oblong to elliptic in outline, readily separating into 2 mericarps; marginal flowers of umbel ± similar to the central ones, without sepals and with petals not over 2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carum 37b. Ultimate segments of leaf blades linear to ovate or obovate, wider than 1 mm, often toothed; mature schizocarps rank-scented or weakly aromatic (strongly aromatic in Myrrhis) 43a. Schizocarps 15–25 mm long; primary branches of umbel dimorphic (especially in fruit), those bearing only staminate flowers thinner and shorter than those bearing bisexual flowers . . . . . Myrrhis 43b. Schizocarps 1.5–10 mm long; primary branches of umbel ± monomorphic 44a. Schizocarp with a smooth body, lacking ribs or wings (only the apical beak ribbed in A. sylvestris) [Fig. 345]; bractlets at base of umbellets narrow-ovate (linear in A. cerefolium), with conspicuously ciliate or fimbriate margins . . . . . . Anthriscus 44b. Schizocarp with a ribbed and/or winged body; bractlets setaceous to lanceolate, not or only minutely ciliate, or absent 45a. Plants bearing vegetative bulbils on the upper parts, these subtended by broad-based, acuminate-tipped bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cicuta 45b. Plants without bulbils 46a. Stems spotted with red-purple; bractlets subtending the umbellets lanceolate to ovate or triangular, conspicuously wider than the pedicels they subtend . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conium 46b. Stems not spotted with red-purple; bractlets setaceous to narrow-lanceolate or narrow-oblanceolate, scarcely, if at all, wider than the pedicels they subtend (narrow-lanceolate to lanceolate and somewhat wider in Peucedanum)

Ap i ac e a e   3 37

47a. Petals yellow to green-yellow; schizocarp somewhat compressed perpendicular to the commissure, with thin ribs narrower than the intervals between them . . . . . . . . . . . . . . . . . . . . . . . . . Petroselinum 47b. Petals white; schizocarp subterete or conspicuously compressed parallel to the commissure, the thick ribs wider than the intervals between them 48a. Primary branches of umbel very unequal, the outer branches 2–3 times (or more) longer than the central branches; outer flowers of umbel zygomorphic; schizocarp subterete . . . . . . . . Aethusa 48b. Primary branches of umbel only somewhat unequal, the outer branches shorter than to as much as 2 times as long as the central branches; outer flowers of umbel not or scarcely zygomorphic; schizocarp somewhat to conspicuously compressed parallel to the commissure 49a. Schizocarp suborbicular in outline, with low, blunt abaxial ribs; bractlets subtending the umbellets narrow-lanceolate to lanceolate; stems hollow . . . . . . . . . . . . . . . . . . . . . Peucedanum 49b. Schizocarp elliptic to oblong or ovate-oblong in outline, with prominently raised, thin-edged, abaxial ribs; bractlets subtending the umbellets linear to narrow-lanceolate; stems solid 50a. Branches of inflorescence glabrous; schizocarp (4.5–) 5–6 (–8) mm long; native plants of swamps, stream banks, and riparian forests . . . . . . . . . . . . . . . . . . . . . . . . Conioselinum 50b. Branches of inflorescence puberulent on the angles; schizocarp 3–4 mm long; non-native plants of waste areas and yards . . . . . . Selinum

Aegopodium 1. Aegopodium podagraria L. E Fig. 344 bishop’s goutweed. Aegopodium podagraria L. var. variegata Bailey • CT, MA, ME, NH, RI, VT. Roadsides, lawn edges, forests, especially riparian types, abandoned lots.

Aethusa 1. Aethusa cynapium L. E fool’s-parsley. CT, MA, ME, RI. Roadsides, waste areas, gardens.

Anethum 1. Anethum graveolens L. E dill. CT, MA, ME, RI. Roadsides, waste areas, gardens.

Angelica 1a. Upper leaves with the sheath shorter than the blade; lateral ribs of schizocarp thick and corky; plants growing at upper edge of or just behind Atlantic coast beaches . . . . . . . A. lucida

Fig. 344  Schizocarp with recurved styles of Aegopodium podagraria.

33 8   tricolpate s

1b. Upper leaves with the sheath longer than the blade or the blade ± absent; lateral ribs of schizocarp thin and wing-like; plants not of seashores 2a. Ovary puberulent; ultimate segments of leaf blades 2–6 (–7) cm long, with an obtuse apex; pericarp tightly adherent to seed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. venenosa 2b. Ovary glabrous; ultimate segments of leaf blades (4–) 8.5–12 (–15) cm long, with an acute apex; pericarp loose from seed at maturity . . . . . . . . . . . . . . . . . . . . . . A. atropurpurea 1. Angelica atropurpurea L. N purple-stemmed Angelica. CT, MA, ME, NH, RI, VT. Wetland margins, river banks, fields, roadsides; most common in regions of high-pH bedrock. 2. Angelica lucida L. N sea coast Angelica. Coelopleurum actaeifolium (Michx.) Coult. & Rose; C. gmelinii (DC.) Ledeb.; C. lucidum (L.) Fern.; C. lucidum (L.) Fern. ssp. gmelinii (DC.) A. & D. Löve • CT, MA, ME, NH, RI. Upper margin of sea beaches, fields and forest edges adjacent to coastal beaches. Frequently confused with Ligusticum scoticum in regional collections. Angelica lucida has puberulent primary umbel branches, obsolete sepals, and leaf blade segments toothed nearly to base, whereas L. scoticum has glabrous primary umbel branches that are sometimes glandularwarty near apex, evident sepals, and basally entire leaf blade segments. 3. Angelica venenosa (Greenway) Fern.

NC

hairy Angelica. Angelica villosa (Walt.) B.S.P. • CT, MA. Dry-mesic to xeric, sandy or rocky soils of fields, woodlands, and slopes.

Anthriscus 1a. Umbel with 2–6 pubescent primary branches; beak of schizocarp prominent, narrower than and mostly 25–33% as long as the body; bracteoles linear; plants annual; ultimate segments of leaf blades mostly 5–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. cerefolium 1b. Umbel with (4–) 6–15 glabrous primary branches; beak of schizocarp short, nearly as thick as and mostly 16–25% as long as the body [Fig. 345]; bracteoles narrow-ovate; plants biennial or perennial; ultimate segments of leaf blades mostly 15–50 mm long . . . . . . . . . . A. sylvestris 1. Anthriscus cerefolium (L.) Hoffmann E garden chervil. Cerefolium cerefolium (L.) Britt.; Scandix cerefolium L. • CT, VT. Roadsides, gardens, waste areas. 2. Anthriscus sylvestris (L.) Hoffmann E Fig. 345 wild chervil. Chaerophyllum sylvestre L. • CT, MA, ME, RI, VT. Roadsides, field margins, disturbed areas. Fig. 345  Schizocarp of Anthriscus sylvestris.

Apium 1. Apium graveolens L. E celery. MA. Dumps, waste areas.

Aralia 1a. Plants scapose, both the leafless peduncle arising from near ground level and the single leaf arising from a very short, woody stem at the surface of the ground . . . . . . . A. nudicaulis 1b. Plants with evident, leafy stems, the peduncles arising from well above the surface of the ground 2a. Plants shrubs or small trees, up to 10 m tall, armed with stout prickles; styles distinct (sometimes connate to near the middle in A. elata)

A p i ac e a e   3 3 9

3a. Leafules (or leafulets on tripinnately compound leaves) ± glabrous abaxially, with veins anastomosing before reaching the marginal teeth, with petiolules longer than 2 mm; branchlets light brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. spinosa 3b. Leafules (or leafulets on tripinnately compound leaves) pubescent abaxially, with veins essentially running into the marginal teeth, subsessile; branchlets light gray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. elata 2b. Plants herbaceous, up to 2 m tall, unarmed or armed with slender bristles near the base of the plant; styles connate about half their length 4a. Inflorescence composed of a few to several umbels in a cluster resembling a corymb; stems armed near the base with slender bristles; plants commonly of dry, sandy, and/or sterile soils; ultimate leaf segments rounded to cuneate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. hispida 4b. Inflorescence composed of numerous umbels in a cluster resembling a panicle; stems unarmed; plants of rich, mesic soils; ultimate leaf segments cordate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. racemosa 1. Aralia elata (Miq.) Seem. E Japanese angelica-tree. Dimorphanthus elatus Miq. • CT, MA, NH. Roadsides, field edges, stream banks, abandoned homesteads. 2. Aralia hispida Vent. N bristly sarsaparilla. CT, MA, ME, NH, RI, VT. Roadsides, sandy fields, open balds, woodlands. 3. Aralia nudicaulis L. N wild sarsaparilla. CT, MA, ME, NH, RI, VT. Forests, woodlands. 4. Aralia racemosa L. ssp. racemosa N American spikenard. CT, MA, ME, NH, RI, VT. Forests, usually mesic, deciduous types. 5. Aralia spinosa L. E Hercules’-club. CT, MA, RI. Forest edges and fragments. Some collections formerly identified as this species actually belong to Aralia elata.

Astrantia 1. Astrantia major L. E greater masterwort. MA. Fields, waste areas.

Bifora 1. Bifora radians Bieb. E wild bishop. RI. Waste areas, yards.

Bupleurum 1a. Leaf blades linear, sessile; bracts at base of umbel present; bractlets at base of umbellets lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. odontites 1b. Leaf blades oblong-lanceolate or ovate to suborbicular, perfoliate; bracts at base of umbel absent; bractlets at base of umbellets oblanceolate or elliptic to suborbicular 2a. Primary branches of inflorescence numbering (3–) 5–10; bractlets at base of umbellets oblanceolate or elliptic to ovate or obovate; fruit smooth; leaf blades usually elliptic-ovate to suborbicular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. rotundifolium

340 tricolpates

2b. Primary branches of inflorescence usually numbering 2 or 3 (–5); bractlets at base of umbellets suborbicular; fruit tuberculate; leaf blades usually ovate to oblong-lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. lancifolium 1. Bupleurum lancifolium Hornem. E lance-leaved thorough-wax. Bupleurum protractum Hoffm. & Link • CT, MA. Fields, roadsides, waste areas. 2. Bupleurum odontites L. E narrow-leaved thorough-wax. Bupleurum fontanesii Guss. ex Caruel • MA. Fields, roadsides, waste areas. 3. Bupleurum rotundifolium L. E round-leaved thorough-wax. NH; also reported from VT by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, waste areas. Reports of Bupleurum rotundifolium from CT and MA were based on specimens of B. lancifolium (specimens at CONN; image seen!, MASS!, and NEBC!).

Carum 1. Carum carvi L. E caraway. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, gardens.

Cicuta 1a. Axils of the upper leaves bearing bulbils; ultimate leaf segments linear, up to 5 mm wide; schizocarps 1.5–2 mm long; styles becoming 0.5–0.7 mm long . . . . . . . . . . . . . . . . . . C. bulbifera 1b. Axils of the leaves without bulbils; ultimate leaf segments linear to lanceolate, (5–) 6–25 (–40) mm wide [Fig. 346]; schizocarps (2–) 3–4 mm long; styles becoming 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. maculata 1. Cicuta bulbifera L. N bulblet-bearing water-hemlock. CT, MA, ME, NH, RI, VT. Shorelines, marshes, shallow water. 2. Cicuta maculata L. var. maculata N Fig. 346 spotted water-hemlock. Cicuta curtissii Coult. & Rose; C. maculata L. var. curtissii (Coult. & Rose) Fern. • CT, MA, ME, NH, RI, VT. Shorelines, marshes, shallow water, wet ditches.

Conioselinum 1. Conioselinum chinense (L.) B.S.P. N Fig. 346  Ultimate segment of leaf of Cicuta maculata showing some veins directed to the sinuses between teeth.

Chinese hemlock-parsley. CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Stream banks, swamps, riparian forests.

Conium 1. Conium maculatum L. E poison-hemlock. CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas.

Coriandrum 1. Coriandrum sativum L. E coriander. CT, MA, ME, RI. Roadsides, waste areas.

A p i ac e a e   3 41

Cryptotaenia 1. Cryptotaenia canadensis (L.) DC. N Fig. 347 Canada honewort. Deringa canadensis (L.) Kuntze • CT, MA, ME, NH, RI, VT; disjunct and rare in northeastern New England. Rich, mesic forests, high-terrace floodplain forests.

Cuminun 1. Cuminum cyminum L. E cumin. MA. Roadsides, waste areas.

Daucus 1. Daucus carota L. E Fig. 348 wild carrot. CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns.

Eleutherococcus

Fig. 347  Inflorescence of Cryptotaenia canadensis.

1. Eleutherococcus pentaphyllus (Sieb. & Zucc.) Nakai E five-leaved-aralia. Acanthopanax sieboldianus Makino; Aralia pentaphylla Sieb. & Zucc. • CT, MA. Forest edges, thickets, and roadsides in areas of habitation.

Eryngium Eryngium are perennial plants from thick, cylindric or turnip-shaped taproots. 1a. Bracts subtending inflorescence relatively short, often not projecting beyond width of inflorescence, mostly entire; leaf blades unlobed, linear to narrow-lanceolate, parallelveined; inflorescence white to green-white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. yuccifolium 1b. Bracts subtending inflorescence conspicuous, usually projecting well beyond the width of the inflorescence, mostly spinulose-toothed [Fig. 349]; leaf blades lobed or unlobed and then elliptic-oblong to elliptic-ovate and net-veined; inflorescence usually blue

Fig. 348  Inflorescence and pinnately lobed bracts of Daucus carota.

2a. Lower leaf blades bipinnatifid; upper stem leaves clasping . . . . . . . . . E. amethystinum 2b. Lower leaf blades simple; upper stem leaves shortly petiolate or sessile . . . . E. planum 1. Eryngium amethystinum L. E amethyst eryngo. MA. Roadsides, waste areas. 2. Eryngium planum L. E Fig. 349 plains eryngo. CT, MA, VT. Roadsides, waste areas. 3. Eryngium yuccifolium Michx. var. yuccifolium E button eryngo. CT; southwestern portion of state. Roadsides, waste areas.

Falcaria 1. Falcaria vulgaris Bernh. E sickleweed. Falcaria soides (Wibel) Aschers. • MA. Roadsides, waste areas.

Foeniculum 1. Foeniculum vulgare P. Mill. E sweet fennel. Foeniculum foeniculum (L.) Karst. • CT, MA, RI; also reported from ME by Campbell et al. (1995), but specimens are unknown. Roadsides, waste areas, gardens.

Fig. 349  Inflorescence of Eryngium planum.

342   tricolpate s

Hedera 1. Hedera helix L. E English-ivy. MA. Forest edges, thickets, and yards in areas of habitation.

Heracleum 1a. Umbel with mostly 50–150 primary branches, up to 50 cm wide; leaflets of principal leaves up to 130 cm long; schizocarp on a pedicel 15–40 mm long, with oil tubes 0.8 mm wide or wider that extend (50–) 65–75% of the way from apex to base of mature fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. mantegazzianum 1b. Umbel with mostly 15–45 primary branches, rarely exceeding 20 cm wide; leaflets of principal leaves 5–30 (–60) cm long; schizocarp on a pedicel 6–20 mm long, with oil tubes up to 0.5 (–0.8) mm wide that extend 50–60 (–70%) of the way from apex to base of mature fruit 2a. Leaves with 3 ternately arranged, sharply serrate leaflets [Fig. 350]; umbels with 5–10 bracts; larger leaflets 10–30 (–60) cm long; stylopodium not, or only slightly, extending above a deep notch in the mericarp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. maximum 2b. Leaves with 3–7 pinnately arranged, often bluntly toothed, leaflets; umbels with fewer than 5 bracts, or lacking bracts altogether; leaflets mostly 5–10 cm long; stylopodium extending above a notch in the mericarp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. sphondylium 1. Heracleum mantegazzianum Sommier & Levier E giant cow-parsnip. CT, MA, ME, VT. Fields, forest openings, abandoned homesteads. 2. Heracleum maximum Bartr. N Fig. 350 American cow-parsnip. Heracleum lanatum Michx.; H. sphondylium L. ssp. montanum (Schleich. ex Gaudin) Briq.; H. sphondylium L. var. lanatum (Michx.) Dorn • CT, MA, ME, NH, RI, VT. River banks, riparian terraces, meadows, roadside ditches, fields, subalpine meadows. 3. Heracleum sphondylium L. E

Fig. 350  Leaf with expanded sheath of Heracleum maximum.

European cow-parsnip. CT, MA, ME, VT. Fields, roadsides, waste areas. Our plants need study to determine which subspecies is represented are New England. Some plants show the white petals of ssp. sphondylium, but also show the scarcely enlarged marginal corollas of ssp. sibericum (L.) Simonkai.

Hydrocotyle 1a. Leaf blades basifixed (i.e., the petiole attached to the base of the blade at the sinus between the two basal lobes); umbels sessile or subsessile; schizocarp 1.5–1.9 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. americana 1b. Leaf blades peltate (i.e., the petiole attached to the center of the abaxial surface of the blade) [Fig. 351]; umbels borne on peduncles; schizocarp 2–4 mm wide 2a. Inflorescence usually a single umbel at the summit of the peduncle (rarely as many as 3 whorls), with (10–) 15–35 (–100) flowers in the lowest whorl; schizocarp 2–3 mm wide, deeply notched or cordate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. umbellata 2b. Inflorescence appearing to be a spike or raceme with 2–12 whorls of flowers, each whorl with 2–7 (–20) flowers [Fig. 351]; schizocarp (2–) 3–4 mm wide, rounded or shallowly emarginate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. verticillata 1. Hydrocotyle americana L. N American marsh-pennywort. CT, MA, ME, NH, RI, VT. Marshes, low fields, wet lawns, stream margins. 2. Hydrocotyle umbellata L. N many-flowered marsh-pennywort. CT, MA, RI. Pond shores, outlet margins.

Ap i ac e a e   3 43

3. Hydrocotyle verticillata Thunb.

N C Fig. 351

whorled marsh-pennywort. Hydrocotyle verticillata Thunb. var. triradiata (A. Rich.) Fern. • CT, MA, RI; coastal plain and the islands. Low, wet, often boggy, depressions and seeps, pond shores, outlet margins.

Kalopanax 1. Kalopanax septemlobus (Thunb.) Koidz. E castor-aralia. Acanthopanax ricinifolius (Sieb. & Zucc.) Seem.; Kalopanax pictus (Thunb.) Nakai • CT, MA. Forest edges and fragments in areas of habitation.

Levisticum 1. Levisticum officinale W.D.J. Koch E garden lovage. Hipposelinum levisticum (L.) Britt. & Rose • CT, MA, ME, VT. Fields, gardens, waste areas.

Ligusticum 1. Ligusticum scoticum L. ssp. scoticum N Scotch wild lovage. CT, MA, ME, NH, RI. Coastal beaches and strands.

Lilaeopsis 1. Lilaeopsis chinensis (L.) Kuntze N eastern grasswort. Lilaeopsis lineata (Michx.) Greene • CT, MA, ME, NH, RI. Brackish-tidal river shores, often on mud or silty mud substrate.

Myrrhis 1. Myrrhis odorata (L.) Scop. E anise. Scandix odorata L. • MA. Compost heaps, gardens, waste areas.

Osmorhiza 1a. Bractlets at base of umbellets absent (infrequently with a reduced, solitary bractlet); styles at maturity strongly outcurved, 0.3–1 mm long [Fig. 352]; flowers all bisexual 2a. Schizocarp 12–22 mm long, concavely tapered to the apex and forming a terminal beak 1–2 mm long [Fig. 352]; branches of inflorescence and pedicels ascending . . . . . O. berteroi 2b. Schizocarp 10–15 mm long, straight or convexly tapered to the apex, without a beak [Fig. 353]; branches of inflorescence and pedicels widely spreading, the pedicels often divaricately so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. depaupertata 1b. Bractlets present (though some or all may be deciduous during and after maturation of the fruit); styles at maturity nearly straight, 1–4 mm long; flowers bisexual or staminate 3a. Styles 1–1.5 mm long; roots and foliage with a slight odor and taste of anise; bractlets of umbellets linear; umbellets with 4–8 (–10) flowers (note: include the withered staminate ones for counts made after anthesis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. claytonii 3b. Styles 2–4 mm long; roots and foliage with a strong odor and taste of anise; bractlets of umbellets lanceolate; umbellets with (6–) 9–18 flowers . . . . . . . . . . . . . . . . . O. longistylis

Fig. 351  Leaves and inflorescence of Hydrocotyle verticillata.

344   tricolpate s

1. Osmorhiza berteroi DC. N Fig. 352 mountain sweet-cicely. Osmorhiza chilensis Hook. & Arn.; O. divaricata (Britt.) Suksdorf; Washingtonia divaricata Britt. • ME, NH, VT. Mesic, deciduous and mixed evergreen-deciduous forests in north-temperate and boreal areas. See Kartesz and Gandhi (1993) for rationale of Osmorhiza berteroi as the correct name for this taxon (vs. O. chilensis). 2. Osmorhiza claytonii (Michx.) C.B. Clarke N bland sweet-cicely. Washingtonia claytonii (Michx.) Britt. • CT, MA, ME, NH, RI, VT. Mesic, deciduous forests. 3. Osmorhiza depauperata Phil.

N C Fig. 353

blunt-fruited sweet-cicely. Osmorhiza obtusa (Coult. & Rose) Fern.; Washingtonia obtusa Coult. & Rose • ME, VT. Mesic, deciduous forests in north-temperate and boreal areas. 4. Osmorhiza longistylis (Torr.) DC. N Fig. 352  Schizocarp of Osmorhiza berteroi.

long-styled sweet-cicely. Osmorhiza longistylis (Torr.) DC. var. villicaulis Fern.; Washingtonia longistylis (Torr.) Britt. • CT, MA, ME, NH, RI, VT. Mesic, deciduous forests.

Panax 1a. Leaflets mostly 6–15 cm long, with long petiolules, acuminate at the apex; plants 2–6 dm tall, in anthesis in July; flowers mostly bisexual; drupe red; root elongate and fusiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. quinquefolius 1b. Leaflets mostly 4–8 cm long, sessile or nearly so, obtuse to acute at the apex; plants 1–2 dm tall, in anthesis from late May through mid-June; flowers often unisexual and staminate; drupe yellow; root globular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. trifolius 1. Panax quinquefolius L. N American ginseng. CT, MA, ME, NH, RI, VT. Rich, mesic forests, often on rocky slopes and near cliff bases, rarely in wet-mesic forests that are influenced by high-pH bedrock. 2. Panax trifolius L. N dwarf ginseng. CT, MA, ME, NH, RI, VT. Mesic, deciduous forests. Fig. 353  Schizocarp of Osmorhiza depauperata.

Pastinaca 1. Pastinaca sativa L. E Fig. 354 wild parsnip. Pastinaca sativa L. var. pratensis Pers. • CT, MA, ME, NH, RI, VT. Fields, roadsides.

Petroselinum 1. Petroselinum crispum (P. Mill.) Nyman ex A.W. Hill E parsley. Apium petroselinum L. • CT, MA, RI, VT. Fields, roadsides, waste areas, gardens.

Peucedanum 1. Peucedanum palustre (L.) Moench E milk-parsley. Calestania palustris (L.) K.-Pol. • MA. Roadsides.

Fig. 354  Cross-section of schizocarp of Pastinaca sativa showing compression parallel to commissure.

Pimpinella 1a. Lower leaf blades pinnately divided; schizocarps glabrous, 2–2.5 mm long . . . . . P. saxifraga 1b. Lower leaf blades simple, with coarse, irregular teeth and incisions; schizocarps puberulent, 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. anisum

Ap i ac e a e   3 45

1. Pimpinella anisum L. E anise burnet-saxifrage. MA. Fields, roadsides, waste areas. 2. Pimpinella saxifraga L. ssp. saxifraga E solid-stemmed burnet-saxifrage. CT, MA, ME, VT. Fields, roadsides, waste areas.

Ptilimnium 1. Ptilimnium capillaceum (Michx.) Raf. N Atlantic mock bishop-weed. Ammi capillaceum Michx. • CT, MA, RI. Saline and brackish marshes.

Sanicula Plants of Sanicula bear two types of flowers—those that are bisexual (i.e., have stamens and carpels) with hooked bristles over the outer surface of the ovary and those that are unisexual and staminate. The distribution of these types of flowers is important. All species in New England have umbellets with both types of flowers, but some species also have umbellets with only staminate flowers. Reference: Pryer and Phillippe (1989). 1a. Styles much exceeding the bristles of the fruit [Fig. 356], usually more than twice as long as the calyx; umbellets dimorphic—some with staminate and bisexual flowers intermixed and some with only staminate flowers, those with both types of flowers bearing 12–25 (–117) staminate flowers per umbellet; plants perennial 2a. Sepals 1–2 mm long, rigid, narrow-subulate; schizocarps subsessile, slightly exceeded by the persistent, staminate flower remnants . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. marilandica 2b. Sepals 0.5–1 mm long, soft, triangular; schizocarps stipitate, slightly exceeding the persistent, staminate flower remnants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. odorata 1b. Styles exceeded by the bristles of the fruit [Fig. 355], usually not exceeding the length of the calyx (as long as 1.5 times the calyx length in S. canadensis var. grandis); umbellets usually monomorphic, all bearing bisexual and staminate flowers, the staminate flowers numbering 1–8 (–15) per umbellet (S. canadensis var. grandis sometimes with umbellets containing only staminate flowers); plants biennial 3a. Sepals 2–2.5 mm long, lanceolate, connivent into a beak and conspicuous in fruit, exceeding the bristles of the schizocarp; carpellate flowers sessile . . . . . . . . . . . S. trifoliata 3b. Sepals 0.6–1.5 mm long, subulate, inconspicuous in fruit, shorter than the bristles of the schizocarp; carpellate flowers on pedicels 0.5–1 mm long . . . . . . . . . . . . . . . . . S. canadensis 1. Sanicula canadensis L. N C Fig. 355 Canada sanicle. 1a. Sanicula marilandica L. var. canadensis (L.) Torr. • CT, MA, NH, VT. Rich, mesic forests, dry-mesic forests on sandy soils.

Fig. 355  Umbellet of Sanicula canadensis var. canadensis.

1a. Umbellets of only one type usually present—those with both bisexaul and staminate flowers, those with only staminate flowers rare; umbellets with both types of flowers bearing 1–3 staminate flowers borne on pedicels up to 2 mm long [Fig. 355]; styles shorter than or equal in length to the calyx; triad of fruits mostly 7–9 mm wide . . . . . . 1a. S. canadensis var. canadensis 1b. Umbellets of both types sometimes present—those with both bisexual and staminate flowers and those with only staminate flowers; umbellets with both types of flowers bearing 3–15 staminate flowers borne on pedicels 2–3 mm long; styles ca. 1.5 times as long as the calyx; triad of fruits mostly 10–15 mm wide . . . . . . . . . . . . . 1b. S. canadensis var. grandis Fern. Variety canadensis is known from CT, MA, NH, RI, VT. Variety grandis is known from CT, MA, VT. Both varieties are of conservation concern. 2. Sanicula marilandica L. N Fig. 356 Maryland sanicle. Sanicula canadensis L. var. marilandica (L.) A.S. Hitchc.; S. marilandica L. var. borealis Fern. • CT, MA, ME, NH, RI, VT. Mesic forests, often associated with rich soils and/or rocky slopes.

Fig. 356  Umbellet of Sanicula marilandica.

346  tricolpates

3. Sanicula odorata (Raf.) K.M. Pryer & L.R. Phillippe N clustered sanicle. Sanicula gregaria Bickn.; Triclinium odoratum Raf. • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Rich, mesic forests, including upland and riparian types. 4. Sanicula trifoliata Bickn. N large-fruited sanicle. CT, MA, ME, NH, VT; also reported from RI by Gould et al. (1998), but specimens are unknown. Rich, mesic forests, including uplants and riparian types.

Scandix 1. Scandix pecten-veneris L. E shepherd’s-needle. MA, RI. Ballast, waste areas.

Selinum 1. Selinum carvifolia (L.) L. E little-leaved-angelica. Seseli carvifolia L. • MA. Waste areas, yards.

Sium Sium sisarum L. was reported from RI by Tucker (2006) as a “weed in vegetable garden.” This is not here considered evidence of naturalization in New England. 1. Sium suave Walt. N Fig. 357 water-parsnip. Sium carsonii Dur. ex Gray; S. cicutifolium Schrank • CT, MA, ME, NH, RI, VT. Shorelines, swamps, marshes, shallow water of lakes and rivers.

Fig. 357  Cross-section of schizocarp of Sium suave.

Taenidia 1. Taenidia integerrima (L.) Drude

NC

yellow-pimpernel. Smyrnium integerrimum L.; Zizia integerrima (L.) DC. • CT, RI, VT. River banks, lake shores and headlands, primarily in regions of high-pH bedrock.

Thaspium 1. Thaspium trifoliolatum (L.) Gray var. trifoliolatum E purple meadow-parsnip. Thaspium aureum (L.) Nutt. var. trifoliolatum (L.) Coult. & Rose • RI. Forests.

Torilis 1a. Umbel with 2 or 3 primary branches; prickles of schizocarp wide-spreading, only slightly ascending, up to 1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. leptophylla 1b. Umbel with (3–) 5–10 primary branches [Fig. 358]; prickles of schizocarp ascending, up to 0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. japonica 1. Torilis japonica (Houtt.) DC. E Fig. 358 erect hedge-parsley. CT, MA. Roadsides, fields, waste areas. Fig. 358  Inflorescence of Torilis japonica.

2. Torilis leptophylla (L.) Reichenb. f. E bristle-fruited hedge-parsley. MA. Roadsides, fields, waste areas.

A p i ac e a e   3 47

Zizia 1a. Basal and sometimes the lower stem leaves with simple, triangular-ovate to oblong-ovate blades with a cordate base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Z. aptera 1b. Basal and lower stem leaves with mostly twice ternately divided blades . . . . . . . . Z. aurea 1. Zizia aptera (Gray) Fern.

NC

heart-leaved golden Alexanders. Zizia cordata W.D.J. Koch ex DC. • CT, RI. Open to partially forested sites on dry-mesic to mesic soils, frequently near river banks and often associated with circumneutral soils, including those influenced by shell middens. 2. Zizia aurea (L.) W.D.J. Koch N common golden Alexanders. CT, MA, ME, NH, RI, VT. Riparian forests, river banks, fields, meadows, open rights-of-way.

Apocynaceae 1a. Flowers with a corona [Figs. 362, 363, 366]; carpels connate only by the thickened stylehead; pollen coherent in masses; seeds always comose 2a. Plants lianas; corolla lobes densely villous adaxially; corona composed of 5 short, bifid, antepetalous lobes alternating with 5 narrow, thread-like lobes 4–8 mm long . . . . . Periploca 2b. Plants herbs or vines; corolla lobes glabrous or minutely pubescent adaxially; corona either a whorl of 5 tubular- or scoop-shaped hoods or a fleshy, lobed cup 3a. Stems climbing or twining (at least the apical part of the plant); calyx and corolla lobes ascending to spreading [Fig. 366]; corona a fleshy, lobed cup [Fig. 366] . . . . . . Cynanchum 3b. Stems upright, neither climbing nor twining; calyx and corolla lobes reflexed [Figs. 362, 363]; corona a whorl of hoods bearing internally near the base a slender, horn-like appendage (the horns absent in A. viridiflora) [Figs. 362, 363] . . . . . . Asclepias 1b. Flowers with only sepals and petals, lacking a corona [Figs. 359, 360]; carpels apically connate with a common style and stigma; pollen grains not cohering in masses; seeds with or without a coma 4a. Flowers solitary from leaf-bearing nodes; corolla 20–50 mm wide in life; trailing plants with upright branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vinca 4b. Flowers in terminal or both terminal and axillary cymes [Fig. 359]; corolla up to 10 mm wide in life; upright plants without trailing stems 5a. Leaves alternate or irregularly scattered; petals blue; seeds lacking a coma of hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amsonia 5b. Leaves opposite; petals white, tinged or striped with pink, green-white, or yellow; seeds comose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apocynum

Amsonia 1. Amsonia tabernaemontana Walter var. tabernaemontana E eastern bluestar. Amsonia amsonia (L.) Britt.; A. glaberrima Woods. • MA. Lake shores.

348   tricolpate s

Apocynum Reference: Woodson (1930). 1a. Corolla pink or white and pink-striped, 6–10 mm long, its lobes spreading or recurving [Fig. 359]; flowers spreading to nodding; seeds 2.5–3 mm long; leaf blades wide-spreading to drooping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. androsaemifolium 1b. Corolla white, green-white, or yellow, 3–6 mm long, its lobes erect or slightly spreading [Fig. 360]; flowers ascending to spreading; seeds 4–6 mm long; leaf blades erect to spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. cannabinum 1. Apocynum androsaemifolium L. N Fig. 359 Fig. 359  Flowers of Apocynum androsaemifolium.

spreading dogbane. Apocynum ambigens Greene; A. androsaemifolium L. var. glabrum Macoun; A. pumilum (Gray) Greene • CT, MA, ME, NH, RI, VT. Fields, roadsides, woodlands, forest edges. 1‌ × 2. Apocynum ×floribundum Greene is an uncommon hybrid that has also been known by the name A. × ‌medium Greene. It is known from CT, MA, ME, NH, RI, VT. It is marked by a white or pink-tinged corolla 4–7 mm long, ascending to spreading flowers, seeds 3–4 mm long, and wide-spreading leaf blades. 2. Apocynum cannabinum L. N Fig. 360

Fig. 360  Flowers of Apocynum cannabinum.

hemp dogbane. Apocynum cannabinum L. var. hypericifolium (Ait.) Gray; A. cannabinum L. var. nemorale (G.S. Mill.) Fern.; A. cannabinum L. var. pubescens (Mitchell ex R. Br.) Woods.; A. hypericifolium Ait.; A. sibiricum Jacq.; A. sibiricum Jacq. var. cordigerum (Greene) Fern. • CT, MA, ME, NH, RI, VT. River shores, fields, shaded roadsides, woodlands. Apocynum cannabinum is a variable taxon. Those collections with sessile or subsessile leaves and rounded to cordate bases were called A. sibiricum (vs. A. cannabinum sensu stricto with petiolate leaves with cuneate to rounded leaf bases).

Asclepias Reference: Woodson (1954). 1a. Leaves alternate; flowers yellow-orange to orange; latex watery . . . . . . . . . . . . . A. tuberosa 1b. Leaves predominantly or entirely opposite or whorled; flowers white to pink, red, purple, or green; latex milky 2a. Corona with hoods only, the horn lacking [Fig. 365]; umbels sessile or on peduncles up to 1 (–2) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. viridiflora 2b. Corona with horns within the hoods [Figs. 362, 363]; umbels borne on peduncles 1–10 cm long [Figs. 361, 364] 3a. Leaf blades narrow-linear, 1–3 (–5) mm wide, in whorls of 3–6 (some scattered) [Fig. 364] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. verticillata 3b. Leaf blades narrow-lanceolate or narrow-oblong to broad-oblong, broad-elliptic, or ovate, the principal ones 20–140 mm wide, opposite or with 1 or 2 whorls of 4 leaves 4a. Corona hoods shorter than to ± equaling the height of the gynostegium, with conspicuously exserted horns [Fig. 362] 5a. Corolla pink to red; corona hoods 2–3 mm long, the lateral margins upwardly divergent (i.e., not parallel), therefore, the hoods scoop-shaped; follicles on erect pedicels; stems branched or, less commonly, simple . . . . . . . . . . . . . . . A. incarnata 5b. Corolla white, green, green-purple, or pale purple; corona hoods 4–6 mm long, the lateral margins parallel, therefore, the hoods tube-shaped [Fig. 362]; follicles on declined pedicels; stems usually unbranched 6a. Leaf blades cuneate at base, acuminate at apex, with petioles 10–20 mm long; margins of corona hoods prolonged into a short, erect tooth 1–1.5 mm long; umbels 2–4 from both the stem summit and upper nodes . . . . . . . . . A. exaltata

A p o c yn ac e a e   3 49

6b. Leaf blades broad-rounded to cordate at the base, obtuse to rounded at apex, sessile or with short petioles to 5 mm long [Fig. 361]; margins of corona hoods not prolonged, the hood summit truncate; umbels solitary (rarely paired) from the summit of the stem [Fig. 361] . . . . . . . . . . . . . . . . . . . . . A. amplexicaulis 4b. Corona hoods taller than the gynostegium, the horns shorter than to equaling the length of the hoods [Fig. 363] 7a. Corolla white to pink; pedicels 1–2.5 cm long; leaf blades abaxially glabrous or with hairs along the midvein 8a. Stems with usually 3 or 4 leaf-bearing nodes, the upper and lower nodes with opposite leaves, the middle 1 or 2 nodes with a whorl of 4 leaves (rarely all the leaves opposite); corolla lobes 4.5–6 mm long; lateral margins of corona hoods bearing a tooth near the middle [Fig. 363]; leaf blades gradually acuminate at apex; follicles on erect pedicels . . . . . . . . . . . . . . . . . . A. quadrifolia 8b. Stems with 2–5 leaf-bearing nodes, all with opposite leaves; corolla lobes 7–9 mm long; lateral margins of corona hoods without a tooth; leaf blades obtuse to abruptly acuminate at apex; follicles on declined pedicels . . . . . . A. variegata 7b. Corolla purple-green to light purple or red-purple to purple; pedicels 2–5 cm long; leaf blades abaxially tomentulose 9a. Corona hoods 5–7 mm long, red-purple to purple, without marginal lobes; follicles smooth; corolla red-purple to purple . . . . . . . . . . . . . . . A. purpurascens 9b. Corona hoods 4–5 mm long, light purple, each with a prominent marginal lobe near the center; follicles covered with conic processes; corolla purplegreen to light purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. syriaca

Fig. 361  Inflorescence and leaves of Asclepias amplexicaulis.

1. Asclepias amplexicaulis J.E. Smith N Fig. 361 clasping milkweed. CT, MA, NH, RI, VT. Woodlands, sandy fields. 2. Asclepias exaltata L. N Fig. 362 poke milkweed. Asclepias bicknellii Vail; A. phytolaccoides Pursh • CT, MA, ME, NH, RI, VT. Forest openings and edges, roadsides. 3. Asclepias incarnata L. N swamp milkweed. 3b. Asclepias incarnata L. var. neoscotica Fern.; A. incarnata L. ssp. pulchra (Ehrh. ex Willd.) Woods.; A. pulchra Ehrh. ex Willd. • CT, MA, ME, NH, RI, VT. Marshes, wet fields, borders of swamps, shorelines. 1a. Stems often repeatedly branched, sparsely pubescent to glabrous; leaf blades lanceolate to narrow-oblong, usually tapering to the base, glabrous or nearly so on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. A. incarnata var. incarnata

Fig. 362  Flower of Asclepias exaltata.

1b. Stems simple to few-branched, short-pilose; leaf blades broad-lanceolate to elliptic, usually rounded at the base, short-pilose on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. A. incarnata var. pulchra (Ehrh. ex Willd.) Pers. Variety incarnata is known from CT, MA, ME, NH, VT. Variety pulchra is known from CT, MA, ME, NH, RI, VT. 4. Asclepias purpurascens L.

NC

purple milkweed. CT, MA, NH, RI; also reported from VT by Seymour (1982), but specimens are unknown. Forest edges, roadsides, dry fields. The report by Magee and Ahles (1999) for Cumberland County, ME, is based on an erroneously determined Asclepias syriaca (specimen at NHA!). 5. Asclepias quadrifolia Jacq. N Fig. 363 four-leaved milkweed. CT, MA, NH, RI, VT. Forests and woodlands, often associated with rich soils and/or circumneutral bedrock.

Fig. 363  Flower of Asclepias quadrifolia.

35 0   tricolpate s

6. Asclepias syriaca L. N common milkweed. Asclepias intermedia Vail; A. syriaca L. var. kansana (Vail) Palmer & Steyermark • CT, MA, ME, NH, RI, VT. Fields, roadsides, open rights-of-way. 7. Asclepias tuberosa L.

NC

butterfly milkweed. 7a. Asclepias tuberosa L. var. interior (Woods.) Shinners • CT, MA, ME, NH, RI, VT. Dry fields, sand plains, roadsides, disturbed areas. 1a. Leaf blades narrow-oblong to lanceolate or oblong-ovate, widest below the middle, ± truncate to cordate at the base, gradually tapering to the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7a. Asclepias tuberosa ssp. interior Woods. 1b. Leaf blades oblong to obovate or oblanceolate, broadest above the middle, cuneate to rounded at the base, abruptly tapering to the apex . . 7b. Asclepias tuberosa ssp. tuberosa Fig. 364  Inflorescences and leaves of Asclepias verticillata.

Subspecies interior is known from CT and is of conservation concern. Subspecies tuberosa is known from CT, MA, ME, NH, RI, VT. 8. Asclepias variegata L.

NC

red-ringed milkweed. Biventraria variegata (L.) Small • CT. Dry slopes and thickets. 9. Asclepias verticillata L. N Fig. 364 whorled milkweed. CT, MA, RI, VT. Rocky woodlands, open glades, balds, cliff bases. 10. Asclepias viridiflora Raf.

N C Fig. 365

green milkweed. Acerates viridiflora (Raf.) Pursh ex Eat.; A. viridiflora (Raf.) Pursh ex Eat. var. linearis Gray; Asclepias viridiflora Raf. var. lanceolata Torr.; A. viridiflora Raf. var. linearis (Gray) Fern. • CT. Sandy fields.

Cynanchum Fig. 365  Flower of Asclepias viridiflora.

1a. Corolla purple-black, minutely pubescent on the adaxial surface, with triangular lobes 1.5–3 mm long [Fig. 366] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. louiseae 1b. Corolla paler, usually pink to maroon or yellow-white, glabrous, with lanceolate lobes 2.5–4.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rossicum 1. Cynanchum louiseae Kartesz & Gandhi E Fig. 366 black swallowwort. Cynanchum nigrum (L.) Pers.; Vincetoxicum nigrum (L.) Pers. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest edges, abandoned homesteads, banks. 2. Cynanchum rossicum (Kleopow) Borhidi E pale swallowwort. Cynanchum medium, auct. non R. Br.; Vincetoxicum medium, auct. non (R. Br.) Dcne.; Vincetoxicum rossicum (Kleopow) Barbarich • CT, MA, ME, NH. Fields, roadsides, forest edges, abandoned homesteads, banks.

Fig. 366  Flower of Cynanchum louiseae.

Periploca 1. Periploca graeca L. E silkvine. CT; also reported from RI by Magee and Ahles (1999), but specimens are unknown. Roadsides, areas of habitation.

Vinca 1a. Leaf blades ovate to triangular-ovate, 3–7 cm long, ciliate; calyx lobes 6–12 (–15) mm long, ciliate; corolla with a connate tube 12–20 mm long expanding distally to a limb 35–50 mm wide in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. major

A p o c yn ac e a e   3 5 1

1b. Leaf blades oblong or lanceolate to narrow-elliptic or ovate, 1.5–5 cm long, eciliate; calyx lobes 2–3 (–4) mm long, eciliate; corolla with a connate tube 8–12 mm long expanding distally to a limb 20–30 mm wide in life 2a. Leaf blades smooth, coriaceous, the larger ones 15–23 mm wide; flowering stems erect from creeping horizontal stems; peduncles 11–22 (–28) mm long . . . . . . . . . . . . . . . . V. minor 2b. Leaf blades scabrous, subherbaceous, the larger ones 7–16 mm wide; flowering stems erect or arching; peduncles 20–40 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. herbacea 1. Vinca herbacea Waldst. & Kit. E herbaceous periwinkle. MA. Fields, roadsides, waste areas. 2. Vinca major L. E greater periwinkle. Vinca major L. var. variegata Loud. • MA. Fields, roadsides, waste areas. 3. Vinca minor L. E lesser periwinkle. CT, MA, ME, NH, RI, VT. Roadsides, fields, cemeteries, areas of habitation.

Aquifoliaceae Ilex 1a. Perianth 4- or 5-merous; sepals minute, deciduous in fruit; petals linear to linear-oblong, yellow-white, distinct; stamens free; leaves without stipules; branchlets and winter buds purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. mucronata 1b. Perianth 4- to 8-merous; sepals evident, persistent in fruit; petals obovate, white, connate at the base; stamens adnate to the corolla tube for a short distance; leaves with a stipular vestige on each side; branchlets and winter buds gray to brown 2a. Leaf blades thick, coriaceous, evergreen 3a. Leaf blades with a spine-tip and often with 2 or more remote, spine-tipped teeth per margin, (30–) 50–100 mm long; carpellate flowers 4-merous; drupe red (rarely yellow), 7–10 mm thick 4a. Leaf blades dull or barely sublustrous; flowers and fruits solitary or in solitary, peduncled cymes on the branchlets of the current season . . . . . . . . . . . . . . . . . I. opaca 4b. Leaf blades highly lustrous, especially adaxially; flowers and fruits usually clustered, produced from the axils on branches of the previous season . I. aquifolium 3b. Leaf blades merely toothed, without spines [Fig. 367], 15–50 mm long; carpellate flowers 6- to 8-merous (4-merous in I. crenata); drupe black, 4–5 (–8) mm thick 5a. Leaf blades with 1–3 teeth per margin [Fig. 367], the teeth usually restricted to the apical ⅓ of the blade; flowers 6- to 8-merous; drupe 4–5 mm thick . . . . . . . . I. glabra 5b. Leaf blades with (3–) 4–10 teeth per margin, the teeth usually not restricted to the apical portion of the blade; flowers 4-merous; drupe 6–8 mm thick . . . . . . . . I. crenata 2b. Leaf blades thinner, herbaceous, deciduous 6a. Sepals eciliate [Fig. 368]; staminate flowers on peduncles 8–16 mm long; drupe usually orange-red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. laevigata 6b. Sepals ciliate (also sometimes pubescent on the abaxial surface) [Fig. 369]; staminate flowers borne on peduncles 1–4 mm long; drupe usually deep red

35 2   tricolpate s

7a. Carpellate flowers 4- or 5 (–6)-merous, with ciliate-margined petals [Fig. 369]; pyrenes striate-ribbed on outer surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. montana 7b. Carpellate flowers 5- to 8-merous, with entire or inconspicuously erose-margined petals; pyrenes smooth on outer surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. verticillata 1. Ilex aquifolium L. E English holly. MA. Forests and borders of swamps in areas of habitation. 2. Ilex crenata Thunb. E Japanese holly. Celastrus adenophyllus Miq.; Ilex crenata Thunb. var. latifolia Goldring • MA. Forest fragments and borders in areas of habitation.

3. Ilex glabra (L.) Gray N Fig. 367 Fig. 367  Leaves of Ilex glabra.

evergreen winterberry. CT, MA, ME, NH, RI. Swamps, lake shores, bogs. 4. Ilex laevigata (Pursh) Gray N Fig. 368 smooth winterberry. CT, MA, ME, NH, RI, VT. Swamps, lake shores, wetland borders. Ilex laevigata can be difficult to separate from I. verticillata when the two do not bear flowers or fruits. The former tends have dimorphic branches—short shoots and long shoots. The latter has essentially monomorphic branches on a given shrub. 5. Ilex montana Torr. & Gray ex Gray

Fig. 368  Flower of Ilex laevigata.

N C Fig. 369

big-leaved holly. Ilex ambigua (Michx.) Torr. var. montana (Torr. & Gray ex Gray) Ahles; I. ambigua (Michx.) Torr. var. monticola (Wood) Wunderlin & Poppleton; I. amelanchier M.A. Curtis ex Chapman var. monticola Wood; I. monticola Gray • CT, MA; western New England. Deciduous and mixed evergreen-deciduous forests, often associated with rocky slopes and/or wooded areas around ponds. 6. Ilex mucronata (L.) M. Powell, Savol., & S. Andrews N

Fig. 369  Flower of Ilex montana.

mountain holly. Nemopanthus mucronatus (L.) Loes.; Vaccinium mucronatum L. • CT, MA, ME, NH, RI, VT. Swamps, peatlands, lake shores, openings and trail edges in boreal and subalpine forests. 7. Ilex opaca Ait. var. opaca N American holly. MA, RI; also reported from CT by Kartesz (1999) and from ME by Gleason and Cronquist (1991), but specimens are unknown; on the coastal plain. Wet-mesic to mesic forests, occasionally found on drier sites. 8. Ilex verticillata (L.) Gray N common winterberry. Ilex fastigiata Bickn.; I. verticillata (L.) Gray var. cyclophylla B.L. Robins.; I. verticillata (L.) Gray var. fastigiata (Bickn.) Fern.; I. verticillata (L.) Gray var. padifolia (Willd.) Torr. & Gray ex S. Wats.; I. verticillata (L.) Gray var. tenuifolia (Torr.) S. Wats. • CT, MA, ME, NH, RI, VT. Swamps, lake shores, wetland borders, laggs.

Asteraceae Brickellia grandiflora (Hook.) Nutt. was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species had questionable naturalization in RI. 1a. Capitula composed entirely of bisexual ray flowers [Figs. 395, 435, 471]; plants usually with a milky latex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 1 1b. Capitula composed entirely of disk flowers or both ray and disk flowers, the ray flowers, when present, restricted to the margin of the capitulum and unisexual or sterile [Figs. 376, 431, 447]; plants commonly without a milky latex

Ast e r ac e a e   3 5 3

2a. Capitula without marginal, zygomorphic flowers that bear a ray, all the flowers tubular and actinomorphic, with or without apical teeth or lobes [Figs. 374, 390, 431] 3a. Pappus composed of capillary bristles (at least in large part), each bristle smooth, barbellate, or plumose [Fig. 391] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group 2 3b. Pappus composed entirely of scales (each scale divided at the apex into many bristles in Dyssodia), or awns, a short crown, or entirely absent [Fig. 382] . . . . . . Group 3 2b. Capitula with zygomorphic ray flowers near the periphery, the rays sometimes minute and inconspicuous in drying [Figs. 401, 416, 464] 4a. Rays of various colors, but not yellow or orange . . . . . . . . . . . . . . . . . . . . . . . . Group 4 4b. Rays largely or entirely yellow or orange (sometimes yellow only at the base and white distally in Glebionis coronaria and Layia platyglossa; the rays with a darker color near the base in a few species) 5a. Pappus composed partly or entirely of capillary bristles (short scales may also be present; ray flowers sometimes lacking pappus) [Figs. 438, 468] . . . . . . . . . . . . Group 5 5b. Pappus composed entirely of scales (each scale divided at the apex into many bristles in Dyssodia), or awns, or a crown, or completely absent . . . . . . . . . . . . Group 6

Group 1 1a. Pappus absent; plants annual; rays yellow 2a. Involucral bracts membranaceous to herbaceous, not prominently keeled; peduncles not swollen; stems with foliaceous leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lapsana 2b. Involucral bracts becoming enlarged with an indurate, keeled midrib after anthesis; peduncles conspicuously swollen; stem with minute, bracteal leaves . . . . . . . . . . . Arnoseris 1b. Pappus present in some form; plants annual, biennial, or perennial; rays of various colors 3a. Pappus composed partly or entirely of scales (bristles may also be present) 4a. Pappus composed entirely of scales, the scales numerous; rays blue (rarely pink or white); involucre 9–15 mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cichorium 4b. Pappus of 5 scales and 5–10 slender bristles; rays yellow; involucre 4–7 mm tall 5a. Pappus composed of two different elements—5 or 10 flat scales (these sometimes inconspicuous) along with 5–35 capillary bristles [Fig. 428]; involucre composed of bracts of ± equal length, 4–14 mm tall; plants annual or perennial . . . . . . . . . . . . . Krigia 5b. Pappus of 5 or fewer bristle-tipped scales; involucre composed of two dissimilar series of bracts, the outer ones much shorter than the inner ones, 7–16 mm tall; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microseris 3b. Pappus composed entirely of slender bristles 6a. Pappus bristles smooth or barbellate, but not pinnately branched [Fig. 424] 7a. Cypsela body terete or several-angled (note: this character can often be successfully assessed with the flowering ovary) 8a. Cypsela body muricate, at least in the apical portion, tipped by a long, slender beak [Fig. 472] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taraxacum 8b. Cypsela without sharp projections, with or without an apical beak [Figs. 422, 423] 9a. Flowers pink, purple, white, yellow-white, or green-white; capitula with 5–16 (–26) flowers [Fig. 436] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nabalus 9b. Flowers yellow, orange, or red-orange; capitula with 8–100 flowers [Figs. 426, 427]

35 4   tricolpate s

10a. Plants taprooted annuals or biennials; pappus bristles white, relatively soft; involucre composed of 2 series of principal bracts, the outer much shorter than the inner . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crepis 10b. Plants fibrous-rooted perennials, from short or long rhizomes or a caudex; pappus bristles sordid white to light brown, relatively stiff; involucre variable, either composed of 2 series of principal bracts (i.e., a long inner series and a short, outer series) or with 3 or more series of principal bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hieracium 7b. Cypsela body evidently compressed 11a. Cypsela body without an enlarged disk at the apex where the pappus attaches, also lacking a beak; capitula with 80–250 flowers . . . . . . . . . . . . Sonchus 11b. Cypsela with an enlarged disk at the apex where the pappus attaches, with or without a beak [Figs. 429, 430]; capitula with 5–56 flowers 12a. Capitula with 5 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mycelis 12b. Capitula with 13–55 flowers 13a. Plants annual or biennial, without rhizomes; involucre 6–12 mm long in flower, becoming 12–15 mm long in fruit (except L. hirsuta, with involucres 15–22 mm long in fruit and cypsela bodies with a single nerve on each face); cypsela body with 1 or 3–9 nerves on each face . . . . . . . . . . . . . . . . . . . Lactuca 13b. Plants perennial, from deep-seated rhizomes; involucre 12–15 mm long in flower, becoming 15–20 mm in fruit; cypsela body with 4–6 nerves on each face . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mulgedium 6b. Pappus bristles plumose (i.e., pinnately branched) [Fig. 391] 14a. Involucre composed of 1 series of involucral bracts of equal length; leaf blades long and slender, grass-like; cypsela body 10–25 mm long excluding the beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tragopogon 14b. Involucre calyculate or with bracts of differing lengths; leaf blades relatively wider, not grass-like; cypsela body 2–7.5 mm long excluding the beak 15a. Stems resembling a scape, without leaves or with few, very small leaves; cypsela body 4–7.5 mm long excluding the apical beak 16a. Receptacle with chaff; cypselas muricate, the inner terminated by a slender beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypochaeris 16b. Receptacle without chaff; cypsela body rugulose, without a beak 17a. Reproductive stem with (1–) 2–7 capitula, with (7–) 10 or more scale-like bracts, these especially numerous just below the involucres; pubescence of plant strictly simple (i.e., of unbranched hairs); pappus in 1 series of plumose hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorzoneroides 17b. Reproductive stem with 1 (–2) capitulum, with 0–3 scale-like bracts; pubescence of plant, at least in part, compound, with 2 or more branches; pappus of inner flowers in 2 series, the outer series not plumose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leontodon 15b. Stems not resembling a scape, definitely leafy; cypsela body 2–4 (–6) mm long excluding the apical beak (when present) 18a. Outer series of involucral bracts numbering (8–) 13 or more, lanceolate to broad-linear, 0.5–1.5 (–3) mm wide; cypsela body apically narrowed but without a beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Picris 18b. Outer series of involucral bracts numbering (3–) 5, ovate to broadlanceolate, 3–5 (–8) mm wide; cypsela body terminated by a slender beak nearly as long as or longer than the body proper . . . . . . . . . . . Helminthotheca

Ast e r ac e a e  3 5 5

Group 2 1a. Leaf blades spiny-margined [Fig. 385]; involucral bracts (at least some of them) with a simple spine tip (the spine tip sometimes short in Cirsium muticum and C. palustre) [Fig. 390]; receptacle densely bristly setose between the disk flowers (except in Onopordum) 2a. Leaf blades streaked or blotched with white; lower part of free portion of filaments connate into a tube; pappus dimorphic—an outer series of minutely barbellate, narrow (i.e. bristle-like) scales and an inner series of minute, smooth bristles . . . . . . . . . . . . Silybum 2b. Leaf blades not streaked or blotched with white (though sometimes entirely or strongly suffused with white in Cirsium arvense); free portion of filaments distinct; pappus monomorphic, consisting of minutely barbellate or plumose bristles 3a. Pappus bristles plumose (i.e., pinnately branched); stem and branches with or without prickly, decurrent wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cirsium 3b. Pappus bristles minutely barbellate; upper stem and branches with conspicuous, prickly and undulate-margined wings formed by decurrent leaf bases 4a. Receptacle flat, fleshy, deeply pitted with polygonal depressions that are dentate or short setose around the margins, lacking elongate bristles between the disk flowers; pappus pale red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Onopordum 4b. Receptacle flat to conic, not fleshy, shallowly pitted, densely bristly setose between the disk flowers; pappus ± white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carduus 1b. Leaf blades without spines; involucral bracts not spine-tipped (except Symphyotrichum ericoides) or the bracts with palmately or pinnately branched spines [Figs. 370, 387, 405]; receptacle lacking bristle-like setae (except Centaurea) 5a. Receptacle densely bristly setose; most or all of the involucral bracts tipped with an erose or a fimbriate- to pectinate-fringed appendage, the appendage sometimes spinose (entire in Amberboa); cypselas attached laterally or obliquely to the receptacle 6a. Apical appendage of the involucral bracts erose or fimbriate- to pectinate-fringed, the appendage sometimes spinose; cypsela body with an entire apex, lacking a conspicuous rim around the attachment scar; pappus absent or of minutely barbellate bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Centaurea 6b. Apical appendage of the involucral bracts entire; cypsela body with a minutely denticulate apex, with a conspicuous rim around the attachment scar; pappus absent or of slender scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amberboa 5b. Receptacle lacking bristle-like setae; involucral bracts with entire or ciliate margins; cypselas attached basally to the receptacle 7a. Pappus double—an inner series of elongate, slender bristles and an outer series of short, scale-like bristles; style branches slender, gradually tapering to the apex, minutely pubescent on the outer surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vernonia 7b. Pappus single, composed entirely of elongate, slender bristles (pappus double in Crupina, the inner series of black-brown bristles); style branches truncate, rounded, clavate-thickened, or shortly tapering to apex, mostly glabrous or minutely papillose, infrequently pubescent and then the hairs restricted to an apical tuft or an apical appendage 8a. Plants pubescent with white or gray tomentum or sericeo-tomentum, at least on the abaxial leaf surface (often also on the stem; some species becoming glabrate late in growing season) 9a. Stems resembling a scape, scaly bracteate, appearing before the leaves; foliage leaves all basal, 5–40 cm wide, palmately lobed . . . . . . . (in part) Petasites 9b. Stems resembling a scape or not, in either case appearing after the leaves; leaves basal or produced on a stem, 0.2–5.5 cm wide, entire

35 6   tricolpate s

10a. Plants dioecious or monoecious, none of the flowers bisexual (note: central flowers of carpellate Anaphalis appearing bisexual but have an undivided style with an abortive ovary and are functionally staminate) 11a. Plants stoloniferous, with conspicuous rosettes of basal leaves, mostly 4–35 cm tall; staminate pappus apically clavate; carpellate pappus connate at the base and falling together; stems subscapose, with reduced, bract-like leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Antennaria 11b. Plants not stoloniferous, without conspicuous rosettes of basal leaves, mostly 20–100 cm tall; pappus slender at the apex and distinct, falling separately; stems leafy, the leaf blades 7.5–12 cm long . . . . . . . . . . Anaphalis 10b. Plants polygamous—the numerous outer flowers unisexual and carpellate, the few inner flowers bisexual 12a. Receptacle with chaff; outer flowers of capitulum (i.e., the carpellate ones) without pappus 13a. Chaff of carpellate flowers blunt at the apex, ± saccate, enclosing a disk flower; receptacle obovoid to abruptly expanded at the apex, 0.4–1.6 times as tall as wide; inner chaff spreading in fruit; cypselas dimorphic—the outer longer than the inner . . . . . . . . . . . . . . . . . . . . . Logfia 13b. Chaff of carpellate flowers acuminate to aristate at the apex, not saccate, enfolding (but not enclosing) a disk flower; receptacle clavate, 5–15 times as tall as wide; inner chaff erect to ascending in fruit; cypselas ± monomorphic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Filago 12b. Receptacle without chaff; both the outer carpellate flowers and the inner bisexual flowers with pappus 14a. Involucral bracts yellow-white to sordid white; capitulescence corymb-like, with elongate lower branches; outer disk flowers yellow when fresh . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudognaphalium 14b. Involucral bracts light brown, yellow-brown, or green-brown to brown (sometimes pink to purple); capitulescence not corymb-like, either tall and slender (i.e., spike-like), subcapitate, or composed of axillary clusters [Figs. 413, 415, 440]; outer disk flowers purple or at least apically purple when fresh 15a. Capitulescence composed of small axillary clusters of capitula [Fig. 415]; involucre 2–3 mm tall; axis of plant often profusely branched; pappus bristles distinct but joined at the base by short, interlocking cilia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gnaphalium 15b. Capitulescence spike-like or subcapitate, in either case terminal [Figs. 413, 440]; involucre 3–7 mm tall; axis of plant simple to sparingly branched; pappus bristles either connate at the base into a ring or distinct and without interlocking cilia 16a. Capitulescence consisting of a terminal cluster of capitula 10–15 mm in diameter, subtended by 3–5 leaf-like bracts, sometimes also with some smaller axillary clusters; stem leaves subclasping, with revolute margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euchiton 16b. Capitulescence spike-like, tall and slender (subcapitate in depauperate plants), the axillary capitula subtended by a solitary leaf-like bract; stem leaves not clasping, with ± plane margins 17a. Body of cypsela glabrous; involucre 3–5 mm tall; pappus bristles basally connate into a ring, the entire set falling as a unit;

Ast e r ac e a e   3 57

body of cypsela 0.4–0.9 mm long; plants annual or biennial, of low elevations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gamochaeta 17b. Body of cypsela sparsely strigose; involucre 5–7 mm tall; pappus bristles basally connate or not; body of cypsela 1–1.5 mm long; plants perennial, boreal or alpine . . . . . . . . . . . Omalotheca 8b. Plants glabrous or pubescent, but not conspicuously tomentose 18a. Leaves opposite or whorled (those of the upper stem sometimes alternate) [Fig. 433] 19a. Plants vines; involucre with 4 principal bracts . . . . . . . . . . . . . . . . . . . Mikania 19b. Plants herbs; involucre with more than 4 principal bracts 20a. Leaves both opposite and borne on distinct petioles (rarely the upper leaves sessile) 21a. Involucral bracts all of similar length (1 or 2 outer, very small bracts may also be present) [Fig. 370]; pappus bristles tapering to a fine point; adaxial (i.e., inner) surface of corolla lobes papillose (i.e., with a layer of projecting cells) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ageratina 21b. Involucral bracts of (2–) 3 or more conspicuously different lengths; pappus bristles blunt at the apex; adaxial surface of corolla lobes smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Eupatorium 20b. Leaves opposite and ± sessile (i.e., the petioles, if present, ill defined), or whorled 22a. Leaves in whorls of 3–7 with blades (15–) 20–150 (–180) mm wide; corollas, and often also the involucral bracts, pink to purple; involucres usually cylindric in flower, the margins parallel or slightly upwardly flared . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrochium 22b. Leaves usually opposite with blades 0.5–40 (–60) mm wide (whorled in E. hyssopifolium and E. torreyanum, but then with blades only 0.5–10 (–17) mm wide); corollas white (pink to purple in rare forms of E. perfoliatum); involucral bracts variously colored, usually with green and white, but not pink to purple; involucres usually with a distinct upward flare in flower, obviously narrower near base compared with apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Eupatorium 18b. Leaves alternate throughout the stem 23a. Capitulum with 3–5 flowers, usually only the inner 1 fertile and producing fruit; pappus of fertile flowers double—an outer series of short, triangularlanceolate scales and an inner series of elongate, black-brown bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crupina 23b. Capitulum with 5 or more flowers, producing more than 1 fruit; pappus single, all of elongate bristles, white to brown or purple 24a. Capitula 3–5 cm in diameter, with 200–450 disk flowers; involucre 10–20 mm long, with bracts that spread from the base during flowering (these becoming deflexed during fruiting . . . . . . . . . . . . . . . . . . . Xerochrysum 24b. Capitula narrower than 3 cm, with (4–) 6–110 florets; involucre 4–17 mm long, with bracts that are erect to loosely ascending from the base (the apices spreading to recurving in some species) 25a. Plants woody, often scurfy-pubescent on the capitulescence branches and involucral bracts, dioecious—the carpellate plants with capitula containing only slender, tubular corollas lacking apical teeth, the

35 8   tricolpate s

staminate plants with apically 5-lobed corollas and abortive ovaries with frequently connate style branches . . . . . . . . . . . . . . . . . . . . . . . . . . . Baccharis 25b. Plants herbaceous, glabrous or pubescent but not scurfy, either synoecious and with all of the disk corollas of the capitulum with apical lobes or polygamous and then with unlobed, tubular corollas near the periphery and apically lobed corollas near the center of the capitulum 26a. All flowers of the capitula bisexual; plants perennial, or annual in most Senecio 27a. Corollas red-purple (rarely white); capitulescence tall and slender, resembling a raceme; involucre consisting of 3 or more series of bracts of distinctly unequal lengths [Fig. 431] . . . . . . Liatris 27b. Corollas yellow to orange or green-white to yellow-white (rarely pink-tinged); capitulescence resembling a panicle or corymb; involucre consisting of a single series of long bracts, sometimes calyculate as well [Fig. 449] . . . . . . . . . . . . . . . . . . . . . (in part) Senecio 26b. At least the marginal flowers of the capitula unisexual and carpellate; plants annual 28a. Involucre evidently pubescent with short, multicelled, glandular-viscid hairs, arranged in several, overlapping series of dissimilar length bracts, commonly tinged with anthocyanin; anthers filiform-tailed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pluchea 28b. Involucre not glandular-pubescent, in 1 or more series of long, ± equal length bracts (sometimes with a very short, outer series of bracts in Erechtites), green; anthers cuneate to sagittate at the base, but not tailed 29a. Involucre 10–15 mm tall, in 1 series of long, nearly equal length bracts, sometimes calyculate, turbinate-cylindric, conspicuously swollen at the base before anthesis [Fig. 399]; leaf blades sharply serrate and sometimes also irregularly lobed; cypsela body 2–5.5 mm long . . . . . . . . . . . . . . . . . . . . . . . Erechtites 29b. Involucre 5–11 mm tall, in 3 or 4 series of nearly equal length bracts, without a basal swelling; leaf blades entire; cypsela body up to 2 (–2.2) mm long . . . . . . . . . (in part) Symphyotrichum

Group 3 1a. Leaf blades spiny margined; involucral bracts tipped by a spine (the spine pinnately branched in Centaurea [Fig. 387]) 2a. Capitula 1-flowered, aggregated in clusters forming a false, ± spherical, secondary capitulum that is subtended by a common involucre [Fig. 398]; true involucre subtended by a tuft of capillary bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Echinops 2b. Capitula with more than 1 flower, not conspicuously aggregated in secondary capitula; involucre not subtended by a tuft of bristles 3a. Pappus composed of 10 outer, longer, minutely barbellate awns and 10 inner, shorter, ciliate awns; cypsela body ± tetete in cross-section; inner involucral bracts tipped by a pectinately fringed and spine-tipped appendage or the apical spine pinnately or palmately branched [Fig. 387] . . . . . . . . . . . . . . . . . . . . . . . . (in part) Centaurea 3b. Pappus absent or composed of many rows of linear scales; cypsela body quadrangular in cross-section; inner involucral bracts lacking fringed appendages and tipped by a simple spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carthamus

Ast e r ac e a e   3 5 9

1b. Leaf blades without spiny margins, though spines may occur on other portions of the plant; involucral bracts not tipped by spines (except Ageratum; though they may be tipped by a hooked appendage or be covered by hooked prickles in some species) 4a. Receptacle bristly setose, hairy, or chaffy, at least near the margin of the capitulum; corollas anthocyanic, cyanic, yellow to orange, green-yellow, green-white, or white (or lacking on some flowers) 5a. Pappus of some form present, at least on the inner flowers 6a. Most or all of the leaves opposite [Fig. 380]; pappus of 2–6 awns, these usually retrorsely barbellate (rarely antrorsely barbellate or smooth or absent) [Figs. 381, 382, 383]; receptacle with flattened scales . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Bidens 6b. Leaves alternate; pappus of bristles or scales; receptacle densely bristly 7a. Involucral bracts with an entire, attenuate, hooked tip [Fig. 377]; leaf blades 15–70 cm wide, usually simple, rounded to cordate at the base . . . . . . . . . . Arctium 7b. Involucral bracts tipped by an erose or a fimbriate to pectinate-fringed appendage, not hooked [Figs. 386, 388, 389]; leaf blades up to 6 cm wide, cuneate at the base or pinnately lobed or both . . . . . . . . . . . . . . . . . . . . . . (in part) Centaurea 5b. Pappus absent 8a. Receptacle densely bristly setose or long-hairy or naked 9a. Most or all the involucral bracts tipped by an erose or a fimbriate- to pectinate-fringed appendage [Figs. 386, 388, 389]; involucre 10–25 mm tall; cypselas attached laterally to the receptacle; style branches with a thickened, often pubescent, ring, the texture of the branches changing to papillate distal to the ring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Centaurea 9b. Involucral bracts with entire margins; involucre 1–7.5 mm tall; cypselas attached basally to the receptacle; style branches truncate and penicillate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Artemisia 8b. Receptacle with scale-like chaff 10a. Staminate and carpellate flowers in separate capitula, the staminate capitula usually the uppermost and possessing an undivided style; involucre armed with tubercles, spines, or prickles [Fig. 478]; carpellate flowers lacking a corolla 11a. Staminate involucre of distinct bracts; carpellate involucre a conspicuous, prickly bur, 8–40 mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthium 11b. Staminate involucre of connate bracts; carpellate involucre with 1 or more series of tubercles or spines [Fig. 371], 3–10 mm tall . . . . . . . . (in part) Ambrosia 10b. Flowers bisexual or unisexual and then the staminate and carpellate flowers in the same capitulum; involucre unarmed; all the flowers with a corolla (except in Cyclachaena xanthiifolia, in which the carpellate flowers sometimes lack a corolla) 12a. Plants woody; leaf blades white-tomentose, evergreen, 3-dimensionally pinnately dissected . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Santolina 12b. Plants herbaceous or usually woody in Iva frutescens; leaf blades glabrous or pubescent, but not white-tomentose, deciduous, simple to 3-times pinnately divided, relatively plane 13a. Involucre with 2 series of dimorphic bracts—the outer series larger, herbaceous to foliaceous, the inner series smaller, membranaceous, and usually striate [Fig. 381]; pappus of 2–6 awns, these usually retrosely barbellate (rarely antrorsely barbellate or smooth or absent) [Figs. 381, 382, 383] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Bidens

36 0   tricolpate s

13b. Involucre with 1 series of monomorphic, herbaceous or subherbaceous bracts; pappus none 14a. Leaves alternate; inner flowers of capitulum (i.e., the disk flowers) bisexual, with a bifid style; plants annual, 1–10 dm tall . . . . . (in part) Madia 14b. Leaves opposite (the uppermost sometimes alternate); inner flowers of capitulum functionally unisexual, the ovary abortive and the style undivided; plants perennial or coarse annuals, 5–30 dm tall 15a. Larger leaves with pinnately lobed and toothed blades [Fig. 445]; involucre 6–11 mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Polymnia 15b. Leaves with simple blades that are toothed or double toothed; involucre 1.5–4 mm tall 16a. Capitulescence (or the principal branches in robust plants) raceme-like or spike-like; peduncles 0–3 mm long, many subtended by leaf-like bracts; carpellate flowers with a corolla 0.5–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Iva 16b. Capitulescence panicle-like; peduncles 1–6 (–12) mm long, few (if any) subtended by leaf-like bracts; carpellate flowers with a corolla 0.1–0.5 mm long, or the corolla absent . . . . . . Cyclachaena 4b. Receptacle without bristles or chaff (the persistent floral stipes of Cotula sometimes misinterpreted as chaff); corollas yellow (or usually blue to lavender in Ageratum or lacking in the carpellate flowers of Ambrosia trifida) 17a. Leaves whorled [Fig. 448]; capitula solitary at the summit of the stem [Fig. 448]; amphibious plants usually found growing in shallow water . . . . . . . . . . . . . . . . Sclerolepis 17b. Leaves alternate or opposite; capitula usually more than one in the axils of leaves, at the tips of branches, and/or at the summit of the stem; terrestrial plants 18a. Leaves opposite 19a. Plants monoecious (i.e., with separate staminate and carpellate capitula); pappus absent; usually at least some leaf blades palmately lobed [Fig. 372] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Ambrosia 19a. Plants synoecious (i.e., with capitula that have bisexual flowers); pappus present; leaf blades simple to dissected, but none palmately lobed 20a. Leaf blades dissected into narrow segments; pappus composed of 10–20 scales that are cleft into 5–10 bristles at the apex; inner involucral bracts rounded to obtuse at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Dyssodia 20b. Leaf blades simple merely toothed; pappus composed of (4–) 5 (–6) scales that are fringed along the margins and terminate in a single, long awn; involucral bracts narrowed to an awn-like point 0.5–2 mm long . . . . Ageratum 18b. Leaves alternate 21a. Pappus of 2–8 firm, but deciduous, awns; involucral bracts squarrose and heavily resinous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Grindelia 21b. Pappus of scales, a short crown, or none; involucral bracts neither squarrose nor resinous 22a. Capitulescence a solitary capitulum at the tips of branches or in the axils of leaves; outermost series of flowers lacking a corolla;; low, procumbent or trailing herbs 23a. Capitula at the tips of branches; inner flowers stipitate, the stipes persistent on the receptacle; leaf blades (10–) 20–30 (–70) mm long; cypselas with two evident ribs or wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cotula

Ast e r ac e a e  3 6 1

23b. Capitula mainly in the axils of leaves; inner flowers not stipitate; leaf blades 5–15 mm long; cypselas with ridged but not winged . . . . . Centipeda 22b. Capitulescence with multiple capitula, all the flowers with a corolla; none of the flowers stipitate; erect or ascending herbs or shrubs 24a. Capitulescence resembling a spike, raceme, or panicle [Figs. 378, 379]; pappus none . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Artemisia 24b. Capitulescence resembling a corymb or cyme; pappus of scales or a short crown 25a. Pappus of 4–10 scales, the longer scales 2–6 mm wide 26a. Corolla lobes narrow-lanceolate to linear, mostly more than 2 times as long as wide; all corollas actinomorphic . . . . . . . . . . . . . . . . . . Palafoxia 26b. Corolla lobes lanceolate to triangular, mostly 1–2 times as long as wide; outer corollas enlarged and zygomorphic. . . . . . . . . . . . Chaenactis 25b. Pappus a short crown 0.2–0.4 mm long 27a. Leaf blades toothed, sometimes with a few basal lobes as well; cypselas with 5–8 rib-like nerves . . . . . . . . . . . . . . . . (in part) Balsamita 27b. Leaf blades 1- to 3-times pinnatifid; cypselas with 5 or fewer rib-like nerves 28a. Receptacle flat or low-convex [Fig. 470]; disk corollas 5-lobed; plants 4–15 dm tall . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Tanacetum 28b. Receptacle high-convex and pointed [Fig. 432]; disk corollas 4-lobed; plants 0.5–4 dm tall . . . . . . . . . . . . . . . . . (in part) Matricaria

Group 4 1a. Pappus composed of capillary bristles (also with an additional series of minute, slender scales in some Erigeron, with a short, outer crown in Callistephus) [Figs. 438, 468] 2a. Plants subdioecious, each capitulum composed almost entirely of unisexual flowers; stems scaly bracteate; well-developed leaves all basal, the blades palmately lobed [Fig. 442] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Petasites 2b. Plants polygamous, each capitulum with the ray flowers unisexual and carpellate and the disk flowers bisexual; stems with leaves; leaves various, but neither all basal nor with palmately lobed blades 3a. Rays up to 2 mm long, shorter than to scarcely exceeding the pappus, often inconspicuous in drying 4a. Involucral bracts glabrous and eciliate; ± glabrous saltmarsh plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Symphyotrichum 4b. Involucral bracts pubescent or ciliate or both; plants often with pubescent stems and/or leaf blades, at least with marginal cilia on the leaf blades, not occurring in saltmarshes 5a. Involucre with 3 or 4 series of foliaceous bracts of ± equal length; style appendages acute to acuminate . . . . . . . . . . . . . . . . . . . . (in part) Symphyotrichum 5b. Involucre with green but not at all foliaceous bracts, the bracts of similar or dissimilar length; style appendage acute to, more commonly, obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Erigeron

362  tricolpates

3b. Rays 2–35 mm long, exceeding the pappus, evident even in drying [Figs. 451, 462] 6a. Leaf blades pinnately lobed, each lobe terminating in a spine; involucral bracts ± spinulose-margined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Perezia 6b. Leaf blades entire to toothed (sometimes lobed in Callistephus), not spiny; involucral bracts without spinulose margins (though the apex sometimes with a spine tip) 7a. Surface of ovary with minute stalked glands [Fig. 438]; capitula nodding in bud; leaves reduced in size toward the base, the lowest scale-like . . . . . . . . . Oclemena 7b. Surface of ovary without glands; capitula erect in bud (spreading or sometimes nodding in Ionactis); leaves larger toward the stem base 8a. Capitula frequently in glomerules, with 3–8 ray flowers [Fig. 451]; foliage leaves with sessile glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sericocarpus 8b. Capitula usually borne singly at the ends of peduncles, with 7–100 or more ray flowers [Figs. 462, 466]; foliage leaves without glands (Oligoneuron album with obscure resin glands, and the bracteal leaves of some Symphyotrichum and Eurybia with stipitate glands) 9a. Basal leaves gradually tapering to an elongate petiole, the blade and petiole combined commonly 30–40 cm long; involucral bracts usually with a thin anthocyanic stripe along each margin; rare escape in New England . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aster 9b. Basal leaves shorter than 30 cm, lacking petioles, and/or abruptly contracted to a petiole; involucral bracts rarely with marginal anthocyanic stripes, anthocyanin, if present, usually distributed near the center and/or apex of the bract (rarely throughout); primarily native plants, though some species weedy 10a. Involucral bracts in three series—the outer series foliaceous and green ± throughout, the inner series membranaceous to scarious; capitula solitary and the ends of branches, 6–8 cm in diameter; involucre double—outer series a short, membranous crown, inner series of minutely barbellate bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callistephus 10b. Involucral bracts variable, usually not foliaceous (except in a few Symphyotrichum); capitula solitary, in glomerules, or few to many along branches; involucre single or double, but not with an outer, membranous crown 11a. Involucral bracts of ± equal length (sometimes with some very small bracts near the base of the involucre), green throughout or in large part, but not foliaceous; style appendage acute to, more commonly, obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Erigeron 11b. Involucral bracts of dissimilar lengths, usually pale at the base with an apically dilated green midzone [Figs. 462, 464, 466], less commonly foliaceous or anthocyanic (green throughout in Doellingeria); style appendages acute to acuminate 12a. Pappus bristles of 2 distinctly uneven lengths—a very short outer series and 1 or 2 series of elongate bristles of nearly even length [Fig. 438]; involucral bracts neither foliaceous nor with a distinct, green apical zone [Fig. 396] 13a. Longer series of pappus bristles thickened at the apex; involucral bracts with raised midvein, but not keeled; rays not coiling; disk flowers abruptly expanded apically; ovaries terete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Doellingeria

Ast e r ac e a e   3 63

13b. Longer series of pappus bristles slender at the apex; involucral bracts somewhat keeled; rays coiling; disk flowers tubular; ovaries compressed . . . . . . . . . . . . . . . . . . . . . . . . . Ionactis 12b. Pappus bristles all elongate, in 2 or 3 series of nearly even length [Fig. 468]; involucral bracts with a distinct green apical zone or entirely foliaceous in a few species (anthocyanic pigments sometimes also present) [Figs. 462, 464, 466] 14a. Disk corollas white; leaf blades with sessile resin glands (these sometimes obscure); pappus bristles thickened at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Oligoneuron 14b. Disk corollas yellow or becoming purple, red, or red-brown; leaf blades without sessile resin glands (though the bracteal leaves stipitate-glandular in a few species); pappus bristles slender at the apex or thickened in some Eurybia 15a. Rays 2–3 mm long [Figs. 452, 459]; capitulescence very slender, resembling a thyrse, sometimes with 1 or more elongate, slender, ascending branches resembling the main axis; disk flowers persistently pale yellow . . . . . . . . (in part) Solidago 15b. Rays 3–30 mm long [Figs. 462, 465]; capitulescence resembling a panicle or corymb or infrequently composed of 1 or few capitula; disk flowers yellow, becoming purple, red, or red-brown in age 16a. Outer and middle involucral bracts less than 2.5 times as long as wide, rounded to obtuse at the apex, densely ciliate along the margins, with a thumbnail to rhombic shaped chlorophyllous zone at the tip [Fig. 407]; capitulescence corymb-like; pappus bristles sometimes thickened at the apex; ovary terete . . . . . . . . . . . . . . . Eurybia 16b. Involucral bracts more than 3 times as long as wide, obtuse to acuminate at the apex, eciliate or sparingly ciliate along the margins, with a rhombic to basally tapering chlorophyllous zone at the tip or entirely foliaceous [Figs. 462, 464, 465]; capitulescence commonly panicle-like when well formed; pappus bristles slender at the apex; ovary compressed in most species . . . . (in part) Symphyotrichum 1b. Pappus composed entirely of scales, awns, or a short crown, or absent 17a. Leaves opposite (the upper may be alternate) 18a. Leaf blades 10–30 cm long, pinnately lobed with broad lobes [Fig. 445]; disk flowers functionally staminate, the ovary abortive and with an undivided style; pappus absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Polymnia 18b. Leaf blades 2–12 cm long, simple or pinnately dissected into linear to linear-filiform segments; disk flowers bisexual with a bifid style; pappus present, at least on the disk flowers, though reduced to a tiny crown in some species (absent in Sigesbeckia) 19a. Leaf blades pinnately dissected into numerous linear to linear-filiform segments narrower than 1.5 mm; rays 5–40 mm long; cypsela body tapering to a slender, apical beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cosmos 19b. Leaf blades simple or infrequently with a few basal lobes in Coreopsis rosea; rays up to 13 mm long; cypsela body not taping to an elongate beak 20a. Rays 8–13 mm long; disk 5–10 mm wide; leaf blades linear . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Coreopsis

36 4   tricolpate s

20b. Rays 1–2 mm long; disk 3–6 mm wide in flower; leaf blades narrow-lanceolate to ovate [Fig. 412] 21a. Involucre biseriate and dimorphic—an outer series of longer, spreading to reflexed bracts and an inner series of shorter, erect bracts—conspicuously stipitate-glandular; leaf blades resin-dotted abaxially . . . . (in part) Sigesbeckia 21b. Involucre uniseriate or biseriate, with none of the bracts spreading to reflexed, stipitate-glandular or not; leaf blades not resin-dotted abaxially 22a. Disk flowers yellow; capitula with 3–6 rays [Fig. 412], commonly 5; pappus of fimbriate scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Galinsoga 22b. Disk flowers white; capitula with numerous ray flowers; pappus a tiny crown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eclipta 17b. Leaves alternate throughout the stem or all basal 23a. Receptacle with spine-like setae; lobes of disk corollas villous-tomentose; pappus composed of 6–10 conspicuously awned scales . . . . . . . . . . . . . . . . . . . (in part) Gaillardia 23b. Receptacle either naked, with scale-like chaff (though the chaff may be spinescent at the tip), or bristly; lobes of the disk corollas glabrous; pappus various, but not composed of awned scales 24a. Receptacle chaffy, at least toward the middle [Fig. 397] 25a. Involucral bracts herbaceous, without a scarious margin, spreading to reflexed; leaf blades simple and ± entire; chaff with a stout, spinescent tip . . . . . . . Echinacea 25b. Involucral bracts dry, scarcely or not at all herbaceous, scarious-margined (without scarious margins in Parthenium), appressed; leaves pinnately divided (except Achillea ptarmica with simple leaves); chaff without a spine-like tip 26a. Pappus usually composed of a pair of scales; disk flowers functionally staminate, the ovary abortive and the style undivided . . . . . . . . . . . . Parthenium 26b. Pappus absent or represented by a minute crown; disk flowers bisexual, fertile, with a bifid style 27a. Rays 5–14 mm long, numbering 10–16 per capitulum [Fig. 376]; disk 5–10 mm wide; disk flowers yellow; capitulescence not resembling a corymb, the capitula located at the tips of branches . . . . . . . . . . . . . . . . . . . . . Anthemis 27b. Rays 2–5 mm long, mostly numbering 4–10 per capitulum (up to 15 in cultivated forms of A. ptarmica); disk 2–8 mm wide; disk flowers white capitulescence resembling a corymb . . . . . . . . . . . . . . . . . . . . . . . . . . . . Achillea 24b. Receptacle without chaff 28a. Stems scapose, the leaves all basal; capitulescence a solitary capitulum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bellis 28b. Stems leafy; capitulescence with multiple capitula or solitary in Arctanthemum, Arctotis, Leucanthemum, and Nipponanthemum 29a. Pappus of disk flowers composed of 5–8 oblong to ovate, hyaline scales 0.5–4 mm long; cypsela with a conspicuous basal tuft of long, silky hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctotis 29b. Pappus of disk flowers absent, a short crown, or dimorphic and composed of barbellate awns and shorter awns or flat scales; cypsela lacking a long basal tuft of silky hairs 30a. Pappus of disk flowers composed of 2–4 long, barbellate awns and a variable number of shorter bristles or flat scales; style branches not truncate at apex, lacking an apical tuft of hairs

Ast e r ac e a e   3 6 5

31a. Leaves with simple and entire blades, not clasping the stem; pappus of 2 or 3 awns and 7–12 shorter bristles or scales . . . . . . . . . . . . . . Boltonia 31b. Leaves with subpalmately lobed blades, narrowed to a winged petiole, expanded at the base and clasping the stem; pappus 2–4 awns and 2–4 scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calotis 30b. Pappus a short crown or absent; style branches truncate and penicillate 32a. Leaf blades twice pinnatifid, with linear to filiform ultimate segments; receptacle convex, rounded, or pointed 33a. Receptacle conic, acute at the apex [Fig. 432]; cypsela with an oblique attachment scar near base, the body with 3–5 raised, but not wing-like, ribs, lacking apical resin glands; plants pleasantly aromatic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Matricaria 33b. Receptacle dome-shaped, rounded at the apex; cypsela with basal attachment scar, the body with 3, prominently thickened and almost wing-like ribs, with apical resin glands; plants nearly inodorous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tripleurospermum 32b. Leaf blades toothed, subpalmately lobed, or 1- to 2-times pinnatifid into oblong-elliptic to ovate primary segments; receptacle flat or low-convex 34a. Plants subshrubs, with deciduous lower leaves, so most of the stems leaves are clustered near the apex of the shoot; leaf blades somewhat succulent, evergreen [Fig. 437] . . . . . . . . Nipponanthemum 34b. Plants herbaceous, the leaves not clustered near the stem apex; leaf blades not succulent, deciduous 35a. Pappus present, a short crown; capitula 6–20 mm across in life, arranged in corymb-like clusters; disk 4–9 mm wide 36a. Leaf blades crenate-dentate, sometimes the larger with a few basal lobes, the lower blades 10–25 cm long. . . (in part) Balsamita 36b. Leaf blades with 3–9 oblong-elliptic to ovate segments that may be again lobed, none of the blades exceeding 8 cm in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Tanacetum 35a. Pappus absent (rarely some ray flowers with wall tissue prolonged to appear as a short crown in Leucanthemella and Leucanthemum); capitula 20–60 mm across in life, solitary at the tips of branches or arranged in corymb-like clusters; disk 10–25 mm wide 37a. Ray flowers sterile, not producing fruits; leaf blades glandular-punctate; disk corolla tubes provided with resin glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucanthemella 37b. Ray flowers fertile, producing fruits; leaf blades without glandular-dots; disk corolla tubes lacking resin glands 38a. Disk flowers compressed and winged near base; body of cypsela with 10 ribs; pericarp of cypsela with resin canals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucanthemum 38b. Disk flowers neither compressed nor winged near base; body of cypsela with 5–8 ribs; pericarp of cypsela without resin canals 39a. Leaf blades glabrous or subglabrous, the lower flabellate to spatulate, nearly or fully as wide as long; capitulescence consisting of a solitary capitulum; rays white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arctanthemum

36 6   tricolpate s

39b. Leaf blades gray-pubescent, lanceolate or elongateelliptic to ovate, clearly longer than wide; capitulescence consisting of 1 to several capitula in a corymb-like array; rays commonly yellow, pink, purple, red, or white (but other color forms also exist) . . . . . . . . . . (in part) Chrysanthemum

Group 5 1a. Anthers sagittate-tailed at the base; involucres with 3–7 series of involucral bracts 2a. Leaf blades lanceolate or elliptic to ovate, 12–200 mm wide; involucres 8–20 mm in diameter; rays 15–40 mm long, conspicuously surpassing the involucres; body of cypsela not contracted just below the attachment of the pappus bristles; pappus bristles 6–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Inula 2b. Leaf blades linear to narrow-lanceolate, 1–3 (–10) mm wide; involucres 4–7 mm in diameter; rays 4–7 mm long, not or scarcely exceeding the involucres; body of the cypsela abruptly contracted just below the attachment of the pappus bristles; pappus bristles 3–4 (–5) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dittrichia 1b. Anthers cuneate to sagitate at the base, but not tailed; involucres with 1 or 2 series of involucral bracts or with 3–5 series in Chrysopsis, Euthamia, Heterotheca, Oligoneuron, Pityopsis, Solidago 3a. Rays usually yellow with a white tip; receptacle with chaff; disk corollas puberulent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Layia 3b. Rays entirely yellow; receptacle without chaff; disk flowers ± glabrous (sometimes sparsely pubescent in Heterotheca) 4a. Middle stem leaves with a short, broadly winged petiole that prominently expands and clasps the stem; upper stem leaves cordate-clasping 5a. Lower leaf blades broad-ovate to nearly orbicular, (5–) 7–14 cm wide; involucres 22–40 mm tall; pappus of disk flowers consisting of only bristles; plants perennial, from rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Doronicum 5b. Lower leaf blades ovate to elliptic or lanceolate, 0.6–5.5 cm wide; involucres 4–8 (–10) mm tall; pappus of disk flowers consisting of both scales and bristles; plants annual or biennial, from taproots . . . . . . . . . . . . . . . . . . . . . (in part) Heterotheca 4b. Middle stem leaves sessile or petioled, but without an expanded, clasping petiole; upper stem leaves not cordate-clasping 6a. Leaves opposite, except sometimes the upper; rays 15–20 mm long . . . . . . Arnica 6b. Leaves alternate throughout the stem; rays shorter than 10 mm (except Chrysopsis with rays 10–20 mm long) 7a. Stems with only small, bract-like leaves (i.e., scapose) [Fig. 475], appearing and flowering before the cordate-suborbicular blades of the basal leaves are produced; disk flowers sterile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tussilago 7b. Stems with leaves, these present during flowering [Fig. 444]; disk flowers fertile 8a. Involucre composed of a single series of long bracts, sometimes also calyculate (i.e., with a short, outer series of bracts) [Fig. 449] 9a. Plants perennial, with rhizomes and fibrous roots; leaves basally disposed (i.e., prominent clusters of basal leaves present, the stem leaves rapidly reduced in size upwards) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Packera 9b. Plants annual or biennial (rarely short-lived perennial), mostly with evident taproots; leaves chiefly cauline (i.e., prominent clusters of basal leaves absent, the stem leaves gradually, if at all, reduced upwards)

Ast e r ac e a e  3 67

10a. Rays 4–8 mm long; leaf blades relatively more divided, usually 2- or 3-times pinnatifid; cypsela bodies from near margin of capitulum glabrous, the inner ones pubescent . . . . . . . . . . . . . . . . . . . . . . . . . Jacobaea 10b. Rays absent or up to 2 mm long [Fig. 449]; leaf blades relatively less divided, usually toothed to pinnatifid; usually all the cypsela bodies of the capitulum similar, either all pubescent or all glabrous in glandular-hairy S. viscosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Senecio 8b. Involucre with (2–) 3–5 series of bracts of distinctly unequal lengths [Figs. 439, 444] 11a. Capitulescence not at all flat-topped, resembling a panicle or thyrse or consisting of axillary clusters of capitula [Figs. 452, 458] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Solidago 11b. Capitulescence or its divisions flat-topped, resembling a corymb 12a. Pappus monomorphic—composed of elongate and slender bristles 13a. Leaves chiefly cauline, the blades linear to narrow-lanceolate, parallel-veined, resinous-punctate [Fig. 409]; involucral bracts with a single, median nerve; lower petiole bases not persistent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euthamia 13b. Leaves basally disposed, the blades elliptic or broad-lanceolate to broad-ovate, pinnately veined, not resinous-punctate; involucral bracts longitudinally few-striate [Fig. 439]; lower petiole bases often persistent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Oligoneuron 12b. Pappus dimorphic—an inner series of elongate, slender bristles and an outer series of short, course bristles or narrow scales 14a. Leaf blades linear, arcuate, parallel-veined, sessile, entire [Fig. 444]; cypsela body narrow-fusiform; involucre white-pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pityopsis 14b. Lower leaf blades lanceolate or oblanceolate to ovate, ± straight, pinnately veined, ± petiolate, toothed to entire; cypsela body obovoid to obconic; involucre stipitate-glandular 15a. Pappus monomorphic, similar on both ray and disk flowers; cypselas monomorphic, all compressed; rays 8–11 × 2–3 mm; stems sericeous to sericeo-tomentose . . . . . . . . . . . . . Chrysopsis 15b. Pappus dimorphic—absent on ray flowers and present on disk flowers; cypselas dimorphic, those of the ray flowers triangular in cross-section, those of the disk flowers compressed; rays 3–7 (–9) × 1–2 mm; stems hirsute to pilose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Heterotheca

Group 6 1a. Receptacle naked 2a. Capitula tiny, sessile and aggregated in head-like glomerules, each with a single ray flower; involucre up to 1 mm in diameter, with usually 2 bracts; upper leaf blades sessile and often connate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Flaveria 2b. Capitula larger, some or all peduncled, not aggregated in glomerules, each with (1–) 3 or more ray flowers; involucre wider than 1 mm in diameter, with 3 or more bracts; upper leaf blades sessile or petioled, but not basally connate 3a. Leaves opposite and with simple, entire blades . . . . . . . . . . . . . . . . . . . . . . . . . Lasthenia

36 8   tricolpate s

3b. Leaves alternate or opposite, the blades toothed to lobed or divided (blades usually entire in Helenium and sometimes in Calendula, but then the leaves alternate) 4a. Leaves opposite (at least the lower), the blades once or twice pinnately divided or -lobed; involucral bracts gland-dotted 5a. Pappus monomorphic, composed of 10–20 scales that are cleft into 3–10 divisions or bristles at the apex; involucre biseriate—the outer series with 3–9 bracts, the inner series with 8–22 bracts 6a. Capitula with 5–8 ray flowers; inner series of 8 distinct involucral bracts; pappus scales apically cleft into 5–10 bristles . . . . . . . . . . . . . . (in part) Dyssodia 6b. Capitula with 10–21 ray flowers; inner series of 12–22 involucral bracts that are connate 80% or more of their length; pappus scales apically cleft into 3–5 divisions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Thymophylla 5b. Pappus scales dimorphic, none of which are divided at the apex into bristles; involucre either uniseriate or biseriate and then with only 2 outer bracts 7a. Capitula with (1–) 3–8 (up to 100+ in cultivars) ray flowers, the ray 1–18 (–25) mm long; pappus of 3–7 scales—1 or 2 longer scales and 2–5 shorter scales; involucre uniseriate, bracts largely connate . . . . . . . . . . . . . . . . . . . Tagetes 7b. Capitula with (0–) 1 ray flower, the ray 0.8–1.2 mm long; pappus of 8 scales that usually are alternately unequal; involucre biseriate, the bracts distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schkuhria 4b. Leaves alternate, simple or 1–3 times pinnately lobed; involucral bracts not gland-dotted (except in Helenium and Hymenoxys) 8a. Pappus a tiny crown or completely absent 9a. Cypsela bodies tuberculate-rugose on the outer surface (i.e., the surface facing the margin of the capitulum); disk flowers functionally staminate with an abortive ovary, not producing fruit; capitula 4–7 (–10) cm wide in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calendula 9b. Cypsela bodies not tuberculate-rugose on any surface; disk flowers bisexual or unisexual (and then carpellate), producing fruit; capitula up to 4 (–5) cm wide in life (2.5–20 cm in diameter in Chrysanthemum) 10a. Capitulescence consisting of (10–) 20–70 (–100) capitula in a compact, corymb-like array . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Tanacetum 10b. Capitulescence consisting of 1–3 capitula 11a. Herbage glabrous or nearly so; cypselas dimorphic—those of the ray flowers 3-angled with 2 or 3 wing-margins, those of the disk flowers ± cylindrical; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glebionis 11b. Herbage gray-pubescent (sometime glabrate in late season); cypselas monomorphic, angled or ribbed, but without wing-angles; plants perennial 12a. Rays longer than 10 mm; leaf blades prominently toothed to lobed; introduced plants of settled areas . . . . . . . . . (in part) Chrysanthemum 12b. Rays up to 4 mm long; leaf blades 2- to 3-times pinnately compound; native plants of northern, ice-scoured river shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Tanacetum 8b. Pappus composed of distinct awns or scales 13a. Inner series of involucral bracts connate 80% or more of their length; pappus of apically cleft scales . . . . . . . . . . . . . . . . . . . . . . . . (in part) Thymophylla 13b. Involucral bracts distinct or connate only at base; pappus of awns or simple scales

Ast e r ac e a e  3 6 9

14a. Involucral bracts resinous; pappus composed of 2–8 firm, but deciduous, awns; capitula with 25–40 rays . . . . . . . . . . . . . . (in part) Grindelia 14b. Involucral bracts resin-dotted; pappus composed of 5–10 lanceolate to ovate or obovate, awn-tipped scales; capitula with 5–21 rays 15a. Involucral bracts reflexed in fruit; mature receptacles ± globose [Fig. 416]; stems winged by decurrent leaf bases (not so in H. amarum); leaf blades simple (rarely pinnately lobed in H. amarum) . . . . . . Helenium 15b. Involucral bracts spreading to erect in fruit; mature receptacles hemispherical; stems unwinged; larger leaf blades usually pinnately lobed into very narrow segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hymenoxys 1b. Receptacle bristly or chaffy, at least toward the margin of the capitulum 16a. Leaves regularly alternate throughout the stem 17a. Receptacle with spine-like setae; lobes of disk corollas villous-tomentose; pappus composed of 6–10 conspicuously awned scales . . . . . . . . . . . . . . . . . . . (in part) Gaillardia 17b. Receptacle chaffy with flattened scales; lobes of disk corollas not villoustomentose; pappus various but not composed of awned scales 18a. Involucral bracts in 1 series, laterally compressed and enfolding a ray cypsela; receptacle chaffy only near the margin; plants aromatic, with stipitate glands (at least in the capitulescence) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Madia 18b. Involucral bracts in 1 or more series, flat, not enfolding a ray cypsela; receptacle chaffy throughout; plants aromatic or not, but lacking abundant stipitate glands 19a. Capitula with 20–40 ray flowers; involucral bracts dry and scarious-margined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cota 19b. Capitula with 2–21 ray flowers [Fig. 447]; involucral bracts herbaceous or foliaceous, commonly without scarious margins 20a. Receptacle convex to low-conic; cypsela body with evident lateral wings (very rarely the wings lacking); leaves either auriculate-dilated at base or decurrent on the stem as conspicuous wings . . . . . . . . . . . . . . . . . . . . . Verbesina 20b. Receptacle conic to columnar [Fig. 447]; cypsela body lacking wings or with narrow wing margins; leaves neither abruptly and conspicuously broadened at base nor decurrent as prominent wings 21a. Ray flowers subtended by chaffy scales; chaff velutinous at tip; cypsela body compressed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ratibida 21b. Ray flowers not subtended by chaff; chaff not velutinous at tip; cypsela body ± quadrangular. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rudbeckia 16b. Leaves opposite or whorled, except sometimes the upper, which are alternate [Fig. 418] 22a. Inner involucral bracts 5- to 7-ribbed, each rib bearing 1 or 2 rows of hooked prickles 1–2 mm long, maturing into a bur in fruit; annuals with prostrate stems commonly rooting from the lower nodes . . . . . . . . . . . . . . . . . . . . . . . . . Acanthospermum 22b. None of the involucral bracts bearing prickles or maturing as a bur; annual or perennial plants, mostly with ascending stems (except the aquatic Bidens beckii) 23a. Leaves, at least the upper, connate-perfoliate; stem square; disk flowers unisexual and staminate, with an undivided style and abortive ovary . . . . . . . Silphium 23b. Leaves not connate-perfoliate; stem terete or angled; disk flowers bisexual, the style divided and the ovary fertile

370  tricolpates

24a. Plants aquatic, with flaccid stems; leaves (except for the emersed ones) whorled, finely dissected into narrow, flaccid segments; capitulescence commonly a solitary capitulum [Fig. 380]; cypselas terete . . . . . . . . . . . . . . . . . . . (in part) Bidens 24b. Plants terrestrial or of wetlands, with erect stems; leaves opposite, entire or divided into broad, firmer segments; capitulescence commonly composed of 2 or more capitula; cypselas compressed, quadrangular, or ± triangular 25a. Involucral bracts conspicuously stipitate glandular, biseriate, dimorphic— the outer longer and spreading, the inner shorter and erect; leaf blades resin-dotted abaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Sigesbeckia 25b. Involucral bracts not stipitate-glandular (though with sessile glands or rarely with glands on short stipes near the margins in Deinandra), arranged in 1 or more series, when biseriate the outer involucral bracts not both longer than the inner and spreading (except in some Bidens); leaf blades not resindotted abaxially (except in Deinandra and Heliomeris and some Helianthus) 26a. Disk corollas densely tomentose basally; involucral bracts uniseriate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Guizotia 26b. Disk corollas glabrous basally; involcural bracts in 2 or more series (in 1 series in Deinandra, but that genus with only 6 disk flowers, rather than 100+ in Guizotia) 27a. Chaff in a single series between the ray and disk flowers; pappus composed of 5–12 lanceolate to oblong or linear scales 1–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Deinandra 27b. Chaff not restricted to a single series ; pappus absent, of 2–4 toothlike scales, or of 2–4 (–5) awns that are commonly minutely barbellate; involucre with 2 or more series of bracts 28a. Ray flowers carpellate and fertile, becoming chartaceous in fruit and persistent on the triangular cypsela [Fig. 421] . . . . . . . . . . Heliopsis 28b. Ray flowers neutral and lacking carpels (with carpels in Heterosperma, but that genus with pinnately lobed leaves), deciduous at or before maturity of the compressed or quadrangular cypsela 29a. Involcural bracts biseriate or triseriate, ± monomorphic [Fig. 417]; chaff of the receptacle partially enfolding the disk flowers; cypsela bodies compressed at right angles to the involucral bracts 30a. Pappus present, of 2 narrow scales (1–) 1.2–5 mm long positioned at the principal angles and 0–4 shorter scales (note: the scales are readily deciduous); cypsela body glabrous or pubescent; leaf blades 1.2–40 cm wide, at least the larger serrate or serrulate to subentire (rarely entire) and usually 3-nerved (1-nerved in H. maximiliani) . . . . . . . . . . . . . . Helianthus 30b. Pappus absent; cypsela body glabrous; leaf blades 0.2–2 cm wide, entire, 1 (–3)-nerved . . . . . . . . . . . . . . . . . . . . . . . . . . Heliomeris 29b. Involucral bracts biseriate, dimorphic [Fig. 381]; chaff flat or nearly so, not or only slightly enfolding the disk flowers; cypsela bodies compressed parallel to the involucral bracts 31a. Cypsela body tapering to a slender, apical beak; leaf blades pinnately lobed or dissected into linear or linear-filiform segments up to 1.5 (–3) mm wide (lanceolate an up to 5 mm wide in the rare introduction Cosmos sulphureus); capitula usually with 3–20 disk flowers

Ast e r ac e a e   37 1

32a. Ray flowers 1–3, with rays usually 1–2 mm long; disk flowers 3–10; leaf blades 1–4 cm long . . . . . . . . Heterosperma 32b. Ray flowers usually 8, with rays 15–50 mm long; disk flowers 10–20; leaf blades 5–12 (–25) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cosmos 31b. Cypsela body not terminated by a beak; leaf blades simple, lobed, or divided into linear to ovate segments mostly wider than 1.5 mm (narrower in a few Coreopsis); capitula with (7–) 12–150+ disk flowers 33a. Pappus of 2–6 awns, these usually retrorsely barbellate (rarely antrorsely barbellate or smooth or absent) [Figs. 381, 382, 383]; outer involucral bracts foliaceous in most species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Bidens 33b. Pappus of 2 short, tooth-like scales or none; cypsela wing-margined; outer involucral bracts ± herbaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Coreopsis

Acanthospermum 1. Acanthospermum australe (Loefl.) Kuntze E Paraguay starburr. Melampodium australe Loefl. • MA. Wool waste, gardens, waste areas.

Achillea 1a. Rays yellow; stems glandular-punctate 2a. Stems villous; ultimate segments of leaf blades 1.5–8 mm wide; involucral bracts sparsely to moderately pubescent (i.e., the surface not concealed by hairs) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. filipendulina 2b. Stems villous-tomentose; ultimate segments of leaf blades usually narrower than 1 mm; involucral bracts moderately to densely pubescent abaxially (i.e., the surface largely concealed by hairs) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. tomentosa 1b. Rays white to pink; stems not glandular-punctate 3a. Leaf blades subentire to serrate with short teeth; rays numbering (5–) 8–10 (–15) per capitulum, 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ptarmica 3b. Leaf blades pinnately dissected into narrow segments; rays numbering (4–) 5 (–6) per capitulum, 1–3 (–4) mm long 4a. Leaves near the middle of the stem usually with 15 or more primary divisions, the blades lanceolate to linear, the divisions usually not plane . . . . . . . . . . . . . A. millefolium 4b. Leaves near the middle of the stem with 10 or fewer lobes, the blades elliptic to ovate, with plane lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ligustica 1. Achillea filipendulina Lam. E Fern-leaved yarrow. VT. Gardens, waste areas. 2. Achillea ligustica All. E Ligurian yarrow. VT. Gardens, waste areas. 3. Achillea millefolium L. ssp. lanulosa (Nutt.) Piper N common yarrow. Achillea borealis Bong.; A. lanulosa Nutt.; A. millefolium L. ssp. borealis (Bong.) Breitung; A. millefolium L. var. lanulosa (Nutt.) Piper; A. millefolium L. var. nigrescens E. Mey.; A. millefolium L. ssp. occidentalis (DC.) Hyl.; A. millefolium L. var. occidentalis DC.;

372  tricolpate s

A. nigrescens (E. Mey.) Rydb. • CT, MA, ME, NH, RI, VT. Fields, roadsides, open areas, coastal headlands, river beaches, alpine plateaus. Collections from New England appear to largely represent our native subspecies (lanulosa). Introduced material of the Old World ssp. millefolium appear to be confined to garden settings. It is distinguished by its leaf blades with broader segments ± oriented in the same plane (vs. narrow segments oriented in more than 1 plane in ssp. lanulosa). More work is needed to confirm its presence as a naturalized plant. Plants identified as Achillea borealis from New England merely represent individuals that show dark margins to the involucral bracts (an ecological response to exposure). 4. Achillea ptarmica L. E pearl yarrow. CT, MA, ME, NH, RI, VT. Roadsides, fields, abandoned homesteads, gravel beaches. 5. Achillea tomentosa L. E woolly yarrow. MA. Wool waste.

Ageratina Reference: Clewell and Wooten (1971). 1a. Leaf blades firm, 3–7 (–10) × 2–5 (–5.5) cm, obtuse to acute at the apex, with 7–12 well-developed, rounded to bluntly pointed teeth per margin, 4 or more times as long as the associated petiole; petioles 1–15 (–20) mm long (rarely lacking altogether); roots 0.8–1 (–1.5) mm thick; capitula with 10–19 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . A. aromatica 1b. Leaf blades herbaceous, (5–) 6–18 × 3–11 (–12) cm, short-acuminate to acuminate at the apex, with 9–25 well-developed, pointed teeth per margin, 1.5–5 times as long as the petiole; petioles of principal leaves 20–80 mm long; roots 0.3–0.8 (–1) mm thick; capitula with (9–) 12–25 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. altissima 1. Ageratina altissima (L.) King & H.E. Robins. var. altissima N Fig. 370

Fig. 370  Capitulescence of Ageratina altissima showing involucre of similar length bracts.

white snakeroot. Ageratum altissimum L.; Eupatorium rugosum Houtt.; Eupatorium rugosum Houtt. var. chlorolepis Fern.; Eupatorium rugosum Houtt. var. tomentellum (B.L. Robins.) Blake; Eupatorium rugosum Houtt. var. villicaule (Fern.) Blake • CT, MA, ME, NH, RI, VT. Mesic, deciduous forests, especially rich types, riparian forests. 2. Ageratina aromatica (L.) Spach

NC

lesser snakeroot. Eupatorium aromaticum L. • CT, MA, RI. Dry-mesic to mesic, often rocky, forests and forested slopes.

Ageratum 1a. Peduncles pubescent, in part, with glandular hairs; involucral bracts narrow-lanceolate, gradually tapering to an awn-like point 0.8–2 mm long, the abaxial surface stipitate-glandular, eciliate or sparsely ciliate on the margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. houstonianum 1b. Peduncles pubescent with eglandular hairs; involucral bracts oblong-lanceolate, abruptly tapering to an awn-like point 0.5–1 mm long, the abaxial surface eglandular, ciliate on the margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. conyzoides 1. Ageratum conyzoides L. E tropical whiteweed. CT. Ballast, areas of habitation. 2. Ageratum houstonianum P. Mill. E Houston’s whiteweed. MA; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Roadsides, waste areas, areas of habitation.

Amberboa 1. Amberboa moschata (L.) DC. E sweet sultan. Centaurea moschata L. • ME. Roadsides, fields, areas of habitation.

Ast e r ac e a e  373

Ambrosia Reference: Strother (2006a). 1a. Staminate capitula sessile or subsessile, in a solitary, spike-like capitulescence; staminate involucres prolonged into a long, retrorse, lanceolate to triangular-ovate, hooded, bristly tooth; leaf blades lanceolate to narrow-lanceolate, entire to lobed, when lobed with 1 or 2 pairs of pinnately arranged lobes that are confined to the base of the blade . . . . A. bidentata 1b. Staminate capitula peduncled, borne in 1 or more raceme-like capitulescences; staminate involucres without a prolonged, retrorse lobe, actinomorphic or nearly so; leaf blades lanceolate or elliptic to ovate or triangular, 1- or 2-times pinnately lobed, or entire to palmately lobed in A. trifida, but that species with some broad-lanceolate to ovate, unlobed blades on the plant 2a. Plants annual, 0.5–5 m high; leaves opposite throughout, the blades entire to palmately lobed with 3 (–5) lobes [Fig. 372]; receptacle without chaff; staminate involucre with 3 evident nerves on one side; carpellate involucre 5–10 mm long in fruit . . . . . . . . . . A. trifida 2b. Plants annual or perennial, 0.2–1 (–2.5) m high; leaves usually opposite below and alternate above, once- or twice-pinnatifid; receptacle with chaff; staminate involucre inconspicuously nerved (i.e., the faint nerves of ± equal prominence in all directions); carpellate involucre mostly 3–5 mm long in fruit 3a. Plants usually perennial from a creeping rootstock; leaf blades usually oncepinnatifid, relatively thicker; carpellate involucre with 4 tubercles near the apex, these sometimes short and inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. psilostachya 3b. Plants annual; leaf blades once-, or more commonly, twice-pinnatifid, relatively thinner; carpellate involucre with 4–7 sharp spines near or above the middle [Fig. 371] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. artemisiifolia 1. Ambrosia artemisiifolia L. N Fig. 371 common ragweed. Ambrosia artemisiifolia L. var. elatior (L.) Descourtils; A. artemisiifolia L. var. paniculata (Michx.) Blank.; A. elatior L.; A. monophylla (Walt.) Rydb.; A. paniculata Michx. • CT, MA, ME, NH, RI, VT. Roadsides, fields, areas of cultivation, disturbed soil. ‌1 × 4. Ambrosia ×helenae Rouleau is an extremely rare ragweed hybrid known only from CT in New England. It ismost likely to be confused with A. trifida due to the palmately or subpalmately lobed leaf blades. However, the upper leaves (i.e., those subtending branches of the capitulescence) are frequently alternate, the lobes on the blades are often with additional lobes, and the staminate involucres have relatively weak veins (vs. leaves opposite throughout, lobes on leaf blades without additional lobes, and staminate involucres with 3 (–4) strong veins on one side in A. trifida).

Fig. 371  Carpellate involucre with small spines of Ambrosia artemisiifolia.

2. Ambrosia bidentata Michx. E lance-leaved ragweed. CT. Roadsides, fields, disturbed soil. 3. Ambrosia psilostachya DC. E perennial ragweed. Ambrosia coronopifolia Torr. & Gray; A. psilostachya DC. var. coronopifolia (Torr. & Gray) Farw. • CT, MA, ME, NH, VT. Roadsides, fields, railroads, disturbed soil. 4. Ambrosia trifida L. var. trifida N Fig. 372 giant ragweed. Ambrosia trifida L. var. integrifolia (Muhl. ex Willd.) Torr. & Gray • CT, MA, ME, NH, RI, VT. Edges of cultivated fields, barnyards, railroads, disturbed soil.

Anaphalis 1. Anaphalis margaritacea (L.) Benth. & Hook. f. N pearly everlasting. Anaphalis margaritacea (L.) Benth. & Hook. f. var. angustior (Miq.) Nakai; A. margaritacea (L.) Benth. & Hook. f. var. intercedens Hara; A. margaritacea (L.) Benth. & Hook. f. var. occidentalis Greene; A. margaritacea (L.) Benth. & Hook. f. var. subalpina Gray; Gnaphalium margaritaceum L. • CT, MA, ME, NH, RI, VT. Roadsides, forest openings, river beaches, fields.

Fig. 372  Leaf blade of Ambrosia trifida.

374 tricolpates

Antennaria Diploid species of Antennaria—A. neglecta and A. plantaginifolia—routinely produce staminate individuals within populations, whereas polyploid species—A. howellii and A. parlinii—do not, or do so with great rarity. Therefore, noting the sex of plants observed in populations on herbarium labels can be useful for later reviewers. Also, whole plant collections, which include the stolons and their rosettes (if formed) are much more valuable than single-stem collections. Reference: Bayer (2006). 1a. Rosette leaves smaller, 2–15 (–21) mm wide, 1-veined or sometimes with 2 additional, evanescent, lateral veins, the lateral veins (when present) located ⅔ to ¾ the distance from the midrib to the margin (i.e., near the margin) 2a. Middle and upper stem leaves blunt- to aristate-tipped [Fig. 373], only the leaves of the capitulescence with a scarious appendage . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) A. howellii 2b. Middle and upper stem leaves tipped by a flat or involute-margined, scarious appendage [Fig. 374] 3a. New rosette leaves bright green and promptly glabrous on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) A. howellii 3b. New rosette leaves white or gray-green and tomentose on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. neglecta 1b. Rosette leaves larger, (7–) 15–55 mm wide, with 3 or 5 prominent veins, the pair of lateral veins closest to the midrib located ⅓ to ½ the distance from the midrib to the margin (i.e., near the center of each half of the leaf blade) [Fig. 375] 4a. Carpellate involucre 5–7 mm tall; central corollas 2.5–4.3 mm long; body of cypselas 1–1.5 mm long; mature pappus 4–5.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . A. plantaginifolia 4b. Carpellate involucre (6–) 7–10 mm tall; central corollas 4.5–7 mm long; body of cypselas 1.3–2.2 mm long; mature pappus 6–9 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. parlinii

Fig. 373  Capitulescence and upper leaf of Antennaria howellii ssp. dioica.

1. Antennaria howellii Greene N Fig. 373 small pussytoes.  1a. Antennaria canadensis Greene; A. neglecta Greene var. canadensis (Greene) Cronq.; A. neodioica Greene ssp. canadensis (Greene) Bayer & Stebbins;  1b. Antennaria neglecta Greene var. neodioica (Greene) Cronq.; A. neodioica Greene; A. neodioica Greene var. attenuata Fern.; A. neodioica Greene var. chlorophylla Fern.; A. neodioica Greene var. grandis Fern.; A. neodioica Greene var. rupicola (Fern.) Fern.; A. rupicola Fern.; 1c. Antennaria neglecta Greene var. petaloidea (Fern.) Cronq.; A. neglecta Greene var. subcorymbosa Fern.; A. neodioica Greene ssp. petaloidea (Fern.) Bayer & Stebbins; A. neodioica Greene var. petaloidea Fern.; A. petaloidea (Fern.) Fern.; A. petaloidea (Fern.) Fern. var. scariosa Fern.; A. petaloidea (Fern.) Fern. var. subcorymbosa (Fern.) Fern. • CT, MA, ME, NH, RI, VT. Dry fields, roadsides, woodlands, rock balds, ledges. 1a. Middle and upper stem leaves tipped by a flat or involute-margined, scarious appendage [Fig. 374]; new rosette leaves bright green and promptly glabrous on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. A. howellii ssp. canadensis (Greene) Bayer 1b. Middle and upper stem leaves blunt- to aristate-tipped [Fig. 373], only the leaves of the capitulescence with a scarious appendage; new rosette leaves white or gray-green and tomentose on the adaxial surface or sometimes bright green and promptly glabrous in forms of A. howellii var. neodioica 2a. Stolons and basal offshoots short, leafy, terminated by rosettes; rosette leaves tending to have defined petioles . . . . . . . . . . . . . . . . . . 1b. A. howellii ssp. neodioica (Greene) Bayer 2b. Stolons elongate, cord-like, with few, small leaves, only tardily developing terminal rosettes; rosette leaves tending to have ill-defined petioles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1c. A. howellii ssp. petaloidea (Fern.) Bayer Subspecies canadensis is known from CT, MA, ME, NH, RI, VT. Subspecies neodioica is known from CT, MA, ME, NH, RI, VT. Subspecies petaloidea is known from CT, MA, ME, NH, RI, VT, but is relatively rare in southern New England.

Ast e r ac e a e  37 5

2. Antennaria neglecta Greene N Fig. 374 field pussytoes. Antennaria campestris Rydb.; A. neglecta Greene var. campestris (Rydb.) Steyermark • CT, MA, ME, NH, RI, VT. Dry fields, roadsides, woodlands, rock balds, ledges. 3. Antennaria parlinii Fern. N Fig. 375 Parlin’s pussytoes. 3a. Antennaria ambigens (Greene) Fern.; A. brainerdii Fern.; A. fallax Greene; A. fallax Greene var. calophylla (Greene) Fern.; A. munda Fern.; A. parlinii Fern. var. farwellii (Greene) Boivin; A. plantaginifolia (L.) Richardson var. ambigens (Greene) Cronq. 3b. Antennaria arnoglossa Greene; A. parlinii Fern. var. arnoglossa (Greene) Fern.; A. plantaginifolia (L.) Richardson var. arnoglossa (Greene) Cronq.; A. plantaginifolia (L.) Richardson var. parlinii (Fern.) Cronq. • CT, MA, ME, NH, RI, VT. Dry fields, roadsides, woodlands, rock balds. 1a. New rosette leaves gray-green and tomentose on the adaxial surface; reproductive stems usually lacking stipitate glands; reproductive stems and involucral bracts pale to green, rarely purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. A. parlinii ssp. fallax (Greene) Bayer & Stebbins 1b. New rosette leaves bright green and promptly glabrous or glabrate on the adaxial surface; reproductive stems often with purple glandular hairs near the apex; reproductive stems and involucral bracts often purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. A. parlinii ssp. parlinii

Fig. 374  Capitulescence and upper leaf of Antennaria neglecta.

Subspecies fallax is known from CT, MA, ME, NH, RI, VT. Subspecies parlinii is known from CT, MA, ME, NH, RI, VT. 4. Antennaria plantaginifolia (L.) Richardson N plantain-leaved pussytoes. Antennaria petiolata Fern.; A. plantaginifolia var. petiolata (Fern.) Heller; Gnaphalium plantaginifolium L. • CT, MA, ME, NH, RI, VT. Dry fields, roadsides, woodlands, rock balds.

Anthemis 1a. Ray flowers staminate or neutral; receptacle chaffy only near the middle; cypselas glandular-tuberculate; plants ill scented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. cotula 1b. Ray flowers carpellate; receptacle chaffy throughout; cypselas not tuberculate; plants not ill scented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. arvensis 1. Anthemis arvensis L. E

Fig. 375  Leaf blade of Antennaria parlinii.

corn chamomile. Anthemis arvensis L. var. agrestis (Wallr.) DC. • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil. 2. Anthemis cotula L. E Fig. 376 stinking chamomile. Maruta cotula (L.) DC. • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil.

Arctanthemum 1. Arctanthemum arcticum (L.) Tzvelev ssp. polare (Hultén) Tzvelev E arctic-daisy. Chrysanthemum arcticum L. var. polare (Hultén) Boivin; Dendranthema arcticum (L.) Tzvelev ssp. polare (Hultén) Heywood; D. hultenii (A. & D. Löve) Tzvelev • MA. Fields, roadsides, gardens.

Arctium Reference: Keil (2006a). 1a. Capitulescence resembling a raceme [Fig. 377]; capitula sessile or with a short peduncle up to 30 (–90) mm long [Fig. 377]; petioles weakly angled; leaf blades narrow- to broad-ovate, acute at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. minus

Fig. 376  Capitulum of Anthemis cotula.

376  tricolpates

1b. Capitulescence resembling a corymb; capitula with a peduncle mostly (15–) 25–120 mm long; petioles strongly angled; leaf blades broad-ovate to orbicular-ovate, rounded at the apex 2a. Involucre 15–25 mm wide, the involucral bracts arachnoid-tomentose; corollas minutely glandular-pubescent; petioles, particularly the lower, often hollow . . . . . . . A. tomentosum 2b. Involucre 25–45 mm wide, the involucral bracts glabrous to thinly arachnoidpubescent; corollas glabrous; petioles usually solid . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. lappa 1. Arctium lappa L. E great burdock. CT, MA, ME, NH, RI, VT. Fields, roadsides, vacant lots. 2. Arctium minus Bernh. E Fig. 377 common burdock. Lappa minor Hill • CT, MA, ME, NH, RI, VT. Fields, roadsides, vacant lots. Plants called Arctium nemorosum Lej. & Court. have been reported from New England (e.g., Kartesz 1999, Magee and Ahles 1999). Keil (2006a) noted that no North American plants display the characters for this species, specifically the broad, middle involucral bracts. Most plants referred to as A. nemorosum are merely large-headed forms of A. minus. 3. Arctium tomentosum P. Mill. E woolly burdock. CT, MA, ME, NH, VT. Fields, roadsides, vacant lots. Fig. 377  Capitulescence of Arctium minus.

Arctotis 1. Arctotis stoechadifolia Berg. E African-daisy. Arctotis grandis Thunb.; A. stoechadifolia Berg. var. grandis (Thunb.) Less. • MA. Waste areas, railroads.

Arnica 1. Arnica lanceolata Nutt. ssp. lanceolata

NC

lance-leaved arnica. Arnica mollis Hook. var. petiolaris Fern. • ME, NH, VT; northern portion of states. Alpine ravines, gullies, and brooks, sometimes found on middle elevation stream banks whose waters drain high alpine areas, also known from northern, ice-scoured river shores.

Arnoseris 1. Arnoseris minima (L.) Schweig. & Koerte E lamb-succory. Hyoseris minima L. • ME, NH. Roadsides, fields, disturbed soil.

Artemisia Reference: Shultz (2006). 1a. Plants definitely woody; capitula homogamous—all the flowers bisexual and with 5-lobed corollas; leaves persistent, the principal ones with blades narrow-obtriangular with three apical teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. tridentata 1b. Plants herbaceous, suffrutescent, or ± woody; capitula heterogamous—the marginal flowers carpellate with 2-lobed corollas, the inner flowers bisexual or functionally staminate with 5-lobed corollas; leaves usually deciduous, the principal ones with pinnatifid blades (A. dracunculus often and A. ludoviciana usually with simple leaves) 2a. Central flowers of disk functionally staminate, the gynoecium of these flowers with an abortive ovary and undivided style that is included within the short-tubular to subglobose corolla

Ast e r ac e a e   37 7

3a. Leaf blades 2- or 3-times pinnatifid into narrow segments [Fig. 379]; herbage scarcely odorous, if at all . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. campestris 3b. Leaf blades entire or sometimes the lower with 2–5 lobes; herbage faintly to strongly tarragon-scented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. dracunculus 2b. Central flowers of disk bisexual, the gynoecium of these flowers with a fertile ovary and divided style that is often exserted beyond the tubular to funnelform corolla 4a. Receptacle densely and conspicuously hairy between the disk flowers 5a. Reproductive stems 4–6 (–10) dm tall; leaf blades 3–8 cm long, the ultimate segments oblong to oblanceolate, 1.5–4 mm wide . . . . . . . . . . . . . . . . . . A. absinthium 5b. Reproductive stems 1–4 dm tall; leaf blades 0.5–1.5 (–2.5) cm long, the ultimate segments linear, up to 1 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. frigida 4b. Receptacle without hairs 6a. Plants annual or biennial, arising from a taproot; leaf blades glabrous on both surfaces; involucral bracts glabrous 7a. Capitulescence dense, resembling a spike, 2–4 cm wide, with inconspicuous peduncles and erect or ascending capitula; involucre 2–3 mm tall; plants inodorous or nearly so; leaf blades 1- or 2-times pinnatifid . . . . . . . . . . . . . . . . . . . . . . A. biennis 7b. Capitulescence open, resembling a panicle, 10–20 cm wide, with conspicuous peduncles and nodding capitula [Fig. 378]; involucre 1–2 mm tall; plants sweetscented; leaf blades 2- or 3-times pinnatifid . . . . . . . . . . . . . . . . . . . . . . . . . . . A. annua 6b. Plants perennial, arising from a rhizome or woody caudex; leaf blades tomentose on one or both surfaces, sometimes sparsely so; involucral bracts tomentose, at least sparsely so 8a. Plants suffrutescent or ± woody, without rhizomes; leaves 2- or 3-times pinnatifid into linear or filiform segments 0.5–1.5 mm wide; involucre 2–3.5 (–4) mm tall 9a. Leaf blades 3–6 cm long, thinly tomentose on the abaxial surface, glabrous on the adaxial surface; plants ± shrubby, 5–20 dm tall . . . . . . . . . . A. abrotanum 9b. Leaf blades 1–3 cm long, tomentose on both surfaces; plants suffrutescent, 4–10 dm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pontica 8b. Plants herbaceous, dying back to the ground each year, with rhizomes; leaves entire or 1- or 2-times pinnatifid into lobes wider than 1.5 mm (the ultimate lobes often narrower than 1.5 mm in A. carruthii); involucre 2.5–7.5 mm tall 10a. Leaf blades entire or toothed, the lower sometimes inconspicuously lobed, but the primary lobes without additional teeth or lobes . . . . . . . A. ludoviciana 10b. Principal leaf blades 1- or 2-times pinnatifid, the primary lobes either entire or with additional teeth or lobes 11a. Leaf blades concolorous, densely white-tomentose on both faces; involucre 6–7.5 mm tall; disk corollas 3.2–4 mm long . . . . . . . . . A. stelleriana 11b. Leaf blades bicolorous, white-tomentose on the abaxial surface, green and glabrous or sparsely pubescent on the adaxial surface; involucre 2.5–4.5 mm tall; disk corollas 1–2.8 mm long 12a. Reproductive stems (4–) 6–19 dm tall; leaf blades with ultimate segments wider than 2 mm; capitula with 7–10 marginal carpellate flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. vulgaris 12b. Reproductive stems 1.5–4 (–7) dm tall; leaf blades with ultimate segments narrower than 2 mm; capitula with 1–5 marginal carpellate flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. carruthii

378  tricolpate s

1. Artemisia abrotanum L. E southern wormwood. Artemisia procera Willd. • CT, MA, ME, VT; also reported from NH by Kartesz (1999), but specimens are unknown. Waste areas. 2. Artemisia absinthium L. E oldman wormwood. CT, MA, ME, NH, VT; also reported from RI by Gould et al. (1998), but specimens are unknown. Fields, roadsides, disturbed soil. 3. Artemisia annua L. E Fig. 378 annual wormwood. CT, MA, ME, NH, VT. Roadsides, disturbed soil, gardens. 4. Artemisia biennis Willd. E biennial wormwood. CT, MA, ME, NH, RI, VT. Roadsides, railroads, fields, disturbed soil. Fig. 378  Capitulescence and upper leaves of Artemisia annua.

5. Artemisia campestris L. n C Fig. 379 field wormwood.  5a. Artemisia campestris L. var. caudata (Michx.) Palmer & Steyermark; A. caudata Michx.; A. caudata Michx. var. calvens Lunell; Oligosporus campestris (L.) Cass. ssp. caudatus (Michx.) W.A. Weber; O. caudatus Poljakov; 5b. Artemisia campestris L. var. canadensis (Michx.) Welsh; A. canadensis Michx.; 5c. Oligosporus campestris (L.) Cass. • CT, MA, ME, nh, ri, VT. Beaches, dunes, sandy areas on the coastal plain, cliffs, talus, river shore ledges, ridges. 1a. Plants biennial from a taproot, usually with a solitary stem; disk corollas 1.4–2 (–2.2) mm long; native plants of coastal plain sand and gravel (rarely of inland cliffs and outcrops) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5a. A. campestris ssp. caudata (Michx.) Hall & Clements 1b. Plants perennial from a branching caudex, usually with multiple reproductive stems; disk corollas (1.8–) 2–3 mm long; native plants of rocky substrate or introduced plants of the Atlantic coastal plain 2a. Capitula with (21–) 23–45 flowers; involucre 3.5–5 mm wide; native plants of northern New England cliffs, talus, and river shore ledges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5b. A. campestris ssp. canadensis (Michx.) Scoggan

Fig. 379  Capitulescence and upper leaves of Artemisia campestris ssp. caudata.

2b. Capitula with 10–22 (–28) flowers; involucre 1.8–2.9 mm wide; rare introduction in southern New England . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5c. A. campestris ssp. campestris Subspecies caudata is native and known from CT, MA, ME, NH, RI, VT. Subspecies canadensis is native, of conservation concern, and known from ME, VT. Subspecies campestris is nonnative and known from CT, MA. Reports of ssp. campestris in New England are based on Hall and Clements (1923). Some of the collections were referred to this subspecies through examination of above-ground collections. It may be that these collections are merely forms of ssp. caudata with open-branched capitulescences. 6. Artemisia carruthii Wood ex Carruth E Carruth’s wormwood. Artemisia carruthii Wood ex Carruth var. wrightii (Gray) Blake; A. kansana Britt.; A. vulgaris L. ssp. wrightii (Gray) Hall & Clements • RI. Roadsides, fields, waste areas. 7. Artemisia dracunculus L. ssp. glauca (Pallas ex Willd.) Hall & Clemants E dragon wormwood. Artemisia dracunculoides Pursh; A. dracunculus L. ssp. glauca (Pallas ex Willd.) Hall & Clements; A. dracunculus L. var. glauca (Pallas ex Willd.) Bess.; A. glauca Pallas ex Willd.; A. glauca Pallas ex Willd. var. dracunculina (S. Wats.) Fern.; Oligosporus dracunculus (L.) Poljakov; O. dracunculus (L.) Poljakov ssp. glaucus (Pallas ex Willd.) A. & D. Löve • CT, MA. Roadsides, fields, waste areas. 8. Artemisia frigida Willd. E prairie wormwood. CT, MA. Roadsides, fields, waste areas. Reports of this species in VT (e.g. Seymour 1982) were based on a collection of Artemisia abrotanum—17 Sep 1898, Kennedy s.n. (NEBC). 9. Artemisia ludoviciana Nutt. ssp. ludoviciana E white wormwood. Artemisia gnaphalodes Nutt.; A. ludoviciana Nutt. var. americana (Bess.) Fern.; A. ludoviciana Nutt. var. gnaphalodes (Nutt.) Torr. & Gray; A. ludoviciana Nutt. var. latifolia (Bess.) Torr. & Gray; A. vulgaris L. ssp. ludoviciana (Nutt.) Hall & Clements; A. vulgaris L. var. ludoviciana (Nutt.) Kuntze • CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas, abandoned lots.

Ast e r ac e a e   379

10. Artemisia pontica L. E Roman wormwood. CT, MA, ME, NH, RI, VT. Roadsides, fields, railroads. 11. Artemisia stelleriana Bess. E beach wormwood. CT, MA, ME, NH, RI, VT. Dunes and other sandy areas near the coast, areas of habitation. 12. Artemisia tridentata Nutt. ssp. tridentata E big sagebrush. Artemisia angusta Rydb.; A. tridentata Nutt. var. angustifolia Gray; Seriphidium tridentatum (Nutt.) W.A. Weber • MA. Fields, waste areas. 13. Artemisia vulgaris L. var. vulgaris E common wormwood. Artemisia vulgaris L. var. glabra Ledeb.; A. vulgaris L. var. latiloba Ledeb. • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil, vacant lots, railroads.

Aster 1. Aster tataricus L. f. E Tatarian aster. CT, MA, RI. Roadsides, waste ares.

Baccharis 1. Baccharis halimifolia L. N eastern false willow. Baccharis halimifolia L. var. angustior DC. • CT, MA, RI. Saline and brackish marshes, coastal beaches, disturbed sites near the coast.

Balsamita 1. Balsamita major Desf. E costmary. Balsamita major Desf. var. tanacetoides (Boiss.) Moldenke; Chrysanthemum balsamita (L.) Baill.; C. balsamita (L.) Baill. var. tanacetoides Boiss.; Pyrethrum majus (Desf.) Tzvelev; Tanacetum balsamita L. • CT, MA, ME, NH, RI, VT. Roadsides, fields, vacant lots, abandoned homesteads.

Bellis 1. Bellis perennis L. E Lawn-daisy. CT, MA, ME, NH, RI, VT. Lawns.

Bidens Several species of Bidens have been divided into varieties on the basis of differences in awn morphology (e.g., number, length, presence and orientation of barbs). Studies have revealed that variation in awn morphology can be observed within populations or even on a given plant (Weedon 1974, Roberts 1982). Therefore, taxa based solely on these differences are not recognized here. Recent phylogenetic work showed that Bidens is not monophyletic and that it is nested within Coreopsis (Kimball and Crawford 2004). This indicates the need for taxonomic realignment and potential merging (in some fashion) of these two genera. References: Weedon (1974), Haines (2003b). 1a. Plants aquatic, with flaccid stems; leaves whorled, finely dissected into narrow, flaccid segments (except the emersed ones); capitulescence commonly a solitary capitulum [Fig. 380]; cypselas nearly terete to weakly 4-angled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. beckii

38 0   tricolpate s

1b. Plants terrestrial or of wetlands, with erect stems; leaves opposite, entire or divided into broad, firmer segments; capitulescence commonly composed of 2 or more capitula; cypselas compressed to quadrangular 2a. Principal leaf blades 1- to 3-times divided into distinct leaflets, at least the terminal leaflet with a thin petiolule 3a. Ultimate segments of leaf blades linear to narrow-lanceolate, (0.5–) 2–8 (–12) mm wide; marginal flowers of capitulum with well-developed rays 10–30 mm long (except B. tenuisecta, with rays absent or short) 4a. Marginal flowers of capitulum lacking rays or with short rays up to 6 mm long; inner cypselas quadrangular in cross-section, 8–15 mm long . . . . . . . . . B. tenuisecta 4b. Marginal flowers of capitulum with well-developed rays 10–30 mm long; inner cypselas ± flat, (4–) 5–9 mm long 5a. Cypsela body narrow-oblanceolate to cuneate-oblong in outline, 2.5–4 times as long as wide, 0.9–2.5 mm wide, merely antrorsely ciliate on the margin; peduncles glabrous; pappus of 2 setose, very stout awns or awn-scales (rarely absent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. trichosperma 5b. Cypsela body obovate to elliptic-obovate, 1.5–2 (–2.5) times as long as wide, 2.5–5.2 mm wide, with a thin, antrorsely ciliate and often erose-notched wing margin; peduncles pubescent; pappus of 2 (–4) antrorsely or restrorsely barbed awns (rarely absent) 6a. Outer, herbaceous series of involucral bracts numbering 8–12 (–16), (4–) 5–7 (–12) mm long, usually shorter than the inner, non-herbaceous series of bracts, eciliate or ciliate on the margin . . . . . . . . . . . . . . . . . . . . . . . . B. aristosa 6b. Outer, herbaceous series of involucral bracts numbering 12–21, (6–) 8–12 (–20) mm long, usually longer than the inner, non-herbaceous series of bracts, coarsely ciliate on the margin . . . . . . . . . . . . . . . . . . . . . . . . . B. polylepis 3b. Ultimate segments of leaf blades lanceolate or oblanceolate to ovate, (5–) 7–40 mm wide; marginal flowers of capitulum lacking rays or with short rays up to 5 mm long (except B. alba, with well-developed, white rays) 7a. Leaf blades 2- or 3-times pinnately divided; cypsela bodies at maturity strongly surpassing (i.e., some 2–3 times as long as) the inner, non-herbaceous series of involucral bracts of intact capitula [Fig. 381]; pappus of (2–) 3 or 4 awns [Fig. 381]; rays pale yellow (rarely white) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. bipinnata 7b. Leaf blades once pinnately divided; cypsela bodies at maturity not or only slightly exceeding (i.e., up to 1.5 times as long as) the inner, non-herbaceous series of involucral bracts of intact capitula; pappus of 2 (–5) awns; rays, when present, white (rarely short yellow rays present) 8a. Cypsela body quadrangular in cross-section, linear in outline, 5–9 times as long as wide; marginal flowers of the capitulum with or without well-developed rays longer than 5 mm 9a. Capitulum with 5–8 ray flowers with well-developed white rays 5–16 mm long; outer involucral bracts (8–) 12 (–16); pappus consisting of 2 awns. . . . . . . . . B. alba 9b. Capitulum lacking ray flowers or these present and with short rays up to 3 mm long; outer involucral bracts 7–10; pappus consisting of 2 or 3 (–5) awns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. pilosa 8b. Cypsela body definitely compressed, oblanceolate to obovate, 2–3.5 times as long as wide; marginal flowers of capitulum lacking rays or with short rays up to 5 mm long

Ast e r ac e a e   3 8 1

10a. Capitula with 3–5 outer, herbaceous involucral bracts; involucral bracts eciliate or only sparsely ciliate; cypsela body 3–6.5 mm long, with awns 0–2.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. discoidea 10b. Capitula with 5–21 outer, herbaceous involucral bracts; involucral bracts ciliate-margined; cypsela body 5.3–11.3 mm long, with awns 2–9.5 mm long 11a. Capitula with 5–10 outer, herbaceous involucral bracts; disk corollas abruptly ampliate above the herbaceous base, the limb delicate and the lobes prominently pigmented with yellow to yellow-orange, after anthesis collapsing around the exserted anthers and weakly attached to the summit of ovary [Fig. 382] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. frondosa 11b. Capitula with 10–21 outer, herbaceous involucral bracts; disk corollas gradually expanded apically, more herbaceous throughout and the lobes only weakly pigmented with yellow, after anthesis becoming chartaceous, scarcely collapsing, with included anthers, and firmly attached to the summit of ovary [Fig. 384] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. vulgata 2b. Leaf blades simple or lobed into 3–7 segments, the segments not divided completely to the midrib and, when lobed, with broadly winged petiolules 12a. Leaf blades sessile, or sometimes the lowermost with a short, narrowed base; cypselas either with a convex, cartilaginous apex or with a truncate apex in B. tripartita 13a. Disk corollas gradually expanded apically, more herbaceous throughout and the lobes only weakly pigmented with yellow, after anthesis becoming chartaceous, scarcely collapsing, with included anthers, and firmly attached to the summit of ovary [Fig. 384]; anthers pale brown; summit of the fruiting ovary with a truncate apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) B. tripartita 13b. Disk corollas abruptly ampliate above the stramineous or herbaceous base, the limb delicate and the lobes prominently pigmented with yellow to yellow-orange, after anthesis collapsing around the exserted anthers and weakly attached to the summit of ovary [Fig. 382]; anthers dark brown to black; summit of fruiting ovary with a convex, cartilaginous apex 14a. Outer, herbaceous involucral bracts ascending to erect; disk corollas usually 4-lobed; capitula erect; body of cypsela conspicuously marked with 7–15 longitudinal striations on each surface . . . . . . . . . . . . . . . . . . . . . B. hyperborea 14b. Outer, herbaceous involucral bracts spreading; disk corollas 5-lobed; capitula arched or nodding in age; body of cypsela not or only inconspicuously marked with longitudinal striae 15a. Rays absent or 2–15 (–18) mm long; chaff yellow near apex; body of inner cypselas with a pale, thickened, cartilaginous margin, those of the outer ones (3–) 5–6 mm long, those of the inner ones 4–8 mm long . . . . . . . . . . . . B. cernua 15b. Rays 15–30 mm long; chaff yellow-orange to orange-red near apex; body of cypselas lacking a pale, thickened margin, those of the outer ones 6–8 mm long, those of the inner ones 8–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . B. laevis 12b. Leaves with distinct petioles 1–4 cm long; cypselas with a truncate apex 16a. Disk corollas gradually expanded apically, more herbaceous throughout and the lobes only weakly pigmented with yellow, after anthesis becoming chartaceous, scarcely collapsing, with included anthers, and firmly attached to the summit of ovary; anthers pale brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) B. tripartita 16b. Disk corollas abruptly ampliate above the herbaceous base, the limb delicate and the lobes prominently pigmented with yellow to yellow-orange, after anthesis collapsing around the exserted anthers and weakly attached to the summit of ovary; anthers dark brown to black

38 2   tricolpate s

17a. Capitula cylindric to narrow-funnelform, each with (7–) 12–25 (–35) flowers; body of cypsela ± flat, usually striate on each face, lacking tuberculae [Fig. 383]; plants restricted to tidal river shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. eatonii 17b. Capitula funnelform to hemispheric, each with 20–60 (–150) flowers; body of cypsela compressed-quadrangular (varying to almost flat), commonly with few or no striations (immature fruits with more evident striations), usually with minute tubercles [Fig. 382]; plants of varied habitats, including tidal river shores, but primarily of inland wetlands and shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. connata Fig. 380  Capitulescence and emersed leaves of Bidens beckii.

1. Bidens alba (L.) DC. E white beggar-ticks. Bidens alba (L.) DC. var. radiata (Schultz-Bip.) Ballard ex T.E. Melchert; B. pilosa L. var. radiata Schultz-Bip. • CT, MA. Wool waste, disturbed soil. 2. Bidens aristosa (Michx.) Britt. E midwestern beggar-ticks. Bidens aristosa (Michx.) Britt. var. mutica (Gray) Gattinger; B. aristosa (Michx.) Britt. var. retrorsa (Sherff) Wunderlin; Coreopsis aristosa Michx. • CT, MA, ME, NH. Fields, pastures, waste areas. 3. Bidens beckii Torr. ex Spreng. N Fig. 380 Beck’s water-marigold. Megalodonta beckii (Torr. ex Spreng.) Greene; M. nudata Greene • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving water of lakes and rivers. Multiple phylogenetic studies have shown that Bidens beckii is best treated as part of Bidens (i.e., separation of this taxon into the genus Megalodonta is artificial; e.g., Kimball and Crawford 2004)

Fig. 381  Fruiting capitulum of Bidens bipinnata.

4. Bidens bipinnata L. E Fig. 381 Spanish needles beggar-ticks. Bidens bipinnata L. var. biternatoides Sherff • CT, MA; also reported from RI by George (1992), but specimens are unknown. Fields, roadsides, disturbed soil. 5. Bidens cernua L. N nodding beggar-ticks. Bidens cernua L. var. dentata (Nutt.) Boivin; B. cernua L. var. elliptica Wieg.; B. cernua L. var. integra Wieg.; B. cernua L. var. minima (Huds.) Pursh; B. cernua L. var. oligodonta Fern. & St. John • CT, MA, ME, NH, RI, VT. Shorelines, wetland edges, margins of pools and beaver flowages. 6. Bidens connata Muhl. ex Willd. N Fig. 382

Fig. 382  Cypsela and disk corolla of Bidens connata.

purple-stemmed beggar-ticks. Bidens connata Muhl. ex Willd. var. anomala Farw.; B. connata Muhl. ex Willd. var. fallax (Warnst.) Sherff; B. connata Muhl. ex Willd. var. gracilipes Fern.; B. connata Muhl. ex Willd. var. inundata Fern.; B. connata Muhl. ex Willd. var. petiolata (Nutt.) Farw.; B. connata Muhl. ex Willd. var. submutica Fassett; B. heterodoxa (Fern.) Fern. & St. John var. heterodoxa; B. heterodoxa (Fern.) Fern. & St. John var. orthodoxa Fern. & St. John • CT, MA, ME, NH, RI, VT. Shorelines, including fresh-tidal rivers, wetland edges, margins of pools and beaver flowages. ‌6 × 8. Bidens ×multiceps Fassett is a rare beggar-ticks hybrid known from fresh-tidal river shores in MA. Collections show variation in cypsela cross-section ranging from compressed-quadrangular to flat (displaying the variation between both parental taxa). 7. Bidens discoidea (Torr. & Gray) Britt. N small beggar-ticks. CT, MA, ME, NH, RI, VT; becoming rare in northeastern New England. Shorelines, wetland margins, swamps. 8. Bidens eatonii Fern.

Fig. 383  Cypsela of Bidens eatonii.

N C Fig. 383

Eaton’s beggar-ticks. Bidens eatonii Fern. var. fallax Fern.; B. eatonii Fern. var. interstes Fassett; B. eatonii Fern. var. kennebecensis Fern.; B. eatonii Fern. var. major Fassett; B. eatonii Fern. var. mutabilis Fassett; B eatonii Fern. var. simulans Fassett; B. heterodoxa (Fern.) Fern. & St. John var. atheistica Fern. • CT, MA, ME. Fresh to brackish-tidal river shores. The report of this species from RI by Magee and Ahles (1999) is based on a specimen of Bidens connata— 23 Sep 1929, Leland s.n. (NEBC!). Bidens heterodoxa has been a problematic taxon. Study revealed that it was comprised of two different, previously described taxa (as to New England

Ast e r ac e a e   3 83

plants) amalgamated together into one artificial taxon (Roberts 1982). Collections from the Kennebec River in ME are referable to B. eatonii. Collections from Pocotopaug Lake in CT are referable to B. tripartita. Bidens eatonii infrequently has a low, ± yellow dome at the summit of the mature ovary (not as strongly developed as in B. cernua and relatives). 9. Bidens frondosa L. N Devil’s beggar-ticks. Bidens frondosa L. var. anomala Porter ex Fern.; B. frondosa L. var. caudata Sherff; B. frondosa L. var. stenodonta Fern. & St. John • CT, MA, ME, NH, RI, VT. Shorelines, margins of wetlands, wet depressions and ditches. 10. Bidens hyperborea Greene

NC

northern beggar-ticks. Bidens hyperborea Greene var. cathancensis Fern.; B. hyperborea Greene var. colpophila (Fern. & St. John) Fern.; B. hyperborea Greene var. laurentiana Fassett; B. hyperborea Greene var. svensonii Fassett • MA, ME, NH. Fresh to somewhat brackish-tidal river shores. 11. Bidens laevis (L.) B.S.P. N smooth beggar-ticks. Bidens elegans Greene; Helianthus laevis L. • CT, MA, NH, RI; also reported from VT by Magee and Ahles (1999), but specimens are unknown. Lake shores, edges of marshes, river shores, including fresh-tidal ones. Reports of this species in ME (e.g., Kartesz 1999) are based on a misidentified specimen of Bidens cernua—23 Aug 1904, Bean s.n. (NEBC!). 12. Bidens pilosa L. E hairy beggar-ticks. Bidens odorata Cav.; B. pilosa L. var. minor (Blume) Sherff • CT. Ballast, waste areas. 13. Bidens polylepis Blake E Ozark beggar-ticks. Bidens involucrata (Nutt.) Britt.; Bidens polylepis Blake var. retrorsa Sherff; Coreopsis involucrata Nutt. • MA, RI. Hydric soil of marshes, wetland margins, and open rights-of-way. 14. Bidens tenuisecta Gray E slim-lobed beggar-ticks. MA. Fields, roadsides, disturbed soil. 15. Bidens trichosperma (Michx.) Britt. N crowned beggar-ticks. Bidens coronata (L.) Britt.; B. coronata (L.) Britt. var. brachyodonta Fern.; B. coronata (L.) Britt. var. tenuiloba (Gray) Sherff; B. coronata (L.) Britt. var. trichosperma (Michx.) Fern. • CT, MA, RI. Low meadows, shorelines, wet ditches, margins of swamps. Bidens coronata (L.) Britt., the name long used for this plant, is a homonym and, therefore, illegitimate. 16. Bidens tripartita L. ssp. comosa (Gray) A. Haines N Fig. 384 three-lobed beggar-ticks. Bidens comosa (Gray) Wieg.; Bidens heterodoxa (Fern.) Fern. & St. John var. agnostica Fern. • CT, MA, ME, NH, RI, VT. Shorelines, margins of wetlands, wet depressions and ditches. This species is often confused with Bidens connata in regional herbaria. In addition to characters used in the key, B. tripartita differs from B. connata in its smooth cypsela bodies that lack tubercles (vs. usually with tubercles). See B. eatonii for comments concerning B. heterodoxa. North American populations of B. tripartita are distinct from European ones. Those from North America have sparser and weaker cilia on the margins of the foliaceous involucral bracts (cilia 0.1–0.4 mm long, numbering 0–2 (–3) per mm, the bases not confluent to form a continuous cartiladgenous band vs. cilia 0.1–0.8 mm long (the longer usually exceeding 0.3 mm, numbering 1–5 per mm, the bases often confluent to form a continuous cartiladgenous band). 17. Bidens vulgata Greene N tall beggar-ticks. Bidens frondosa L. var. puberula Wieg.; B. puberula Wieg.; B. vulgata Greene var. puberula (Wieg.) Greene • CT, MA, ME, NH, RI, VT. Roadsides, ditches, shorelines.

Fig. 384  Cypsela and disk corolla of Bidens tripartita.

38 4   tricolpate s

Boltonia 1. Boltonia asteroides (L.) Ľ Hér. E white doll’s-daisy. 1a. Boltonia latisquama Gray; 1b. Boltonia latisquama Gray var. microcephala Fern. & Grisc.; B. latisquama Gray var. occidentalis Gray; B. latisquama Gray var. recognita Fern. & Grisc.; B. recognita (Fern. & Grisc.) G.N. Jones • CT, MA, ME, RI, VT. Roadsides, fields, waste areas. 1a. Outer and middle involucral bracts oblanceolate to rhombic or spatulate, obtuse to rounded at the apex, 1–2 mm wide; leaf blades near base of capitulescence mostly 1–2 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. B. asteroides var. latisquama (Gray) Cronq. 1b. Involucral bracts linear to narrow-oblong, acute at the apex, 0.5–1.3 mm wide; leaf blades near base of capitulescence mostly wider than 4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. B. asteroides var. recognita (Fern. & Grisc.) Cronq. Variety latisquama is known from CT, MA, ME, RI, VT. Variety recognita is known from CT, MA, ME, RI.

Calendula 1. Calendula officinalis L. E pot-marigold. CT, MA, ME, NH. Roadsides, gardens, waste areas.

Callistephus 1. Callistephus chinensis (L.) Nees E Chinese-aster. Aster chinensis L.; Callistemma chinense (L.) Skeels; Callistephus hortensis Cass.; Diplopappus chinensis (L.) Less. • CT, MA, ME. Waste areas, pond shores.

Calotis 1. Calotis cuneifolia R. Br. E Australian wedgeleaf. MA. Roadsides, waste areas.

Carduus Reference: Keil (2006b). 1a. Involucres 20–60 mm wide; involucral bract appendages 2–7 mm wide; capitula often nodding; peduncles 2–30 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nutans 1b. Involucres 10–25 mm wide; involucral bract appendages 0.5–1 mm wide; capitula erect [Fig. 385]; peduncles up to 4 (–10) cm long 2a. Abaxial leaf blade surface with dimorphic hairs—longer, curling septate hairs on the veins and sparse to dense tomentum across the surface; involucres 12–17 mm tall; disk corollas 11–16 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. crispus 2b. Abaxial leaf blade surface with only curling, septate hairs; involucres 14–20 mm tall; disk corollas 13–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. acanthoides 1. Carduus acanthoides L. ssp. acanthoides E Fig. 385 spiny plumeless-thistle. CT, MA, ME, RI, VT. Roadsides, fields, pastures. 2. Carduus crispus L. E Fig. 385 Capitulescence and upper stem of Carduus acanthoides.

curly plumeless-thistle. RI, VT. Roadsides, fields, pastures. The distribution of Carduus crispus has long been erroneous due to reliance on non-diagnostic characters. Keil (2006b) presented C. crispus as the more widespread species in New England, but this is incorrect. Carduus acanthoides is more frequent and is responsible for most reports of C. crispus in New England.

Ast e r ac e a e  3 8 5

3. Carduus nutans L. E nodding plumeless-thistle. Carduus nutans L. ssp. leiophyllus (Petrovic) Stojanov & Stef.; C. nutans L. var. leiophyllus (Petrovic) Arènes • CT, MA, NH, RI. Roadsides, fields, pastures.

Carthamus 1a. Leaf blades usually entire or spinose-toothed; stems ± glabrous; pappus absent, or rarely present on cypselas from near middle of capitulum and then consisting of short, narrow scales; corollas orange, orange-red, or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tinctorius 1b. Leaf blades usually pinnately lobed and spinose-toothed; stems white-tomentose; pappus present, of narrow, ciliate scales; corollas yellow (rarely white) . . . . . . . . . . . . . . . . . . . C. lanatus 1. Carthamus lanatus L. ssp. lanatus E woolly safflower-thistle. MA. Fields, roadsides, gardens. 2. Carthamus tinctorius L. E common safflower-thistle. MA. Fields, roadsides, gardens.

Centaurea Centaurea is a difficult genus that is plagued by misapplication of names (e.g., C. maculosa, C. pratensis). Report of Centaurea bovina Velen. in New England by Kartesz (1999) was probably based on a collection of C. diffusa. The report of Plectocephalus americanus (Nutt.) D. Don in Sweet by George (1992) was based on a specimen of C. nigra (Fernald 1922a). Reference: Keil and Ochsmann (2006). 1a. Involucral bracts tipped by 1 or more spines [Fig. 387]; disk corollas yellow, white to pink, or purple 2a. Capitula sessile, closely subtended and partially concealed by a series of non-spinetipped, foliaceous bracts; pappus composed of 2 series of awns—the outer series smooth or minutely barbellate and 9–10 mm long, the inner series minutely pubescent with spreading hairs and 2–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. benedicta 2b. Capitula sessile or borne on peduncles, not concealed by foliaceous bracts; pappus of capillary bristles or absent 3a. Apical spine of involucral bracts 2–3 (–5) mm long; involucre 4–5 mm wide; flowers lacking pappus; apical appendage of involucral bracts decurrent on body of bract, therefore the base of the appendage concave; disk corollas usually white to pink or pale purple; stems angled but not winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. diffusa 3b. Apical spine of involucral bracts 5–25 (–30) mm long; involucre 7–12 (–15) mm wide; at least the central flowers of a capitulum with pappus; apical appendage of involucral bracts not decurrent on body of bract, therefore the base of the appendage ± truncate; disk corollas yellow or varying shades of purple; stems with decurrent wings (except C. calcitrapa) 4a. Disk corollas pale purple, pink-purple, or purple; stem leaves not decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. calcitrapa 4b. Disk corollas yellow; stem leaves long-decurrent 5a. Apical spine of involucral bracts 10–15 (–30) mm long; marginal flowers of capitulum lacking pappus; disk corollas eglandular; pappus of inner flowers mostly 3–5 mm long, ca. twice as long as cypsela body . . . . . . . . . . . . . . . . . . C. solstitialis 5b. Apical spine of involucral bracts 5–8 mm long; marginal flowers of capitulum with pappus; disk corollas glandular; pappus of inner flowers mostly 1.5–3 mm long, ca. as long as cypsela body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. melitensis

38 6   tricolpate s

1b. Involucral bracts without spine-tips; disk corollas white to pink, red, or purple, or blue 6a. Apical appendage of involucral bracts decurrent on body of bract, therefore, the base of the appendage concave [Fig. 389]; principal stem leaf blades entire or conspicuously pinnatifid with linear to oblong lobes 7a. Plants annual; stem leaf blades linear . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cyanus 7b. Plants perennial; stem leaf blades oblong or lanceolate to ovate 8a. Appendage of involucral bracts decurrent nearly to the base of the bract; leaf blades entire (rarely the lower ones obscurely dentate); involucre 10–15 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. montana 8b. Appendage of involucral bracts decurrent only in the apical half of the bract [Fig. 389]; leaf blades pinnatifid; involucre narrower than 10 mm or wider than 15 mm 9a. Involocural bracts lacking prominent, longitudinal veins; pappus (3–) 4– 5 (–6) mm long; involucre 18–25 mm wide; cypsela bodies 4–5.5 mm long; corollas of fertile flowers 20–25 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. scabiosa 9b. Involucral bracts with several, prominent longitudinal veins [Fig. 389]; pappus 1–2.5 (–3) mm long (rarely absent); involucre 6–8 mm wide; cypsela bodies 2–3.5 mm long; corollas of fertile flowers 12–15 mm long . . . . . . . . . . . . . . C. stoebe 6b. Apical appendage of involucral bracts not decurrent on body of bract, therefore the base of the appendage ± truncate [Fig. 388]; principal stem leaf blades entire to toothed (infrequently with a few small lobes on larger leaf blades) 10a. Apical appendage of involucral bracts often recurved, long-tapering to a filiformfringed tip, the rachis of the appendage 0.2–0.4 mm wide on the middle bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. phrygia 10b. Apical appendage of involucral bracts ascending to appressed (rarely some with a loosely spreading apex), obtuse to acute at tip, the rachis of the appendage 0.5–3.5 mm wide on the middle bracts [Figs. 386, 388] 11a. Apical appendage of involucral bracts light brown to brown, those of the middle and outer series irregularly lacerate, those of the inner series often bifid; pappus absent; outer flowers of capitulum usually enlarged and falsely appearing as ray flowers (but the corollas still actinomorphic) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. jacea 11b. Apical appendage of involucral bracts brown to black, those of the middle and outer series regularly pectinate-fringed [Fig. 388], usually none of them bifid; pappus absent or present and then consisting of unequal bristles mostly 0.5–1 mm long; outer flowers of capitulum enlarged or not 12a. Apical appendage of involucral bracts 1–2 (–2.2) mm long, with 5–8 fringe segments on each margin, relatively narrow, the appendage not completely obscuring the distal portion of adjacent involucral bracts; involucre definitely taller than wide in life, ca. 1.5 times as tall as wide; outer flowers of capitulum sometimes enlarged and falsely appear as ray flowers . . . . . . . . . . . . C. nigrescens 12b. Apical appendage of involucral bracts longer than 2 mm, the larger ones (3–) 4–6 mm long, with 7–15 fringe segments on each margin [Fig. 388], relatively broad, obscuring the distal portion of the adjacent involucral bracts; involucre nearly as wide or wider than tall in life; outer flowers of capitulum not enlarged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nigra 1. Centaurea benedicta (L.) L. E blessed knapweed. Cnicus benedictus L. • CT, MA. Roadsides, fields, waste areas. 2. Centaurea calcitrapa L. E purple knapweed. Centaurea adulterina Morreti • MA. Roadsides, fields, waste areas.

Ast e r ac e a e   3 87

3. Centaurea cyanus L. E garden knapweed. Centaurea segetum Hill; C. pulchra DC.; Leucacantha cyanus (L.) Nieuwl. & Lunell • CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas, rubbish heaps. Centaurea cyanus often shows involucral bracts that change colors from base to apex of the involucre. The visible portion of the basal ones are frequently white to pale, the middle ones green (in life) to brown (in drying), and the apical ones tinged with or wholly pink. 4. Centaurea diffusa Lam. E white knapweed. Acosta diffusa (Lam.) Soják; Acrolophus diffusus (Lam.) A. & D. Löve • CT, MA, NH, RI. Roadsides, fields, waste areas, abandoned homesteads. 4 ‌ × 13. Centaurea ×psammogena Gáyer is a rare knapweed hybrid in New England that is frequently confused with C. diffusa. The hybrid, known from MA, is morphologically variable. Involucral bract appendages are usually brown to black (rarely light brown), and appendage spines are absent or short (spine mostly 1 mm long or shorter). However, an evident pappus always crowns the ovary, and the marginal flowers of the capitulum are falsely radiant (pappus absent and marginal flowers not enlarged in C. diffusa). 5. Centaurea jacea L. E brown knapweed. Centaurea debeauxii Gren. & Godr. ssp. thuillieri Dostál; C. jacea L. ssp. decipiens (Thuill.) Čelak.; C. jacea L. ssp. pratensis Čelak.; C. pratensis Thuill.; C. thuillieri (Dostál) J. Duvign. & Lambinon; Cyanus jacea (L.) P. Gaertn.; Jacea pratensis Lam. • CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas. ‌5 × 8. Centaurea ×moncktonii C.E. Britton [Fig. 389] has been known by the illegitimate name C. pratensis Thuill. It is known from CT, MA, ME, NH, RI, VT. It is a variable hybrid, with marginal corollas usually enlarged and falsely appearing as ray flowers (but sometimes not; usually enlarged in C. jacea, usually not enlarged in C. nigra), either lacking pappus or with vestigial bristles, and involucral bracts with brown apical appendages, those on the middle involucral bracts becoming irregularly pectinate (these irregularly lacerate in C. jacea and regularly pectinate in C. nigra). The involucral bract appendage often conceals the distal portion of adjacent involucral bracts but can be highly variable in later-generation hybrids (i.e., they can approach one or the other parent). The involucres are usually at least as wide as tall in life.

Fig. 386  Involucre of Centaurea ×moncktonii.

6. Centaurea melitensis L. E Fig. 387 Maltese knapweed. Calcitrapa melitensis (L.) Soják; Triplocentron melitense (L.) Cass. • MA. Roadsides, fields, waste areas. 7. Centaurea montana L. E

Fig. 387  Involucre and spinetipped bracts of Centaurea melitensis.

mountain knapweed. ME, NH, VT. Gardens, roadsides, fields. 8. Centaurea nigra L. E Fig. 388 black knapweed. Jacea nigra (L.) Hill • CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas. 9. Centaurea nigrescens Willd. E short-fringed knapweed. Centaurea dubia Suter; C. dubia Suter ssp. nigrescens (Willd.) Hayek; C. dubia Suter ssp. vochinensis (Bernh. ex Reichenb.) Hayek; C. transalpina Schleich. ex DC.; C. vochinensis Bernh. ex Reichenb.; Jacea nigrescens (Willd.) Soják • CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas. Plants identified as Centaurea transalpina Schleich. ex DC. in regional herbaria are referable to C. nigrescens. 10. Centaurea phrygia L. E wig knapweed. Centaurea austriaca Willd. • VT. Roadsides, fields, waste areas. 11. Centaurea scabiosa L. E greater knapweed. Acrocentron scabiosa (L.) A. & D. Löve; Colymbada scabiosa (L.) Holub • CT, ME, NH. Roadsides, fields, waste areas.

Fig. 388  Involucral bracts of Centaurea nigra showing truncate-based apical appendages.

38 8   tricolpate s

12. Centaurea solstitialis L. E yellow knapweed. Calcitrapa solstitialis (L.) Lam.; Cyanus solstitialis (L.) Baumg.; Leucantha solstitialis (L.) A. & D. Löve • CT, MA, NH, RI. Roadsides, fields, waste areas. 13. Centaurea stoebe L. ssp. micranthos (Gugler) Hayek E Fig. 389 spotted knapweed. Centaurea biebersteinii DC.; C. maculosa, auct. non Lam.; C. maculosa Lam. ssp. micranthos Gugler • CT, MA, ME, NH, RI, VT. Fields, railroads, open rights-of-way, sandy or gravelly banks. This taxon has long been known as Centaurea maculosa Lam.; however, this name properly applies to a strictly biennial knapweed with larger capitula (and is included in C. stoebe L. ssp. stoebe).

Fig. 389  Involucral bracts of Centaurea stoebe showing concave-based apical appendages.

Centipeda 1. Centipeda minima (L.) A. Braun & Aschers. E spreading-sneezeweed. Artemisia minima L. • MA. Gardens, waste areas.

Chaenactis 1. Chaenactis glabriuscula DC. var. glabriuscula E yellow pincushion. Chaenactis glabriuscula DC. var. tenuifolia (Nutt.) Hall; C. tenuifolia Nutt. • MA. Wool waste, roadsides, fields.

Chrysanthemum 1. Chrysanthemum morifolium Ramat. E florist’s daisy. Anthemis grandiflora Ramat.; Dendranthema grandiflorum (Ramat.) Kitam.; D. morifolium (Ramat.) Tzvelev • MA. Gardens, roadsides, fields. This plant is suspected to be of hybrid origin. Chrysanthemum indicum L. and C. japonicum Makino are often implicated as the progenitor taxa.

Chrysopsis 1. Chrysopsis mariana (L.) Ell.

NC

Maryland golden-aster. Chrysopsis mariana (L.) Ell. var. macradenia Fern.; Heterotheca mariana (L.) Shinners; Inula mariana L. • RI; Block Island. Sandy fields and grasslands.

Cichorium 1a. Vegetative leaf blades pubescent, at least on the abaxial midrib; bracteal leaves subtending the capitula usually shorter than capitula; ray flowers blue, white, or pink . . . . . . . . . . C. intybus 1b. Vegetative leaf blades glabrous; bracteal leaves subtending capitula usually exceeding capitula; ray flowers purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. endivia 1. Cichorium endivia L. E cultivated endive. CT, ME. Gardens, roadsides, fields. 2. Cichorium intybus L. E chicory. CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas.

Cirsium Given the polymorphic nature of Cirsium and its ability to form hybrids that can reproduce sexually (and backcross), not all collections will be referable to a species by the following key. Careful comparison with museum specimens and technical descriptions may be necessary. References: Moore and Frankton (1974), Keil (2006c).

Ast e r ac e a e   3 8 9

1a. Upper stem and branches with conspicuous, prickly and undulate-margined wings formed by decurrent leaf bases 2a. Involucre 25–40 mm tall; involucral bracts (or most of them) with prominent spine-tips 2–6 mm long; leaf blades adaxially with abundant, appressed spines . . . . . . . . . . C. vulgare 2b. Involucre 10–15 (–20) mm tall; involucral bracts lacking spine tips or with short, vestigial spines up to 1 mm long; leaf blades adaxially with sparse, septate hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. palustre 1b. Neither the stem nor the branches with decurrent wings 3a. Involucre 10–20 mm tall; capitula with, or with mostly, unisexual flowers by abortion; plants perennial, colonial from deep-seated, creeping roots, the reproductive stems not arising from a rosette of leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. arvens 3b. Involucre 20–50 mm tall; capitula with bisexual flowers; plants biennial or short-lived perennial, not colonial, the reproductive stems arising from a rosette of leaves 4a. Capitula closely invested by series of narrow, spiny-margined leaves that form a false involucre sometimes overtopping the true involucre [Fig. 390]; corollas yellow (rarely purple); scarious apex of innermost (i.e., longer) involucral bracts slender and not expanded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. horridulum 4b. Capitula either without subtending leaves or with 1 or 2 approximate, reduced leaves; corollas purple to pink (rarely white); scarious apex of innermost involucral bracts expanded or not 5a. Leaf blades persistently white-tomentose on the abaxial surface 6a. Adaxial surface of leaf blades thinly tomentose (rarely glabrate); stem thinly, uniformly, and persistently tomentose; pappus bristles 20–30 mm long; plants from taproots and also with horizontal roots that produce new shoots . . . . . C. flodmanii 6b. Adaxial surface of leaf blades glabrate or sparsely villous with septate hairs; stem glabrate, infrequently with thin tomentum apically; pappus bristles 12–25 mm long; plants from taproots and sometimes also a cluster of coarse, fibrous roots, but without prominent horizontal roots 7a. Stem leaf blades with flat margins, unlobed or with short lobes separated by shallow sinuses (rarely with deep sinuses and then with broad-triangular lobes); tips of innermost involucral bracts usually dilated and erose to serrulatae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. altissimum 7b. Stem leaf blades with ± revolute margins, evidently pinnately lobed with narrow-lanceolate lobes; tips of innermost involucral bracts slender, not dilated, entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. discolor 5b. Leaf blades abaxially thinly tomentose or arachnoid-pubescent when young, becoming glabrate in age 8a. Outer involucral bracts tipped with a coarse spine 3–6 mm long; cypsela body (3–) 3.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pumilum 8b. Outer involucral bracts lacking a spine tip or tipped with a vestigial spinule up to 0.5 (–1) mm long; cypsela body 4.5–5.5 mm long . . . . . . . . . . . . . . . . . . C. muticum 1. Cirsium altissimum (L.) Spreng. E tall thistle. Carduus altissimus L. • CT, MA, VT. Fields, roadsides, disturbed soil. 2. Cirsium arvense (L.) Scop. E creeping thistle. Breea arvensis Less.; Carduus arvensis (L.) Robson; Cirsium arvense (L.) Scop. var. horridum Wimmer & Grab.; C. arvense (L.) Scop. var. integrifolium Wimmer & Grab.; C. arvense (L.) Scop. var. mite Wimmer & Grab.; Serratula arvensis L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings.

39 0   tricolpate s

3. Cirsium discolor (Muhl. ex Willd.) Spreng. N field thistle. Carduus discolor (Muhl. ex Willd.) Nutt. • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil, often on mesic to wet-mesic soil. 4. Cirsium flodmanii (Rydb.) Arthur E Flodman’s thistle. Carduus flodmanii Rydb. • VT. Fields, pastures. 5. Cirsium horridulum Michx. var. horridulum

N C Fig. 390

yellow thistle. Carduus spinosissimus Walt.; Cirsium spinosissimum, auct. non (L.) Scop. • CT, MA, ME, NH, RI. Fields, clearings, and meadows near the coast, including upper edges of saltmarshes. 6. Cirsium muticum Michx. N Fig. 390  Capitulescence and subtending upper leaves of Cirsium horridulum.

swamp thistle. Carduus muticus (Michx.) Pers.; Cirsium bigelovii DC.; C. muticum Michx. var. monticola Fern.; C. muticum Michx. var. subpinnatifidum (Britt.) Fern. • CT, MA, ME, NH, RI, VT. Wet depressions, margins of wetlands, shorelines, upper edges of saltmarshes. 7. Cirsium palustre (L.) Scop. E marsh thistle. Carduus palustris L. • MA, NH. Marshes, swamp edges, low clearings. 8. Cirsium pumilum (Nutt.) Spreng. var. pumilum N pasture thistle. Carduus pumilus Nutt.; Cirsium odoratum (Muhl. ex W. Bart.) Petrak • CT, MA, ME, NH, RI, VT. Fields, roadsides, pastures, clearings. 9. Cirsium vulgare (Savi) Ten. E Fig. 391 common thistle. Carduus lanceolatus L.; C. vulgaris Savi; Cirsium lanceolatum (L.) Scop. • CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings, edges of lawns.

Coreopsis Reference: Strother (2006b).

Fig. 391  Cypsela and plumose pappus bristles of Cirsium vulgare.

1a. Disk corollas mostly 4-lobed at apex; outer series of involucral bracts much shorter than the inner series [Fig. 394]; style branch apex blunt or short-conical 2a. Rays pink to white; disk corollas yellow; cypsela body lacking wing-like edges; principal leaf blades usually simple (rarely with 2 or 3 lobes) [Fig. 394]; plants perennial, from rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rosea 2b. Rays yellow distally with a red-brown base; disk corollas dark red; cypsela body narrowly to broadly winged (infrequently wingless); principal leaf blades 1- to 2-times pinnatifid; plants annual or biennial, from taproots . . . . . . . . . . . . . . . . . . . . . . . . . . C. tinctoria 1b. Disk corollas mostly 5-lobed at apex; outer series of involucral bracts nearly or fully as long as inner series (shorter in C. tripteris); style branch apex acute, long-conical, or cuspidate 3a. Leaf blades simple or pinnately lobed to compound with 3–9 (or more) segments [Fig. 392]; chaff long-attenuate at apex; rays apically with prominent tooth-like lobes 4a. Disk corollas dark red; principal leaf blades pinnately lobed or pinnately compound with oblong to broad-ovate segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. basalis 4b. Disk corollas yellow; principal leaf blades simple or with a pair of basal lobes or pinnatifid with linear to narrow-lanceolate lobes 5a. Leaves confined to lower portion of stem; peduncles elongate, scape-like, mostly 20–40 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lanceolata 5b. Leaves borne throughout the stem; peduncles relatively shorter, mostly 5–20 cm long 6a. Principal leaf blades pinnatifid into linear to narrow-lanceolate segments; pappus often wanting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. grandiflora

Ast e r ac e a e  3 9 1

6b. Principal leaf blades simple, elliptic to obovate or ovate, sometimes with a pair of smaller lobes at the base; pappus often of 2, short, scale-like teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pubescens 3b. Principal leaf blades palmately or ternately lobed or divided (the central segment sometimes again palmately or ternately lobed or divided) [Fig. 393]; chaff rounded to acute at apex; rays apically entire or shortly toothed 7a. Leaf blades borne on evident, narrow petioles that are clearly much narrower than the associated leaf segments; outer involucral bracts 2–3 mm long, ca. ⅓ to ½ as long as the inner involucral bracts; reproductive stems 10–30 dm tall . . . . . . . . . . . . C. tripteris 7b. Leaf blades sessile or borne on petiole-like structures that are nearly or fully as wide as the associated leaf segments; outer involucral bracts (3–) 4–12 mm long, subequal in length to the inner involucral bracts; reproductive stems 3–10 dm tall 8a. Ultimate segments of leaf blades linear to filiform, 0.3–1.5 mm wide; cypsela body 3–5 × 1–1.7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. verticillata 8b. Ultimate segments of leaf blades linear to oval, 2–25 (–30) mm wide; cypsela body 4.5–6.5 × 1.8–4 mm 9a. Leaf blades trifurcating well above the base (i.e., the blades appearing to be on a broadly winged petiole) into linear to narrow-oblong segments 2–7 mm wide [Fig. 393]; internodes of stem usually glabrous . . . . . . . . . . . . . . . . . . . . . . . C. palmata

Fig. 392  Leaf of Coreopsis lanceolata.

9b. Leaf blades trifurcating at the base (i.e., the leaf blades definitely sessile) into lanceolate to oval segments (5–) 12–25 (–30) mm wide; internodes of stem usually short-pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. major 1. Coreopsis basalis (A. Dietr.) Blake E golden mane tickseed. Calliopsis basalis A. Dietr.; Coreopsis drummondii (D. Don) Torr. & Gray • CT. Fields, roadsides, waste areas. 2. Coreopsis grandiflora Hogg ex Sweet E large-flowered tickseed. Coreopsis grandiflora Hogg ex Sweet var. pilosa Sherff • CT, RI, VT; also reported from ME by Campbell et al. (1995), but specimens are unknown. Fields, roadsides, waste areas, dumps. Coreopsis grandiflora is sometimes confused in regional herbaria and collections thought to be this species are actually C. lanceolata. In addition to the characters used in the key, these two species can be separated by disk corolla length and node number to first peduncle. Coreopsis grandiflora has disk corollas 3.3–4.8 mm long and stems with 6–10 (or more) leaf-bearing nodes below the first peduncle. Coreopsis lanceolata has disk corollas 6–7.5 mm long and stems with 1–3 (–5) leaf-bearing nodes below the first peduncle.

Fig. 393  Leaf of Coreopsis palmata.

3. Coreopsis lanceolata L. E Fig. 392 lance-leaved tickseed. Coreopsis heterogyna Fern.; C. lanceolata L. var. villosa Michx. • CT, MA, ME, NH, RI, VT. Fields, roadsides, dry clearings. 4. Coreopsis major Walt. E greater tickseed. MA. Fields, roadsides, waste areas. 5. Coreopsis palmata Nutt. E Fig. 393 stiff tickseed. MA. Railroads, fields, roadsides. 6. Coreopsis pubescens Ell. E hairy tickseed. Coreopsis pubescens Ell. var. robusta Gray ex Eames • CT, MA. Fields, roadsides, clearings. 7. Coreopsis rosea Nutt. N Fig. 394 pink tickseed. MA, RI. Pond shores, swales, and boggy depressions on the coastal plain, sometimes in shallow water.

Fig. 394  Capitula of Coreopsis rosea.

39 2   tricolpate s

8. Coreopsis tinctoria Nutt. var. tinctoria E golden tickseed. Coreopsis cardaminifolia (DC.) Torr. & Gray; C. tinctoria Nutt. var. imminuta Sherff • CT, MA, ME, RI, VT. Fields, roadsides, waste areas, dumps. 9. Coreopsis tripteris L. E tall tickseed. Coreopsis tripteris L. var. smithii Sherff; C. tripteris L. var. subrhomboidea Sherff • CT, MA. Fields, roadsides, clearings. This species was reported from RI by Kartesz (1999),

based on George (1992); however, George (1999) stated this species had questionable naturalization in RI. 10. Coreopsis verticillata L. E thread-leaved tickseed. CT, MA. Fields, roadsides, clearings.

Cosmos 1a. Ultimate segments of leaf blades lanceolate to elliptic, 2–5 mm wide; rays pale to bright yellow; cypsela body 15–30 mm long including beak . . . . . . . . . . . . . . . . . . . . . . . . . C. sulphureus 1b. Ultimate segments of leaf blades linear to filiform, narrower than 1.5 mm; rays usually white to pink to red; cypsela body 7–16 mm long including beak 2a. Rays 15–40 mm long; margins of ultimate leaf segments entire . . . . . . . . . C. bipinnatus 2b. Rays 5–15 mm long; margins of ultimate leaf segments spinulose-ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. parviflorus 1. Cosmos bipinnatus Cav. E garden cosmos. CT, MA, ME, NH, RI. Roadsides, waste areas, gardens. 2. Cosmos parviflorus (Jacq.) Pers. E southwestern cosmos. MA, ME, RI. Roadsides, waste areas, wool waste. 3. Cosmos sulphureus Cav. E sulphur cosmos. CT, MA. Fields, roadsides, waste areas.

Cota 1. Cota tinctoria (L.) J. Gay E golden-chamomile. Anthemis tinctoria L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil.

Cotula Cotula australis (Sieber) Hook. f. was reported from ME by several recent sources, these originally based on Abrams and Ferris (1960), but specimens are unknown. 1. Cotula coronopifolia L. E common brassbuttons. MA. Saline, brackish, and freshwater marshes, stream banks.

Crepis Reference: Bogler (2006a). 1a. Cypselas of 2 kinds—those borne near the margin of the capitulum stout, 7–9 mm long, with a short beak and those from the interior of the capitulum slender, 12–17 mm long, and with a long, slender beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. foetida 1b. Cypselas ± uniform, 1.4–7.5 mm long

Ast e r ac e a e   3 9 3

2a. Inner series of involucral bracts glabrous on the adaxial (i.e., inner) surface; cypsela body 1.4–2.5 mm long (up to 4 mm long in the rarely collected C. nicaeensis) 3a. Abaxial surface of the involucral bracts with usually 2 rows of black, stipitate glands; involucre 5–8 (–9) mm tall; cypsela body 1.4–2.5 mm long . . . . . . . . . . . . . . . . . C. capillaris 3b. Abaxial surface of the involucral bracts glabrous or glabrate; involucre 8–10 mm tall; cypsela body 2.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nicaeensis 2b. Inner series of involucral bracts pubescent on the adaxial (i.e., inner) surface; cypsela body (2.5–) 3–8 (–9) mm long 4a. Capitula with 10–20 florets; involucral bracts coarsely hispid abaxially; cypsela tapered to a short beak 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. setosa 4b. Capitula with 30–100 florets; involucral bracts tomentose to hispidulous abaxially, sometimes also with stipitate glands in C. vesicaria; cypsela narrowed to summit of ovary and without a beak or with a prominent beak 2–5 mm long 5a. Cypsela with a slender beak 2–5 mm long; outer involucral bracts (i.e., the smaller ones) reflexed at maturity, sometimes with stipitate glands on the abaxial surface; abaxial surface of rays red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. vesicaria 5b. Cypsela narrowed to summit of ovary but without a beak; outer involucral bracts spreading-ascending to erect, usually without stipitate glands on the abaxial surface; rays entirely yellow 6a. Involucre (8–) 10–13 mm tall; cypsela body 4–7.5 mm long, yellow to red-brown, with (10–) 13–20 smooth, longitudinal ribs; pappus 5–7 mm long; stem leaf blades oblanceolate or oblong to spatulate, with plane margins . . . . . . . . . . . . . . C. biennis 6b. Involucre 6–9 mm tall; cypsela body (2.5–) 3–4 (–4.5) mm long, red to purplebrown, with 10 minutely scabrous, longitudinal ribs; pappus 4–5 mm long; stem leaf blades linear to lanceolate or oblanceolate, with revolute margins . . . . C. tectorum 1. Crepis biennis L. E rough hawk’s-beard. VT. Fields, roadsides, disturbed soil. 2. Crepis capillaris (L.) Wallr. E Fig. 395 smooth hawk’s-beard. CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil. 3. Crepis foetida L. E stinking hawk’s-beard. MA. Fields, roadsides, disturbed soil. 4. Crepis nicaeensis Balbis ex Pers. E Turkish hawk’s-beard. MA, VT. Fields, roadsides, disturbed soil. 5. Crepis setosa Haller f. E bristly hawk’s-beard. CT, VT. Fields, roadsides, disturbed soil. 6. Crepis tectorum L. E narrow-leaved hawk’s-beard. CT, MA, ME, RI, VT. Fields, roadsides, disturbed soil. 7. Crepis vesicaria L. ssp. haenseleri (Boiss. ex DC.) P.D. Sell E beaked hawk’s-beard. Crepis vesicaria L. ssp. taraxacifolia (Thuill.) Thellung; C. vesicaria L. var. taraxacifolia (Thuill.) Boivin • CT. Fields, roadsides, disturbed soil.

Crupina 1. Crupina vulgaris Cass. E common crupina. MA. Fields, roadsides, disturbed soil.

Fig. 395  Capitula and bract of Crepis capillaris.

39 4   tricolpate s

Cyclachaena 1. Cyclachaena xanthiifolia (Nutt.) Fresen. E carelessweed. Iva xanthiifolia Nutt. • CT, MA, ME, NH, RI, VT. Fields, railroads, barnyards, disturbed soil.

Deinandra 1. Deinandra fasciculata (DC.) Greene E clustered moonshine-daisy. Hemizonia fasciculata DC.; H. fasciculata ssp. ramosissima (Benth.) Keck & D.A. Johans.; H. ramosissima Benth. • MA. Wool waste.

Dittrichia 1. Dittrichia graveolens (L.) W. Greuter E stinkwort. Erigeron graveolens L.; Inula graveolens (L.) Desf. • CT. Open, disturbed, sandy or gravelly soil.

Doellingeria 1a. Body of cypsela short-hairy; involucre 3–5 mm tall; upright stems borne from elongate creeping rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. umbellata 1b. Body of cypsela glabrous; involucre 4.5–7 mm tall; upright stems mostly solitary from a fibrous root crown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. infirma 1. Doellingeria infirma (Michx.) Greene

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Appalachian white-aster. Aster infirmus Michx. • CT, MA, RI. Dry-mesic forests and woodlands, often with Quercus, Carya, and/or Pinus in the overstory, sparsely forested talus, clearings. 2. Doellingeria umbellata (P. Mill.) Nees var. umbellata N Fig. 396 tall white-aster. Aster umbellatus P. Mill. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest edges, meadows, margins of wetlands.

Doronicum Fig. 396  Involucre of Doellingeria umbellata with bracts that lack a distinct, green apical zone.

1. Doronicum pardalianches L. E greater false leopard’s bane. MA. Gardens, roadsides, fields.

Dyssodia 1. Dyssodia papposa (Vent.) A.S. Hitchc. E fetid-marigold. Boebera papposa (Vent.) Rydb. ex Britt.; Tagetes papposa Vent. • MA, VT; also reported from ME by Campbell et al. (1995), but specimens are unknown. Wool waste, fields, disturbed soil.

Echinacea 1a. Lower leaf blades broad-lanceolate to elliptic or broad-ovate, (1–) 5–12 cm wide, mostly 1.5–5 times as long as wide, usually toothed, abruptly tapered to the petiole; plants from fribrous roots and a weak taproot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. purpurea 1b. Lower leaf blades narrow-lanceolate to narrow-elliptic, 1–4 cm wide, mostly 5–20 times as long as wide, usually entire, gradually tapered to an often ill-defined petiole; plants from a strong taproot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. pallida

Ast e r ac e a e   3 9 5

1. Echinacea pallida (Nutt.) Nutt. E pale purple coneflower. Brauneria pallida (Nutt.) Britt.; Rudbeckia pallida Nutt. • CT, MA, ME. Fields, clearings. 2. Echinacea purpurea (L.) Moench E Fig. 397 eastern purple coneflower. Brauneria purpurea (L.) Britt. • MA, VT; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Fields, clearings, forest borders.

Echinops 1. Echinops sphaerocephalus L. E Fig. 398 greater globe-thistle. CT, MA, ME, NH, VT. Fields, roadsides, waste areas.

Fig. 397  Disk corollas and chaff of Echinacea purpurea.

Eclipta 1. Eclipta prostrata (L.) L. E false daisy. Eclipta alba (L.) Hassk.; Verbesina prostrata L. • CT, MA. Waste areas, disturbed places, trail edges. This species recently collected in the Connecticut River drainage of CT along the edge of a trail through a fresh-tidal wetland.

Erechtites 1. Erechtites hieraciifolius (L.) Raf. ex DC. N Fig. 399 American burnweed.  1a. Erechtites hieraciifolius (L.) Raf. ex DC. var. intermedius Fern.; E. hieraciifolius (L.) Raf. ex DC. var. praealtus (Raf.) Fern.; Senecio hieraciifolius L.; 1b. Erechtites megalocarpus Fern. • CT, MA, ME, NH, RI, VT. Dry-mesic to wet-mesic fields, clearings, and disturbed soil, also sandy sea beaches and edges of saline marshes.

Fig. 398  Capitula of Echinops sphaerocephalus.

1a. Cypsela body 2–3 mm long, with mostly 10–12 longitudinal ribs; inner (i.e., longer) series of involucral bracts attenuate at apex, 0.5–1.5 mm wide at the scarcely widened base; denuded receptacle 5–8.5 mm wide; leaf blades herbaceous; plants primarily of inland areas (casually found in coastal and near coastal habitats) . . . . . . . . . 1a. E. hieraciifolius var. hieraciifolius 1b. Cypsela body 4–5.5 mm long, with mostly 16–20 longitudinal ribs; inner involucral bracts obtuse to rounded at apex, 1–3 mm wide at the prominently widened base; denuded receptacle 9–12 mm wide; leaf blades somewhat fleshy; plants restricted to coastal habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. E. hieraciifolius var. megalocarpus (Fern.) Cronq. Variety hieraciifolius is known from CT, MA, ME, NH, RI, VT. Variety megalocarpus is known from CT, MA, RI.

Erigeron Phylogenetic work by Noyes (2000) has altered the circumscription of the genus Erigeron as treated by some recent authors. Both Conyza and Trimorpha have been shown to be nested within Erigeron. Therefore, these genera are here subsumed under Erigeron. Reference: Nesom (2006a). 1a. Involucre 5–12 mm tall; peduncles with glandular hairs; capitula with a series of rayless, carpellate flowers within the series of ray flowers; pappus elongating in fruit and conspicuously surpassing the involcure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. acris 1b. Involucre 2–7 mm tall; peduncles without glandular hairs; capitula with carpellate ray flowers and bisexual disk flowers; pappus not or scarcely elongating in fruit, not or only somewhat exceeding the involucre

Fig. 399  Involucre of Erechtites hieraciifolius var. hieraciifolius.

39 6   tricolpate s

2a. Rays 0.5–1 (–2) mm long [Fig. 400]; involucre 2–5 mm wide (sometimes wider in pressed specimens); disk 1–3 (–4) mm wide in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. canadensis 2b. Rays 4–10 mm long (very rarely wanting) [Fig. 401]; involucre 5–20 mm wide; disk (3–) 5–20 mm wide in life 3a. Leaves linear to linear-oblanceolate, 1–5 mm wide; stems tufted and bearing axillary, sterile, leafy branches [Fig. 401]; capitula with 20–50 ray flowers [Fig. 401]; capitula numbering 1–4 (–5) per stem, often solitary . . . . . . . . . . . . . . . . . . . . . . . . . . E. hyssopifolius 3b. Leaves linear to suborbicular, 2.5–70 mm wide; stems not tufted, usually without sterile, axillary branches; capitula with 50–400 ray flowers; capitula numbering 3–30 (–100) per stem (only 1–4 (–5) in E. pulchellus) 4a. Stem leaves gradually tapering to the base; pappus of 2 types—the inner of long, slender bristles, the outer of short scales; pappus of ray flowers shorter than 1 mm 5a. Principal stem leaves linear to lanceolate, entire or subentire, mostly 2.5– 10 (–15) mm wide; pubescence of mid-stem 0.1– 0.4 (–0.8) mm long, usually ascending to appressed (longer and more spreading in var. septentrionalis); rays 4–6 mm long; basal leaf blades often persistent through flowering . . . . E. strigosus 5b. Principal stem leaves broad-lanceolate to ovate, toothed, mostly 10– 35 (–70) mm wide; pubescence of mid-stem 0.5–1.2 mm long, spreading; rays 4–10 mm long; basal leaf blades often withering prior to flowering . . . . . E. annuus 4b. Stem leaves rounded to clasping at the base; pappus of 1 type—long, slender bristles; pappus of all the flowers much longer than 1 mm 6a. Capitula with 50–100 ray flowers, the rays 0.8–1.7 mm wide; disk corollas (4–) 4.5–6 mm long; body of cypsela 1.3–1.8 mm long; plants with slender, elongate, superficial rhizomes; capitula numbering 1–4 (–5) per stem . . . . . . . . . . E. pulchellus 6b. Capitula with 150–400 ray flowers, the rays up to 0.5 mm wide; disk corollas (2.1–) 2.5–3.2 mm long; body of cypsela 0.6–1.1 mm long; plants without superficial rhizomes; capitula numbering 3–30 (–40) per stem . . . . . . . . . . . E. philadelphicus 1. Erigeron acris L. ssp. kamtschaticus (DC.) Hara

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bitter fleabane. Erigeron acris L. var. kamtschaticus (DC.) Herder; E. angulosus Gaudin var. kamtschaticus (DC.) Hara; E. kamtschaticus DC.; Trimorpha acris (L.) Nesom var. kamtschatica (DC.) Nesom • ME; mainly in northern portion of state. Forest clearings, river banks. 2. Erigeron annuus (L.) Pers. N Fig. 400  Capitula of Erigeron canadensis.

annual fleabane. Erigeron annuus (L.) Pers. var. discoideus (Victorin & Rouss.) Cronq.; Stenactis annua (L.) Nees • CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings, disturbed soil. 3. Erigeron canadensis L. N Fig. 400 Canada fleabane. Conyza canadensis (L.) Cronq.; C. canadensis (L.) Cronq. var. pusilla (Nutt.) Cronq.; C. parva Cronq.; Erigeron canadensis L. var. pusillus (Nutt.) Boivin; E. pusillus Nutt.; Leptilon canadense (L.) Britt.; L. pusillum (Nutt.) Britt. • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, disturbed soil. 4. Erigeron hyssopifolius Michx. N Fig. 401 hyssop-leaved fleabane. ME, VT; also reported from NH by Kartesz (1999), but specimens are unknown. Cliffs, gorges, and river shore ledges in regions of high-pH bedrock. 5. Erigeron philadelphicus L.

Fig. 401  Capitulum and upper leaves of Erigeron hyssopifolius.

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Philadelphia fleabane. 5a. Erigeron philadelphicus L. var. glaber Henry; E. philadelphicus L. var. scaturicola (Fern.) Fern.; E. purpureus Ait.; 5b. E. philadelphicus L. ssp. provancheri (Victorin & Rouss.) J.K. Morton; E. provancheri Victorin & Rouss. • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, clearings; also basic, river shore ledges.

Ast e r ac e a e  3 97

1a. Reproductive stems 20–70 cm tall; basal leaves usually withering prior to anthesis; stems, leaf blades, and involucral bracts usually pubescent; ray flowers pink or white tinged with pink; capitula 18–25 mm wide . . . . . . . . . . . . . . . . . . . 5a. E. philadelphicus var. philadelphicus 1b. Reproductive stems 4–18 (–30) cm tall; basal leaves persisting through flowering; stems, leaf blades, and involucral bracts glabrous or sparsely pubescent; ray flowers white to pale pink; capitula 10–15 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . 5b. E. philadelphicus var. provancheri (Victorin & Rouss.) Boivin Variety philadelphicus is known from CT, MA, ME, NH, RI, VT. It occurs in open, frequently human-disturbed, habitats. Variety provancheri is known from VT. It occurs on high-pH, river shore ledges and is of conservation concern. 6. Erigeron pulchellus Michx. var. pulchellus N Robin’s plantain fleabane. CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, clearings. 7. Erigeron strigosus Muhl. ex Willd. N rough fleabane.  7a. Erigeron strigosus ssp. septentrionalis (Fern. & Wieg.) Wagenitz;  7b. Erigeron annuus (L.) Pers. ssp. strigosus (Muhl. ex Willd.) Wagenitz; E. ramosus (Walt.) B.S.P.; E. ramosus (Walt.) B.S.P. var. beyrichii (Fisch. & C.A. Mey.) Gray; E. strigosus Muhl. ex Willd. var. beyrichii (Fisch. & C.A. Mey.) Torr. & Gray; E. strigosus Muhl. ex Willd. var. discoideus Robbins ex Gray; E. strigosus Muhl. ex Willd. var. eligulatus Cronq.; Stenactis strigosa (Muhl. ex Willd.) DC. • CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings, and other open places. 1a. Involucral bracts pubescent with flattened hairs 0.5–1.2 mm long; mid-stem pubescent with appressed to spreading hairs 0.5–1 mm long; basal leaf blades usually dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7a. E. strigosus var. septentrionalis (Fern. & Wieg.) Fern. 1b. Involucral bracts pubescent with hairs that are not conspicuously flattened and are 0.1–0.5 mm long; mid-stem pubescent with appressed to ascending hairs 0.1–0.4 (–0.8) mm long; basal leaf blades usually entire to subentire . . . . . . . . . . . 7b. E. strigosus var. strigosus Variety septentrionalis is known from CT, MA, ME, NH, VT. It is transitional in some aspects (e.g., stem hair length and orientation, leaf blade margin) to Erigeron annuus. Variety strigosus is known from CT, MA, ME, NH, RI, VT.

Euchiton 1. Euchiton involucratus (G. Forst.) Holub E common creeping-cudweed. Gnaphalium involucratum Forst. • MA. Gardens, lawns, fields.

Eupatorium The traditionally defined genus Eupatorium has been dismantled into smaller, homogeneous, and monophyletic groups based on morphological and phylogenetic evidence (see also the genera Ageratina and Eutrochium). Most authors’ treatments of Eupatorium (including the one referenced here) suffer from poor taxonomy—they repeatedly treat hybrid-derived taxa as a variety of one of the putative parents, a system that creates obvious conflicts due to its nonmonophyletic and arbitrary nature (i.e., which parent should the hybrid be allied to, and on what basis?). Hybrid-derived taxa are here treated as species because they are reproductively competent and can be found outside of the range of parental co-occurence. Reference: Siripun and Schilling (2006). 1a. Capitula with 9–23 flowers; leaves with either evident petioles longer than 1 cm or the blades connate at the base 2a. Leaf blades narrowly to broadly tapering to a distinct petiole . . . . . . . . . . . E. serotinum 2b. Leaf blades broad-based, usually connate and surrounding the stem . . E. perfoliatum 1b. Capitula with (3–) 5 (–9) flowers; leaves sessile or subsessile, not connate at the base

39 8   tricolpate s

3a. Leaf blades linear to elliptic or oblanceolate, cuneate (i.e., tapering) at the base 4a. Involucre 8–11 mm tall; involucral bracts glabrous abaxially or sparsely pubescent near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. album 4b. Involucre 4–7 mm tall; involucral bracts moderately to densely pubescent over the abaxial surface 5a. Involucral bracts acute to acuminate and prominently white-scarious at the apex [Fig. 403] 6a. Principal leaf blades 8–15 (–20) mm wide, usually ± evenly serrate with narrow, sharp teeth; capitula with 5–7 (–9) disk flowers; body of cypsela 2–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. novae‑angliae 6b. Principal leaf blades (10–) 20–40 mm wide, coarsely and sometimes irregularly serrate to crenate-serrate with broad, pointed to blunt teeth [Fig. 404]; capitula with (4–) 5 (–6) disk flowers; body of cypsela 3–4 mm long . . . . . . . . . . (in part) E. pilosum 5b. Involucral bracts rounded to obtuse (sometimes acute) at the apex, with only a thin, white-scarious border near apex [Fig. 405] 7a. Leaf blades 8–30 mm wide, lanceolate to elliptic-oblong, mostly 2.5–7 times as long as wide, opposite; nodes without axillary fascicles or these poorly developed and with consisting of few leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. altissimum 7b. Leaf blades 0.5–10 (–17) mm wide, linear to lanceolate, mostly 6–40 times as long as wide, whorled in 3s or 4s at each node (sometimes opposite or alternate at the upper nodes); nodes often with well-developed axillary fascicles containing many small leaves 8a. Leaves usually 4 per node, sometimes only 3 (rarely fewer), with blades 0.5–5 (–10) mm wide, entire or irregularly few-toothed, 10–40 times as long as wide, the principal leaves with blades 2–7 cm long [Fig. 402]; reproductive stems 30–100 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. hyssopifolium 8b. Leaves usually 3 per node, sometimes 2 or 4 per node (rarely only 1), with blades 5–10 (–17) mm wide, serrate to nearly lacerate-toothed, 6–15 times as long as wide, the principal leaves with blades 6–11 cm long; reproductive stems mostly 75–150 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. torreyanum

Fig. 402 Capitulescence and leaves of Eupatorium hyssopifolium.

3b. Leaf blades lanceolate or oblong to suborbicular, broad-cuneate to cordate at the base [Figs. 404, 406] 9a. Leaf blades pinnately veined, lanceolate to oblong, acuminate to subacuminate at the apex; stem glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. sessilifolium 9b. Leaf blades triple-veined from near the base, oblong to suborbicular, rounded to acute at the apex; stem pubescent 10a. Leaf blades oblong or oblong-lanceolate to narrow-ovate, each margin with 3–12 coarse teeth, sometimes the blades more incised with lobe-like teeth near the base [Fig. 404]; upper leaves and lower branches of capitulescence often alternate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) E. pilosum 10b. Leaf blades oblong-ovate to suborbicular, each margin with 8–25 teeth [Fig. 406]; upper leaves and lower branches of capitulescence usually opposite 11a. Leaf blades triangular-ovate to suborbicular, mostly 1–1.5 (–1.7) times as long as wide, rounded to obtuse at the apex, usually with crenate margins; principal pair of lateral veins originating above base of leaf blade . . . . . . . E. rotundifolium

Fig. 403  Involucral bract of Eupatorium novae-angliae showing acuminate apex and prominent white-scarious border.

11b. Leaf blades oblong-ovate to ovate, mostly 1.5–2 times as long as wide, obtuse to acute at apex, usually with serrate margins [Fig. 406]; principal pair of lateral veins originating at the base of leaf blade . . . . . . . . . . . . . . . . . . . . . . . . E. pubescens

Ast e r ac e a e  3 9 9

1. Eupatorium album L. var. album

NC

white thoroughwort. Eupatorium album L. var. glandulosum (Michx.) DC.; E. glandulosum Michx.; E. petaloideum Britt. ex Small • CT. Dry-mesic woodlands, forest edges, and fields. 2. Eupatorium altissimum L. N tall thoroughwort. Eupatorium saltuense Fern. • CT, MA. Railroads, roadsides, clearings. 3. Eupatorium hyssopifolium L. N Fig. 402 hyssop-leaved thoroughwort. Eupatorium hyssopifolium L. var. calcaratum Fern. & Schub. • CT, MA, RI. Dry-mesic to wet-mesic, often sandy soils of fields, roadsides, pond shores, clearings, and borrow pits. 4. Eupatorium novae-angliae (Fern.) V. Sullivan ex A. Haines & Sorrie

N C Fig. 403

New England thoroughwort. Eupatorium leucolepis (DC.) Torr. & Gray var. novae-angliae Fern. • MA, RI. Coastal plain pond shores. This species is derived through hybridization of Eupatorium paludicola E.E. Schill. & LeBlond and E. perfoliatum.

Fig. 404  Leaf of Eupatorium pilosum.

5. Eupatorium perfoliatum L. N boneset thoroughwort. Eupatorium perfoliatum L. var. colpophilum Fern. & Grisc. • CT, MA, ME, NH, RI, VT; throughout. Shorelines, marshes, swamps, wetland margins, ditches. Plants from tidal shores show narrower leaves and shorter, sparser pubescence on the stems. These forms have been called var. colpophilum, but they intergrade with typical forms as tidal influence decreases. 6. Eupatorium pilosum Walt. N Fig. 404 ragged thoroughwort. Eupatorium rotundifolium L. var. saundersii (Porter ex Britt.) Cronq. • CT, MA, RI. Mesic to wet-mesic or seasonally hydric, sandy soils of pond shores, fields, and

depressions. 7. Eupatorium pubescens Muhl. ex Willd. N Fig. 405, 406 hairy thoroughwort. Eupatorium rotundifolium L. var. ovatum (Bigelow) Torr. • CT, MA, ME, NH, RI. Dry-mesic, often sandy, woodlands, fields, and clearings. This species is thought to have been derived from hybridization between Eupatorium rotundifolium and E. sessilifolium. 8. Eupatorium rotundifolium L.

NC

Fig. 405  Involucral bract of Eupatorium pubescens showing rounded apex and narrow whitescarious border.

round-leaved thoroughwort. CT, MA; also reported from RI by Kartesz (1999), but specimens are unknown. Pond shores, clearings, fields. 9. Eupatorium serotinum Michx. E late thoroughwort. CT, MA; also reported from RI by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, disturbed soil. 10. Eupatorium sessilifolium L. N upland thoroughwort. Eupatorium sessilifolium L. var. brittonianum Porter • CT, MA, ME, NH, RI, VT; southern portion of northern states. Rocky forests and woodlands, edges of rock balds. 11. Eupatorium torreyanum Short & Peter

NC

Torrey’s thoroughwort. Eupatorium hyssopifolium L. var. laciniatum (Torr. & Gray) Small • CT. Dry, open areas, such as fields, roadsides, and road cuts.

Eurybia Reference: Brouillet (2006a). 1a. Basal leaf blades both cordate and borne on petioles 2a. Leaf blades minutely stipitate-glandular (especially on distal portions); branches of capitulescence with minute stipitate glands; rays purple or at least purple-tinged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. macrophylla

Fig. 406  Leaf of Eupatorium pubescens.

40 0 tricolpate s

2b. Leaf blades eglandular; branches of capitulescence without glands (sometimes with sparse, tiny glands in E. schreberi); rays white 3a. Plants with vegetative tufts of leaves produced separately from the flowering stem; involucre slender-cylindric, 5.5–7.5 mm tall; leaves at base of plant relatively firmer, often with rectangular basal sinuses, with mostly 15–30 teeth per margin . . . . . . . E. schreberi 3b. Plants without separate vegetative tufts of leaves; involucre funnelform, 5–6.5 mm tall; leaves at base of plant relatively thinner, with a V-shaped basal sinus, with mostly 6–15 teeth per margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. divaricata 1b. Basal leaf blades not cordate, sessile or petiolate [Fig. 408] 4a. Stipitate glands present on involucral bracts and branches of capitulescence; leaves basally disposed; plants of dry, sandy soils along the coastal plain . . . . . . . . . E. spectabilis 4b. Stipitate glands lacking; leaves chiefly cauline; plants of peatlands, shores, and moist soils of forests, more common in north-temperate, boreal, and cool, maritime climates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. radula 1. Eurybia divaricata (L.) Nesom N white wood-aster. Aster castaneus Burgess; A. divaricatus L.; A. excavatus Burgess; A. tenebrosus Burgess • CT, MA, ME, NH, RI, VT; in ME confined to southern portion of state. Deciduous forests. 2. Eurybia macrophylla (L.) Cass. N Fig. 407 Fig. 407  Involucral bract of Eurybia macrophylla showing abundant cilia and broad, apical green zone.

large-leaved wood-aster. Aster ianthinus Burgess; A. macrophyllus L.; A. macrophyllus L. var. apricensis Burgess; A. macrophyllus L. var. excelsior Burgess; A. macrophyllus L. var. ianthinus (Burgess) Fern.; A. macrophyllus L. var. pinguifolius Burgess; A. macrophyllus L. var. velutinus Burgess • CT, MA, ME, NH, RI, VT. Deciduous and mixed evergreen-deciduous forests. ‌2 × 5. Eurybia ×herveyi (Gray) Nesom is an uncommon hybrid typically occurring on the coastal plain of CT, MA, RI. It has basally disposed leaves with elliptic to broad-ovate leaf blades that are contracted to a definite petiole, appressed to slightly squarrose and stipitate-glandular involucral bracts, and purple rays. 3. Eurybia radula (Ait.) Nesom N Fig. 408 rough wood-aster. Aster radula Ait.; A. radula Ait. var. strictus (Pursh) Gray; A. strictus Pursh • CT, MA, ME, NH, RI, VT. Shorelines, wetland edges, openings in and trail edges through evergreen and mixed evergreen-deciduous forests. 4. Eurybia schreberi (Nees) Nees N

Fig. 408  Capitulescence and leaves of Eurybia radula.

Schreber’s wood-aster. Aster curvescens Burgess; A. glomeratus (Bernh. ex Nees) Burgess; A. schreberi Nees; Eurybia glomerata Bernh. ex Nees • CT, MA, ME, NH, RI, VT; restricted to southern ME and southern half of NH. Upland and riparian forests, forest fragments. 5. Eurybia spectabilis (Ait.) Nesom N purple wood-aster. Aster commixtus (Nees) Kuntze; A. spectabilis Ait.; A. spectabilis Ait. var. suffultus Fern.; Eurybia commixta Nees • CT, MA, RI. Sandy soils of open woodlands, roadsides, fields, and clearings.

Euthamia Leaf blade measurements in the following key are for those leaves produced along the main axis of the stem. Leaves from within the branches of the capitulescence (i.e., bracts), which are frequently (and incorrectly) referenced for measurements, will yield smaller results. Reference: Haines (2006). 1a. Leaf blades with numerous and prominent resin glands [Fig. 409], 1–3 (–6) mm wide; plants ± glabrous on the stem and leaf surfaces; axillary fascicles often well developed and conspicuous throughout much of the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. caroliniana

Ast e r ac e a e   40 1

1b. Leaf blades with few and/or inconspicuous resin glands, usually 3–12 mm wide; plants varying from glabrous to conspicuously pubescent on the stem and abaxial leaf surfaces; axillary fascicles, if present, usually poorly developed and sparse . . . . . . . . . . . . E. graminifolia 1. Euthamia caroliniana (L.) Greene ex Porter & Britt. N Fig. 409 coastal plain grass-leaved-goldenrod. Euthamia galetorum Greene; E. microcephala Greene; E. microphylla Greene; E. remota Greene; Solidago tenuifolia Pursh; S. tenuifolia Pursh var. pycnocephala Fern. • CT, MA, ME, NH, RI; also reported from VT by Seymour (1982), but specimens are unknown. Coastal plain pond shores, peaty depressions, wet borrow pits. 2. Euthamia graminifolia (L.) Nutt. N common grass-leaved-goldenrod. Chrysocoma graminifolia L.; Euthamia graminifolia (L.) Nutt. var. major (Michx.) Moldenke; E. graminifolia (L.) Nutt. var. nuttallii (Greene) W. Stone; Solidago graminifolia (L.) Salisb. var. major (Michx.) Fern.; S. graminfolia (L.) Salisb. var. nuttallii (Greene) Fern.; S. graminifolia (L.) Salisb. var. septentrionalis Fern.; S. lanceolata L. • CT, MA, ME, NH, RI, VT; throughout. Fields, shorelines, roadsides, wetland edges.

Fig. 409  Leaf of Euthamia caroliniana with detail of numerous resin glands.

Eutrochium Reference: Lamont (2006). 1a. Capitula with (8–) 9–22 flowers; capitulescence or its divisions flat-topped [Fig. 411]; stems typically green with anthocyanic spots or streaks . . . . . . . . . . . . . . . . . . . . . E. maculatum 1b. Capitula with (4–) 5–9 (–10) flowers; capitulescence or its divisions slightly to strongly convex [Fig. 410]; stems spotted with anthocyanin or anthocyanic only in bands at the nodes or uniformly anthocyanic throughout the stem 2a. Leaf blades triple-veined, with the lowest pair of lateral veins more conspicuous and more prolonged than the other lateral veins, abruptly contracted to the petiole; stems spotted or streaked with anthocyanin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. dubium 2b. Leaf blades pinnately veined or gradually tapered to the petiole or both; stems usually with anthocynin concentrated in nodal bands or uniformly distributed throughout the stem 3a. Stems usually with anthocyanin concentrated in nodal bands 1–2 cm long, not or only scarcely glaucous, solid or developing a slender central cavity near the base; plants typically occurring in upland, deciduous woodlands and in open, well-drained habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. purpureum 3b. Stems usually anthocyanic throughout (though often green in deep shade and sometimes spotted when young), prominently glaucous, developing a large central cavity; plants typically occurring in wetlands, riparian communities, and other low, hydric habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. fistulosum 1. Eutrochium dubium (Willd. ex Poir.) E.E. Lamont N coastal plain Joe-Pye weed. Eupatoriadelphus dubius (Willd. ex Poir.) King & H.E. Robins.; Eupatorium dubium Willd. ex Poir.; E. ternifolium Ell. • CT, MA, ME, NH, RI; restricted in ME to southern portion of state. Pond and stream shores, edges of pools, low meadows, wet fields. Reports of this species in VT are based primarily on collections of Eutrochium maculatum. 2. Eutrochium fistulosum (Barratt) E.E. Lamont N Fig. 410 hollow Joe-Pye weed. Eupatoriadelphus fistulosus (Barratt) King & H.E. Robins.; Eupatorium fistulosum Barratt; E. laevigatum Torr. in Eat. • CT, MA, ME, NH, RI; restricted in ME to sourthern portion of state. Fields, edges of swamps, riparian forests, low meadows. Reports of this species in VT (e.g., Seymour 1982) were based on misidentified collections of Eutrochium maculatum. Hollow stems do not uniquely distinguish this species because E. maculatum sometimes has a hollow stem. Fig. 410  Capitulescence of Eutrochium fistulosum.

402 tricolpate s

3. Eutrochium maculatum (L.) E.E. Lamont var. maculatum N Fig. 411 spotted Joe-Pye weed. Eupatoriadelphus maculatus (L.) King & H.E. Robins.; Eupatorium maculatum L.; E. maculatum L. var. foliosum (Fern.) Wieg.; E. purpureum L. var. maculatum (L.) Darl. • CT, MA, ME, NH, RI, VT; rare or absent from part of the coastal plain in MA and RI. Fields, edges of marshes, swamps, wet ditches, shorelines; ranging further north and ascending to higher elevations than other members of the genus in New England. 4. Eutrochium purpureum (L.) E.E. Lamont var. purpureum N Fig. 411  Capitulescence of Eutrochium maculatum.

purple Joe-Pye weed. Cunigunda purpurea (L.) Lunell; Eupatoriadelphus purpureus (L.) King & H.E. Robins.; Eupatorium amoenum Pursh; E. falcatum Michx.; E. purpureum L.; E. trifoliatum L.; E. verticillatum Muhl. ex Willd. • CT, MA, NH, RI, VT. Woodlands, forests, fields, pastures.

Filago 1. Filago vulgaris Lam. E common cotton-rose. Filago germanica L. • MA. Dry-mesic, sandy soils of fields, roadsides, and waste areas.

Flaveria 1. Flaveria trinervia (Spreng.) C. Mohr E clustered yellowtops. Oedera trinervia Spreng. • MA. Disturbed areas, frequently near water and often with saline influence.

Gaillardia 1a. Rays entirely yellow or yellow with a purple base; setae-like chaff evidently exceeding the mature cypsela bodies; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. aristata 1b. Rays entirely red-purple or red-purple with a yellow apex; setae-like chaff about equaling to shortly exceeding the mature cypsela bodies; plants annual to short-lived perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. pulchella 1. Gaillardia aristata Pursh E common blanket-flower. CT, MA, NH. Fields, roadsides, disturbed soil. 2. Gaillardia pulchella Foug. var. pulchella E rosering blanket-flower. Gaillardia drummondii (Hook.) DC.; G. picta D. Don; G. pulchella Foug. var. picta (D. Don) Gray • CT, MA, ME, NH, VT. Fields, roadsides, disturbed soil, areas of cultivation.

Galinsoga 1a. Pappus of the disk flowers with a short awn-tip; pappus of ray flowers well-developed and equaling the length of the connate, tubular portion of the ray flower; cypselas pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. quadriradiata 1b. Pappus of the disk flowers without an awn-tip; pappus of ray flowers absent or poorly developed; cypselas sparsely pubescent or glabrous . . . . . . . . . . . . . . . . . . . . . . . . . G. parviflora 1. Galinsoga parviflora Cav. E Fig. 412 lesser quickweed. Galinsoga parviflora Cav. var. semicalva Gray; G. semicalva (Gray) St. John & White • CT, MA, ME, NH, RI, VT. Gardens, cultivated fields, barnyards, waste areas, roadsides. 2. Galinsoga quadriradiata Ruiz & Pavón E Fig. 412  Capitula and leaves of Galinsoga parviflora.

common quickweed. Galinsoga aristulata Bickn.; G. bicolorata St. John & White; G. ciliata (Raf.) Blake; G. caracasana (DC.) Schultz-Bip. • CT, MA, ME, NH, RI, VT. Gardens, cultivated fields, barnyards, waste areas, roadsides.

Ast e r ac e a e   403

Gamochaeta Reference: Nesom (2006b). 1a. Stem leaf blades obovate-spatulate, loosely tomentose to glabrate abaxially, weakly bicolored (i.e., adaxial and abaxial surfaces of nearly same color); hairs on adaxial surface of leaf blades slender at base; capitulescence usually interrupted; outer involucral bracts acuminate-apiculate at apex; inner involucral bracts 3–3.5 mm long, not apiculate; receptacles deeply concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. pensylvanica 1b. Stem leaf blades usually oblanceolate, closely tomentose abaxially, strongly bicolored; hairs on adaxial surface of leaf blades expanded at base; capitulescence usually continuous; outer involucral bracts acute at apex; inner involucral bracts (3.5–) 4–5 mm long, usually with a minute apiculus; receptacles shallowly concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. purpurea 1. Gamochaeta pensylvanica (Willd.) Cabrera E Pennsylvania cudweed. Gnaphalium pensylvanicum Willd.; G. purpureum Willd. var. spathulatum (Lam.) Baker • MA. Dumps, disturbed soil. 2. Gamochaeta purpurea (L.) Cabrera

N C Fig. 413

purple cudweed. Gnaphalium purpureum L. • CT, MA, ME, RI. Sandy soils of fields, grasslands, woodland margins, and beaches.

Fig. 413  Capitulescence of Gamochaeta purpurea.

Glebionis 1a. Rays with 2 or more colors, frequently yellow with a dark red band just above the base, though the apical color of the ray may be various shades of red, orange, pink, or white; outer involucral bracts keeled; cypselas ± flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. carinata 1b. Rays yellow; outer involucral bracts not keeled; cypselas compressed-triangular to compressed-columnar or columnar 2a. Principal leaf blades toothed to pinnatifid, the rachis of the blade (i.e., excluding the teeth or lobes) wider than 6 mm, clasping the stem (except the lowest leaves which are narrowed to petiole-like base) [Fig. 414]; cypsela bodies of center of capitilum subterete, the angles with a prominent rib . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. segetum 2b. Principal leaf blades bipinnatifid, the rachis of the blade narrower than 6 mm, not clasping the stem; cypsela bodies of center of capitulum subterete, the angles with a prominent rib, and also with a minor rib between them . . . . . . . . . . . . . . . . . . . . G. coronaria 1. Glebionis carinata (Schousb.) Tzvelev E tricolor daisy. Chrysanthemum carinatum Schousb. • MA. Roadsides, rubbish heaps.

Fig. 414  Capitula and leaves of Glebionis segetum.

2. Glebionis coronaria (L.) Cass. ex Spach E crown daisy. Chrysanthemum coronarium L. • MA, ME, VT. Fields, roadsides, gardens, compost heaps. 3. Glebionis segetum (L.) Fourr. E Fig. 414 corn daisy. Chrysanthemum segetum L. • CT, MA, ME. Fields, roadsides, gardens, compost heaps.

Gnaphalium Gnaphalium has been segregated into several smaller genera on the basis of morphology, base chromosome number, and phylogeny. See also Euchiton, Gamochaeta, Omalotheca, and Pseudognaphalium. 1. Gnaphalium uliginosum L. E Fig. 415 brown cudweed. Filaginella uliginosa (L.) Opiz • CT, MA, ME, NH, RI, VT. Fields, roadsides, yards, disturbed soil.

Fig. 415  Axillary capitulescence of Gnaphalium uliginosum.

404  tricolpate s

Grindelia 1a. Leaf blades toothed with mostly acute teeth, the apex of each tooth apiculate to setose (leaves rarely subentire); middle involcural bracts terminated by green appendages (1.2–) 2.5–5.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. squarrosa 1b. Leaf blades toothed with obtuse to rounded teeth, the apex of each tooth with a prominent resin gland (leaves rarely subentire); middle involucral bracts terminated by green appendages 1–1.7 (–3) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. hirsutula 1. Grindelia hirsutula Hook. & Arn. E hairy gum-weed. Grindelia squarrosa (Pursh) Dunal var. integrifolia (Nutt.) Boivin; G. squarrosa (Pursh) Dunal var. quasiperennis Lunell • CT. Fields, yards. The specimens vouchering this species are problematic in that the leaf blades are subentire (i.e., details of the leaf teeth are unavailable for confirming the identity of the collection). Further, other collections from the same site are clearly referable to Grindelia squarrosa. 2. Grindelia squarrosa (Pursh) Dunal E curly-top gum-weed. Donia squarrosa Pursh; Grindelia squarrosa (Pursh) Dunal var. serrulata (Rydb.) Steyermark; G. serrulata Rydb. • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, waste areas.

Guizotia 1. Guizotia abyssinica (L. f.) Cass. E niger-seed. Polymnia abyssinica L. f. • CT, MA. Fields, roadsides, waste areas.

Helenium 1a. Leaves not decurrent on stems and branches, the blades linear to linear-filiform, 1–2 (–4) mm wide; plants annual, from taproots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. amarum 1b. Leaves decurrent on stems and branches, the blades broad-linear to oblanceolate, elliptic, or narrow-ovate, 5–40 mm wide; plants perennial, from fibrous roots 2a. Ray flowers carpellate, numbering mostly 13–21 per capitulum [Fig. 416]; disk flowers yellow; larger leaf blades coarsely toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. autumnale 2b. Ray flowers neutral, numbering mostly 8–13 per capitulum, rarely absent; disk flowers red-brown to purple-brown; leaf blades entire or subentire . . . . . . . . . . . . . . . . H. flexuosum 1. Helenium amarum (Raf.) H. Rock var. amarum E narrow-leaved sneezeweed. Helenium tenuifolium Nutt. • CT, MA. Fields, roadsides, waste areas. 2. Helenium autumnale L. N Fig. 416

Fig. 416  Capitulum of Helenium autumnale.

fall sneezeweed. Helenium autumnale L. var. canaliculatum (Lam.) Torr. & Gray; H. autumnale L. var. parviflorum (Nutt.) Fern.; H. canaliculatum Lam.; H. parviflorum Nutt. • CT, MA, ME, RI, VT. Shorelines, river banks, fields, ditches. 3. Helenium flexuosum Raf. N purple-headed sneezeweed. Helenium nudiflorum Nutt. • CT, MA, ME, NH, RI, VT. River banks, fields, pastures, roadsides.

Helianthus Identification of Helianthus is complicated by phenotypic plasticity, polyploidy, and occasional hybridization. Micromorphological characters are a great asset within this genus and are used extensively in the key (e.g., anther appendage and style branch colors, disk corolla indument, cypsela size and indument). Reference: Schilling (2006).

Ast e r ac e a e  40 5

1a. Plants annual, with fibrous roots; leaves alternate (except the lowermost); receptacle flat or nearly so; disk corollas red-purple (sometimes yellow in H. annuus) 2a. Involucral bracts narrow-oblong to ovate, (3–) 5–8 mm wide, abruptly narrowed to the apex, conspicuously long-ciliate; cypselas usually glabrous except near the pubescent apex, 4–8 mm wide; style branches yellow; leaf blades 5–25 cm wide, dentate; plants (0.5–) 1–3 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. annuus 2b. Involucral bracts lanceolate, 1–4 (–5) mm wide, gradually narrowed to the apex, either eciliate or ciliate with short hairs of similar length to the hairs of the abaxial surface; cypselas pubescent, 1.2–2.5 mm wide; style branches red (rarely yellow in H. debilis); leaf blades 1.5–9 cm wide, entire to undulate-dentate; plants 0.4–1 (–1.5) m tall 3a. Chaff near the center of the disk inconspicuously short-pubescent; cypselas mottled, pubescent with spreading-ascending hairs; stems frequently mottled; leaf blades dark green and scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. debilis 3b. Chaff near the center of the disk conspicuously pubescent near the apex with long, white hairs; cypselas not mottled, pubescent with appressed-ascending hairs; stems not mottled; leaf blades pale green and strigose . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. petiolaris 1b. Plants perennial, with rhizomes, stolons, tuberous roots, and/or tough, overwintering bases; leaves opposite (except often the upper); receptacle usually convex to some degree; disk corollas yellow (red-purple in H. pauciflorus) 4a. Reproductive stems with 3–5 (–8) nodes bearing highly reduced leaves (rarely the lower 2–4 nodes with well-formed blades), often the upper 50% of the stem lacking leaves; plants with a basal rosette of leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. occidentalis 4b. Reproductive stems with (5–) 6–15 or more leaf-bearing nodes, the blades gradually, if at all, descreasing in size upward, only the upper 25% or less of the stem lacking leaves; plants without a basal rosette of leaves, during anthesis the leaves chiefly cauline (except sometimes in H. pauciflorus) 5a. Disk corollas red-purple (sometimes only the lobes red-purple and the remainder of the limb yellow); involucral bracts acute to obtuse at the apex, 3–5 mm wide, appressed, of several, conspicuously different lengths [Fig. 420] . . . . . . . . . . . . . . . . . . H. pauciflorus 5b. Disk corollas yellow; involucral bracts acuminate to attenuate (rarely acute) at the apex, 1.2–3.5 mm wide, at least some merely ascending, of several, inconspicuously different lengths [Fig. 417] 6a. Stem and axis of capitulescence conspicuously pubescent, not glaucous 7a. Stem, abaxial surface of leaf blades, and involucral bracts densely pubescent with short, soft hairs; leaf blades sessile, subcordate to cordate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. mollis 7b. Stem, abaxial surface of leaf blades, and involucral bracts with sparser and/ or coarser pubescence; leaf blades short- to long-petioled (sometimes sessile in H. giganteus), tapering to rounded at the base (sometimes truncate in H. hirsutus) 8a. Leaf blades (4–) 5–12 (–15) cm wide, with a petiole (15–) 20–80 mm long; roots forming tubers later in growing season; cypsela body 5–7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. tuberosus 8b. Leaf blades 1–4 (–8) cm wide, with a petiole up to 20 mm long; roots fibrous or fleshy, merely thickened; cypsela body 3–5 mm long 9a. Stem strigose; leaf blades sometimes folded and falcate-arched, especially in drying, pinnately veined near the base [Fig. 419] . . . . . . . . . . . . H. maximiliani 9b. Stem spreading-hirsute; leaf blades flat, usually with 3 prominent, parallel nerves near the base

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10a. Leaf blades cuneate at base; petioles 0–12 mm long; roots fleshy and somewhat thickened; involucral bracts linear, 1.2–2 mm wide; anther appendages dark brown . . . . . . . . . . . . . . . . . . . . . . . . . (in part) H. giganteus 10b. Leaf blades rounded to truncate at base; petioles 4–20 mm long; roots fibrous; involucral bracts lanceolate, 2.5–3.5 mm wide; anther appendages usually yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. hirsutus 6b. Stem glabrous (or very nearly so) and often glaucous, only the axis of the capitulescence pubescent 11a. Leaves sessile or with short petioles to 5 (–10) mm long [Fig. 418] 12a. Leaf blades lanceolate to oblong-lanceolate, widest near middle, narrowly tapering to the base, usually alternately arranged on the upper portion of the stem; disk 15–25 mm wide; anther appendages dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) H. giganteus 12b. Leaf blades lanceolate to triangular-ovate, widest near the base, truncate to broadly rounded at the base [Fig. 418], usually oppositely arranged throughout the stem; disk 10–15 mm wide; anther appendages yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. divaricatus 11b. Well-developed leaves with petioles (5–) 10–60 mm long 13a. Disk 4–10 mm wide; capitula with 5–8 ray flowers . . . . . H. microcephalus 13b. Disk 15–35 mm wide; capitula with 8–20 ray flowers 14a. Leaf blades at least somewhat triple-veined, those near the middle of the stem 25–100 mm wide, broad-lanceolate to ovate, usually less than 3 times as long as wide; capitula with 8–15 ray flowers 15a. Involucral bracts spreading, evidently surpassing the disk [Fig. 417]; lobes of the disk corollas pubescent; petioles 15–60 mm long; leaves relatively thin, membranaceous to herbaceous, prominently serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. decapetalus 15b. Involucral bracts ascending, approximately equaling the height of the disk; lobes of the disk corollas glabrous; petioles 5–30 mm long; leaves thick and firm, subentire to shallowly serrate . . . . . . . H. strumosus 14b. Leaf blades pinnately veined, those near the middle of the stem 10–40 (–90) mm wide, lanceolate to narrow-ovate, usually 3–8 times as long as wide; capitula with 10–20 ray flowers 16a. Lobes of the disk corollas glabrous or glabrate; anther appendages yellow; leaf blades slightly, if at all, scabrous on the adaxial surface and pubescent on the abaxial surface with hairs up to 0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. grosseserratus 16b. Lobes of the disk corollas pubescent; anther appendages red-brown to dark brown; leaf blades strongly scabrous on the adaxial surface and pubescent on the abaxial surface with hairs 1 mm long or longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) H. giganteus 1. Helianthus annuus L. E common sunflower. Helianthus annuus L. var. lenticularis (Dougl. ex Lindl.) Steyermark; H. annuus L. var. macrocarpus (DC.) Cockerell; H. annuus L. var. texanus (Heiser) Shinners • CT, MA, ME, NH, RI, VT. Roadsides, fields, yards, waste areas. 2. Helianthus debilis Nutt. ssp. cucumerifolius (Torr. & Gray) Heiser E cucumber-leaved sunflower. Helianthus cucumerifolius Torr. & Gray; H. debilis Nutt. var. cucumerifolius (Torr. & Gray) Gray • CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas.

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3. Helianthus decapetalus L. N Fig. 417 thin-leaved sunflower. Helianthus tracheliifolius P. Mill. • CT, MA, ME, NH, RI, VT. Riparian forests, mesic, deciduous forests, forest edges. 4. Helianthus divaricatus L. N Fig. 418 woodland sunflower. Helianthus divaricatus L. var. angustifolius Kuntze • CT, MA, ME, NH, RI, VT. Woodlands, forest edges, dry fields and glades. 4 ‌ × 6. Helianthus ×divariserratus R.W. Long is an extremely rare sunflower hybrid in New England known from CT. It has a glabrous and glaucous stem bearing opposite leaves that are triple-veined (those of H. grosseserratus are pinnately veined), rounded at the base, and borne on short petioles 5–10 mm long (those of H. divaricatus truncate to broadly rounded at the base and sessile). The involucral bracts are 10–15 mm long (6–12 mm long in H. divaricatus, 10–14 mm long in H. grosseserratus).

Fig. 417  Involucre and ray flowers of Helianthus decapetalus.

5. Helianthus giganteus L. E tall sunflower. Helianthus alienus E.E. Wats.; H. borealis E.E. Wats.; H. giganteus L. ssp. alienus (E.E. Wats.) R.W. Long; H. giganteus L. var. subtuberosus Britt.; H. subtuberosus (Britt.) Britt.; H. validus E.E. Wats. • CT, MA, ME, VT; also reported from RI by George (1997), but specimens are unknown. Fields, borders of brackish marshes, disturbed soil. 5 ‌ × 6. Helianthus ×luxurians E.E. Wats. is a very rare hybrid sunflower known from CT, MA. It closely resembles its parental species. It shows ± glabrous and glaucous stems, and variably toothed leaf blades (subentire to coarsely toothed) that are usually 5–10 times as long as wide and borne on short petioles 5–10 mm long.

Fig. 418  Leaves of Helianthus divaricatus.

6. Helianthus grosseserratus Martens E saw-toothed sunflower. Helianthus grosseserratus Martens var. hypoleucus Gray • CT, MA, ME, NH. Roadsides, fields, disturbed soil. ‌ × 8. Helianthus ×intermedius R.W. Long is a very rare sunflower hybrid known from 6 MA, ME. It has sparsely strigose stems, petioles 5–15 mm long, and flat leaf blades that are strongly scabrous adaxially. ‌6 × H. salicifolius A. Dietr. Helianthus ×kellermanii Britt. is a rare and local hybrid known from MA, ME. It is similar to H. grosseserratus, differing primarily in the linear to narrow-lanceolate principal leaf blades that are 10–20 mm wide and taper to the petiole and lobes of the disk corollas that are white-pubescent on the abaxial surface (vs. principal leaf blades lanceolate to narrow-ovate, 12–90 mm wide, tapering or rounded to the petiole, and disk corolla lobes glabrous or nearly so). 7. Helianthus hirsutus Raf. E hairy sunflower. Helianthus hirsutus Raf. var. stenophyllus Torr. & Gray; H. stenophyllus (Torr. & Gray) E.E. Wats. • CT. Roadsides, fields, disturbed soil. 8. Helianthus maximiliani Schrad. E Fig. 419 Maximilian’s sunflower. CT, MA, ME. Roadsides, fields, disturbed soil. 9. Helianthus microcephalus Torr. & Gray E

Fig. 419  Leaf blade of Helianthus maximiliani.

small-headed sunflower. CT. Gardens, waste areas. 10. Helianthus mollis Lam. E ashy sunflower. Helianthus mollis Lam. var. cordatus S. Wats. • CT, MA, ME, RI. Roadsides, fields, disturbed soil. 11. Helianthus occidentalis Riddell ssp. occidentalis E naked-stemmed sunflower. MA. Roadsides, fields, disturbed soil. 12. Helianthus pauciflorus Nutt. E Fig. 420 stiff sunflower.  12a. Helianthus laetiflorus Pers. var. rigidus (Cass.) Fern.; H. rigidus (Cass.) Desf.; 12b. Helianthus laetiflorus Pers. var. subrhomboideus (Rydb.) Fern.; H. pauciflorus

Fig. 420  Involucre of Helianthus pauciflorus.

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Nutt. var. subrhomboideus (Rydb.) Cronq.; H. rigidus (Cass.) Desf. ssp. subrhomboideus (Rydb.) Heiser; H. rigidus (Cass.) Desf. var. subrhomboideus (Rydb.) Cronq.; H. subrhomboideus Rydb. • CT, MA, ME, NH, RI; also reported from VT by Kartesz (1999), but specimens are unknown. Roadsides, fields, disturbed soil. 1a. Reproductive stems 8–20 dm tall, with 9–15 leaf-bearing nodes below the capitulescence; leaves usually alternate on upper portion of stem, the blades oblong-lanceolate to narrowovate, 8–27 cm long, and acuminate at the apex . . . . . . 12a. H. pauciflorus ssp. pauciflorus 1b. Reproductive stems 5–12 dm tall, with 5–10 leaf-bearing nodes below the capitulescence; leaves usually opposite throughout the stem, the blades rhombic-ovate to narrow-lanceolate, 5–12 cm long, and acute to obtuse at the apex . . . . . . . . . . . . . . . . . . 12b. H. pauciflorus ssp. subrhomboideus (Rydb.) O. Spring & E. Schilling Subspecies pauciflorus is known from CT, MA, ME, NH, RI. Subspecies subrhomboideus is known from CT, MA, ME, NH. Clevenger and Heiser (1963) gave detailed accounts of crossing studies that suggested Helianthus × ‌laetiflorus is best treated as a hybrid (see below) and that ssp. pauciflorus may have arisose through through hybridization between H. pauciflorus ssp. subrhomboideus and H. tuberosus with later character segregation such that the stabilized entity closely resembles the former parent. Introgression with the latter species would explain the taller stems, longer leaves that are often alternate above, and longer petioles. ‌12 × 15. Helianthus ×laetiflorus Pers. is a rare hybrid sunflower that frequently occurs in the absence of its parents. It is known from CT, MA, ME, NH, RI, VT. It is similar to H. pauciflorus in regard to involucre characteristics and often disk corolla color (though the corollas can also be yellow), but the involucral bracts are oblong-lanceolate with an acuminate apex, are sometimes sparsely short-pubescent abaxially, and 7–12 mm long (vs. elliptic to oblong-ovate, acute to obtuse, ± glabrous abaxially, and 6–10 mm long; lanceolate with an acuminae apex, hispid abaxially, and 8.5–15 mm long in H. tuberosus). It also occasionally shows the branching tubers of H. tuberosus. 13. Helianthus petiolaris Nutt. ssp. petiolaris E prairie sunflower. CT, MA, ME, VT. Roadsides, fields, disturbed soil. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species had questionable occurrence in RI and was unaware of any collections. 14. Helianthus strumosus L. N pale-leaved sunflower. CT, MA, ME, NH, RI, VT. Deciduous forests, riverbanks, fields, roadsides, open rights-of-way. 15. Helianthus tuberosus L. E tuberous sunflower. Helianthus tomentosus Michx. • CT, MA, ME, NH, RI, VT. Stream banks, riparian forests, fields, abandoned homesteads, areas of cultivation. Helianthus tuberosus is often considered an introduced species. However, it behaves as a native species in many riparian forests (Schilling 2006). The species may be best considered as native to North America with introduced occurrences (the species was frequently planted for its starchy tubers).

Heliomeris 1. Heliomeris multiflora Nutt. var. multiflora E showy false goldeneye. Gymnolomia multiflora (Nutt.) Benth. & Hook. f.; Viguiera multiflora (Nutt.) Blake • MA. Wool waste. This species sometimes shows a mixture of leaf arrangements, ranging from alternate to opposite (the latter character state most prevalent).

Heliopsis Fig. 421  Fruiting capitulum of Heliopsis helianthoides with persistent rays.

1. Heliopsis helianthoides (L.) Sweet E Fig. 421 sunflower-everlasting.  1a. Buphthalmum helianthoides L.; Heliopsis helianthoides (L.) Sweet var. solidaginoides (L.) Fern.; 1b. Heliopsis helianthoides (L.) Sweet ssp. occidentalis T.R. Fisher;

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H. helianthoides (L.) Sweet var. occidentalis (T.R. Fisher) Steyermark; H. helianthoides (L.) Sweet ssp. scabra (Dunal) T.R. Fisher; H. scabra Dunal • CT, MA, ME, NH, RI, VT; also reported from RI by Kartesz (1999), but specimens are unknown. Fields, roadsides, dumps, disturbed soil. 1a. Leaf blades thin, smooth on both surfaces or slightly scabrous adaxially, 4–8 (–12) cm wide; stem smooth, glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. H. helianthoides var. helianthoides 1b. Leaf blades firm, harshly scabrous adaxially and somewhat scabrous abaxially, 2–5 cm wide; stem often scabrous, not glaucous . . . . . 1b. H. helianthoides var. scabra (Dunal) Fern. Variety helianthoides is known from CT, MA, VT. Variety scabra is known from CT, MA, ME, NH, VT.

Helminthotheca 1. Helminthotheca echioides (L.) Holub E bristly ox-tongue. Picris echioides L. • CT, MA, ME, VT. Fields, roadsides, waste areas.

Heterosperma 1. Heterosperma pinnatum Cav. E wingpetal. MA. Gardens, wool waste.

Heterotheca 1. Heterotheca subaxillaris (Lam.) Britt. & Rusby var. latifolia (Buckl.) Gandhi & Thomas E camphor false golden-aster. Heterotheca latifolia Buckl.; H. subaxillaris (Lam.) Britt. & Rusby ssp. latifolia (Buckl.) Semple • CT. Fields, roadsides, vacant lots, disturbed soil.

Hieracium Sell (1987) argued convincingly, on morphological grounds, for the segregation of Pilosella from Hieracium. Gaskin and Wilson (2007) showed that Pilosella is a monophyletic group. However, they also showed that recognition of Pilosella creates a paraphyletic Hieracium unless the genus Stenotheca, a genus of western hemisphere plants, is also recognized. The morphological distinction between Hieracium and Stenotheca is relatively weak and does not seem to justify generic separation. Therefore, the genus Hieracium is recognized here in the broad sense until such time as study elucidates morphological separation of Stenotheca from Hieracium. References: Voss (1996), Strother (2006c). 1a. Cypsela body 1.2–2 (–2.2) mm long, each rib on the surface of the ovary projecting to form a crenulate apex [Fig. 422]; pappus bristles mostly 25–40 in 1 series [Fig. 422]; rays yellow, yellow with an red abaxial stripe, or orange-red; stems scapose, without leaves or with few and reduced leaves; leaves tapering to an ill-defined petiole; plants with a well-developed rosette of basal leaves at anthesis, commonly with stolons [Fig. 426] 2a. Reproductive stems bearing a solitary capitulum or a capitulescence with 2–4 (–6) capitula on elongate peduncles (5–) 15–150 mm long [Fig. 426]; leaf blades 2–4 (–6) times as long as wide; plants 3–25 (–40) cm tall 3a. Capitulescence with 1 or 2 (–3) capitula [Fig. 426]; involucres 7.5–9 (–10) mm tall; leaf blades densely pubescent with stellate hairs on the abaxial surface, the hairs usually contiguous and concealing the surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. pilosella 3b. Capitulescence with (1–) 2–4 (–6) capitula; involucres (9–) 10–13 mm tall; leaf blades moderately pubescent with stellate hairs on the abaxial surface, the hairs not so numerous as to be contiguous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. flagellare 2b. Reproductive stems bearing (3–) 5–30 (–50) capitula, these usually in compact, corymb-like capitulescences on short peduncles 1–15 (–28) mm long; leaf blades 3–8 times as long as wide; plants (10–) 20–100 cm tall

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4a. Ray flowers orange-red (drying dark red); involucral bracts 1.5–3 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. aurantiacum 4b. Ray flowers yellow; involucral bracts 0.5–1.25 mm wide 5a. Involucres 7.5–9 mm tall; pappus bristles 4–5 (–6) mm long; ray corollas 8–12 mm long; leaf blades green, with abundant setae adaxially; plants with short or, more commonly, long rhizomes and often also short, prostrate stolons . . . . . H. caespitosum 5b. Involucres 5–6 (–7) mm tall; pappus bristles 3–4 mm long; ray corollas 6–9 mm long; leaf blades glaucous, sparsely and unevenly setose to subglabrous adaxially; plants usually with short, praemorse rhizomes and also usually lacking prostrate stolons (sometimes with divergent to ascending branches) 6a. Peduncles lacking stellate hairs or with very few stellate hairs; basal leaf blades lacking stellate-hairs on the abaxial surface, either glabrous or with simple, eglandular hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. piloselloides 6b. Peduncles moderately to densely pubescent with stellate hairs; basal leaf blades usually with minute stellate-hairs on the abaxial surface (longer, simple, eglandular ones may be present as well) . . . . . . . . . . . . . . . . . . . . . . . . . H. praealtum 1b. Cypsela body (2–) 2.5–5 mm long, each rib apically confluent to form an obscure ring [Fig. 424]; pappus bristles mostly 30–60 (or more) in 2 series; rays yellow; stems scapose or evidently leafy; lower leaves usually tapering to a ± distinct petiole; plants with a basal rosette of leaves or these withered at anthesis, never with stolons 7a. Reproductive stems with (4–) 6–50 leaves; basal leaves mostly absent or withered at anthesis, not forming a conspicuous rosette 8a. Involucres 6–7 mm tall; pappus bristles 4–5 mm long; ray corollas 5–8 mm long; capitula with 8–20 (–30) flowers; peduncles very slender, 0.2–0.3 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. paniculatum 8b. Involucres 7–10 (–11) mm tall; pappus bristles (4.5–) 5–7 mm long; ray corollas 8–18 mm long; capitula with 20–100 flowers; peduncles stouter, 0.3–1.1 mm thick 9a. Cypsela body tapering to an expanded, circular apex where the pappus bristles attach [Fig. 423]; capitula with 20–40 flowers; capitulescence panicle-like, cylindrical in outline, with short lower branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. gronovii 9b. Cypsela body truncate or scarcely narrowed to the circular apex [Fig. 424]; capitula with (30–) 40–100 flowers; capitulescence corymb-like or umbel-like, with elongate lower branches (sometimes narrow and panicle-like in H. scabrum) 10a. Peduncles with abundant stipitate glands [Fig. 427]; leaf blades entire or remotely denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. scabrum 10b. Peduncles with very few or no stipitate glands; at least the lower leaf blades usually dentate 11a. Hairs of the lower stem and leaf surfaces simple, firm, and bulbous-based, short compound hairs usually absent from the leaf surfaces . . . . H. sabaudum 11b. Hairs of the lower stem and leaf surfaces simple or compound, but not bulbous-based, that of the leaves sometimes compound 12a. Involucral bracts tapering to an acuminate apex; reproductive stems 10–35 cm tall, with 4–10 leaves . . . . . . . . . . . . . . . . . . . . . (in part) H. robinsonii 12b. Involucral bracts tapering to a narrow-rounded to acute apex; reproductive stems 15–60 (–150) cm tall, with 6–50 leaves 13a. Leaf blades linear to lanceolate or narrow-oblong, the principal ones 3–15 (–19) mm wide, mostly 4–12 times as long as wide, provided with abundant stout, rigid, conical hairs, at least toward the margin of the blade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. umbellatum

Ast e r ac e a e   41 1

13b. Leaf blades lanceolate to oblong-lanceolate or narrow-ovate, the principal ones (7–) 12–40 mm wide, mostly 2–5 times as long as wide, lacking rigid, conical hairs or these very sparse . . . . . . . . . . . . . . . . H. kalmii 7b. Reproductive stems with 0–5 (–10) leaves, these often reduced and bract-like; basal leaves present at anthesis, forming a conspicuous rosette 14a. Leaf blades entire or merely denticulate, adaxially green to glaucous-green and usually with red to purple coloration that closely follows the outline of the veins; ray corollas 7–11 mm long; mid-portion of reproductive stem glabrous or nearly so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. venosum 14b. At least the lower leaf blades with conspicuous, sharp teeth, adaxially green, red coloration, when present, either more randomly patterned (i.e., mottled) or merely suffusing a region of the blade; ray corollas 12–18 mm long; mid-portion of reproductive stem with setose or glandular hairs or both 15a. Basal leaf blades truncate to cordate at the base; reproductive stems with 0–2 leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. murorum 15b. Basal leaf blades tapering to the base; reproductive stems with 2–10 leaves 16a. Leaf blades 7–20 mm wide; peduncles lacking stipitate glands; plants 10–35 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) H. robinsonii 16b. Leaf blades 15–50 mm wide; peduncles stipitate-glandular; plants 30–60 (–100) cm tall 17a. Leaf blades with red to red-purple blotches, streaks, and/or spots [Fig. 425]; stem leaves mostly numbering 2–4; involucres with numerous stipitate glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. maculatum 17b. Leaves blades green, without red to red-purple coloration; stem leaves numbering mostly 2–8; involucres with few to numerous stipitate glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. lachenalii 1. Hieracium aurantiacum L. E orange hawkweed. Pilosella aurantiaca (L.) F.W. Schultz & Schultz-Bip. • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns. 2. Hieracium caespitosum Dumort. E Fig. 422 yellow hawkweed. Hieracium pratense Tausch; Pilosella caespitosa (Dumort.) P.D. Sell & C. West • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, shorelines.

Fig. 422  Cypsela of Hieracium caespitosum.

‌2 × Hieracium lactucella Wallr. Hieracium ×floribundum Wimmer & Grab. is a rare hawkweed hybrid known from CT, MA, ME, NH, RI, VT. It is relatively tall (20–80 cm) and has 3–30 (–50) capitula borne in a compact, corymb-like capitulescence with short peduncles. The leaf blades are glaucous and nearly glabrous adaxially. This hybrid hawkweed most notably differs from H. piloselloides in its early production of prostrate stolons and elongate rhizomes (vs. not producing stolons or these divergent to ascending, but not prostrate, and with short, praemorse rhizomes). 3. Hieracium flagellare Willd. E whip hawkweed. Hieracium flagellare Willd. var. pilosius Lepage; Pilosella flagellaris (Willd.) P.D. Sell. & C. West • CT, MA, ME, NH, VT. Fields, roadsides, lawns. This plant is sometimes treated as the hybrid of Hieracium caespitosum and H. pilosella. Sell and West (1976) point out that it has larger capitula than either parent and it has a widespread distribution with little variation. These facts suggest it is better treated as an orthospecies. 4. Hieracium gronovii L. N Fig. 423 beaked hawkweed. CT, MA, RI. Fields, open woodlands, clearings, usually on dry-mesic soils. Hieracium gronovii and H. scabrum are sometimes confused, especially in flower considering H. scabrum sometimes shows a tall, cylindrical capitulescence (like H. gronovii). The two species

Fig. 423  Cypsela of Hieracium gronovii.

412   tricolpate s

can be separated by flower number per capitulum and peduncle thickness. Hieracium gronovii has 20–40 flowers per capitulum and peduncles 0.3–0.5 (–0.6) mm thick. Hieracium scabrum has 40–100 flowers per capitulum and peduncles (0.4–) 0.5–0.9 mm thick. Reports of this species from me were based on a collection of Hieracium scabrum—1882, Furbish s.n. (nebc!) ‌ × 17. Hieracium ×marianum Willd. is a rare hawkweed hybrid in New England known 4 from CT, MA, NH, VT. It is like H. gronovii in having a leafy stem, usually (2–) 3–6 leaves on the stem, but the capitulescence is corymb-like (rather than cylindrical and panicle-like). This nothospecies is further characterized by a persistent, loose, basal rosette of leaves present at anthesis, the leaf blades green or with a faint tinge of red on the veins, and cypsela bodies sometimes slightly tapered to the apex. 5. Hieracium kalmii L. N Fig. 424 Canada hawkweed. Hieracium canadense Michx.; H. canadense Michx. var. fasciculatum (Pursh) Fern.; H. canadense Michx. var. kalmii (L.) Scoggin, nom. illeg.; H. kalmii L. var. fasciculatum (L.) Lepage; H. umbellatum L. var. canadense (Michx.) Breitung • CT, MA, ME, NH, RI, VT. Fields, sand plains, roadsides, forest edges, cliffs, grasslands, shorelines.

Fig. 424  Cypsela of Hieracium kalmii.

‌5 × 15. Hieracium ×fernaldii Lepage is an extremely rare hawkweed hybrid in New England. In general aspect it resembles H. kalmii but without large leaf blade teeth (which are sometimes present in H. kalmii) and with some of the peduncles of the plant bearing 1 or a few stipitate glands. 6. Hieracium lachenalii K.C. Gmel. E common hawkweed. Hieracium vulgatum Fries • CT, MA, ME; also reported from NH by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, trail edges, waste areas. This species has been attributed to NH, in part, on the basis of Hodgdon 7834 (NHA; image seen!) and Steele 1417 (NHA; image seen!). The former is Hieracium robinsonii and the latter is H. venosum. 7. Hieracium maculatum Sm. E Fig. 425 spotted hawkweed. ME, VT. Fields, roadsides, shorelines, clearings. 8. Hieracium murorum L. E wall hawkweed. CT, MA, ME, NH, VT. Fields, roadsides, lawns, yards. 9. Hieracium paniculatum L. N panicled hawkweed. CT, MA, ME, NH, RI, VT. Mesic to dry-mesic, deciduous forests and woodlands, trail edges.

Fig. 425  Leaf blade of Hieracium maculatum.

10. Hieracium pilosella L. E Fig. 426 mouse-ear hawkweed. Hieracium pilosella L. var. niveum Muell.-Arg.; Pilosella officinarum F.W. Schultz & Schultz-Bip. • CT, MA, ME, NH, RI, VT. Open areas such as lawns, fields, and roadsides. 11. Hieracium piloselloides Vill. E glaucous hawkweed. Hieracium florentinum All.; Pilosella piloselloides (Vill.) Soják; • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, waste areas. 12. Hieracium praealtum Vill. ex Gochnat E tall hawkweed. Hieracium praealtum Vill. ex Gochnat var. decipiens W.D.J. Koch; Pilosella praealta (Vill. ex Gochnat) F.W. Schultz & Schultz-Bip. • CT, MA, ME, NH, VT. Fields, roadsides, railroads, waste areas. 13. Hieracium robinsonii (Zahn) Fern.

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Robinson’s hawkweed. Hieracium ungavense Lepage • ME, NH; northern portion of states. River shore outcrops, usually of high pH. 14. Hieracium sabaudum L. E Savoy hawkweed. CT, MA, ME, VT. Fields, roadsides, vacant lots, trail edges. 15. Hieracium scabrum Michx. N Fig. 427 Fig. 426  Habit of Hieracium pilosella showing stolon.

rough hawkweed. Hieracium scabrum Michx. var. tonsum Fern. & St. John • CT, MA, ME, NH, RI, VT. Woodlands, forest openings, fields, clearings.

Ast e r ac e a e   4 13

16. Hieracium umbellatum L.

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narrow-leaved hawkweed. Hieracium scabriusculum Schwein.; H. scabriusculum Schwein. var. saximontanum Lepage • NH, VT. Forests, logging trails, clearings. 17. Hieracium venosum L. N rattlesnake hawkweed. Hieracium venosum L. var. nudicaule (Michx.) Farw. • CT, MA, ME, NH, RI, VT. Woodlands, clearings.

Hymenoxys 1. Hymenoxys odorata DC. E poison rubberweed. Actinella odorata Gray; Picradenia odorata (DC.) Britt. • ME. Waste areas, yards.

Hypochaeris The fruits of Hypochaeris vary depending on their placement on the capitulum. Those from the margin are shorter and either are short-beaked or lack beaks altogether compared with the inner cypselas, which are longer and have slender, apical beaks. 1a. Leaf blades glabrous or puberulent; involucre 8–10 mm tall during anthesis (elongating to 17 mm in fruit); rays relatively inconspicuous, not much exceeding the involucre (if at all) and ca. 2 times as long as wide; body of inner cypselas 6–8.5 (–13.5) mm long; plants annual from a taproot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. glabra 1b. Leaf blades hispid; involucre 10–15 mm tall during anthesis (elongating to 25 mm in fruit); rays evident, exceeding the involucre and ca. 4 times as long as wide; body of inner cypselas 8–17 mm long; plants perennial from a caudex and fibrous roots (sometimes 1 or more roots thickened as well) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. radicata 1. Hypochaeris glabra L. E smooth cat’s-ear. MA, ME. Fields, road and trail edges, disturbed soil. 2. Hypochaeris radicata L. E hairy cat’s-ear. CT, MA, ME, NH, RI, VT. Fields, road and trail edges, disturbed soil.

Inula 1a. Basal and lower stem leaves 1.2–2 cm broad; rays 10–15 mm long; disk corollas 5–7 mm long; outer involucral bracts 1.5–2.5 mm wide; involucre 8–12 (–20) mm in diameter; body of cypsela 1.5–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. salicina 1b. Basal and lower stem leaves 10–20 cm broad; rays (10–) 20–30 mm long; disk corollas 7–11 mm long; outer involucral bracts 6–8 (–20) mm wide; involucre (20–) 30–40 mm in diameter; body of cypsela 2–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. helenium 1. Inula helenium L. E horse yellowhead. CT, MA, ME, NH, RI, VT. Roadsides, fields, pastures, mostly on dry-mesic to wet-mesic substrate. 2. Inula salicina L. E willow-leaved yellowhead. MA. Roadsides, waste areas. Collected in 1879 near a carpet factory in Danvers.

Ionactis 1. Ionactis linariifolia (L.) Greene N flax-leaved stiff-aster. Aster linariifolius L.; A. linariifolius L. var. victorinii Fern. • CT, MA, ME, NH, RI, VT. Roadsides, dry fields, sand plains, open woodlands, river shore outcrops, open balds.

Fig. 427  Capitula and peduncles of Hieracium scabrum showing stipitate-glands.

414   tricolpate s

Iva 1a. Leaf blades glabrous; axis of capitulescence glabrous; petioles 3–10 mm long; peduncles 1–3 mm long; plants suffrutescent perennials of coastal marshes and shores . . . . I. frutescens 1b. Leaf blades adaxially strigose-scaberulous; axis of capitulescence pubescent; petioles 5–20 (–30) mm long; peduncles 0–1 mm long; plants annual of moist, often disturbed, inland soils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. annua 1. Iva annua L. E rough marsh-elder. Iva ciliata Willd. • MA, ME. Roadsides, dumps, waste areas. 2. Iva frutescens L. N maritime marsh-elder. Iva frutescens L. ssp. oraria (Bartlett) R.C. Jackson; I. frutescens L. var. oraria (Bartlett) Fern. & Grisc. • CT, MA, ME, NH, RI. Saline marshes, most commonly near limit of high tide.

Jacobaea 1. Jacobaea vulgaris Gaertn. E tansy ragwort. Senecio jacobaea L. • MA, ME. Fields, pastures, roadsides.

Krigia 1a. Pappus of 10 inconspicuous, hyaline scales and 20–35 longer, slender bristles; involucre 7–14 mm tall; basal leaf blades 10–50 mm wide, glabrous; plants perennial from a short caudex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. biflora 1b. Pappus of 5 evident, flat scales alternating with 5 longer, slender bristles [Fig. 428]; involucre 3–7 mm tall; basal leaf blades 1–12 mm wide, villous-hirsute or glandular-hirsute to glabrous; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. virginica 1. Krigia biflora Walt. var. biflora

nC

two-flowered dwarf-dandelion. Hyoseris amplexicaulis Michx.; H. biflora Walt.; Krigia amplexicaulis (Michx.) Nutt.; K. biflora Walt. ssp. glandulifera (Fern.) Iltis • CT, ME; also reported from MA by Smith (1899), but specimens are unknown and the given ecology is unusual. Sandy roadsides and fields, woodlands. This species is native to CT and introduced in ME (Portland City, Cumberland County, 1891). 2. Krigia virginica (L.) Willd. N Fig. 428 Virginia dwarf-dandelion. Hyoseris virginica L. • CT, MA, ME, NH, RI, VT. Open woodlands, rock balds, fields, clearings.

Lactuca Fig. 428  Cypsela of Krigia virginica showing dimorphic pappus.

Lactuca floridana (L.) Gaertn. was attributed to New England based on a collection from Eastham, MA (Collins 1774 NEBC!). The specimen is L. biennis with apically pale pappus bristles (the basal half of the pappus bristles, the portion concealed by the involucral bracts, is light brown; L. floridana would have white pappus bristles). Lactuca saligna L. was reported from MA and ME by Strother (2006d), but specimens are unknown. Reference: Strother (2006d). 1a. Basal leaf blades of commonly cultivated forms ovate to orbicular, 12–25 cm wide; plants rare escapes from cultivation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sativa 1b. Basal leaf blades linear to ovate, 0.6–12 cm wide; plants native or introduced, but not generally cultivated 2a. Cypsela apically beakless or with a short, stout beak 0.1–0.5 mm long [Fig. 429], the body compressed but not flat; pappus light brown to light olive-brown; capitula with (15–) 20–30 (–55) flowers; corollas blue to white or infrequently yellow . . . . . . . . . L. biennis

Ast e r ac e a e  4 15

2b. Cypsela apically with a slender, elongate beak 1–6 mm long [Fig. 430], the body ± flat; pappus ± white; capitula with 6–24 (–27) flowers; corollas yellow (sometimes blue in age or drying) 3a. Cypsela body gray or yellow-gray to light brown, with (3–) 5–9 prominent nerves on each face; leaf blades often prickly setose on the abaxial midrib . . . . . . . . . . . L. serriola 3b. Cypsela body brown to dark brown, with 1 prominent nerve on each face, sometimes with an additional pair of faint nerves [Fig. 430]; leaf blades not prickly setose 4a. Involucre 10–15 mm tall in fruit; cypsela body 4.5–6 mm long including the beak; pappus 5–7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. canadensis 4b. Involucre 15–22 mm tall in fruit; cypsela body 7–10 mm long including the beak; pappus 8–12 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. hirsuta 1. Lactuca biennis (Moench) Fern. N Fig. 429 tall blue lettuce. Lactuca spicata (Lam.) A.S. Hitchc. var. integrifolia (Torr. & Gray) Britt.; Mulgedium spicatum (Lam.) Small var. integrifolium (Torr. & Gray) Small; Sonchus biennis Moench • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest edges and clearing

Fig. 429  Cypsela of Lactuca biennis.

‌1 × 2. Lactuca ×morssii B.L. Robins. is a rare lettuce hybrid known from MA, ME. It is intermediate between the parental taxa or sometimes combines characteristics. It usually has blue ray flowers and off-white pappus bristles, the cypsela bodies intermediate or as in each parent (see key for differences in beak length and nerving of faces). 2. Lactuca canadensis L. N Fig. 430 tall lettuce. Lactuca canadensis L. var. integrifolia (Bigelow) Torr. & Gray; L. canadensis L. var. latifolia Kuntze; L. canadensis L. var. longifolia (Michx.) Farw.; L. canadensis L. var. obovata Wieg.; L. sagittifolia Ell. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest edges and clearings. 3. Lactuca hirsuta Muhl. ex Nutt.

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tall hairy lettuce. Lactuca hirsuta Muhl. ex Nutt. var. sanguinea (Bigelow) Fern. • CT, MA, ME, NH, RI, VT. Dry fields, roadsides, forest edges and clearings, woodlands. 4. Lactuca sativa L. E cultivated lettuce. MA, ME. Gardens, compost heaps, fields. Lactuca sativa is further characterized by a yellow corolla, white pappus, involucral bracts erect in fruit, and a flat cypsela body that has 5- to 9-nerved faces, the body terminated by an elongate, filiform beak. 5. Lactuca serriola L. E prickly lettuce. Lactuca scariola L.; L. scariola L. var. integrata Gren. & Godr.; L. scariola L. var. integrifolia (Bogenh.) G. Beck • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas.

Lapsana 1. Lapsana communis L. E common nipplewort. CT, MA, ME, RI, VT. Fields, roadsides, trail edges, disturbed soil.

Lasthenia 1a. Involucre 5–10 mm tall; anthers of disk flowers triangular at apex; styles of disk flowers triangular at apex; pappus of 1–7 awns or awn-tipped scales or absent; leaf blades entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. californica 1b. Involucre 4–6 mm tall; anthers of disk flowers ovate to elliptic at apex; styles of disk flowers triangular to dome-shaped at apex; pappus of 2 or 3 narrow-lanceolate to lanceolate scales intermixed with shorter, truncate, fringed scales or absent; leaf blades entire to toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. minor

Fig. 430  Cypsela of Lactuca canadensis.

416  tricolpate s

1. Lasthenia californica DC. ex Lindl. E California goldfields. Baeria chrysostoma Fisch. & C.A. Mey.; B. chrysostoma Fisch. & C.A. Mey. var. gracilis (DC.) Hall; Lasthenia chrysostoma (Fisch. & C.A. Mey.) Greene • MA. Fields, roadsides, waste areas. 2. Lasthenia minor (DC.) Ornduff E coastal goldfields. Baeria minor (DC.) Ferris • MA. Fields, roadsides, waste areas.

Layia 1. Layia platyglossa (Fisch. & C.A. Mey.) Gray E coastal tidytips. Layia platyglossa (Fisch. & C.A. Mey.) Gray var. breviseta Gray; L. platyglossa (Fisch. & C.A. Mey.) Gray ssp. campestris Keck • MA. Wool waste.

Leontodon Leontodon has been dismantled because it was diphyletic (i.e., the former definition of the genus contained two distinct, evolutionay entities; Samuel et al. 2006). Our common species (L. autumnalis) has been transferred to Scorzoneroides. 1a. Pappus of inner flowers with an inner series of plumose bristles and an outer series of scales that may be tipped with a scabrous awn, that of the outer flowers reduced to a short, laciniate crown; involucre 6–11 mm tall during anthesis, calyculate . . . . . . . . . . . . . . . L. saxatilis 1b. Pappus of inner flowers and outer flowers alike, well-developed, with an inner series of plumose bristles and an outer series of slender scales or barbellate bristles; involucre 10–18 mm tall during anthesis, not calyculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. hispidus 1. Leontodon hispidus L. ssp. hispidus E bristly hawkbit. Apargia hispida (L.) Willd.; Leontodon hastilis L. var. vulgaris W.D.J. Koch • CT, RI. Fields, roadsides, lawns, disturbed soil. 2. Leontodon saxatilis Lam. ssp. saxatilis E little hawkbit. Leontodon leysseri (Wallr.) G. Beck; L. nudicaulis Mérat; L. nudicaulis Mérat ssp. taraxacoides (Vill.) Schinz & Thellung; L. taraxacoides (Vill.) Mérat • CT, MA; also reported from RI and VT by Bogler (2006b), but specimens are unknown. Fields, roadsides, lawns, disturbed soil.

Leucanthemella 1. Leucanthemella serotina (L.) Tzvelev E giant-daisy. Chrysanthemum serotinum L.; Leucanthemum serotinum (L.) Stankov; Tanacetum serotinum (L.) Schultz-Bip. • CT; also reported from MA by Strother (2006e), but specimens are unknown. Gardens, fields, roadsides, vacant lots.

Leucanthemum 1. Leucanthemum vulgare Lam. E ox-eye daisy. Chrysanthemum leucanthemum L.; C. leucanthemum L. var. pinnatifidum Lecoq & Lamotte; Leucanthemum leucanthemum (L.) Rydb.; L. vulgare Lam. var. pinnatifidum (Lecoq & Lamotte) Moldenke • CT, MA, ME, NH, RI, VT; throughout. Fields, roadsides, lawns, logging roads, river beaches. Leucanthemum lacustre (Brot.) Samp. × L. maximum (Ramond) DC. Leucanthemum ‌×superbum (Berg. ex J.W. Ingram) Berg. ex Kent is a rarely collected plant that has escaped from cultivation in ME. It is one species that is commonly referred to as the Shasta-daisy (along with L. maximum). It has toothed leaf blades, capitula mostly 5.1–12.7 cm in diameter, and stems usually 60–90 cm tall (with lobed lower leaf blades, capitula mostly 2.5–5.1 cm in diameter, and stems 10–30 (–100) cm tall in L. vulgare).

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Liatris Reference: Nesom (2006c). 1a. Capitula borne mostly on short peduncles 10–50 mm long, with (21–) 25–80 flowers [Fig. 431]; disk flowers longer than 11 mm, pubescent adaxially (i.e., inside) near the base of the connate tube; involucre 9–17 mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. novae‑angliae 1b. Capitula ± sessile, with 5–10 (–14) flowers; disk flowers 7–11 mm long, glabrous adaxially; involucre 7–11 mm tall 2a. Outer involucral bracts obtuse to rounded at apex, erect; axis of capitulescence usually glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. spicata 2b. Outer involucral bracts acute to short-acuminate at apex, squarrose; axis of capitulescence usually hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. pycnostachya 1. Liatris novae-angliae (Lunell) Shinners var. novae-angliae

N C Fig. 431

northern blazing star. Lacinaria scariosa (L.) Hill var. novae-angliae Lunell; Liatris borealis, auct. non Nutt. ex J. McNab; L. scariosa (L.) Willd. var. novae-angliae (Lunell) Gandhi, S.M. Young, & P. Somers • CT, MA, ME, NH, RI. Woodlands, sandplains, dry fields, sandy sea beaches, roadsides, railroads, limestone outcrops. Liatris novae-angliae has been treated as a variety of L. scariosa by some authors (along with the primarily midwestern and Great Lakes taxon L. nieuwlandii). However, L. novae-angliae has larger capitula with more disk flowers, taller stems, and more leaves per stem than does the L. scariosa. Treating L. novae-angliae as a species distinct from L. scariosa was done by Shinners (1943) and Gaiser (1946), and suggested by Nesom (2006c). 2. Liatris pycnostachya Michx. var. pycnostachya E thick-spiked blazing star. Lacinaria pycnostachya (Michx.) Kuntze • MA, ME. Fields, roadsides, rubbish heaps. 3. Liatris spicata (L.) Willd. var. spicata E sessile-headed blazing star. Lacinaria spicata (L.) Kuntze • CT, MA. Fields, roadsides, waste areas.

Logfia 1. Logfia minima (Sm.) Dumort. E little false cotton-rose. Filago minima (Sm.) Pers.; Gnaphalium minimum Sm.; Oglifa minima (Sm.) Reichenb. • MA. Fields, areas of cultivation.

Madia 1a. Leaves 2–7 × 0.1–0.4 cm; stems stipitate-glandular only near the summit; capitula 2–5 mm wide after pressing, with 0–5 ray flowers; rays 1.5–3 mm long . . . . . . . . . . . . . . . . . M. glomerata 1b. Leaves 4–18 × 0.4–1.2 cm; stems conspicuously stipitate-glandular throughout; capitula 6–12 mm wide after pressing, with 5–13 ray flowers; rays 2–7 mm long . . . . . . . . . . . . . M. sativa 1. Madia glomerata Hook. E mountain tarplant. CT, ME; also reported from VT by Magee and Ahles (1999), but specimens are unknown. Cultivated fields, disturbed soil. 2. Madia sativa Molina E Chile tarplant. Madia capitata Nutt.; M. sativa Molina ssp. capitata (Nutt.) Piper; M. sativa Molina var. congesta Torr. & Gray • CT, MA, VT. Cultivated fields, disturbed soil. Reports of this species from ME (e.g., Kartesz 1999) are based, in part, on a collection taken from a cultivated specimen—27 Aug 1888, unknown s.n. (MAINE!).

Fig. 431  Capitula of Liatris novae-angliae.

418  tricolpates

Matricaria 1a. Capitula without ray flowers [Fig. 432]; disk corollas 4-lobed, green-yellow; pappus a short crown; ultimate segments of leaf blades short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. discoidea 1b. Capitula with conspicuous, white ray flowers; disk corollas 5-lobed, yellow; pappus absent or a short crown; ultimate segments of leaf blades longer . . . . . . . . . . . . . . . . . . M. chamomilla 1. Matricaria chamomilla L. E wild chamomile.  1a. Chamomilla chamomilla (L.) Rydb.; Matricaria chamomilla L. var. coronata J. Gay ex Boiss.; 1b. Chamomilla recutita (L.) Rauschert; Matricaria recutita L. • CT, MA, ME, RI. Fields, roadsides, waste areas. 1a. Pappus of a short crown at the summit of the ovary present on ray flowers and sometimes also on the disk flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. M. chamomilla var. chamomilla 1b. Pappus absent on all flowers . . . . . . . . . . . . . . . . . . 1b. M. chamomilla var. recutita (L.) Fiori Variety chamomilla is known from CT, MA, ME, RI. Variety recutita is known from CT, MA, ME. See Applequist (2002) for nomenclature of this species. 2. Matricaria discoidea DC. E Fig. 432 Fig. 432  Capitulum and upper leaves of Matricaria discoidea.

rayless chamomile. Chamomilla suaveolens (Pursh) Rydb.; Lepidanthus suaveolens (Pursh) Nutt.; Matricaria matricarioides, auct. non (Less.) Porter; M. suaveolens (Pursh) Buch.; Tanacetum suaveolens (Pursh) Hook. • CT, MA, ME, NH, RI, VT. Roadsides, yards, waste areas, disturbed soil.

Microseris 1. Microseris douglasii (DC.) Schultz-Bip. ssp. douglasii E Douglas’ silverpuffs. Calais douglasii DC. • MA. Wool waste.

Mikania 1. Mikania scandens (L.) Willd. N Fig. 433 climbing hempvine. Eupatorium scandens L.; Mikania scandens (L.) Willd. var. pubescens (Nutt.) Torr. & Gray; Willoughbya scandens (L.) Kuntze • CT, MA, ME, NH, RI; restricted to the coastal plain in northern New England. Riparian and lacustrine forests and thickets.

Mulgedium Fig. 433  Capitulescence and leaves of Mikania scandens.

1. Mulgedium pulchellum (Pursh) G. Don in R. Sweet E Russia blue lettuce. Lactuca pulchella (Pursh) DC.; L. tatarica (L.) C.A. Mey. ssp. pulchella (Pursh) Stebbins; L. tatarica (L.) C.A. Mey. var. pulchella (Pursh) Breitung; Sonchus pulchellus Pursh • MA, ME. Waste areas, yards, roadsides.

Mycelis 1. Mycelis muralis (L.) Dumort. E wall-lettuce. Lactuca muralis (L.) Fresen. • MA, ME, NH, VT. Fields, roadsides, disturbed soil.

Nabalus Leaf blade outline has been used for identification despite the fact it is unreliable for our species occurring in temperate forests and woodlands. Specimens based solely on vegetative collections should not be considered as adequately vouchered. Pappus bristles frequently

Ast e r ac e a e   419

break off and the bristle fragments can be found about the involucres. These bristles are sometimes misinterpreted as hairs on the involucres, leading to misidentification. The pappus bristles of Nabalus are minutely barbellate, the involucral bract hairs are smooth. Reference: Bogler (2006c). 1a. Involucral bracts sparsely to densely long-hirsute (caution: sometimes only a few hairs are present on the involucre of N. serpentarius, look near the base of the inner involucral bracts) [Figs. 434, 435] 2a. Lower leaf blades simple, broad-oblanceolate to obovate, the middle and upper blades clasping; capitulescence tall and narrow, raceme-like; rays pink to purple (infrequently white); capitula with (9–) 11–29 flowers [Fig. 434] . . . . . . . . . . . . . . . . . . . . . . . . . N. racemosus 2b. Lower leaf blades pinnately or palmately lobed or simple and then ovate to elliptic, the middle and upper blades short-petioled or sessile, but not clasping; capitulescence broader, corymb- or panicle-like; rays yellow to yellow-white; capitula with 8–14 flowers [Fig. 435] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. serpentarius 1b. Involucral bracts glabrous except for minute, apical cilia [Fig. 436] 3a. Each capitulum with 4–6 inner (i.e., longer) involucral bracts and 5 or 6 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. altissimus 3b. Each capitulum with 7–14 inner involucral bracts and 9–20 flowers [Fig. 436] 4a. Pappus deep red-brown; rays white to pink to lavender . . . . . . . . . . . . . . . . . . N. albus 4b. Pappus light brown to pale yellow-brown; rays green-yellow to yellow-white or white 5a. Rays white; axis of capitulescence and often upper portion of stem villouspuberulent; involucral bracts black-green to black, the dark pigmentation resolvable as minute spots or streaks at high magnification (i.e., 30× and higher); lower leaf blades simple, triangular-sagittate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. boottii 5b. Rays green-yellow to yellow-white; axis of capitulescence and stem glabrous; involucral bracts green to black, provided with minute, white, waxy papillae (view at 15× or higher) or with minute, dark spots in exposed situations; lower leaf blades pinnately or palmately lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. trifoliolatus 1. Nabalus albus (L.) Hook. N white rattlesnake-root. Harpalyce alba (L.) D. Don ex Beck; Nabalus integrifolius Cass.; N. serpentarius (Pursh) Hook. var. integrifolius (Cass.) Britt.; Prenanthes alba L. • CT, MA, ME, NH, VT; also reported from RI by Bogler (2006c), but specimens are unknown. Deciduous forests, woodlands, roadsides. 2. Nabalus altissimus (L.) Hook. N tall rattlesnake-root. Harpalyce altissima (L.) D. Don ex Beck; Nabalus altissimus (L.) Hook. var. hispidulus House; Prenanthes altissima L.; P. altissima L. var. cinnamomea Fern.; P. altissima L. var. hispidula Fern. • CT, MA, ME, NH, RI, VT. Deciduous forests, rarely ascending to high elevation. 3. Nabalus boottii DC.

NC

Boott’s rattlesnake-root. Prenanthes boottii (DC.) Gray • ME, NH, VT. Alpine plateaus and ridges. 4. Nabalus racemosus (Michx.) Hook.

N C Fig. 434

glaucous rattlesnake-root. Harpalyce racemosa (Michx.) D. Don ex Beck; Prenanthes racemosa Michx.; P. racemosa Michx. ssp. multiflora Cronq.; P. racemosa Michx. var. multiflora (Cronq.) Cronq. • ME; northern portion of state. Ice-scoured river shores in high-pH bedrock and/or till regions. ‌ × 6. Nabalus ×mainensis (Gray) Heller is a rare hybrid known from only northern Maine 4 river shores within New England. It ± resembles N. racemosus vegetatively (including the clasping upper stem leaves), but the basal leaf blades are more abruptly contracted to a petiole-like base, and the involucral bracts are glabrous (as in N. trifoliolatus).

Fig. 434  Capitulescence of Nabalus racemosus showing raceme-like appearance and pubescent involucral bracts.

420   tricolpate s

5. Nabalus serpentarius (Pursh) Hook.

N C Fig. 435

lion’s-foot rattlesnake-root. Prenanthes serpentaria Pursh; P. serpentaria Pursh forma simplicifolia Fern. • CT, MA, NH, RI. Woodlands, rocky slopes, cliffs, roadsides, powerline rightsof-way, sandplains, clearings. 6. Nabalus trifoliolatus Cass. N Fig. 436

Fig. 435  Capitula of Nablus serpentarius showing pubescent involucral bracts.

three-leaved rattlesnake-root. Nabalus nanus (Bigelow) DC.; Prenanthes alba L. var. nana Bigelow; P. nana (Bigelow) Torr. ex DC.; P. trifoliolata (Cass.) Fern.; P. trifoliolata (Cass.) Fern. var. nana Bigelow • CT, MA, ME, NH, RI, VT. Forests, woodlands, cliffs, ravines, open mountain tops, coastal headlands and islands. Nabalus nanus has been variously treated as a distinct species or variety with limited distribution in northeastern North America. The dark involucral bracts of N. nanus are not diagnostic; specimens of N. trifoliolatus from exposed habitats (e.g., subalpine areas, downeast coastal islands) also possess them. Plant size has also been used to distinguish taxa, but this is certainly a function of environment, and northern specimens of N. trifoliolatus from Quebec and Labrador intergrade with N. nanus. Milstead (1964) arbitrarily designated alpine plants as N. nanus and dismissed the intermediate material as N. trifoliolatus. Field observations on Katahdin in central Maine show that N. trifoliolatus demonstrates clinal changes in morphology correlated with elevation (i.e., lower elevation plants resemble typical N. trifoliolatus and plants gradually change toward N. nanus with increased elevation). Based on this evidence, N. nanus is here treated as a taxonomic synonym of N. trifoliolatus.

Nipponanthemum 1. Nipponanthemum nipponicum (Franch. ex Maxim.) Kitam. E Fig. 437 Nippon-daisy. Chrysanthemum nipponicum (Franch. ex Maxim.) Spreng.; Leucanthemum nipponicum Franch. ex Maxim. • CT. Fields, roadsides, compost heaps, stream shores. Fig. 436  Capitula of Nablus trifoliolatus showing glabrous involucral bracts.

Oclemena 1a. Rays white or tinged with pink; leaves herbaceous, prominently toothed, with flat margins, the blades 15–60 mm wide, those of the stem below the capitulescence mostly numbering 10–22 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. acuminata 1b. Rays pink to purple; leaves firm, entire or nearly so, often with revolute margins, the blades 2–12 mm wide, those of the stem below the capitulescence numbering 40–75 . . . . O. nemoralis 1. Oclemena acuminata (Michx.) Nesom N Fig. 438 sharp-toothed nodding-aster. Aster acuminatus Michx. • CT, MA, ME, NH, RI, VT. Forests. 1‌ × 2. Oclemena ×blakei (Porter) Nesom is a frequent hybrid where the habitat of the two parental species overlap. It is known from CT, MA, ME, NH, RI, VT. It is recognized by its somewhat firm leaf blades 5–24 mm wide, with small teeth, and numbering 20–45 below the capitulescence. 2. Oclemena nemoralis (Ait.) Greene N

Fig. 437  Capitulum and upper leaves of Nipponanthemum nipponicum.

bog nodding-aster. Aster nemoralis Ait. • CT, MA, ME, NH, RI, VT; mainly in eastern New England. Bogs, fens, mossy lake shores, ditches, and meadows.

Oligoneuron The generic placement of species of Oligoneuron is controversial. Recent arguments to include these species within Solidago (e.g., Semple et al. 1999) are inconclusive given that the data used in the analyses were uniparentally inherited (cpDNA) and Oligoneuron is not nested within Solidago as stated by the authors—it was nested within Solidago only because they considered Oreochrysum parryi (Gray) Rydb. to be part of Solidago (see Nesom 2006d for rationale that Oreochrysum is distinct from Solidago).

Ast e r ac e a e  42 1

1a. Rays white, 5–9 mm long; lower leaf blades elliptic or elliptic-oblong to to broadlanceolate or broad-ovate, 2–10 mm wide; involucral bracts with a single, thickened midrib; many pappus bristles expanded at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. album 1b. Rays yellow, 3–5 mm long; lower leaf blades linear to narrow-oblanceolate, 15–100 mm wide; involucral bracts few-striate [Fig. 439]; pappus bristles slender at the apex . . . . . . . . . O. rigidum 1. Oligoneuron album (Nutt.) Nesom

NC

white flat-topped-goldenrod. Aster ptarmicoides (Nees) Torr. & Gray; Doellingeria ptarmicoides Nees; Inula alba Nutt.; Solidago asteroides Semple; S. ptarmicoides (Nees) Boivin; Unamia alba (Nutt.) Rydb. • CT, MA, NH, VT. Deciduous woodlands, cliffs, river shore outcrops, commonly on soils derived from limestone or trap rock. 2. Oligoneuron rigidum (L.) Small var. rigidum

N C Fig. 439

stiff flat-topped-goldenrod. Solidago rigida L. • CT, MA, RI. Forest borders, banks and plains near the coast, woodlands on high-pH bedrock.

Omalotheca

Fig. 438  Cypsela of Oclemena acuminata showing dimorphic pappus bristles and stipitateglands on ovary.

1a. Pappus bristles distinct and falling separately; principal leaf blades 12–30 mm long; alpine plants 2–10 cm tall, growing at elevation exceeding 1000 m . . . . . . . . . . . . . . . . . . . . . . O. supina 1b. Pappus bristles united at the base and falling together; principal leaf blades 25–51 mm long; boreal plants 10–70 cm tall, growing at elevation less than 1000 m . . . . . . . . O. sylvatica 1. Omalotheca supina (L.) DC.

NC

alpine arctic-cudweed. Gnaphalium supinum L. • ME, NH. Alpine summits, plateaus, and ravines. 2. Omalotheca sylvatica (L.) Schultz-Bip. & F.W. Schultz N Fig. 440 woodland arctic-cudweed. Gnaphalium sylvaticum L. • ME, NH, VT; mainly in northern counties. In dry-mesic to wet-mesic soil of fields, roadsides, logging roads, clearings, and lake shores.

Onopordum 1. Onopordum acanthium L. ssp. acanthium E

Fig. 439  Capitula of Oligoneuron rigidum showing striate involucral bracts.

Scotch cotton-thistle. CT, MA, RI, VT. Fields, roadsides, waste areas.

Packera Packera plattensis (Nutt.) W.A. Weber & A. Löve, a species primarily of the Midwest, Great Lakes region, and southern states, continues to be reported from New England (Magee and Ahles 1999). However, the plants responsible for this report were carefully examined and found to be P. paupercula (Barkley 1962). References: Barkley (1962), Trock (2006). 1a. Plants with stolons and shallow rhizomes, forming colonies; leaf blades decurrent on petioles for some distance; involucral bracts abruptly tapering to an acuminate apex from above the middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. obovata 1b. Plants without stolons, or these short and poorly developed, not forming colonies; leaf blades not or very shortly decurrent on petioles; involucral bracts gradually tapering from base or with parallel margins and gradually tapering from near apex 2a. Basal leaf blades tapering to petiole, usually cuneate at the base; plants of gravels, ledges, and balds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. paupercula 2b. Blade of basal leaves abruptly contracted to the petiole, usually truncate to cordate at the base [Fig. 441]; plants of wet-mesic to hydric soils of fields, low forests, and wetlands 3a. Blade of basal leaves usually 1.75–3.5 times as long as wide, sharply and finely toothed, rounded to subcordate at the base, acute to obtuse at the apex [Fig. 441] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. schweinitziana

Fig. 440  Capitula of Omalotheca sylvatica.

422   tricolpate s

3b. Blade of basal leaves usually 0.75–1.5 (–1.75) times as long as wide, crenate, strongly cordate at the base, rounded at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aurea 1. Packera aurea (L.) A. & D. Löve N golden groundsel. Senecio aureus L.; S. aureus L. var. gracilis (Pursh) Hook.; S. aureus L. var. intercursus Fern.; S. gracilis Pursh • CT, MA, ME, NH, RI, VT. Swamps, wetland margins, wet meadows, stream banks. 2. Packera obovata (Muhl. ex Willd.) W.A. Weber & A. Löve N running groundsel. Senecio obovatus Muhl. ex Willd.; S. obovatus Muhl. ex Willd. var. rotundus Britt.; S. rotundus (Britt.) Small • CT, MA, NH, RI, VT. Rich, rocky forests and woodlands, usually in regions of high-pH bedrock. 3. Packera paupercula (Michx.) W.A. Weber & A. Löve N balsam groundsel. Senecio balsamitae Muhl. ex Willd.; S. gaspensis Greenm.; S. pauperculus Michx.; S. pauperculus Michx. var. balsamitae (Muhl. ex Willd.) Fern.; S. pauperculus Michx. var. praelongus (Greenm.) House • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. River shore outcrops and gravels, woodlands, ridges. 4. Packera schweinitziana (Nutt.) W.A. Weber & A. Löve N Fig. 441 New England groundsel. Senecio robbinsii Oakes ex Rusby; S. schweinitzianus Nutt. • ME, NH, VT. Swamps, wetland margins, wet meadows.

Palafoxia Fig. 441  Leaf of Packera schweinitziana.

1. Palafoxia texana DC. var. texana E Texas palafox. MA. Disturbed soil, river banks.

Parthenium 1a. Plants annual; leaf blades 1- or 2-times pinnatifid . . . . . . . . . . . . . . . . . . . . P. hysterophorus 1b. Plants perennial; leaf blades toothed, or some of them lobed near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. integrifolium 1. Parthenium hysterophorus L. E Santa Maria feverfew. Parthenium lobatum Buckl. • CT, MA. Fields, roadsides, waste areas. 2. Parthenium integrifolium L. var. integrifolium E wild feverfew. CT, MA. Fields, areas of cultivation, railroads.

Perezia 1. Perezia multiflora (Humb. & Bonpl.) ssp. sonchifolia (Baker) Vuill. E perezia. Perezia aletes Macbride; P. sonchifolia Baker • MA. Fields, roadsides, waste areas.

Petasites 1a. Capitula usually with marginal ray flowers, the rays 0.1–6.3 mm long; leaf blades prominently ± palmately lobed with deep sinuses, the sinuses extending more than 50% of the distance from the margin to the leaf midrib [Fig. 442] . . . . . . . . . . . . . . . . . . . . . . P. frigidus

Fig. 442  Leaf of Petasites frigidus.

1b. Capitula without marginal ray flowers [Fig. 443]; leaf blades ± dentate, the lobes absent or obscure, the sinuses (when present) extending much less than 50% of the distance from the margin to the leaf midrib

Ast e r ac e a e   4 23

2a. Disk corollas cream-white; bracts of scape usually green, the one subtending the lowermost branch of the capitulescence 17–40 mm wide; leaf blades with (4–) 5 (–6) pairs of lateral veins emerging along the inner margins of the basal sinus . . . . . . . . . P. japonicus 2b. Disk corollas largely purple; bracts of scape usually purple or purple-tinged, the one subtending the lowermost branch of the capitulescence 5–16 mm wide; leaf blades with 2–5 pairs of lateral veins emerging along the inner margins of the basal sinus . . . . . . P. hybridus 1. Petasites frigidus (L.) Fries var. palmatus (Ait.) Cronq. N Fig. 442 northern sweet-coltsfoot. Nardosmia arctica (Porsild) A. & D. Löve; Petasites arcticus Porsild; P. frigidus (L.) Fries ssp. arcticus (Porsild) Cody; P. frigidus (L.) Fries ssp. palmatus (Ait.) Cody; P. palmatus (Ait.) Gray; Tussilago palmata Ait. • CT, MA, ME, NH, RI, VT. Swamps, fens, seeps, often in regions of high-pH bedrock in southern and/or western New England. 2. Petasites hybridus (L.) P.G. Gaertn., B. Mey., & Scherb. E Fig. 443 butterbur sweet-coltsfoot. Petasites officinalis Moench; P. vulgaris Hill • CT, MA, VT. Roadsides, fields, stream banks.

Fig. 443  Capitulescence of Petasites hybridus.

3. Petasites japonicus (Sieb. & Zucc.) Maxim. E Japanese sweet-coltsfoot. Nardosmia japonica Sieb. & Zucc. • ME. Disturbed soil, thickets, edges of lawns.

Picris 1. Picris hieracioides L. E hawkweed oxtongue. CT, MA, ME, VT; also reported from RI by Hultén and Fries (1986), but specimens are unknown. Roadsides, fields, waste areas.

Pityopsis 1. Pityopsis falcata (Pursh) Nutt. N Fig. 444 sickle-leaved silk-grass. Chrysopsis falcata (Pursh) Ell.; Heterotheca falcata (Pursh) Harms • CT, MA, RI. Dry-mesic to xeric fields, plains, and woodland openings. Fig. 444  Capitulum and leaves of Pityopsis falcata.

Pluchea 1. Pluchea odorata (L.) Cass. var. succulenta (Fern.) Cronq. N sweet-scented camphorweed. Pluchea purpurascens (Sw.) DC. var. succulenta Fern. • CT, MA, NH, RI; also reported from ME by Fernald (1942b), but specimens are unknown.

Saline and brackish marshes.

Polymnia 1. Polymnia canadensis L.

N C Fig. 445

white-flowered leaf-cup. Polymnia canadensis L. var. radiata Gray; P. radiata (Gray) Small • CT, VT. Mesic to dry-mesic forests and woodlands on rocky slopes and ridges, also at cliff bases and among talus slopes, usually in regions of circumneutral to basic bedrock.

Pseudognaphalium Reference: Nesom (2001). 1a. Leaf blades decurrent on the stem as thin wings [Fig. 446], acuminate at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. macounii

Fig. 445  Leaf of Polymnia canadensis.

424   tricolpate s

1b. Leaf blades not decurrent on the stem, obtuse to acute at the apex 2a. Stems usually with persistent, white tomentum, sometimes only sparsely so, appearing gray or gray-green, eglandular or infrequently stipitate-glandular near the base; capitula wth 4–8 bisexual flowers and 38–96 carpellate flowers; plants relatively inodorous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. obtusifolium 2b. Stems rapidly losing most of the white tomentum, appearing green, covered with abundant stipitate glands 0.1–0.2 mm tall; capitula with (7–) 11–20 bisexual flowers and 47–78 carpellate flowers; plants fragrant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. micradenium 1. Pseudognaphalium macounii (Greene) Kartesz N Fig. 446 Macoun’s rabbit-tobacco. Gnaphalium decurrens Ives; G. macounii Greene; Pseudognaphalium viscosum, auct. non (Kunth) W.A. Weber • CT, MA, ME, NH, VT. Fields, roadsides, forest clearings, logging roads. 2. Pseudognaphalium micradenium (Weatherby) Nesom

Fig. 446  Leaf with decurrent base in Pseudognaphalium macounii.

NC

Weatherby’s rabbit-tobacco. Gnaphalium helleri Britt. var. micradenium (Weatherby) Mahler; G. obtusifolium L. var. micradenium Weatherby; Pseudognaphalium helleri (Britt.) A. Anderb. ssp. micradenium (Weatherby) Kartesz • MA, ME, NH. Sandy soils of woodlands, clearings, and roadbeds, usually in association with Pinus rigida and/or Quercus ilicifolia. 3. Pseudognaphalium obtusifolium (L.) Hilliard & Burtt N blunt-leaved rabbit-tobacco. Gnaphalium obtusifolium L.; G. obtusifolium L. var. praecox Fern.; Pseudognaphalium obtusifolium (L.) Hilliard & Burtt ssp. praecox (Fern.) Kartesz • CT, MA, ME, NH, RI, VT. Fields, roadsides, dry banks, clearings.

Ratibida Reference: Richards (1968). 1a. Disk ellipsoid to spherical, 10–25 × 10–18 mm; plants from fibrous roots; pappus usually absent (rarely consisting of 1 or 2 tooth-like scales); style branches stigmatic in the apical ½ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. pinnata 1b. Disk cylindric, 10–50 × 7–12 mm; plants from a taproot; pappus usually consisting of 1 or 2 tooth-like scales (rarely absent); style branches stigmatic in the apical ¼ to ⅓ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. columnifera 1. Ratibida columnifera (Nutt.) Woot. & Standl. E prarie Mexican-hat. Ratibida columnaris (Sims) D. Don; Rudbeckia columnifera Nutt.; Lepachys columnifera (Nutt.) J.F. Macbr. • CT, MA. Fields, roadsides, waste areas. 2. Ratibida pinnata (Vent.) Barnh. E gray-headed Mexican-hat. Rudbeckia pinnata Vent. • CT, MA, VT. Fields, roadsides, forest edges, gardens.

Rudbeckia Reference: Urbatsch and Cox (2006). 1a. Pappus absent; style appendages elongate and pointed; chaff sharply acute to acuminate and hispidulous near the apex; leaf blades toothed to entire, but not lobed 2a. Plants biennial or short-lived perennial, with basal tufts of leaves; leaves basally disposed, decreasing in size upwards, the lower borne on evident petioles . . . . . . . . R. hirta 2b. Plants annual, lacking basal tufts of leaves; leaves chiefly cauline, remaining relatively constant in size until near base of capitulescence, all sessile or subsessile . . . . . . R. bicolor

Ast e r ac e a e   425

1b. Pappus present, represented by a minute crown up to 0.2 mm long; style appendages short and blunt; chaff blunt to acute at apex (tipped by a cusp in R. triloba), glabrous or minutely downy-pubescent near apex; lower leaf blades with 3 or more prominent lobes or entire to toothed in R. fulgida 3a. Leaf blades not lobed; chaff 2.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. fulgida 3b. Lower leaf blades with 3 or more prominent lobes; chaff 4–6.5 mm long 4a. Chaff tipped by a glabrous cusp; rays 1–2 (–2.5) cm long; plants biennial or infrequently short-lived perennial, lacking rhizomes and leafy offshoots . . . . . R. triloba 4b. Chaff obtuse to acute at apex, minutely downy-pubescent near apex; rays 2–6 cm long; plants perennial from rhizomes that produce leafy offshoots 5a. Disk flowers yellow; stem ± glabrous; leaf blade surfaces lacking sessile glands; cypsela body flat, obliquely attached to receptacle . . . . . . . . . . . . . . . . . . . . R. laciniata 5b. Disk flowers dark brown to purple (rarely yellow); stem densely short-pilose, at least in the apical half; leaf blade surfaces with sessile glands; cypsela body equilaterally quadrangular, basally attached to receptacle . . . . . . . . R. subtomentosa 1. Rudbeckia bicolor Nutt. E pine-woods coneflower. MA. Fields, roadsides, waste areas. 2. Rudbeckia fulgida Ait. var. speciosa (Wenderoth) Perdue E showy coneflower. Rudbeckia speciosa Wenderoth • CT; also reported from MA by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, waste areas. 3. Rudbeckia hirta L.

nC

black-eyed coneflower.  3a. Rudbeckia brittonii Small; R. hirta L. var. brittonii (Small) Fern.;  3b. Rudbeckia hirta L. var. lanceolata (Bisch.) Core; R. hirta L. var. serotina (Nutt.) Core; R. serotina Nutt. • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, open woodlands, clearings. 1a. Leaves blades coarsely toothed, the basal ones ovate to rhombic-oval, 2.5–7 cm wide, about 2 times as long as wide, those borne on the stem lance-ovate to ovate or pandurate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. R. hirta var. hirta 1b. Leaf blades entire to finely toothed, the basal ones lanceolate to oblanceolate, 1–2.5 (–5) cm wide, (3–) 4–5 times as long as wide, those borne on the stem linear to oblanceolate or spatulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. R. hirta var. pulcherrima Farw. Vareity hirta is known from MA, VT. It is native and of conservation concern. Variety pulcherrima is non-native and known from CT, MA, ME, NH, RI, VT. 4. Rudbeckia laciniata L. var. laciniata n Fig. 447 green-headed coneflower. 4a. Rudbeckia laciniata L. var. hortensis Bailey • CT, MA, ME, NH, RI, VT. Lake and river shores, riparian forests, swamps. 1a. Lower stem leaves once pinnatifid; chaff 4.4–6.1 mm long; cypsela body 4.2–6 mm long; pappus 0.1–0.7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a. R. laciniata var. laciniata 1b. Lower stem leaves twice pinnatifid; chaff 3.1–4.1 mm long; cypsela body 3.5–4 mm long; pappus 0.7–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. R. laciniata var. bipinnata Perdue Variety laciniata is the common form in New England and is known from CT, MA, ME, NH, RI, VT. A cultivated form of this species, with many disk flowers converted to ray flowers, sometimes escapes. It has been collected in CT, MA, ME, and VT (Seymour 1982). Variety bipinnata is known from CT, MA, NH. 5. Rudbeckia subtomentosa Pursh E sweet coneflower. CT, MA. Fields, roadsides, waste areas. 6. Rudbeckia triloba L. var. triloba E three-lobed coneflower. CT, MA, VT. Fields, roadsides, waste areas.

Fig. 447  Capitulum of Rudbeckia laciniata.

426   tricolpate s

Santolina 1. Santolina chamaecyparissus L. E lavender-cotton. MA. Fields, roadsides, waste areas.

Schkuhria 1. Schkuhria pinnata (Lam.) Kuntze var. pinnata E feathery false threadleaf. Pectis pinnata Lam. • MA. Waste areas, gardens. Strother (2006f) noted that this species has persisted after planting in ME; however, specimens documenting incipient naturalization are unknown.

Sclerolepis 1. Sclerolepis uniflora (Walt.) B.S.P.

N C Fig. 448

sclerolepis. Aethulia uniflora Walt.; Sclerolepis verticillata (Michx.) Cass.; Sparganophorus verticillatus Michx. • MA, nh, RI. Shallow water of lakes, sandy and gravelly lake shores. Some occurrences may be introduced (e.g., Wallum Lake, Worcester County, MA), given that the plants are concentrated near the boat landing.

Scorzoneroides Fig. 448  Capitulum and leaves of Sclerolepis uniflora.

1. Scorzoneroides autumnalis (L.) Moench E fall-dandelion.  1a. Apargia autumnalis (L.) Hoffmann; Leontodon autumnalis L.;  1b. Leontodon autumnalis L. ssp. pratensis (Link) Arcang.; L. autumnalis L. var. pratensis (Link) W.D.J. Koch • CT, MA, ME, NH, RI, VT. Lawns, fields, roadsides, waste areas. 1a. Involucre tomentose-puberulent to glabrous, sometimes with a few, longer, pale to dark hairs; reproductive stems 5–60 cm tall, with (1–) 2–7 capitula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1a. S. autumnalis ssp. autumnalis 1b. Involucre spreading-hirsute with many, long, dark hairs in addition to the underlying tomentose-puberulence; reproductive stems 5–25 cm tall, with 1–3 capitula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. S. autumnalis ssp. pratensis (Link) Holub Subspecies autumnalis is the common form and is known from CT, MA, ME, NH, RI, VT. Subspecies pratensis is known from CT, MA, ME, NH.

Senecio Arnoglossum atriplicifolium (L.) H.E. Robins. was reported from New England based on an erroneously determined collection by Faxon from Middlesex County, MA (GH!). This specimen was, in fact, Senecio suaveolens. Senecio eremophilus Richards. var. macdougalii (Heller) Cronq. was reported from MA by Knowlton and Deane (1924), but specimens are unknown. Senecio pseudoarnica Less. was reported from ME by Gleason and Cronquist (1991), but specimens are unknown. See Pelser et al. (2007) for new circumscription of this genus (i.e., rationale for inclusion of Hasteola and segregation of Jacobaea). Reference: Barkley (2006). 1a. Corollas green-white to yellow-white (rarely pink-tinged); lower leaf blades triangularhastate, sharply toothed; cypsela body 5–7 (–9) mm long . . . . . . . . . . . . . . . . . . . . S. suaveolens 1b. Corollas yellow to orange; leaf blades mostly oblong to obovate in outline, coarsely toothed to 1 or more times pinnately lobed; cypsela body 1.5–2.5 mm long 2a. Leaf blades linear to filiform (sometimes parted into linear-filiform lobes); subshrubs with the taproots forming a woody crown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. spartioides

Ast e r ac e a e  427

2b. Leaf blades oblong or oblanceolate to ovate or obovate, toothed or pinnately lobed to divided [Fig. 449]; annuals with taproots lacking a woody crown 3a. Leaf blades and stems densely and conspicuously glandular-pubescent [Fig. 449]; cypsela body glabrous; shorter, outer series of involucral bracts 2–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. viscosus 3b. Leaf blades and stems pubescent to subglabrous, glandular hairs absent or sparse; cypsela body pubescent; shorter, outer series of involucral bracts 1–2 (–2.5) mm long 4a. Rays present, though inconspicuous; involucral bracts without a dark apex; involucre with about 13 principal involucral bracts . . . . . . . . . . . . . . . . . . . . S. sylvaticus 4b. Rays absent; involucral bracts with a dark apex, this dark red to red-brown, becoming black [Fig. 450]; involucre with about 21 principal involucral bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. vulgaris 1. Senecio spartioides Torr. & Gray E broom-like ragwort. Senecio multicapitatus Greenm. ex Rydb.; S. spartioides Torr. & Gray var. multicapitatus (Greenm. ex Rydb.) Welsh • MA. Gardens, wool waste. 2. Senecio suaveolens (L.) Ell.

nC

Fig. 449  Capitulum and leaves of Senecio viscosus.

false Indian-plantain. Cacalia suaveolens L.; Hasteola suaveolans (L.) Pojark.; Synosma suaveolens (L.) Raf. ex Britt. • CT, MA, RI. River banks and rich, mesic forests, introduced to roadsides to rubbish heaps. This species is native to CT but introduced in MA and likely so in RI. 3. Senecio sylvaticus L. E woodland ragwort. ME; reported from MA by Hultén and Fries (1986), but specimens are unknown. Roadsides, waste areas, clearings, forest edges, gardens, coastal beaches and headlands 4. Senecio viscosus L. E Fig. 449 sticky ragwort. CT, MA, ME, NH, RI, VT. Waste areas, railroads, river beaches, coastal headlands. 5. Senecio vulgaris L. E Fig. 450 common ragwort. CT, MA, ME, NH, RI, VT. Roadsides, fields, areas of cultivation, sidewalks, waste areas.

Sericocarpus Reference: Semple and Leonard (2006).

Fig. 450  Capitula of Senecio vulgaris showing dark-tipped involucral bracts.

1a. Principal leaf blades oblanceolate to obovate, 10–45 mm wide, at least the lower toothed; each capitulum with 9–20 disk flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. asteroides 1b. Principal leaf blades linear to narrow-oblong, 2–12 mm wide, entire; each capitulum with 5–10 disk flowers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. linifolius 1. Sericocarpus asteroides (L.) B.S.P. N Fig. 451 toothed white-topped-aster. Aster asteroides (L.) MacMillan; A. paternus Cronq.; Conyza asteroides L. • CT, MA, ME, NH, RI, VT; limited to southern portions of the northern New England states. Woodlands, sand plains, dry clearings. 2. Sericocarpus linifolius (L.) B.S.P. N narrow-leaved white-topped-aster. Aster solidagineus Michx.; Conyza linifolia L. • CT, MA, NH, RI; restricted to southern NH. Woodlands, dry fields and clearings.

Sigesbeckia 1. Sigesbeckia orientalis L. E common St. Paul’s-wort. MA. Waste areas, dumps.

Fig. 451  Capitula of Sericocarpus asteroides showing few ray flowers and squarrose involucral bracts.

428   tricolpate s

Silphium 1. Silphium perfoliatum L. var. perfoliatum E cup-plant rosinweed. CT, MA, ME, VT. Ditches, thickets, trail edges, waste areas.

Silybum 1. Silybum marianum (L.) Gaertn. E blessed milk-thistle. Carduus marianus L.; Mariana mariana (L.) Hill • CT, MA, NH. Roadsides, waste areas, gardens.

Solidago Pubescence of the cypsela body can often be assessed by examining the ovary while plants are in flower. Given that the color of ray flowers sometimes fades on herbarium specimens with age, some collections of Solidago can be difficult to distinguish from small-headed Symphyotrichum. This confusion can be solved by noting the fresh ray flower color on herbarium labels. Hybridization is reported between closely related species but does not appear to be a common event on the New England landscape. References: Semple et al. (1999), Semple and Cook (2006). 1a. Capitulescence nodding at the summit and/or with branches that have secund capitula [Fig. 452] 2a. Leaves basally disposed (i.e., leaves progressively reduced upward, those of the apical portion of the stem smaller, often of different shape, and less petiolate than those of the basal portion) [Fig. 455]; plants usually with basal tufts of leaves 3a. Stems and often the leaves closely and minutely pubescent [Fig. 456]; rays pale yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nemoralis 3b. Stems and leaves glabrous or with long, scattered hairs or scabrous; rays bright yellow 4a. Leaf blades fleshy, entire; capitula with 7–17 ray flowers; plants of Atlantic coast shorelines and saline marshes (inland along heavily salted roadways) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sempervirens 4b. Leaf blades not fleshy, serrate to subentire; capitula with 1–12 ray flowers; plants not of Atlantic coast shores 5a. Leaf blades adaxially strongly scabrous; stems prominently angled, each angle often with a narrow wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. patula 5b. Leaf blades glabrous, pubescent, or slightly scabrous adaxially; stems terete or striate, but without prominent angles 6a. Lower leaf blades abruptly contracted to a petiole 7a. Abaxial surface of leaf blades hirsute; capitulum with 4–7 disk flowers; cypsela body minutely pubescent . . . . . . . . . . . . . . . . . . . . (in part) S. ulmifolia 7b. Abaxial surface of leaf blades glabrous; capitulum with 8–20 disk flowers; cypsela body glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. arguta 6b. Lower leaf blades gradually tapered to the base 8a. Lower leaves conspicuously sheathing, the petiole covering 50–75% of the circumference of the stem; capitulescence taller than wide; capitula with 1–8 ray flowers and 4–8 disk flowers; plants primarily of organic soil wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. uliginosa

Ast e r ac e a e   42 9

8b. Lower leaves not conspicuously sheathing, the petiole covering less than 50% of the circumference of the stem; capitulescence as wide as or wider than tall; capitula with 7–12 ray flowers and 9–14 disk flowers; plants of open or lightly shaded, mineral soils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. juncea 2b. Leaves chiefly cauline (i.e., leaves of the apical portion of the stem of nearly similar shape and not dramatically reduced in size relative to the leaves of the basal portion of the stem) [Fig. 454]; plants lacking tufts of basal leave 9a. Leaf blades triple-veined (i.e., with 3 conspicuous, parallel veins—a midrib and 2 evident and prolonged, lateral veins) [Fig. 453] 10a. Stem glabrous and glaucous below the capitulescence; involucral bracts obtuse to acute at the apex, green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. gigantea 10b. Stem pubescent in at least the apical half, not glaucous; involucral bracts acuminate at the apex, yellow-green 11a. Leaf blades abaxially subglabrous or pubescent only on the midrib and major veins (also pubescent between the veins in var. hargeri), usually sharply serrate and relatively thin; involucre 2–3 mm tall; disk corollas 2.3–2.7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. canadensis 11b. Leaf blades abaxially pubescent on and between major veins, usually subentire to remotely serrate and relatively firm; involucre (2.7–) 3–4 (–5) mm tall; disk corollas 3–3.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. altissima 9b. Leaf blades pinnately veined (i.e., with much-branched lateral veins that are not aligned parallel to the midrib) [Fig. 457] 12a. Leaf blades translucent-punctate, all entire, usually anise-scented when bruised . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. odora 12b. Leaf blades not translucent-punctate, some with teeth, not fragrant when bruised 13a. Plants from a caudex, without creeping rhizomes; lower leaf blades abruptly tapered to a petiole; capitula with 3–6 ray flowers . . . . . . . . . . (in part) S. ulmifolia 13b. Plants with long, creeping rhizomes; leaf blades sessile or nearly so; capitula with 6–11 (–12) ray flowers 14a. Leaf blades abaxially pubescent, the hairs commonly found on the tertiary veins (as well as the midvein and primary lateral veins); stems conspicuously spreading-pubescent (varying to sparsely pubescent in ssp. aspera); widespread plants found throughout most of New England . . . . . . . . . . . . . . . . . . . . S. rugosa 14b. Leaf blades ± glabrous abaxially, hairs, when present, few in number and confined to the midrib and primary lateral veins; stems glabrous or with pubescent lines of decurrence from the leaf blades; plants mainly of southern New England and the coastal plain 15a. Involucre 2.5–4 mm tall; involucral bracts narrow-lanceolate to narrowoblong, mostly 0.3–0.6 mm wide, acute at the apex; disk corollas 2.5–3.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. aestivalis 15b. Involucre 4–6 mm tall; involucral bracts oblong to narrow-ovate, the larger 0.7–1.2 mm wide, obtuse to rounded at the apex; disk corollas 4–5.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. latissimifolia 1b. Capitulescence neither nodding at the summit nor with secund capitula [Fig. 458], either terminal and resembling a thryse or panicle or consisting of clusters of capitula in the axils of well-developed leaves 16a. Leaves chiefly cauline (i.e., leaves of the apical portion of the stem of nearly similar shape and not dramatically reduced in size relative to the leaves of the basal portion of the stem) [Fig. 454]; plants lacking tufts of basal leaves

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17a. Capitula with 30–50 flowers, the ray flowers numbering 10–13; leaves of the stem below the capitulescence numbering 2–4 (–7); plants 5–35 cm tall, of alpine summits and plateaus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. leiocarpa 17b. Capitula with 6–13 flowers, the ray flowers numbering (1–) 3–5; leaves of the stem below the capitulescence numbering 7–60; plants 30–150 cm tall, primarily of temperate forests, woodlands, edges, and fields 18a. Leaf blades entire to obscurely toothed; cypsela body glabrous; capitulescence terminal, only the lower branches subtended by leaves [Fig. 458] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. speciosa 18b. At least the lower leaf blades with prominent teeth; cypsela body pubescent; capitulescence partly or entirely axillary, only the upper portion (if any) not subtended by leaves 19a. Leaf blades elliptic to ovate, 3–10 cm wide, 1.2–2.2 (–2.5) times as long as wide; stem angled, not glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. flexicaulis 19b. Leaf blades lanceolate to narrow-elliptic, 1–3 cm wide, 3–10 times as long as wide; stem terete, glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. caesia 16b. Leaves basally disposed (i.e., leaves progressively reduced upward, those of the apical portion of the stem smaller, often of different shape, and less petiolate than those of the basal portion) [Fig. 455]; plants usually with basal tufts of leaves 20a. Involucre 8–11 mm tall, composed of acuminate- to attenuate-tipped involucral bracts; cypsela body 4–5 mm long; lower leaf blades with an acuminate apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. macrophylla 20b. Involucre 3–9 mm tall, composed of round- to acuminate-tipped involucral bracts; cypsela body shorter than 4 mm long; lower leaf blades usually with an obtuse to acute apex 21a. Stem and often the leaves pubescent with minute, viscidulous hairs [Fig. 456]; involucral bracts narrow, acuminate at the apex, up to 0.5 (–0.75) mm wide at the midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. puberula 21b. Stem and leaves glabrous or pilose, but not copiously pubescent with minute hairs; involucral bracts wider, rounded to acute at the apex, (0.75–) 1–2 mm wide at the midpoint 22a. Leaves conspicuously sheathing, the petioles covering 50–75% of the circumference of the stem; plants primarily of organic soil wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. uliginosa 22b. Leaves not conspicuously sheathing, the petiole covering less than 50% of the circumference of the stem; plants not of organic soil wetlands 23a. Involucral bracts, especially the outer, with squarrose tips [Fig. 459]; involucre 5–9 mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. squarrosa 23b. Involucral bracts appressed to ascending; involucre 3–6 mm tall (up to 9 mm tall in S. leiocarpa) 24a. Leaf blades pilose on one or both surfaces; stems often pilose 25a. Ray flowers white or yellow-white; involucre 3–5 mm long; involucral bracts white-brown or light yellow-brown to light green or white-green at the base, green at the apex, the colors often sharply contrasting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. bicolor 25b. Rays yellow; involucre 4–6 mm long; involucral bracts light green or yellow-green at the base, green at the apex, the colors usually weakly contrasting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hispida

Ast e r ac e a e  43 1

24b. Leaf blades glabrous or nearly so (sometimes scabrous); stems essentially glabrous (though often pubescent in the capitulescence) 26a. Cypsela body glabrous; capitula with 5–9 ray flowers; plants of temperate woodlands, edges, and fields 27a. Capitulescence panicle-like, usually with several ascending branches [Fig. 458]; leaf blades entire or sometimes the lower ones slightly toothed, the middle blades (1.5–) 2–7 cm wide; cypsela body 1.1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. speciosa 27b. Capitulescence thyrse-like, very slender (infrequently with 1 or more prolonged branches); lower leaf blades crenate to crenateserrate, the middle blades 0.5–2 (–2.5) cm wide; cypsela body 2.2–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. erecta 26b. Cypsela body pubescent; capitula with 8–13 ray flowers; plants of mountains, cliffs, and northern river shores 28a. Involucral bracts oblong to lanceolate, not glutinous, the middle ones 1.2–2 mm wide, green in large part; leaves of the stem below the capitulescence numbering 2–4 (–7) . . . . . . . . . . . . . (in part) S. leiocarpa 28b. Involucral bracts linear to narrow-lanceolate, glutinous, the surface shiny as if covered with varnish, the middle ones 0.7–1.2 mm wide, with chartaceous base and sides, green only at the apex and along the central band; leaves of the stem below the capitulescence numbering 5–40 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. simplex 1. Solidago aestivalis Bickn. N swamp wrinkle-leaved goldenrod. Solidago rugosa P. Mill. var. sphagnophila Graves • CT, MA, ME, NH, RI. Swamps, forests and forest edges, and clearings along the coastal plain.

This species tends to flower 20–35 days earlier than Solidago rugosa when the two species grow sympatrically. 2. Solidago altissima L. ssp. altissima N Fig. 452 tall goldenrod. Solidago altissima L. var. procera (Ait.) Fern.; S. canadensis L. var. scabra Torr. & Gray • CT, MA, ME, NH, RI, VT; confined primarily to the coastal plain in ME. Fields, open rights-of-way, roadsides. Solidago lepida DC. was reported from ME based on a number of specimens, mostly from near coastal habitats, that were S. altissima.

Fig. 452  Capitulescence of Solidago altissima showing branches with secund capitula.

3. Solidago arguta Ait. var. arguta N forest goldenrod. CT, MA, ME, NH, RI, VT. Deciduous forests and woodlands, forest edges, clearings. 4. Solidago bicolor L. N white goldenrod. CT, MA, ME, NH, RI, VT. Fields, roadsides, woodlands, disturbed open areas. 5. Solidago caesia L. var. caesia N axillary goldenrod. Solidago axillaris Pursh; S. caesia L. var. axillaris (Pursh) Gray • CT, MA, ME, NH, RI, VT. Mesic, deciduous forests and woodlands, forest borders. 6. Solidago canadensis L. N Fig. 453 Canada goldenrod. CT, MA, ME, NH, RI, VT. Fields, roadsides, open rights-of-way, clearings. 1a. Leaf blades abaxially pubescent on and between the veins, adaxially pubescent or scabrous; stems generally pubescent to the base . . . . . . . . . . . . . . . . . 6a. S. canadensis var. hargeri Fern. 1b. Leaf blades abaxially subglabrous or pubescent only on the midvein and lateral veins, adaxially glabrous or scabrous; stems generally pubescent only in the apical half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6b. S. canadensis var. canadensis Variety hargeri is known from CT, MA, ME, NH, VT. Vareity canadensis is known from CT, MA, ME, NH, RI, VT.

Fig. 453  Triple-veined leaf blade of Solidago canadensis.

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‌ × 14. Solidago ×calcicola Fern. is a rare hybrid known only from northern New 6 England (ME, NH). It demonstrates triple-veined leaf blades (as in S. canadensis), but the capitula are neither secund nor are they borne on nodding or recurved branches (in this trait the hybrid is similar to S. macrophylla). The stems arise from a stout caudex in this nothospecies (rather than elongate rhizomes as in S. canadensis). 7. Solidago erecta Pursh

NC

slender goldenrod. Solidago speciosa Nutt. var. erecta (Pursh) MacM. • CT, MA, RI. Woodlands, roadsides, dry banks. 8. Solidago flexicaulis L. N zig-zag goldenrod. Solidago latifolia L. • CT, MA, ME, NH, RI, VT. Mesic, deciduous forests, stream banks. 9. Solidago gigantea Ait. N Fig. 454 smooth goldenrod. Solidago gigantea Ait. ssp. serotina (Kuntze) McNeill; S. gigantea Ait. var. serotina (Kuntze) Cronq. • CT, MA, ME, NH, RI, VT. Fields, roadsides, riparian forests. Fig. 454  Chiefly cauline leaves of Solidago gigantea.

10. Solidago hispida Muhl. ex Willd. var. hispida N hairy goldenrod. Solidago bicolor L. var. concolor Torr. & Gray; S. bicolor L. var. lanata (Hook.) Seymour; S. hispida Muhl. ex Willd. var. lanata (Hook.) Fern. • CT, MA, ME, NH, RI, VT; mainly in western counties in southern New England states. Ledges, ravines, river shore outcrops, woodlands, often in regions of high-pH bedrock. 11. Solidago juncea Ait. N Fig. 455 early goldenrod. Solidago juncea Ait. var. neobohemica Fern.; S. juncea Ait. var. ramosa Porter & Britt. • CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings, woodlands. 12. Solidago latissimifolia P. Mill. N Elliott’s goldenrod. Solidago elliottii Torr. & Gray; S. elliottii Torr. & Gray var. ascendens Fern.; S. elliottii Torr. & Gray var. pedicellata Fern. • CT, MA, RI. Swamps, thickets, and meadows on the coastal plain. Long known by the name Solidago elliottii in New England literature, the name S. latissimifolia pre-dates it by 73 years (1768 vs. 1841). 13. Solidago leiocarpa DC.

NC

Cutler’s goldenrod. Solidago cutleri Fern.; S. virgaurea L. var. alpina Bigelow • ME, NH, VT. Alpine ridges, plateaus, and ravines. 14. Solidago macrophylla Pursh N Fig. 455  Basally disposed leaves of Solidago juncea.

large-leaved goldenrod. Solidago macrophylla Pursh var. thyrsoidea (E. Mey.) Fern. • MA, ME, NH, VT; restricted to western portion of state in MA. North-temperate, boreal, and subalpine forests, alpine ravines and plateaus. 14 × 19b. This very rare goldenrod hybrid is known from VT. It closely resembles Solidago × ‌calcicola (6 × 14) except that the leaf blades are not triple-nerved (the morphological similarity is evidenced by the fact the collections were initially ‌calcicola determined as S. × by Fernald; specimens at GH!, NEBC!, VT!). 15. Solidago nemoralis Ait. ssp. nemoralis N gray goldenrod. Solidago nemoralis Ait. ssp. haleana (Fern.) G.W. Douglas; S. nemoralis Ait. var. haleana Fern. • CT, MA, ME, NH, RI, VT. Dry fields, ledges, roadsides. 16. Solidago odora Ait. ssp. odora N licorice goldenrod. Solidago suaveolens Schoepf • CT, MA, NH, RI, VT; southern portion of northern New England states. Woodlands, dry fields, roadsides. 17. Solidago patula Muhl. ex Willd. var. patula N rough-leaved goldenrod. CT, MA, NH, VT; western MA and VT, disjunct in Carroll County, NH. Swamps, riparian forests.

Fig. 456  Minute pubescence on stem of Solidago puberula.

18. Solidago puberula Nutt. var. puberula N Fig. 456 downy goldenrod. CT, MA, ME, NH, RI, VT. Fields, sand plains, roadsides, woodlands, clearings.

Ast e r ac e a e   43 3

19. Solidago rugosa P. Mill. N Fig. 457 common wrinkle-leaved goldenrod.  19a. Solidago aspera Ait.; S. celtidifolia Small; S. rugosa P. Mill. var. celtidifolia (Small) Fern.; 19b. Solidago rugosa P. Mill. var. villosa (Pursh) Fern. • CT, MA, ME, NH, RI, VT. Fields, roadsides, swamps, edges of wetlands. 1a. Leaf blades thick and firm, low-serrate or crenate to subentire, subacute to shortacuminate at the apex; involucral bracts usually obtuse or rounded at apex (the innermost sometimes acute); capitula with (5–) 6–8 (–10) ray flowers; plants with relatively shorter and stiffer pubescence, occurring in relatively drier habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19a. S. rugosa ssp. aspera (Ait.) Cronq. var. aspera (Ait.) Fern. 1b. Leaf blades thin to moderately thick, usually sharply serrate, short-acuminate to acuminate at the apex; involucral bracts subacute to acute at apex; capitula with (6–) 8–11 (–13) ray flowers; plants with relatively longer and softer pubescence, occurring in relatively wetter habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19b. S. rugosa ssp. rugosa Subspecies aspera is known from CT, MA, ME, NH, RI. Subspecies rugosa is known from CT, MA, ME, NH, RI, VT. It is much more common on the New England landscape than ssp. aspera. ‌ × 20. Solidago ×asperula Desf. is a rare goldenrod hybrid known from CT, MA, ME, NH. 19 It is known from coastal states where S. sempervirens comes into contact with S. rugosa at the upper limit of coastal marshes and beaches. It can be separated from the former by pubescent stems (at least on the upper portion) and from the latter by leaf blades with shorter teeth and less rugose veins. It can resemble S. latissimifolia, but the hybrid shows more hairs on the stems and ± sheathing petiole bases on lower leaves. 20. Solidago sempervirens L. n seaside goldenrod. 20a. Solidago mexicana L. • CT, MA, ME, NH, RI. Coastal marshes, beaches, and dunes, rarely inland along heavily salted roads and salt storage areas. This species is native to coastal areas, but has been collected from some interior counties (e.g., Berkshire and Worcester County, MA), where it should be considered non-native. 1a. Involucres 3–4 mm tall; capitula with 7–11 ray flowers and 10–16 disk flowers; lower leaf blades linear to oblong-lanceolate, 10–30 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20a. S. sempervirens var. mexicana (L.) Fern. 1b. Involucres 4–7 mm tall; capitula with 12–17 ray flowers and 17–22 disk flowers; lower leaf blades oblanceolate to narrow-ovate, 12–50 (–60) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20b. S. sempervirens var. sempervirens Variety sempervirens is known from CT, MA, ME, nh, RI. It is a relatively more northern race. Variety mexicana is known from CT, MA, RI. It is relatively more southern and reaches its northern limit in southern New England. Involucre measurements need to be made on mature capitula with open flowers (or later stages of development). 21. Solidago simplex Kunth. ssp. randii (Porter) Ringius N Rand’s goldenrod.  21a. Solidago glutinosa Nutt. var. racemosa (Greene) Cronq.; S. racemosa Greene; S. spathulata DC. var. racemosa (Greene) Gleason; 21b. Solidago glutinosa Nutt. ssp. randii (Porter) Cronq.; S. randii (Porter) Britt.; S. randii (Porter) Britt. var. monticola (Porter) Fern.; S. spathulata DC. ssp. randii (Porter) Gleason • MA, ME, NH, VT; western portion of MA. Cliffs, open and/or rocky summits, river shore outcrops. 1a. Basal leaves mostly 7–10 times as long as wide, often subentire, with a slender midrib 0.1–0.2 mm wide; capitulescence more open when well-developed, with longer peduncles 5–15 mm long; plants of river shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21a. S. simplex ssp. randii var. racemosa (Greene) Ringius 1b. Basal leaves mostly 3–8 times as long as wide, often sharply toothed, with a broad midrib 0.7–1 mm wide; capitulescence slender and compact, with shorter peduncles (0–) 1–4 mm long; plants of cliffs, summits, and alpine areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21b. S. simplex ssp. randii var. monticola (Porter) Ringius Variety racemosa is known from ME, NH, VT. Variety monticola is known from MA, ME, NH, VT.

Fig. 457  Pinnately veined leaf blade of Solidago rugosa.

434   tricolpate s

22. Solidago speciosa Nutt. var. speciosa N Fig. 458 showy goldenrod. Solidago conferta P. Mill. • CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings. The collection from ME is anomalous in that it has pubescent ovaries. 23. Solidago squarrosa Muhl. N Fig. 459 squarrose goldenrod. CT, MA, ME, NH, VT; restricted to western part of CT. Roadsides, forest edges, open banks, fields and clearings, trail edges. 24. Solidago uliginosa Nutt. N bog goldenrod. S. neglecta Torr. & Gray; S. purshii Porter; Solidago uliginosa Nutt. var. linoides (Torr. & Gray) Fern.; S. uliginosa Nutt. var. neglecta (Torr. & Gray) Fern.; S. uliginosa Nutt. var. terrae-novae (Torr. & Gray) Fern.; S. uniligulata (DC.) Porter • CT, MA, ME, NH, RI, VT. Bogs, fens, peaty meadows, fen-like river shore seeps. Numerous names have been applied to the various forms of this species, most of these apparently phenotypic responses to different natural communities and latitudes. Fig. 458  Capitulescence of Solidago speciosa showing branches with spirally arranged capitula.

25. Solidago ulmifolia Muhl. ex Willd. var. ulmifolia N elm-leaved goldenrod. CT, MA, ME, RI, VT. Woodlands, dry fields and clearings, forest edges. Reports of this species from NH (e.g., Magee and Ahles 1999) were based on misidentified collections. The Hillsborough County voucher was Solidago rugosa ssp. rugosa—30 Aug 1904, Batchelder s.n. (NHA!). The Strafford County voucher was S. arguta var. arguta—Hodgdon & Woodward 5397 (NHA!).

Sonchus Reference: Hyatt (2006). 1a. Plants perennial, with deep-seated, creeping rhizomes; capitula 3–5 cm wide in flower; involucre 14–22 mm tall in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. arvensis 1b. Plants annual, with taproots; capitula 1.5–2.5 cm wide in flower; involucre 9–13 mm tall in fruit Fig. 459  Involucre of Solidago squarrosa showing squarrose bracts.

2a. Cypsela body with 3 (–5) ribs on each face, otherwise smooth; leaf blades relatively firm, with stiff prickles and rounded basal auricles [Fig. 460]; peduncles glandularpubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. asper 2b. Cypsela body with 5–7 ribs on each face and transversely rugulose; leaf blades relatively soft, with softer prickles and triangular to narrow-triangular basal auricles; peduncles glabrous or infrequently with a few glandular hairs . . . . . . . . . . . . . . S. oleraceus 1. Sonchus arvensis L. E field sow-thistle.  1a. Sonchus arvensis L. ssp. uliginosus (Bieb.) Nyman; S. uliginosus Bieb. • CT, MA, ME, NH, RI, VT. Fields, roadsides, pastures, waste areas. 1a. Peduncles and involucres glabrous (though sometimes with sessile yellow glands on the involucre); longer involucral bracts 10–15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. S. arvensis var. glabrescens Guenth., Grab., & Wimmer 1b. Peduncles and involucres pubescent with yellow, glandular-hairs; longer involucral bracts 14–17 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. S. arvensis var. arvensis Variety glabrescens is known from CT, MA, ME, NH, VT. It is less common than typical variety. Variety arvensis is known from CT, MA, ME, NH, RI, VT. 2. Sonchus asper (L.) Hill E Fig. 460 spiny-leaved sow-thistle. Sonchus oleraceus L. var. asper L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, pastures, waste areas.

Fig. 460  Leaf base of Sonchus asper showing rounded auricle.

3. Sonchus oleraceus L. E common sow-thistle. CT, MA, ME, NH, RI, VT. Fields, roadsides, pastures, waste areas.

Ast e r ac e a e   43 5

Symphyotrichum Though Symphyotrichum is generally regarded as an extremely difficult genus, the vast majority of New England collections are straightforward to assign to species. Once learning the large number of taxa is overcome, infrequent hybridization and rare aberrant forms are some of the few stumbling blocks that botanists will face here in New England. Identification necessitates familiarity with details of the involucral bracts and disk flowers (and the need for magnification to view the details, generally 14× is sufficient for most people). Many hybrid combinations have been reported from New England, but more work is needed to confirm some of these. It is clear that some collections determined as hybrids are not identified ‌gravesii correctly and merely represent atypical forms of species. Symphyotrichum × (Burgess) Nesom is a poorly understood taxon that is believed to be a hybrid involving S. dumosum and S. laeve, though S. cordifolium may be a parent (rather than S. dumosum). More work is needed to understand the origin of this plant from CT. References: Semple et al. (2002), Brouillet et al. (2006). 1a. Annuals from a short taproot; rays very short and inconspicuous, scarcely or not exceeding the mature pappus, or the rays absent altogether [Fig. 461] 2a. Involucral bracts with a chartaceous base and chlorophyllous tip, of several conspicuously different lengths; ray flower corolla longer than the style; saltmarsh plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. subulatum 2b. Involucral bracts, especially the outer, herbaceous, of nearly equal length; ray flower corolla shorter than the style, the style protruding from the corolla; rare introductions in New England 3a. Ray flowers numbering 90–110 or more per capitulum, with a short, but evident, strap-like ray 1.5–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. frondosum 3b. Ray flowers numbering 75–95 (rarely more), lacking rays altogether (note: these flowers are still recognizable as a ray flower due to the lack of lobes at the summit of the corolla tube) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. ciliatum 1b. Perennials from a rhizome, caudex, or crown; rays elongate and conspicuous 4a. Basal leaves both cordate and borne on a petiole (often merely truncate or broadrounded at base of the petiolate blade in S. lowrieanum) [Fig. 469] 5a. Middle and upper stem leaves sessile and conspicuously clasping the stem; leaves usually with low, crenate teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. undulatum 5b. Middle and upper stem leaves petiolate or sessile, generally narrowed to the base and not conspicuously clasping; leaves mostly with prominent, sharp teeth 6a. Peduncles with few or no bracts, very uneven, some generally exceeding 1 cm; rays 7–15 mm long; involucral bracts usually green at apex, the outer 0.5–1 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. ciliolatum 6b. Peduncles copiously bracteate, especially when the peduncles approach or exceed 1 cm in length; rays 4–8 mm long; involucral bracts often with the chlorophyllous tip partly or wholly obscured by anthocyanin, the outer mostly 0.2–0.7 mm wide 7a. Involucral bracts acuminate, with a slender, elongate chlorophyllous zone; branches of capitulescence often strongly ascending . . . . . . . . . . . . S. urophyllum 7b. Involucral bracts obtuse to narrow-acute, with an apically dilated chlorophyllous zone [Fig. 464]; branches of capitulescence spreading to ascending 8a. Petioles scarcely or not at all winged; lower leaves deeply cordate with prominent basal sinuses; leaf blades usually pubescent, not glaucous on the abaxial surface, smooth to scabrous on the adaxial surface . . . . S. cordifolium

436   tricolpate s

8b. Petioles broadly winged; lower leaves usually rounded or truncate to shallowly cordate; leaf blades glabrous, thinly glaucous on the abaxial surface, very smooth on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . S. lowrieanum 4b. Basal leaves not both cordate and borne on a petiole 9a. Leaf blades auriculate- or cordate-clasping 10a. Involucral bracts stipitate-glandular [Fig. 465] 11a. Capitula with 45–100 ray flowers, red-purple; stem leaves abundant and crowded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. novae‑angliae 11b. Capitula with 15–30 ray flowers, commonly blue; stems with remote to approximate, but not crowded, leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. patens 10b. Involucral bracts eglandular [Figs. 462, 464] 12a. Involucral bracts long-acuminate to attenuate at the apex, of nearly equal length; at least the lower portion of the stem usually hispid pubescent with stiff, spreading hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. puniceum 12b. Involucral bracts mostly obtuse to acute at the apex, often of different lengths; stem glabrous to pubescent, when pubescent the hairs often softer and/ or curving toward stem apex 13a. Most or all of the stem leaves conspicuously narrowed to a broadly winged and conspicuously clasping petiolar base; stem often notably flexuous, changing angles at successive nodes . . . . . . . . . . . . . . . . . . . . . S. prenanthoides 13b. Stem leaves not conspicuously narrowed nor with a prominently clasping base; stems straight or weakly flexuous 14a. Stem and leaves below the capitulescence glabrous and glaucous; involucral bracts with a short, rhombic chlorophyllous zone; plants from a short, stout rhizome or woody caudex . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. laeve 14b. Stem and leaves glabrous to pubescent, not glaucous; involucral bracts with an elongate chlorophyllous zone, or the outer involucral bracts entirely green; plants from elongate rhizomes 15a. Leaf blades narrow-lanceolate to oblanceolate, narrowing to the base, flat, herbaceous to fleshy; most peduncles shorter than 3 cm; involucral bracts often loose or somewhat spreading apically, commonly green in large part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. novi‑belgii 15b. Leaf blades tending to be narrow-lanceolate, scarcely narrowing to the base, folded along the midrib, coriaceous; many peduncles longer than 3 cm; involucral bracts usually appressed and with a chartaceous base and green apex (rarely loose, expanded and foliaceous apically) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. anticostense 9b. Leaf blades narrowed to the base 16a. Involucral bracts with a non-chlorophyllous spine tip, with short, stiff cilia on the margins [Fig. 462]; leaf blades linear to oblanceolate, moderately to densely pubescent on both surfaces and provided with stiff, marginal cilia . . . . . . S. ericoides 16b. Involucral bracts lacking a spine tip (though with an involute chlorophyllous tip in S. pilosum that is commonly misinterpreted as a spine), eciliate or softly ciliate on the margins [Fig. 464]; leaf blades not both narrow and pubescent on both surfaces 17a. Many of the involucral bracts with a subulate, involute, chlorophyllous tip [Fig. 466] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pilosum 17b. Involucral bracts without an involute tip [Fig. 464]

Ast e r ac e a e  437

18a. Involucral bracts densely sericeous; leaves densely sericeous on both surfaces (becoming glabrate in age), entire; capitulescence often tall and slender, raceme-like; cypselas densely sericeous, the hairs obscuring the nerves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. concolor 18b. Involucral bracts glabrous to pubescent, but not sericeous; leaves glabrous to pubescent, but not sericeous on both surfaces, entire to toothed; capitulescence variable, though often broad; cypselas usually sparsely strigose, the nerves visible 19a. Glabrous, fleshy saltmarsh plants with narrow, linear leaves rarely wider than 10 mm; involucral bracts chartaceous, sometimes green-tinged in the apical portion but often lacking a sharply defined chlorophyllous zone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tenuifolium 19b. Plants variable in habit, habitat, and leaf morphology, not of saltmarshes except varieties of S. novi-belgii; involucral bracts with a conspicuous chlorophyllous zone in the apical portion or entirely foliaceous 20a. Lobes of the disk corolla comprising (45–) 50–75% of the limb, flaring to recurving [Fig. 463] 21a. Leaf blades abaxially glabrous or pubescent only on the veins; disk flowers yellow, turning purple post anthesis; lobes of the disk corolla comprising 50–75% of the limb; capitula with 8–15 (–25) ray flowers; plants from a branched caudex or short, stout rhizome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lateriflorum 21b. Leaf blades abaxially sparsely to densely pubescent on and between the veins (glabrous in rare forms); disk flowers yellow, turning brown post anthesis; lobes of the disk corolla comprising 45–55 (–65)% of the limb; capitula with (10–) 15–26 ray flowers; plants from slender, elongate rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. ontarionis 20b. Lobes of the disk corolla comprising 15–45% of the limb, erect to ascending [Fig. 468] 22a. Branches and peduncles provided with copious, imbricate, short bracts mostly 1–4 mm long that are conspicuously shorter than the leaves of the main stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. dumosum 22b. Branches and peduncles with relatively fewer bracts, the bracts not crowded, mostly longer than 4 mm, and usually gradually decreasing in size from the leaves of the main stem (abruptly reduced in S. racemosum) 23a. Lobes of disk corolla comprising 15–30% of the limb [Fig. 468]; ray flowers commonly light blue to purple 24a. Leaves with conspicuous veins that connect to form isodiametric aereolae [Fig. 467]; stems glabrous and thinly glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. praealtum 24b. Leaves with fainter veins that connect to form aereolae that are longer than wide; stems glabrous to pubescent, not glaucous 25a. Leaf blades linear to oblong-linear, often with revolute margins, 2–5 (–9) mm wide; base of stem thinner than 2.5 mm; rhizome thinner than 2 mm; plants mostly of high-pH fens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. boreale 25b. Leaf blades narrow-lanceolate to lanceolate, elliptic, or oblanceolate, with plane margins, 5–25 mm wide; base of stem thicker than 2.5 mm; rhizome thicker than 2 mm; plants mostly of lake and river shores, saltmarshes, and moist fields

438   tricolpate s

26a. Leaf blades narrow-lanceolate to oblanceolate, narrowing to the base, flat, herbaceous to fleshy; most peduncles less than 3 cm long; involucral bracts often loose or somewhat spreading apically, commonly green in large part . . . . . . . . . . . . . . . . . . . . . . (in part) S. novi‑belgii 26b. Leaf blades tending to be narrow-lanceolate, scarcely narrowing to the base, folded along the midrib, coriaceous; many peduncles longer than 3 cm; involucral bracts usually appressed and with a chartaceous base and green apex (rarely loose, expanded and foliaceous apically) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. anticostense 23b. Lobes of disk corolla comprising 30–45% of the limb; ray flowers commonly white 27a. Involucre 2.5–3.5 (–4) mm tall; capitula often numerous and secund along the branches 28a. Bracteal leaves usually shorter than 1.5 cm, conspicuously reduced in size from the stem leaves . . . . . . . . . . S. racemosum 28b. Bracteal leaves, at least some of them, usually longer than 1.5 cm, gradually reduced in size from the stem leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. lanceolatum 27b. Involucre 4–6 mm tall; capitula usually neither crowded nor notably secund 29a. Stems 1–6 dm tall; stems 1–3 mm thick at the base; leaf blades 3–10 × 0.3–1 cm; capitula with 15–30 ray flowers; rays 3–8 mm long; plants of rocky shores . . . . . . . . . S. tradescantii 29b. Stems 5–15 dm tall; stems 3–6 mm thick at the base; leaf blades 8–15 × 0.3–3.5 cm; capitula with 20–40 ray flowers; rays 4.5–12 mm long; plants mostly of wet-mesic to hydric fields, ditches, and open swamp edges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. lanceolatum 1. Symphyotrichum anticostense (Fern.) Nesom

NC

Anticosti American-aster. Aster anticostensis Fern. • ME; northern portion of state. Ice-scoured river shores and adjacent, disintegrating ledges. 2. Symphyotrichum boreale (Torr. & Gray) Nesom N rush American-aster. Aster borealis (Torr. & Gray) Prov.; A. junciformis Rydb. • ME, NH, VT. Circumneutral fens. The report of this species in MA (e.g., Sorrie and Somers 1999) is based on a collection of Symphyotrichum novi-belgii var. elodes—Goodale et al. 69563 (MASS!). 3. Symphyotrichum ciliatum (Ledeb.) Nesom E Fig. 461

Fig. 461  Capitulum of Symphyotrichum ciliatum.

rayless annual American-aster. Aster brachyactis Blake; Brachyactis angusta (Lindl.) Britt.; B. ciliata (Ledeb.) Ledeb.; B. ciliata (Ledeb.) Ledeb. ssp. angusta (Lindl.) A.G. Jones; Tripolium angustum Lindl. • VT. Salted roadsides. Reports of this species from York County, ME, are based on Symphyotrichum frondosum. 4. Symphyotrichum ciliolatum (Lindl.) A. & D. Löve N Lindley’s American-aster. Aster ciliolatus Lindl.; A. lindleyanus Torr. & Gray • MA, ME, NH, VT. Forests, forest edges, clearings, fields. This species is frequently confused with Symphyotrichum cordifolium. In addition to the characters used in the identification key, the two can be separated on involucral bract and leaf blade morphology. Symphyotrichum ciliolatum has involucral bracts with a slender, elongate chlorophyllous zone, basal leaf blades that are usually truncate to subcordate, and stem leaves, when petioled, with winged petioles. Symphyotrichum cordifolium has involucral bracts with an apically dilated chlorophyllous zone, basal leaf blades that are usually prominently cordate, and stem leaves, when petioled, with slender petioles.

Ast e r ac e a e   43 9

5. Symphyotrichum concolor (L.) Nesom ssp. concolor

NC

eastern silver American-aster. Aster concolor L.; Lasallea concolor (L.) Semple & Brouillet; Virgulus concolor (L.) Reveal & Keener • MA, RI; coastal plain. Grasslands, sand plains, woodlands. 6. Symphyotrichum cordifolium (L.) Nesom N heart-leaved American-aster. Aster cordifolius L.; A. cordifolius L. var. furbishiae Fern.; A. cordifolius L. var. incisus Britt.; A. cordifolius L. var. polycephalus Porter; A. cordifolius L. ssp. sagittifolius (Wedemeyer ex Willd.) A.G. Jones; A. cordifolius L. var. sagittifolius (Wedemeyer ex Willd.) A.G. Jones; A. sagittifolius Wedemeyer ex Willd; Symphyotrichum cordifolium (L.) Nesom var. furbishiae (Fern.) Nesom; S. cordifolium (L.) Nesom var. polycephalum (Porter) Nesom; S. sagittifolium (Wedemeyer ex Willd.) Nesom • CT, MA, ME, NH, RI, VT. Forests, forest edges, fields, roadsides, waste areas, sometimes weedy and usually found in at least partial shade of trees and/or buildings. 6 × 15. This very rare American-aster hybrid is known from VT. It has adaxially scabrous lower leaves with rounded leaf blade bases that are abruptly contracted to a winged petiole. The capitula have blue rays with wider involucral bracts 0.7–1 mm wide that have a pronounced apical green zone, this sometimes representing 50% of more of the bract length (i.e., some involucral bracts are somewhat foliaceous). ‌6 × 21. Symphyotrichum ×tardiflorum (L.) Greuter, M.V. Agab., & Wagenitz is a rare American-aster hybrid that can be locally frequent. It is known from CT, MA, ME, NH, VT. It has often been associated with Symphyotrichum novi-belgii (usually as a variety under the name Aster novi-belgii L. var. tardiflorus (L.) A.G. Jones). It has broadlanceolate to narrow-ovate leaf blades that are abruptly contracted to a broadly winged, somewhat clasping petiole. The involucral bracts are intermediate between the parental taxa, being more elongate than in S. cordifolium, and the involucre is (5–) 6–8 mm tall. This nothospecies generally shows blue to light purple rays. 7. Symphyotrichum dumosum (L.) Nesom N bushy American-aster. Aster coridifolius Michx.; A. dumosus L. var. coridifolius (Michx.) Torr. & Gray; A. dumosus L. var. gracilentus Torr. & Gray; A. dumosus L. var. dodgei Fern.; A. dumosus L. var. strictior Torr. & Gray; A. dumosus L. var. subulifolius Torr. & Gray; Symphyotrichum dumosum (L.) Nesom var. dodgei (Fern.) Nesom; S. dumosum (L.) Nesom var. strictior (Torr. & Gray) Nesom; S. dumosum (L.) Nesom var. subulifolium (Torr. & Gray) Nesom • CT, MA, ME, NH, RI; also reported from VT by Seymour (1982), but specimens are unknown; most common on the coastal plain. Fields, woodlands, dry, sandy places. 7 × 12. This very rare American-aster hybrid is known from MA. It has the relatively long disk corolla lobes (relative to the length of the corolla limb) as in Symphyotrichum lateriflorum combined with the elongate and copiously bracteate peduncles of S. dumosum. 8. Symphyotrichum ericoides (L.) Nesom var. ericoides N Fig. 462 heath American-aster. Aster ericoides L.; A. ericoides L. var. prostratus (Kuntze) Blake; A. multiflorus Ait.; A. multiflorus Ait. var. prostratus Kuntze; Lasallea ericoides (L.) Semple & Brouillet; Virgulus ericoides (L.) Reveal & Keener • CT, MA, ME, NH, RI, VT. Sandy, open areas, also sometimes rocky turfs near the coast. ‌8 × 14. Symphyotrichum ×amethystinum (Nutt.) Nesom is an uncommon Americanaster hybrid known from CT, MA, RI, VT. It is normally found growing with both parental species and is intermediate in morphology between them. The involucral bracts do not taper into the long apex found in S. novae-angliae, are sparsely stipitate-glandular, and lack a spine tip (with a spine tip in S. ericoides). 9. Symphyotrichum frondosum (Nutt.) Nesom E leafy annual American-aster. Aster frondosus (Nutt.) Torr. & Gray; Brachyactis frondosa (Nutt.) Gray; Tripolium frondosum Nutt. • ME. Wool waste.

Fig. 462  Capitulum of Symphyotrichum ericoides showing spine-tipped involucral bracts.

440  tricolpates

10. Symphyotrichum laeve (L.) A. & D. Löve var. laeve n smooth American-aster. Aster laevis L. • CT, MA, ME, NH, RI, VT; introduced to some counties in ME, but native to York County. Sandy or rocky woodlands, dry fields, roadsides, sandy, open places. 11. Symphyotrichum lanceolatum (Willd.) Nesom ssp. lanceolatum

NC

lance-leaved American-aster.  11a. Aster interior Wieg.; A. lanceolatus Willd. ssp. interior (Wieg.) A.G. Jones; A. lanceolatus Willd. var. interior (Wieg.) Semple & Chmielewski; A. simplex Willd. var. interior (Wieg.) Cronq.; 11b. Aster lanceolatus Willd.; A. lanceolatus Willd. ssp. simplex (Willd.) A.G. Jones; A. lanceolatus Willd. var. simplex (Willd.) A.G. Jones; A. paniculatus Lam. var. simplex (Willd.) Burgess; A. simplex Willd.; A. simplex Willd. var. ramosissimus (Torr. & Gray) Cronq.; Symphyotrichum simplex (Willd.) A. & D. Löve; 11c. Aster lanceolatus Willd. var. latifolius Semple & Chmielewski • CT, MA, ME, NH, RI, VT; common. Wet-mesic to hydric fields, wetland edges, open swamps, ditches. 1a. Involucres usually less than 4 mm tall; capitula crowded on the branches . . . . . . . . . . . . . . . . . . . . . . . 11a. S. lanceolatum ssp. lanceolatum var. interior (Wieg.) Nesom 1b. Involucres 4–6 mm tall; capitula more remote 2a. Leaf blades linear to oblanceolate, mostly 3–12 mm wide, often exceeding 12 times as long as wide, reduced in the capitulescence; rays white to pale violet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11b. S. lanceolatum ssp. lanceolatum var. lanceolatum Fig. 463  Disk corolla of Symphyotrichum lateriflorum showing long lobes relative to the limb.

2b. Leaf blades broadly oblanceolate, mostly 10–35 mm wide, rarely exceeding 11 times as long as wide, only somewhat reduced in the capitulescence; rays white . . . . 11c. S. lanceolatum ssp. lanceolatum var. latifolium (Semple & Chmielewski) Nesom Variety interior is known from VT; also reported from New Hampshire by Kartesz (1999), but specimens are unknown. It is of regional conservation concern. Variety lanceolatum is known from CT, MA, ME, NH, RI, VT. It is the most common variety of this species in New England. Variety latifolium is known from CT, MA, ME, NH, RI, VT. 12. Symphyotrichum lateriflorum (L.) A. & D. Löve N Fig. 463, 464 calico American-aster. Aster hirsuticaulis Lindl. ex DC.; A. lateriflorus (L.) Britt.; A. lateriflorus (L.) Britt. var. angustifolius Wieg.; A. lateriflorus (L.) Britt. var. hirsuticaulis (Lindl. ex DC.) Porter; Solidago lateriflora L. • CT, MA, ME, NH, RI, VT. Mesic to hydric forests, forest edges, and fields.

Fig. 464  Capitulum of Symphyotrichum lateriflorum showing involucral bracts with a distinct, apical green zone.

12 × 21. This uncommon American-aster hybrid is known from MA, ME, VT. It is usually found in wet-mesic forests and usually occurs where the two parental taxa co-occur. It resembles a slender Symphyotrichum puniceum with leaves that do not clasp at the base and less elongate involucral bracts. Examination of the disk corolla lobes will show them to be intermediate in the length of the corolla lobes relative to the corolla limb (see identification key for details of ratios). 13. Symphyotrichum lowrieanum (Porter) Nesom

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Lowrie’s American-aster. Aster cordfolius L. ssp. laevigatus (Porter) A.G. Jones; A. cordifolius L. var. laevigatus Porter; A. lowrieanus Porter • CT. Woodlands and forest edges. Historical, not seen in many years. Possibly the hybrid of Symphyotrichum cordifolium and S. laeve, as suggested by this plant’s morphology. Much over-reported in New England, most specimens actually S. cordifolium or S. laeve. 14. Symphyotrichum novae-angliae (L.) Nesom N Fig. 465

Fig. 465  Capitulum of Symphyotrichum novaeangliae showing involucre with foliaceous and stipitate-glandular bracts.

New England American-aster. Aster novae-angliae L.; Lasallea novae-angliae (L.) Semple & L. Brouillet; Virgulus novae-angliae (L.) Reveal & Keener • CT, MA, ME, NH, RI, VT. Mesic to drymesic soils of open places and edges, such as fields, roadsides, and banks. 15. Symphyotrichum novi-belgii (L.) Nesom N New York American-aster.  15a. Aster elodes Torr. & Gray; A. novi-belgii L. var. elodes (Torr. & Gray) Gray; 15b. Aster johannensis Fern.; A. longifolius Lam.; A. novi-belgii L.; A. novi-belgii L. ssp. johannensis (Fern.) A.G. Jones; A. novi-belgii L. var. johannensis (Fern.) A.G. Jones;  15c. Aster johannensis Fern. var. villicaulis (Gray) Fern.; A. novi-belgii L. var. villicaulis (Gray)

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Boivin • CT, MA, ME, NH, RI, VT. Saltmarshes and open places near the coast, fields and forest edges, riparian and lacustrine shorelines, rarely ascending to high elevation in northern New England. Plants from New England determined as Aster crenifolius and A. foliaceus in regional herbaria represent (in part) environmental forms of Symphyotrichum novi-belgii with elongate, foliaceous outer involucral bracts. Symphyotrichum novi-belgii var. litoreum (Gray) Nesom represents a set of brackish marsh plants with thick, broad blades (relative to length). These have been considered to be the hybrid of S. laeve and S. praealtum (Labrecque and Brouillet 1996); however, this combination does not explain the coastal marsh habitat and foliaceous involucral bracts. The identity of these plants is still in question. 1a. Leaf blades narrow-lanceolate, 10–25 times as long as wide, mostly 4–12 mm wide, barely clasping at the base, the larger blades 4–12 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15a. S. novi-belgii var. elodes (Torr. & Gray) Nesom 1b. Leaf blades oblanceolate to narrow-ovate to oblong, (5–) 7–9 times as long as wide, mostly 9–20 mm wide, usually clasping at the base, the larger blades 6.5–25 mm wide 2a. Stems glabrous to sparsely pubescent; plants widespread, though most frequent in the coastal region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15b. S. novi‑belgii var. novi-belgii 2b. Stems conspicuously pubescent; plants mostly of inland river shores . . . . . . . . . . . . . . . . . . . . . . . . . . 15c. S. novi‑belgii var. villicaule (Gray) Lebrecque & Brouillet Variety elodes is known from CT, MA, ME, RI; also reported from NH and VT by Seymour (1982), but specimens are unknown. Variety novi-belgii is known from CT, MA, ME, NH, RI, VT. Both of these varieties are found in open, coastal, near coastal, and inland natural communities (river banks, fens, boggy shorelines, marsh edges). Variety villicaule is known from ME. It is usually found on rocky shorelines of rivers. 16. Symphyotrichum ontarionis (Wieg.) Nesom

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Ontario American-aster. Aster ontarionis Wieg.; A. ontarionis Wieg. var. glabratus Semple; Symphyotrichum ontarionis (Wieg.) Nesom var. glabratum (Semple) Brouillet & Bouchard • VT; western portion of state. Edges and open places of lacustrine floodplain forests. Rare collections of this plant in New England show nearly or completely glabrous abaxial leaf surfaces. These forms have been distinguished as var. glabratus. Though many collections can be assigned to the pubescent or ± glabrous categories, there exist plants with intermediate hair density, which call into question the validity of the two varieties. The species is treated here as a single, variable entity until further study supports the separation of two forms based on leaf indument density.

Fig. 466  Capitulum of Symphyotrichum pilosum showing involucral bracts that are apically involute.

17. Symphyotrichum patens (Ait.) Nesom var. patens N late purple American-aster. Aster patens Ait.; Lasallea patens (Ait.) Semple & Brouillet; Virgulus patens (Ait.) Reveal & Keener • CT, MA, ME, NH, RI; becoming rare in northern New England. Dry fields and sandy or rocky woodlands. Reports of Symphyotrichum phlogifolium (Muhl. ex Willd.) Nesom from New England are based on specimens of S. patens. 18. Symphyotrichum pilosum (Willd.) Nesom N Fig. 466 awl American-aster.  18a. Aster pilosus Willd.; 18b. Aster ericoides L. var. pringlei Gray; A. pilosus Willd. var. demotus Blake; A. pilosus Willd. var. pringlei (Gray) Blake; A. pringlei (Gray) Britt. • CT, MA, ME, NH, RI, VT. Fields and edges, most frequent in well-drained soils. 1a. Plants sparsely to densely pubescent on stems, branches, and often the leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18a. S. pilosum var. pilosum 1b. Vegetative parts of plant glabrous or essentially so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18b. S. pilosum var. pringlei (Gray) Nesom Variety pilosum is known from CT, MA, ME, RI, VT. Variety pringlei is known from CT, MA, ME, NH, RI, VT. 19. Symphyotrichum praealtum (Poir.) Nesom ssp. angustior (Wieg.) A. Haines N C Fig. 467 willow-leaved American-aster. Aster praealtus Poir. var. angustior Wieg.; Symphyotrichum praealtum (Poir.) Nesom var. angustior (Wieg.) Nesom • CT, MA. Mesic to wet-mesic fields and borders. Much over-reported and very rare in New England.

Fig. 467  Leaf blade of Symphyotrichum praealtum showing veins that create isodiametric aereolae.

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20. Symphyotrichum prenanthoides (Muhl. ex Willd.) Nesom

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crooked-stemmed American-aster. Aster prenanthoides Muhl. ex Willd. • CT, MA, VT. In open areas and along edges of mesic to wet-mesic forests, often riparian types. 21. Symphyotrichum puniceum (L.) A. & D. Löve var. puniceum N Fig. 468

Fig. 468  Disk corolla of Symphyotrichum puniceum showing short lobes relative to the limb.

purple-stemmed American-aster. Aster firmus Nees; A. lucidulus (Gray) Wieg.; A. puniceus L.; A. puniceus L. ssp. firmus (Nees) A.G. Jones; A. puniceus L. var. firmus (Nees) Torr. & Gray; A. puniceus L. var. lucidulus Gray; Symphyotrichum firmum (Nees) Nesom • CT, MA, ME, NH, RI, VT. Swamps, hydric fields, wet ditches, shorelines. Plants called Symphyotrichum firmum, identified by rhizomatous habit, ± glabrous stems and leaf blades, and leafy capitulescences, are not morphologically distinct. Review of specimens from multiple herbaria shows that supposed diagnostic characters vary independently. A recent study supporting the distinction of S. firmum is flawed in that it included only “textbook” specimens of each taxon for analysis (Warners and Laughlin 1999). 22. Symphyotrichum racemosum (Ell.) Nesom N small white American-aster. Aster fragilis, auct. non Willd.; A. racemosus Ell.; A. vimineus Lam. var. subdumosus Wieg.; A. vimineus, auct. non Lam.; Symphyotrichum racemosum var. subdumosum (Wieg.) Nesom • CT, MA, ME, NH, RI, VT. Fields and moist, open sandy areas, including coastal plain pond shores. 23. Symphyotrichum subulatum (Michx.) Nesom var. subulatum N annual saltmarsh American-aster. Aster subulatus Michx.; A. subulatus Michx. var. euroauster Fern. & Grisc.; A. subulatus Michx. var. obtusifolius Fern. • CT, MA, ME, NH, RI; restricted in ME to southern portion of state. Saline marshes, brackish river shores. 24. Symphyotrichum tenuifolium (L.) Nesom var. tenuifolium N perennial saltmarsh American-aster. Aster tenuifolius L. • CT, MA, NH, RI. Saline marshes, brackish river shores. 25. Symphyotrichum tradescantii (L.) Nesom N Tradescant’s American-aster. Aster saxatilis (Fern.) Blanch., nom. illeg.; A. tradescantii L.; A. tradescantii L. var. saxatilis (Fern.) House; A. vimineus Lam. var. saxatilis Fern. • MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Rocky and ledgy shorelines, most frequently rivers. 26. Symphyotrichum undulatum (L.) Nesom N Fig. 469 wavy-leaved American-aster. Aster loriformis (Burgess) Burgess; A. undulatus L.; A. undulatus L. var. loriformis Burgess • CT, MA, ME, NH, RI, VT. Dry fields, forest edges and openings, woodlands, roadsides. 27. Symphyotrichum urophyllum (Lindl.) Nesom

Fig. 469  Leaf of Symphyotrichum undulatum showing clasping petiole base.

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arrow-leaved American-aster. Aster sagittifolius, auct. non Wedemeyer ex Willd.; A. sagittifolius Wedemeyer ex Willd. var. urophyllus (Lindl.) Burgess; A. urophyllus Lindl. • VT; also reported from CT, MA, ME, NH by Brouillet et al. (2006), but specimens are unknown. Fields and open places in high-pH bedrock regions. The name Aster sagittifolius has been misapplied to this plant for some time. The type collection of A. sagittifolius proved to be Symphyotrichum cordifolium; therefore, another name needed to be chosen for this species (Semple et al. 2002).

Tagetes 1a. Involucre 7–12 mm tall, composed of 3–5 involucral bracts; capitula with (1–) 3 ray flowers, the rays 1–2 mm long; disk corollas 3–4 mm long; cypsela body 4.5–7 mm long; pappus of 1 or 2 acuminate scales 2–3 mm long and 3–5 shorter, ovate to lanceolate scales 0.5–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. minuta 1b. Involucre 13–19 mm tall, composed of 7–11 involucral bracts; capitula with 8 or more ray flowers, the rays 10–30 mm long; disk corollas 10–15 mm long; cypsela body 6.5–10 mm long; pappus of 1 or 2 acuminate scales 8–11 mm long and 2 or 3 blunt scales 3–6 mm long

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2a. Peduncles swollen; capitula 6–10 cm wide; corollas yellow or orange . . . . . . . . T. erecta 2b. Peduncles not or only scarcely swollen; capitula 3.5–6 cm wide; corollas partly or entirely red-orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. patula 1. Tagetes erecta L. E Aztec marigold. Tagetes major Gaertn. • MA; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Roadsides, fields, waste areas. 2. Tagetes minuta L. E miniature marigold. CT, MA. Roadsides, fields, waste areas. 3. Tagetes patula L. E French marigold. Tagetes tenuifolia Millsp. • VT. Roadsides, fields, waste areas.

Tanacetum Reference: Watson (2006). 1a. Rays 4–8 mm long, white; cypsela body with 7–10 ribs . . . . . . . . . . . . . . . . . . . T. parthenium 1b. Rays absent or up to 4 mm long and yellow; cypsela body with 5 ribs 2a. Leaf blades glabrous or nearly so; disk 5–10 mm wide; capitulescence with (10–) 20–70 (–100) capitula; rays absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. vulgare 2b. Leaf blades villous; disk 10–18 mm wide; capitulescence with 1–15 (–30) capitula [Fig. 470]; rays present, 1–3 (–4) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. bipinnatum 1. Tanacetum bipinnatum (L.) Schultz-Bip. ssp. huronense (Nutt.) Breitung N C Fig. 470 eastern tansy. Chrysanthemum bipinnatum L. ssp. huronense (Nutt.) Hultén; Tanacetum huronense Nutt.; T. huronense Nutt. var. johannense Fern. • ME; northern portion of state. Ice-scoured river shores, usually on gravel substrate. 2. Tanacetum parthenium (L.) Schultz-Bip. E feverfew tansy. Chrysanthemum parthenium (L.) Bernh.; Matricaria parthenium L.; Pyrethrum parthenium (L.) Sm. • CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas, rubbish heaps. 3. Tanacetum vulgare L. E common tansy. Chrysanthemum uliginosum Pers.; C. vulgare (L.) Bernh. • CT, MA, ME, NH, RI, VT. Fields, roadsides, open river shores, disturbed soil.

Taraxacum Authorities disagree on how best to treat Taraxacum. Numerous microspecies have been distinguished, sometimes on very subtle distinctions. The species are treated here in the broad sense. Taraxacum ceratophorum (Ledeb.) DC. was reported from NH based on an immature specimen collected in Kings Ravine on Mount Adams, Coos County (Pease 1917). However, no mature specimens have ever been collected to verify the identity of the plant. Further, the specimen (at NEBC!) shows narrow and reflexed (in part) outer involucral bracts that are very unlike T. ceratophorum (i.e., the specimen appears misidentified and is likely T. officinale). Taraxacum ceratophorum has also been reported from MA and ME by Magee and Ahles (1999); however, this report was based on incorrectly synonymizing T. latilobum (specimens at NEBC!) with T. ceratorphorum (the two taxa are very distinct are do not comprise the same species). References: Fernald (1933), Brouillet (2006b). 1a. Inner (i.e., longer) involucral bracts with a dark, corniculate appendage at the tip [Fig. 471]; cypsela body at maturity purple-red to brown-red; leaf blades lacerate-lobed with narrow lobes, the medial lateral lobes of leaf blades 1–10 (–12) mm wide at the base . . . . . . . . . . T. laevigatum

Fig. 470  Capitula and leaves of Tanacetum bipinnatum.

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1b. Inner involucral bracts lacking dark, horn-like appendages at the tip; cypsela body at maturity light brown to gray-brown to brown; leaf blades toothed to lobed, the medial lateral lobes of leaf blades, when present, (5–) 10–32 (–38) mm wide 2a. Outer (i.e., shorter) involucral bracts, as well as the inner ones, persistently appressed to ascending; plants 5–10 cm tall; involucres excluding the outer calyculus 12–16 mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. palustre 2b. Outer involucral bracts becoming reflexed [Fig. 471]; plants (1–) 5–75 cm tall; involucres excluding the outer calyculus (13–) 14–25 mm tall 3a. Cypsela with a body (2–) 2.5–2.8 (–4) mm long that is muricate only in the apical half [Fig. 472], with a beak 7–9 mm long; outermost involucral bracts (i.e., the reflexed ones) lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. officinale 3b. Cypsela with a body 2.8–3.8 mm long that is usually muricate also in the basal half (sometimes all the way to the base), with a beak 8–12 mm long; outermost involucral bracts ovate to broad-lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. latilobum 1. Taraxacum laevigatum (Willd.) DC. E Fig. 471 Fig. 471  Involucre of Taraxacum laevigatum showing cucullate involucral bract tips.

red-seeded dandelion. Leontodon erythrospermum (Andrz. ex Bess.) Britt.; Taraxacum erythrospermum Andrz. ex Bess. • CT, MA, ME, NH, RI, VT. Fields, lawns, waste areas, roadsides. The leaf blades of this species tend to have narrower lobes and deeper sinuses than other species of Taraxacum. Once learned, this trait is useful for forming an initial hypothesis concerning the identity of vegetative specimens. 2. Taraxacum latilobum DC.

nC

large-lobed dandelion. Leontodon latiloba (DC.) Britt.; Taraxacum officinale G.H. Weber ex Wiggers ssp. vulgare (Lam.) Schinz & R. Keller; T. vulgare Lam. • MA, ME. Open, often disturbed, soil, dumps. The status of this species in New England is debatable. The MA record is certainly introduced, and those from ME may be as well. 3. Taraxacum officinale G.H. Weber ex Wiggers E Fig. 472 common dandelion. Leontodon taraxacum L. • CT, MA, ME, NH, RI, VT. Lawns, fields, roadsides, waste areas, open shorelines, clearings. 4. Taraxacum palustre (Lyons) Symons E marsh dandelion. Taraxacum turfosum (Schultz-Bip.) Soest • CT, VT; also reported from MA, ME, and NH by Seymour (1982), but specimens are unknown. Lawns, ditches, and other disturbed places in regions of high-pH bedrock. Fig. 472  Cypsela of Taraxacum officinale.

Thymophylla 1. Thymophylla tenuiloba (DC.) Small var. tenuiloba E Dahlburg-daisy. Dyssodia tenuiloba (DC.) B.L. Robins.; Hymenatherum tenuilobum DC. • MA. Fields, roadsides, waste areas.

Tragopogon Reference: Soltis (2006). 1a. Peduncles slender, not enlarged in flower and scarcely enlarged even in fruit; cypsela body including beak 15–25 mm long; involucre 12–24 mm tall in flower, becoming 19–38 mm tall in fruit; involucral bracts shorter than to equaling the ray flowers in length . . . . . . . . . . T. pratensis 1b. Peduncles apically enlarged and fistulous in flower and fruit [Fig. 473]; cypsela body 25–40 mm long; involucre 25–40 mm tall in flower, becoming 40–70 mm tall in fruit; involucral bracts exceeding ray flowers in length 2a. Ray corollas yellow; pappus white to sordid-white; leaf apex usually straight (i.e., not prominently curved) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. dubius

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2b. Ray corollas purple; pappus light brown; leaf apex usually recurved to recoiled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. porrifolius 1. Tragopogon dubius Scop. E fistulous goat’s beard. Tragopogon dubius Scop. ssp. major (Jacq.) Voll.; T. major Jacq. • CT, MA, NH, VT; also reported from ME and RI by Magee and Ahles (1999), but specimens are

unknown. Fields, roadsides, pastures. 2. Tragopogon porrifolius L. E Fig. 473 purple goat’s beard. CT, MA, ME, NH, VT. Fields, roadsides, pastures. 3. Tragopogon pratensis L. E meadow goat’s beard. CT, MA, ME, NH, RI, VT. Fields, roadsides, pastures.

Tripleurospermum Capitula must be neatly pressed for accurate width measurements. 1a. Plants ascending to erect; capitula 3–4 cm wide; ultimate segments of the leaves filiform [Fig. 474], 4–20 mm long; rays 10–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. inodorum

Fig. 473  Involucre and swollen peduncle of Tragopogon porrifolius.

1b. Plants depressed to spreading; capitula 1.5–3 cm wide; ultimate segments of the leaves linear, 1.5–5 mm long; rays 7–12 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. maritimum 1. Tripleurospermum inodorum (L.) Schultz-Bip. E Fig. 474 corn scentless-chamomile. Chamomilla inodora (L.) Gilib.; Matricaria inodora L.; M. maritima L. ssp. inodora (L.) Clapham; M. maritima L. var. agrestis (Knaf) Wilmott; M. perforata Mérat; Tripleurospermum perforatum (Mérat) M. Laínz • CT, MA, ME, NH. Fields, roadsides, waste areas. 2. Tripleurospermum maritimum (L.) W.D.J. Koch ssp. maritimum E scentless-chamomile. Chamomilla maritima (L.) Rydb.; Matricaria maritima L. • CT, MA, ME, NH. Fields, roadsides, waste areas.

Tussilago 1. Tussilago farfara L. E Fig. 475

Fig. 474  Capitula and leaves of Tripleurospermum inodorum.

coltsfoot. CT, MA, ME, NH, RI, VT. Roadsides, stream and river banks, trail edges, open or partially shaded, disturbed soil.

Verbesina 1a. Plants annual, 2–10 dm tall; stems without decurrent wing-angles; capitula with 10–15 carpellate ray flowers; leaf blades coarsely serrate . . . . . . . . . . . . . . . . . . . . . . . . . . V. encelioides 1b. Plants perennial, 10–30 dm tall; stems with prominent wing-angles that are decurrent from the leaf bases; capitula with 2–10 sterile ray flowers; leaf blades serrate to subentire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. alternifolia 1. Verbesina alternifolia (L.) Britt. ex Kearney E wingstem crownbeard. Actinomeris alternifolia (L.) DC.; Coreopsis alternifolia L.; Ridan alternifolia (L.) Britt. • MA. Roadsides, fields, waste areas. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species had questionable naturalization in RI and was unaware of any collections. 2. Verbesina encelioides (Cav.) Benth. & Hook. f. ex Gray ssp. exauriculata (Robins. & Greenm.) J.R. Coleman E golden crownbeard. Verbesina encelioides (Cav.) Benth. & Hook. f. var. exauriculata Robins. & Greenm.; V. exauriculata (Robins. & Greenm.) Cockerell; Ximenesia encelioides Cav. var. exauriculata (Robins. & Greenm.) F.C. Gates; X. exauriculata (Robins. & Greenm.) Rydb. • MA; also reported from RI by Kartesz (1999), but specimens are unknown. Fields, roadsides, waste areas.

Fig. 475  Flowering stem and capitulum of Tussilago farfara.

446 tricolpates

Vernonia Reference: Strother (2006g). 1a. Involucral bracts apically prolonged into a filiform tip 2–8 mm long [Fig. 476] 2a. Leaf blades lanceolate to narrow-lanceolate, (3.3–) 4–6 or more times as long as wide; pappus brown to purple-brown, the 20 outer scales 0.2–0.6 mm long (rarely longer), abruptly transitioning to the longer, slender, inner bristles . . . . . . . . . . . . V. noveboracensis 2b. Leaf blades lanceolate to ovate, 2.5–3.5 (–4) times as long as wide; pappus stramineous or very pale brown to nearly white, the outer 30 scales 0.5–1.5 mm long (rarely longer), gradually transitioning to the longer, slender, inner bristles . . . . . . . . . . . . . . . . . . . . V. glauca 1b. Involucral bracts rounded to acuminate at the apex, but not prolonged 3a. Abaxial surface of leaf blades ± glabrous and conspicuously punctate; capitula with 12–25 (–30) flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. fasciculata 3b. Abaxial surface of leaf blades thinly to densely tomentose, the pubescence usually concealing the puncta; capitula with (30–) 32–55 (–58) flowers . . . . . . . . . . . . . V. missurica 1. Vernonia fasciculata Michx. E prairie ironweed. MA. Fields, roadsides, waste areas. 2. Vernonia glauca (L.) Willd. E broad-leaved ironweed. Serratula glauca L. • MA. Fields, roadsides, waste areas. 3. Vernonia missurica Raf. E Missouri ironweed. MA. Fields, waste areas. 4. Vernonia noveboracensis (L.) Michx. n Fig. 476 New York ironweed. Serratula noveboracensis L. • CT, MA, NH, RI. Fields and meadows, these often wet-mesic to hydric, stream banks, wet ditches. This species is non-native in NH and occurs as a native in the other states. Fig. 476  Capitula of Vernonia noveboracensis showing prolonged tips of involucral bracts.

Xanthium Xanthium is a difficult genus in that it requires careful examination of micromorphological details of the fruiting carpellate involucres (often referred to as “burs”). Accurate determination of many collections is complicated by the fact that they possess flowering or immature fruiting involucres. Tremendous disagreement exists as to the number of species to recognize. A moderate treatment, that of Cronquist (1945), is here followed with reservation (I suspect that earlier works that recognized more species are, in fact, closer to the truth). Though Cronquist’s ideas regarding Xanthium appear to be supported by correlated characters, some distinctive forms with ecological differences are not recognized (e.g., X. echinatum, a species of Atlantic coast beaches, dune hollows, and marsh borders). Reference: Fernald (1946). 1a. Leaf blades lanceolate, tapering to the base, abaxially gray to white and densely strigose; plants with 3-forked, axillary spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. spinosum 1b. Leaf blades ovate to reniform, truncate to cordate at the base, abaxially ± green, hirtellous; plants without axillary spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. strumarium 1. Xanthium spinosum L. E spiny cocklebur. Acanthoxanthium spinosum (L.) Fourr.; Xanthium spinosum L. var. inerme Bel. • CT, MA, ME, NH, RI. Waste areas, disturbed ground, often in the vicinity of old mills.

Fig. 477  Carpellate involucre of Xanthium strumarium var. canadense showing stipitateglandular prickle bases.

2. Xanthium strumarium L. n Fig. 477 rough cocklebur.  2a. Xanthium canadense P. Mill.; X. echinatum Murr.; X. glanduliferum Greene; X. italicum Moretti; X. oviforme Wallr.; X. pensylvanicum Wallr.; X. speciosum Kearney; X. strumarium L. var. pensylvanicum (Wallr.) M.E. Peck; 2b. Xanthium americanum Walt.;

Ast e r ac e a e  447

X. chasei Fern.; X. chinense P. Mill.; X. curvescens Millsp. & Sherff; X. cylindraceum Millsp. & Sherff; X. echinellum Greene; X. globosum Shull; X. inflexum Mackenzie & Bush; X. orientale L. • CT, MA, ME, NH, RI, VT. River banks, river shores, especially those of sand and gravel substrate, lake shores, waste areas, coastal beaches, borders of saline marshes, dune hollows. 1a. Fruiting carpellate involucres (17–) 20–40 mm long and 12–30 mm thick, terminated by 2 beaks 3–11 mm long; prickle bases of carpellate involucres spreading-pubescent and sometimes also stipitate-glandular [Fig. 477] . . . . . . 2a. X. strumarium var. canadense (P. Mill.) Torr. & Gray 1b. Fruiting carpellate involucres 8–20 (–25) mm long and 4–18 mm thick, terminated by 2 beaks 1–7 mm long; prickle bases of carpellate involucres glabrous or puberulent and sometimes also glandular-puberulent and/or with sessile glands 2a. Carpellate involucres terminated by 2 incurved or, less commonly, straight beaks, the fruiting ones usually pale or light brown; petioles usually scabrous; leaf blades usually scabrous adaxially . . . . . . . . . . . . . . . . . . . . . 2b. X. strumarium var. glabratum (DC.) Cronq. 2b. Carpellate involucres terminated by 2 straight beaks, the fruiting ones green to yellowgreen; petioles minutely and softly pilose; leaf blades weakly scabridulous adaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2c. X. strumarium var. strumarium Variety canadense is native (at least in part) and known from CT, MA, ME, NH, RI, VT. It is the most frequently encountered variety in New England. Variety glabratum is native (at least in part) and known from CT, MA, VT; also reported from RI by George (1992), but specimens are unknown. Variety strumarium is non-native and known from MA, VT.

Xerochrysum 1. Xerochrysum bracteatum (Vent.) Tzvelev E bracted strawflower. Helichrysum bracteatum (Vent.) Andr.; Xeranthemum bracteatum Vent. • CT, MA. Roadsides, fields, disturbed soil.

Balsaminaceae Impatiens The flowers of Impatiens are difficult to interpret. There are 3 sepals—a lower one that is petaloid and forms a conical structure prolonged backward into a slender nectary spur and 2 upper ones that are small and sepaloid or petaloid. There are 5 petals—an upper one that is concave and projected upward or forward and 2 lower ones that are each formed from 2 connate petals. 1a. Leaves opposite or whorled, the blades sharply serrate; petioles provided with large, dark glands; Perianth blue or purple to pink or white; capsule obovoid . . . . . . . . . . . . I. glandulifera 1b. Leaves alternate, the blades crenate-serrate; petioles lacking glands or infrequently with slender, dark glands near the apex at the junction of the blade; corolla orange-yellow or pale yellow; capsule slender-clavate in outline 2a. Perianth orange-yellow (rarely bright yellow, pink, or white), spotted with red-brown (rarely unspotted); spur 7–10 mm long, strongly curved, projecting forward; saccate portion of the spurred sepal longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. capensis 2b. Perianth pale yellow (rarely yellow-white or white), sparingly dotted with red-brown; spur 4–6 mm long, curved at a right angle, not projecting forward [Fig. 478]; saccate portion of the spurred sepal wider than long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. pallida

448   tricolpate s

1. Impatiens capensis Meerb. N spotted touch-me-not. Impatiens biflora Walt.; I. noli-tangere L. ssp. biflora (Walt.) Hultén; I. nortonii Rydb. • CT, MA, ME, NH, RI, VT. Mesic to wet-mesic, commonly shaded, soil of deciduous forests, stream banks, ditches, and swamps. Impatiens capensis and I. pallida are often found vegetative or in fruit and, therefore, lack obvious means of identification. Impatiens capensis typically has longer-petioled upper leaves (i.e., petioles longer than 1 cm on all fully expanded leaves) and the leaf blades are flat. Impatiens pallida usually has the upper few leaves shortpetiolate (i.e., petioles shorter than 1 cm), and these blades are usually folded downward about the basal portion of the midrib. Compare these features on flowering individuals to learn the vegetative characters. 2. Impatiens glandulifera Royle E Himalaya touch-me-not. Impatiens roylei Walp. • MA, ME, VT. Mesic to wet-mesic soil of ditches, field edges, banks, and waste areas. 3. Impatiens pallida Nutt. N Fig. 478 pale touch-me-not. CT, MA, ME, NH, RI, VT; most frequent in western New England. Mesic to wetmesic, usually shaded, soil of deciduous forests and stream banks. Fig. 478  Flower of Impatiens pallida.

Berberidaceae 1a. Plants woody, with stipular spines [Fig. 479]; perianth yellow . . . . . . . . . . . . . . . . . . . Berberis 1b. Plants herbaceous, without spines; perianth white to pink or green-yellow, green-purple, purple, or purple-brown (rarely yellow) 2a. Perianth 4-merous; leaves pinnately compound with (2–) 9 leaflets that are asymmetrically cordate at the base; inflorescence a raceme or panicle . . . . . . . Epimedium 2b. Perianth 5- to 9-merous [Fig. 480]; leaves palmately lobed or pinnately compound and then with leaflets that are cuneate, rounded, or truncate at the base (sometimes asymmetrically so); inflorescence a solitary flower or cyme 3a. Leaf blades lobed, the basal ones peltate; inflorescence a solitary flower; perianth white or pink; anthers dehiscing by longitudinal slits; ovary maturing as a berry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podophyllum 3b. Leaf blades 2–4 times pinnately compound, basifixed to the petioles; inflorescence a cyme; perianth green-yellow, green-purple, purple, or purple-brown (rarely yellow); anthers dehiscing by uplifting valves; ovary ruptured by the 2 enlarging seeds, the stalked, blue seeds ripening fully exposed . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caulophyllum

Berberis 1a. Leaves pinnately compound with 5–9 leaflets; plants lacking spines; berries blue, glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. aquifolium 1b. Leaves simple; plants spinose [Fig. 479]; berries red to purple-red, not glaucous 2a. Margins of leaves entire; spines simple; inflorescence a solitary flower or an umbel-like cluster of 2–4 flowers; berries rather dry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. thunbergii 2b. Margins of leaves toothed; some of the spines compound (the upper usually simple in B. koreana); inflorescence a raceme of 10–25 flowers [Fig. 479]; berries fleshy 3a. Berry obloid to obloid-ovoid; branchlets gray . . . . . . . . . . . . . . . . . . . . . . . . . B. vulgaris 3b. Berry subglobose; branchlets red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. koreana

Be r b e r i dac e a e   449

1. Berberis aquifolium Pursh E holly-leaved Oregon-grape. Mahonia aquifolia (Pursh) Nutt. • VT. Gardens, forest edges and fragments. 2. Berberis koreana Palibin. E Korea barberry. VT. Fields, lawns, forest edges. 3. Berberis thunbergii DC. E Japanese barberry. CT, MA, ME, NH, RI, VT. Fields, forests, especially riparian types, disturbed soil, areas of habitation. ‌3 × 4. Berberis ×ottawensis Schneid. is an uncommon barberry hybrid known from CT, MA, ME, VT. It combines characteristics of its parents. The leaf blades are typically entire, but the lower spines of a given branch are often trifid. The inflorescence is a raceme with a short axis. 4. Berberis vulgaris L. E Fig. 479 common barberry. CT, MA, ME, NH, RI, VT. Fields, pastures, forests, roadsides.

Caulophyllum

Fig. 479  Inflorescence and spine of Berberis vulgaris.

References: Haines (2003c), Loconte (1997). 1a. Flowers subprecocious, numbering mostly 4–18 per inflorescence; sepals mostly purple to purple-brown (rarely yellow), 6–9 mm long; filaments 1.5–2.5 mm long; carpels 3–5 mm long; styles 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. giganteum 1b. Flowers nearly coetaneous, numbering mostly 5–70 per inflorescence; sepals yellow to green, often tinged with purple (rarely purple to purple-brown), 3–6 mm long; filaments 0.5–1.5 mm long; carpels 1–3 mm long; styles 0.25–1 mm long . . . . . . . . . . . . . . C. thalictroides 1. Caulophyllum giganteum (Farw.) Loconte & Blackwell N Fig. 480 early blue cohosh. Caulophyllum thalictroides (L.) Michx. var. giganteum Farw. • CT, MA, NH, VT. Mesic, deciduous forests, often rich and rocky, high-terrace floodplain forests. Caulophyllum giganteum flowers ca. 10–15 days ealier than C. thalictroides when both are present at the same site. The two species differ somewhat vegetatively. The ultimate leaf segments of C. giganteum are 5–10 cm long and first leaf is 2- or, more commonly, 3-times pinnately divided. The ultimate leaf segments of C. thalictroides are 3–8 cm long and the first leaf is 3- or, more commonly, 4-times pinnately divided. 2. Caulophyllum thalictroides (L.) Michx. N blue cohosh. CT, MA, ME, NH, RI, VT. Mesic, deciduous forests, often rich and rocky, high-terrace flood plain forests.

Epimedium Epimedium ‌ diphyllum Lodd. × E. grandiflorum Morr. Epimedium ×youngianum Fisch. & C.A. Mey. is a rare Bishop’s-hat hybrid known from MA that has escaped from gardens to lawns and forest edges. It is readily identified by the characters presented in the identification key to the genera.

Podophyllum 1. Podophyllum peltatum L.

nC

May-apple. CT, MA, ME, NH, RI, VT. Deciduous forests, fields, along fence rows and stone walls, stream banks, forest borders. Records of this plant in New England appear to be a mixture of native and introduced occurrences. Native occurrences appear restricted to Litchfield County, CT, Berkshire County, MA, and Addison and Bennington Counties, VT; however, those counties also contain populations that are clearly naturalized. Because it is sometimes difficult to determine which populations that are truly native, conservation of this species is complicated.

Fig. 480  Flower of Caulophyllum giganteum.

45 0   tricolpate s

Betulaceae References: Furlow and Mitchell (1990), Furlow (1997). 1a. Leaf blades abaxially with black glands; mature stems with gray bark and slightly raised, longitudinal ridges; involucre a hastate-shaped, leaf-like bract; winter buds quadrangular in cross-section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carpinus 1b. Leaf blades without black glands (though resin or resin glands may be present); mature stems with smooth or exfoliating bark, variously colored (rarely uniformly gray), without longitudinal ridges; involucre a firm or woody scale, an enclosing husk, or an inflated, bladder-like bract; winter buds ± terete in cross-section 2a. Involucre an inflated, bladder-like bract, the carpellate inflorescence resembling a hop-like cluster; mature stems with gray-brown bark that exfoliates in vertical strips; winter bud scales conspicuously longitudinally striate . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ostrya 2b. Involucre a husk or a flat scale, the carpellate inflorescence resemling a cluster of fruits, an ament, or a cone; mature stems with variously colored bark, but this not exfoliating in vertical strips (though sometimes exfoliating and curling in horizontal strips or patches); winter bud scales not, or scarcely, longitudinally striate 3a. Fruit a nut 10–15 mm long, enclosed in an husk-like involucre 15–70 mm long [Fig. 487]; staminate flowers without perianth; anthers pilose at the tip . . . . . . . . Corylus 3b. Fruit a samara 1.2–4.5 mm long, subtended by, but not enclosed in, flat scales 2.5–13 mm long; staminate flowers with a minute calyx; anthers glabrous 4a. Carpellate scales thin, decicuous with the fruit; androecium with 4 monothecal half-anthers; winter buds sessile; phyllotaxis ⅖ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Betula 4b. Carpellate scales thick, woody, long-persistent after the fruits are shed; androecium with 4 dithecal anthers; winter buds stalked (nearly sessile in A. viridis); phyllotaxis ⅓ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alnus

Alnus Hybridization is known to occur between some species in New England. Steele (1961) described introgressed plants he observed in New Hampshire. Reference: Furlow (1979). 1a. Trees to 20 m tall; leaf blades obovate to suborbicular, rounded or retuse to obcordate at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. glutinosa 1b. Shrubs or small trees to 10 m tall; leaf blades elliptic or narrow-ovate to broad-elliptic, broad-ovate, or obovate, rounded to acute at the apex 2a. Winter buds sessile or on stalks up to 1 mm, with 3–6 scales of unequal sizes, acute to acuminate at the apex, glabrous and glutinous; samara with a thin wing; only the staminate aments emerging from naked buds; peduncles of carpellate infructescences often subtended by leaf-like bracts, 0.5–1.2 mm thick in fruit . . . . . . . . . . . . . . . . . . . . . . . . . A. viridis 2b. Winter buds on stalks 2–4 mm long, with 2 or 3 scales of equal size, rounded to acute at the apex, scurfy-pubescent; samara merely thin-edged; staminate and carpellate aments emerging from naked buds; peduncles of carpellate infructescences usually without subtending bracts, 0.8–1.5 (–2) mm thick in fruit 3a. Leaf blades coarsely double-serrate, narrow-ovate to elliptic, with conspicuous cross-veins on the abaxial surface, slightly or not at all resinous and frequently glaucous later in the season abaxially, broad-cuneate to rounded or subcordate at the base; bark with evident, white lenticels; branchlet supporting the staminate aments usually weakly to moderately diverging from the inflorescence axis [Fig. 481] . . . . . . . . . . . . . . . A. incana

Be t u l ac e a e   45 1

3b. Leaf blades finely and regularly serrulate to somewhat double-serrate, obovate to broad-elliptic, with delicate or inconspicuous cross-veins on the abaxial surface, slightly to moderately resinous and without bloom abaxially, cuneate (rarely to rounded) at the base; bark with fewer, darker lenticels; branchlet supporting the staminate aments usually strongly diverging from the inflorescence axis [Fig. 482] . . . . . . . . . . . A. serrulata 1. Alnus glutinosa (L.) Gaertn. E European alder. Alnus alnus (L.) Britt.; Betula alnus L. var. glutinosa L. • CT, MA, ME, RI, VT. Pond shores, river banks, human-constructed wetlands. This species has recently (and accidentally) been planted in several human-constructed wetlands in ME. It was believed the nusery stock was Alnus incana. 1 × 3. This very rare alder hybrid is known from MA. In habit is resembles Alnus glutinosa (i.e., small trees). However, the leaf blades are rounded or sometimes shortly truncate at the apex, not showing the typical notched apex on many of the leaves. 2. Alnus incana (L.) Moench ssp. rugosa (Du Roi) Clausen N Fig. 481 speckled alder. Alnus incana (L.) Moench var. americana Regel; A. rugosa (Du Roi) Spreng.; A. rugosa (Du Roi) Spreng. var. americana (Regel) Fern.; Betula alnus L. var. rugosa Du Roi • CT, MA, ME, NH, RI, VT. Swamps, shorelines, wetland edges, borders of low fields. ‌2 × 3. Alnus ×fallacina Callier is a rarely collected alder hybrid to be expected wherever the parental taxa co-occur. It is intermediate in morphology; however, given the slight differences between the progenitors, it is subtle and often difficult to determine (especially on herbarium sheets).

Fig. 481  Inflorescence of Alnus incana showing weak divergence of branchlets supporting staminate aments.

3. Alnus serrulata (Ait.) Willd. N Fig. 482 smooth alder. Alnus incana (L.) Moench var. serrulata (Ait.) Boivin; A. noveboracensis Britt.; A. rugosa (Du Roi) Spreng. var. serrulata (Ait.) H.J.P. Winkl.; A. serrulata (Ait.) Willd. var. subelliptica Fern.; Betula serrulata Ait. • CT, MA, ME, NH, RI, VT. Shorelines, swamps. Variety subelliptica may represent introgressed plants with gene exchange from Alnus incana. 4. Alnus viridis (Vill.) Lam. & DC. ssp. crispa (Ait.) Turrill N green alder. Alnus crispa (Ait.) Pursh; A. crispa (Ait.) Pursh var. mollis (Fern.) Fern.; A. mollis Fern.; A. viridis (Vill.) Lam. var. crispa (Ait.) House; Betula crispa Ait. • MA, ME, NH, VT; restricted in MA to the western half of the state. Regenerating clearings, swamps, shorelines, slopes, ravines, talus, common along the ME coastline and ascending high into the mountains.

Betula Measurements of carpellate aments provided in the identification key are from infructescences that are fully mature and beginning to disarticulate. Measurements (especially width) taken from before this stage of maturation will be too narrow and cause problems in the key. Betula platyphylla Sukaczev has been attributed to MA on the basis of a collection from along the banks of the Neponset River—Svenson 11746 (GH!). However, the specimen is clearly misidentified and has leaf blades that are of the wrong shape, are too large, and have too many pairs of lateral veins (among other discrepancies). Though the identity of the specimen is still in question, it is superficially similar to B. papyrifera. 1a. Leaf blades crenate to crenate-dentate with rounded or bluntly pointed teeth, obovate to orbicular, rounded to obtuse at the apex [Fig. 484] 2a. Leaf blades 0.5–3 cm long; branchlets with abundant, large resin glands; scales of carpellate ament with upturned lateral lobes; plants of alpine ridges and plateaus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. glandulosa 2b. Leaf blades mostly 2.5–5 (–7) cm long; branchlets without resin glands or with few, small resin glands, these most abundant near the nodes; scales of carpellate ament with widely divergent lateral lobes; plants of fens and swamps . . . . . . . . . . . . . . . . . . . . . B. pumila

Fig. 482  Inflorescence of Alnus serrulata showing strong divergence of branchlets supporting staminate aments.

45 2   tricolpate s

1b. Leaf blades obscurely to evidently doubly serrate with pointed teeth (the teeth often blunt in B. minor), ovate, rhombic, or triangular to narrow-ovate or oblong-ovate, acute to long-acuminate at the apex 3a. Leaf blades ovate to narrow-ovate or oblong-ovate, the larger with 12–18 pairs of lateral veins; fresh branchlets with wintergreen odor; carpellate aments ± sessile; body of samara wider than the individual wings 4a. Bark exfoliating, yellow to silver; scales of carpellate aments pubescent and/or ciliate, 6–13 mm long; leaf blades somewhat coarsely, and often irregularly, serrate, usually with fewer than 6 teeth per cm; branchlets often pubescent. . . . . . . . . . . . . . . . B. alleghaniensis 4b. Bark smooth, not exfoliating, brown; scales of carpellate aments glabrous, 5–7 mm long; leaf blades finely and regularly serrate, usually with 6 or more teeth per cm; branchlets often glabrous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. lenta 3b. Leaf blades ovate or narrow-ovate to rhombic or triangular, the larger with 2–12 pairs of lateral veins; fresh branchlets lacking wintergreen odor; carpellate aments borne on peduncles 5–15 mm long; body of samara as wide as or narrower than the individual wings (wider than wings in B. nigra and B. pubescens) 5a. Leaf blades triangular or rhombic-triangular to rhombic-ovate, acuminate to longacuminate at apex [Fig. 486], glabrous to sparsely pubescent along and in the axils of major veins; central lobe of carpellate scales much shorter than the lateral lobes 6a. Branches on mature trees usually pendulous; staminate aments mostly in pairs (sometimes solitary or in trios); carpellate aments (19–) 23–40 × (7–) 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. pendula 6b. Branches spreading to ascending; staminate aments mostly solitary (sometimes paired); carpellate aments 10–25 (–30) × 6–8 mm . . . . . . . . . . . . . . . . . . . B. populifolia 5b. Leaf blades ovate to narrow-ovate or rhombic-ovate (to rhombic in B. nigra), obtuse to acute or short-acuminate at the apex, usually sparsely to moderately pubescent along and in the axils of major veins; central lobe of carpellate scales as long as or longer than the lateral lobes 7a. Leaves with rhombic-ovate blades and tomentose petioles; carpellate aments 10–25 mm thick in fruit; body of samara wider than the individual wings . . . . . . B. nigra 7b. Leaves with broad-ovate to narrow-ovate blades (sometimes rhombic-ovate in B. pubescens) and glabrous to pubescent, but not tomentose, petioles; carpellate aments 5–12 mm thick in fruit; body of samara as wide as or narrower than the individual wings 8a. Leaf blades 1.5–5.5 (–6) cm long, obscurely double serrate; mature carpellate aments 10–30 mm long 9a. Native shrubs to 3 m tall of alpine ravines and plateaus; bark dark brown; branchlets glabrous to sparsely pubescent with relatively soft hairs; lateral lobes of carpellate scales ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. minor 9b. Planted trees to 20 m tall, typically near areas of civilization; bark graywhite; branchlets pubescent with short, stiff, erect hairs; lateral lobes of carpellate scales divergent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. pubescens 8b. Leaf blades 5–10 (–14) cm long (sometimes shorter in high-elevation individuals), double serrate; mature carpellate aments 25–55 mm long 10a. Leaf blades cuneate to truncate at the base, with 7–9 pairs of lateral veins; scales of carpellate aments 3.9–6.2 mm long, with divergent, lateral lobes [Fig. 485]; bark of mature trees white (rarely light brown to dark brown) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. papyrifera

Be t u l ac e a e   45 3

10b. Leaf blades cordate at the base, with (8–) 9–12 pairs of lateral veins (with fewer pairs in dwarfed, high-elevation individuals); scales of carpellate aments 5.6–8.7 mm long, with upturned, lateral lobes [Fig. 483]; bark of mature trees pink-white to brown-white or red-brown-white . . . . . . . . . . . . . . . . . B. cordifolia 1. Betula alleghaniensis Britt. N yellow birch. Betula alleghaniensis Britt. var. fallax (Fassett) Brayshaw; B. alleghaniensis Britt. var. macrolepis (Fern.) Brayshaw; B. lutea Michx. f.; B. lutea Michx. f. var. fallax Fassett; B. lutea Michx. f. var. macrolepis Fern. • CT, MA, ME, NH, RI, VT; nearly throughout. Deciduous forests, swamps. 2. Betula cordifolia Regel N Fig. 483 heart-leaved paper birch. Betula alba L. var. cordifolia (Regel) Regel; B. papyrifera Marsh. var. cordifolia (Regel) Fern. • CT, MA, ME, NH, VT. North-temperate to subalpine deciduous and mixed evergreen-deciduous forests, alpine ravines and plateaus, talus slopes, maritime forests. ‌ × 9. Betula ×caerulea Blanch. is an infrequent birch hybrid often referred to as the 2 blue birch. It is a tree attaining heights to 20 m with exfoliating cream-white to pink-white bark. The ovate-triangular leaf blades are 6–10 cm long, have an acuminate apex, and have (6–) 7–9 pairs of lateral veins (vs. (8–) 9–12 in B. cordifolia and 5–7 in B. populifolia). The carpellate aments are 25–50 mm long at maturity (vs. 10–25 (–30) mm in B. populifolia) and have scales with divergent lateral lobes (vs. ascending lobes in B. cordifolia). Collections of this nothospecies were incorrectly referred to as B. pendula and B. pendula var. japonica Rehd. by many early collectors (due to misapplication of names; see Fernald 1922b). This has produced many mislabeled collections in regional herbaria. 3. Betula glandulosa Michx.

Fig. 483  Carpellate scale of Betula cordifolia.

N C Fig. 484

glandular birch. ME, NH. Alpine plateaus, ravines, and snow bank communities. This species was once collected at ca. 485 m elevation in an open field on the slope of a large hill in Coos County, NH. 4. Betula lenta L. N cherry birch. CT, MA, ME, NH, RI, VT. Deciduous or mixed evergreen-deciduous forests. 5. Betula minor (Tuckerman) Fern.

Fig. 484  Leaves of Betula glandulosa.

NC

dwarf birch. Betula papyracea Ait. var. minor Tuckerman; B. pubescens Ehrh. ssp. minor (Tuckerman) A. & D. Löve; B. saxophila Lepage • ME, NH, vt. Alpine plateaus, slopes, gullies, and ravines. This species is sometimes often considered to be a hybrid-derived species of Betula cordifolia and B. glandulosa, given that it is intermediate in morphology and occurs where those species are sympatric. There may be two taxa involved here. One is characterized by glabrous branchlets and leaf blades and samaras 2.5–5 mm wide with individual wings wider than achene body (B. minor s.s.). The other is characterized by pubescent young branchlets and leaf blades and samaras 2–3.5 mm wide with individual wings up to as wide as the achene body (B. borealis Spach sensu Fernald 1950b; but this name misapplied as it consists of material of B. pumila). I am tentatively following Furlow (1997) in treating B. minor as a single, variable taxon. 6. Betula nigra L.

nC

river birch. CT, MA, NH. River banks, riparian forests, lake shores. Introduced to many locations but likely native to some sites in MA and NH, such as along the Merrimack River (MA, NH) and several ponds in Essex County, MA. Reports of this species in VT (e.g., Seymour 1982) are based on a collection taken from a cultivated plant—Charette 2345 (VT). 7. Betula papyrifera Marsh. N Fig. 485 paper birch. Betula alba L. var. commutata Regel; B. papyrifera Marsh. var. commutata (Regel) Fern.; B. papyrifera Marsh. var. macrostachya Fern. • CT, MA, ME, NH, RI, VT. Deciduous and mixed evergreen-deciduous forests, ascending to moderate elevation (but usually replaced by Betula cordifolia at higher elevations).

Fig. 485  Carpellate scale of Betula cordifolia.

45 4   tricolpate s

‌ × 11. Betula ×sandbergii Britt. is a rare hybrid that occurs in VT. It is a shrub or small 7 tree with dark red-brown, close bark. The oval to ovate leaf blades are 2.5–6.5 cm long with serrate margins, acute at the apex and cuneate at the base. The body of the samara is wider than its individual wings. 8. Betula pendula Roth E European weeping birch. Betula verrucosa Ehrh. • CT, MA, VT. Roadsides, forest fragments, areas of habitation. The leaf blades of Betula pendula and the native B. populifolia are very similar; however, with practice they can sometimes be distinguished on subtle differences in shape. Betula pendula frequently has shorter acuminations at the leaf apex and sometimes shows sharp, forward curving, primary teeth. Note that these differences are not displayed on all individuals. Reports of this species from ME and NH (specimens at GH! and MAINE!) are based on collections from cultivated plants. Many early collections of this species were in fact B. ‌caerulea × (the name B. pendula was misapplied by several authors). 9. Betula populifolia Marsh. N Fig. 486 gray birch. CT, MA, ME, NH, RI, VT. Woodlands, swamps, wetland margins, forests, regenerating fields. 10. Betula pubescens Ehrh. ssp. pubescens E downy birch. Betula alba L. var. pubescens (Ehrh.) Spach • CT, MA, ME, VT; also reported from RI by George (1992), but specimens are unknown. Thickets, forest fragments, roadsides. Fig. 486  Leaf blade of Betula populifolia.

11. Betula pumila L. N bog birch. Betula glandulifera (Regel) Butler; B. glandulosa Michx. var. glandulifera (Regel) Gleason; B. pumila L. var. glandulifera Regel; B. pumila L. var. renifolia Fern. • CT, MA, ME, NH. Fens, Evergreen swamps dominated by Thuja occidentalis, fen-like lake and stream shores in high-pH bedrock regions. Reports of this species from VT are based on hybrids with Betula pumila.

Carpinus 1. Carpinus caroliniana Walt. ssp. virginiana (Marsh.) Furlow N American hornbeam. Carpinus betulus L. var. virginiana Marsh.; C. caroliniana Walt. var. virginiana (Marsh.) Fern.; C. virginiana (Marsh.) Sudworth • CT, MA, ME, NH, RI, VT. River banks, seepage swamps, riparian and deciduous forests.

Corylus A collection of Corylus that was originally determined as C. avellana L. or “near it” was taken from near a nursery in Hartford County, CT—9 Jul 1909, Bissell s.n. (GH!). The specimen has an unmounted annotation that reads “=heterophylla”, a statement that, if applying to the synonymy, is untrue. The plant does not appear to be C. heterophylla Fisch. & Trautv. because it lacks the characteristic leaf blade apex of that species (+/- truncate with a central acumination). The plant also does not appear to be C. avellana because it lacks the leaf blade outline (orbicular to broad-obovate) and has acute stipules. The specimen is vegetative and appears equivocal. 1a. Husk-like involucre 4–7 cm long, with a pronounced, tubular beak, bristly with slender spicules; branchlets without red, stipitate glands; staminate aments sessile or on very short peduncles up to 1 mm long; apex of buds acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cornuta 1b. Husk-like involucre 1.5–3 cm long, without a tubular beak [Fig. 487], pubescent but not bristly; branchlets with red, stipitate glands [Fig. 487]; staminate aments on a short, woody peduncles 1–5 mm long; apex of buds obtuse to rounded . . . . . . . . . . . . . . . . . . . . C. americana Fig. 487  Carpellate involucres and stipitate-glandular branchlet of Corylus americana.

1. Corylus americana Walt. N Fig. 487 American hazelnut. Corylus americana Walt. var. indehiscens Palmer & Steyermark • CT, MA, ME, NH, RI, VT. Forests and forest borders, sandy thickets, fields, hedge rows, roadsides. 2. Corylus cornuta Marsh. ssp. cornuta N beaked hazelnut. Corylus rostrata Ait. • CT, MA, ME, NH, RI, VT. Forests and forest borders.

Be t u l ac e a e   45 5

Ostrya 1. Ostrya virginiana (P. Mill.) K. Koch N hop-hornbeam. Carpinus virginiana P. Mill.; Ostrya virginiana (P. Mill.) K. Koch var. lasia Fern. • CT, MA, ME, NH, RI, VT. Deciduous forests and rich, dry-mesic woodlands and rocky slopes.

Bignoniaceae 1a. Leaves whorled, the blades simple, entire or with short lobes; trees with upright stems; corolla largely white or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Catalpa 1b. Leaves opposite, the blades pinnately divided into 5–13 serrate leaflets; lianas with trailing or climbing stems; corolla largely red or orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campsis

Campsis 1. Campsis radicans (L.) Seem. ex Bureau E trumpet-creeper. Bignonia radicans L.; Tecoma radicans (L.) Juss. • CT, MA; also reported from NH by Kartesz (1999), but specimens are unknown. Field edges, forest fragments, waste areas, sometimes near former dwellings. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this species had questionable naturalization in RI and was unaware of any collections.

Catalpa 1a. Corolla yellow, with orange stripes and purple spots on the adaxial limb, up to 2.5 cm long; leaf blades often 3- to 5-lobed, each lobe terminating in a slender point, usually glabrous on the abaxial surface (varying to pubescent); capsule 5–8 mm thick . . . . . . . . . . . . . . . . . . C. ovata 1b. Corolla white, with yellow stripes and purple-brown spots on the adaxial limb, 3.7–5 cm long; leaf blades unlobed, pubescent on the abaxial surface; capsule 6–15 mm thick 2a. Limb of corolla (2–) 3–4 cm wide, the adaxial surface densely spotted with purplebrown; capsule 6–10 mm thick, with a thin placenta; seed wings typically acute to acuminate at the apices and terminated by a narrow tuft of hairs, the width of the tuft usually less than ½ the width of the seed wing [Fig. 488]; leaf blades distinctly and abruptly acuminate at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bignonioides 2b. Limb of corolla 4–6 cm wide, the adaxial surface sparsely spotted with purple-brown; capsule 10–15 mm thick, with a stout, quadrangular placenta; seed wings typically obtuse to rounded or obliquely truncate at the apices, terminated by a broad fringe of hairs, the fringe usually more than ½ the width of the seed wing [Fig. 489]; leaf blades long-acute to short-acuminate at the apex, the acumination (when present) usually more gradual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. speciosa 1. Catalpa bignonioides Walt. E Fig. 488 southern catalpa. Catalpa catalpa (L.) Karst. • CT, MA, VT; also reported from ME by Kartesz (1999), but specimens are unknown. Roadsides, forest fragments, areas of habitation. Reports of this species from RI are based on a mixed collection (neither of the species represented in the collection are Catalpa bignonioides)—Champlin 1301 (Champlin Herb.).

Fig. 488  Seed of Catalpa bignonioides.

2. Catalpa ovata G. Don E Chinese catalpa. CT, MA, VT. Abandoned pastures, river banks, forest fragments, areas of habitation. 3. Catalpa speciosa (Warder) Warder ex Engelm. E Fig. 489 northern catalpa. CT, MA, ME, NH. Roadsides, forest fragments, riparian forests, areas of habitation.

Fig. 489  Seed of Catalpa speciosa.

45 6   tricolpate s

Boraginaceae Reference: Johnston (1924). 1a. Each carpel of the ovary with 2–many ovules; fruit a 2-valved capsule with 1–80 seeds; at least the lower leaf blades lobed or compound (entire to coarsely toothed in some Phacelia) 2a. Flowers solitary in the axils of opposite the leaves; at least the lower leaves opposite; corolla 5–8 mm long, ± equaled or exceeded by the calyx . . . . . . . . . . . . . . . . . . . . . . . . Ellisia 2b. Flowers arranged in helicoid cymes or subdichotomously branched cymes; leaves alternate; corolla 7–12 mm long, exceeding the calyx 3a. Inflorescence a subdichotomously branched cyme, lacking an elongate central axis; plants from fibrous roots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrophyllum 3b. Inflorescence a helicoid cyme, with 1 or more elongate, sympodial axes; plants from taproots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phacelia 1b. Each carpel of the ovary with 2 ovules; fruit a schizocarp separating into (1–) 4 mericarps; leaf blades simple and mostly entire 4a. Ovary unlobed or inconspicuously lobed; style terminating the summit of the ovary; stigma peltate-annular, surmounted by a cone-like process . . . . . . . . . . . . . . . . Heliotropium 4b. Ovary with deep sinuses, appearing as 4 ± distinct carpels; style arising from between the lobes; stigma not peltate-annular 5a. Corolla rotate [Fig. 492]; anthers 5–9 mm long, with conspicuous linear appendages ca. 3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Borago 5b. Corolla tubular, funnelform, salverform, or umbraculiform [Figs. 490, 494, 502]; anthers shorter, without appendages 6a. Corolla weakly zygomorphic [Fig. 494], the tubular, connate portion sometimes also bent 7a. Apex of corolla tube lacking fornices, the tube appearing open; stamens exserted beyond the corolla; mericarps without a stipe-like process, attaching directly to the receptacle; receptacle flat, without a pit . . . . . . . . . . . . . . . . . . Echium 7b. Apex of corolla tube with fornices, these appearing to close off the tube; stamens included within the corolla tube; mericarps attached to the receptacle by a stipe-like process, this fitting into a pit in the receptacle . . . . . . (in part) Anchusa 6b. Corolla actinomorphic [Figs. 491, 502, 503], the tubular portion straight 8a. Ovary and mericarps with barbed prickles, at least on the marginal or abaxial surfaces [Figs. 493, 495] 9a. Plants annual; each flower closely subtended by a bract . . . . . . . . . Lappula 9b. Plants biennial or perennial; all or most of the flowers without closely associated bracts 10a. Corolla 5–8 mm wide; mericarps attached to one another by the apical third of their inner faces; mericarps 3.5–5 mm long . . . . . . . . . . Cynoglossum 10b. Corolla 2–3 mm wide; mericarps attached to one another along the middle third of their inner faces; mericarps 2–3 mm long . . . . . . . . . . Hackelia 8b. Ovary and mericarps smooth or textured, sometimes muricate, but never with barbed prickles 11a. Stems retrorsely prickly hispid [Fig. 491], weak and scrambling; inflorescence axillary, of 1–3 flowers borne on short, recurved pedicels; calyx accrescent in

Bo r ag i n ac e a e   4 57

fruit, becoming 10–20 mm wide, compressed, firm, and evidently reticulate-veiny, with dentate margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asperugo 11b. Stems glabrous to hispid-pubescent, but not retrorsely prickly hispid, weak and sprawling to rigid and erect; inflorescence of 1 or more helicoid cymes or consisting of crowded flowers in the upper leaf axils in Lithospermum and Buglossoides; calyx not notably accrescent in fruit, ± circular in cross-section, not or scarcely veiny, with entire margins 12a. Corolla yellow to orange, the apex of the tube without fornices (though the tube obstructed by pubescent bulges near the orifice in Amsinckia lycopsoides) 13a. Corolla salverform, with wide-spreading, rounded to obtuse lobes [Fig. 490]; style included within tubular portion of corolla [Fig. 500] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amsinckia 13b. Corolla ± tubular, with erect, acuminate lobes [Fig. 500]; style evidently exserted from the corolla [Fig. 500] . . . . . . . . . . . . Onosmodium 12b. Corolla white, yellow-white, pink, red-purple, blue, purple, or brown (pale yellow in Symphytum tuberosum, with a yellow center in some Myosotis), the apex of the tube with fornices (lacking fornices in Buglossoides arvensis and Mertensia virginica) 14a. Lower 1–4 nodes of the stem producing opposite leaves 15a. Mericarp attachment scar grooved on inner median surface, the groove flaring open or forking at the base of the mericap, without a raised keel apical to the attachment scar; basal leaves usually present during early flowering (these early deciduous) . . . . . . . . . . . Cryptantha 15b. Mericarp with a raised attachment scar on the inner median surface, with a raised keel apical to the attachment scar; basal leaves absent, all leaves borne on the stem . . . . . . . . . . . . . . . . . Plagiobothrys 14b. Stem leaves all alternate 16a. Corolla tubular or narrow-umbraculiform, the lobes erect to ascending [Figs. 502, 503] 17a. Leaf blades conspicuously white-spotted; stems with stipitate glands; tube of corolla obstructed by 5 tufts of hairs . . . . Pulmonaria 17b. Leaf blades without white spots; stems lacking stipitate glands; tube of corolla obstructed by 5 scales or lacking fornices altogether (with hairs at the base of the corolla tube in Nonea) 18a. Leaf blades and stems glabrous; mericarps without a stipelike process, attaching directly to the receptacle; receptacle flat, without a pit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mertensia 18b. Leaf blades and stems hispid-hirsute; mericarps attached to the receptacle by a stipe-like process, this fitting into a pit in the receptacle 19a. Style exserted beyond the corolla; fornices present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symphytum 19b. Style not exserted beyond the corolla; fornices absent, the tube of the corolla obstructed by tufts of hairs . . . . Nonea 16b. Corolla salverform or funnelform, the lobes spreading [Figs. 498, 499]

45 8   tricolpate s

20a. Flowers, except sometimes the lower, not subtended by bracts; corolla lobes convolute in bud (i.e., arranged so that one margin is exposed and the other margin covered by the adjacent petal); mericarps smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myosotis 20b. Flowers all or mostly all subtended by bracts; corolla lobes imbricate in bud (i.e., arranged so that the outer members have both margins exposed, the inner members have both margins covered); mericarps wrinkled or pitted, or smooth in Lithospermum 21a. Limb of the corolla 6–11 mm wide; mericarps attached to the receptacle by a stipe-like process, this fitting into a pit in the receptacle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Anchusa 21b. Limb of the corolla up to 4 mm wide; mericarps without a stipe-like process, attaching directly to the receptacle; receptacle flat, without a pit 22a. Plants perennial; corolla 4–5 mm long; leaves with 2 or 3 conspicuous, lateral veins; mericarps lustrous, white to whitebrown, smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lithospermum 22b. Plants annual (rarely biennial); corolla 5–8 mm long; leaves without conspicuous, lateral veins; mericarps dull brown, wrinkled and pitted or roughened . . . . . . Buglossoides

Amsinckia Amsinckia douglasiana A. DC. has been erroneously attributed to Ma and Me due to misapplication of names (i.e., the name A. douglasiana was formerly misapplied to plants of A. menziesii var. intermedia) and misidentifications. This species has actually not been collected in New England. Amsinckia spectabilis Fisch. & C.A. Mey. var. spectabilis has been attributed to Middlesex County, MA, by several sources (e.g., Kartesz 1999, Sorrie and Somers 1999). The reports were based on the following specimen—1878, Alcott s.n. (NEBC!). The specimen was incorrectly determined and is Amsinckia menziesii var. menziesii. Reference: Ganders (1993). 1a. Corolla tube obstructed by pubescent bulges near the orifice; anthers and style included within connate tube of corolla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. lycopsoides 1b. Corolla tube open; anthers and style exserted beyond orifice of connate tube of corolla (but not exserted beyond lobes of corolla) 2a. Corolla 4–11 mm long, the limb 2–10 mm wide at the apex; anthers usually appressed to stigma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. menziesii 2b. Corolla 10–20 mm long, the limb 8–14 mm wide at the apex; anthers not appressed to stigma, the stigma elevated above the anthers . . . . . . . . . . . . . . . . . . . . . . . . . A. eastwoodiae 1. Amsinckia eastwoodiae J.F. Macbr. E Eastwood’s fiddleneck. Amsinckia intermedia Fisch. & C.A. Mey. var. eastwoodiae (J.F. Macbr.) Jepson & Hoover • MA. Waste areas, wool waste. The specimen supporting this record shows atypical calyx pubescence for this species—the hairs are entirely white (they usually are, in part, brown) and finer hairs beneath the course, stiff hairs are essentially lacking (they are usually present). 2. Amsinckia lycopsoides Lehm. E bugloss fiddleneck. Amsinckia barbata Greene; Benthamia lycopsoides (Lehm.) Lindl. ex Druce • CT, MA, NH. Roadsides, waste areas, wool waste. Fig. 490  Inflorescence of Amsinckia menziesii var. intermedia.

3. Amsinckia menziesii (Lehm.) A. Nels. & J.F. Macbr. E Fig. 490 rancher’s fiddleneck.  3a. Amsinckia echinata Gray; A. intermedia Fisch. & C.A. Mey.; A. intermedia Fisch. & C.A. Mey. var. echinata (Gray) Wiggins; 3b. Amsinckia micrantha

Bo r ag i nac e a e   45 9

Suksdorf; A. parviflora Heller; A. retrorsa Suksdorf; A. rugosa Rydb.; Echium menziesii Lehm. • CT, MA, ME. Roadsides, waste areas, yards, wool waste.

1a. Corolla 7–11 mm long, the limb 4–10 mm wide in life and usually with 5 red-orange marks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. A. menziesii var. intermedia (Fisch. & C.A. Mey.) Ganders 1b. Corolla 4–7 mm long, the limb 2–3 mm wide in life and lacking red-orange marks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. A. menziesii var. menziesii Variety intermedia is known from CT, MA. At least some early collections of this plant were later considered to Amsinckia lycopsoides (Chamberlain 1901). Variety menziesii is known from CT, MA, ME, NH.

Anchusa 1a. Corolla zygomorphic, the basal tube bent; mericarps obliquely oriented in life, 1–2 mm tall; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. arvensis 1b. Corolla actinomorphic, the basal tube straight; mericarps obliquely oriented or erect in life, 2–9 mm tall; plants perennial 2a. Limb of corolla 12–20 mm wide in life; mericarps erect, 5–9 mm tall; bracts subtending flowers with a narrow-lanceolate blade; calyx lobes narrow-lanceolate . . . . . . . . . A. azurea 2b. Limb of corolla 5–10 mm wide in life; mericarps obliquely oriented, ca. 2 mm tall; bracts subtending flowers with an ovate to oblong blade; calyx lobes narrow-triangular to lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. officinalis 1. Anchusa arvensis (L.) Bieb. E small bugloss. Lycopsis arvensis L. • CT, MA, ME, NH, RI. Roadsides, railroads, disturbed soil. 2. Anchusa azurea P. Mill. E Italian bugloss. Anchusa italica Retz. • CT, MA, NH. Roadsides, fields, disturbed soil. 3. Anchusa officinalis L. E common bugloss. Anchusa procera Bess. ex Link • CT, MA, ME, NH, RI. Roadsides, fields, dumps, disturbed soil.

Asperugo 1. Asperugo procumbens L. E Fig. 491 German madwort. CT, MA. Fields, roadsides, waste areas.

Fig. 491  Upper stem, leaves, and flowers of Asperugo procumbens.

Borago 1. Borago officinalis L. E Fig. 492 borage. CT, MA, ME, NH, RI, VT. Fields, roadsides, compost heaps, gardens.

Buglossoides 1. Buglossoides arvensis (L.) I.M. Johnson ssp. arvensis E corn-gromwell. Lithospermum arvense L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, waste areas.

Cryptantha 1. Cryptantha ambigua (Gray) Greene E basin cat’s-eye. Krynitzkia ambigua Gray • MA. Wool waste, disturbed soil.

Fig. 492  Flower of Borago officinalis.

460  tricolpates

Cynoglossum 1a. Inflorescence consisting of (1–) 3 (–4) helicoid cymes borne on a common, terminal peduncle (i.e., none borne in the axils of leaves); mericarps ascending, concealing the persistent style at maturity; rounded on the outer surface [Fig. 493], without a raised, marginal rim . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. virginianum 1b. Inflorescences of several helicoid cymes, the lower borne in the axils of the upper leaves and on short axillary branches; mericarps horizontally oriented and not concealing the persistent style at maturity, ± flattened on the outer surface, with a raised, marginal rim 2a. Corolla red-purple (rarely white); calyx 3–5 mm long at anthesis . . . . . . . . . C. officinale 2b. Corolla blue to blue-purple (rarely white); calyx 2–3 mm long at anthesis 3a. Limb of corolla 8–10 mm wide; leaf blades gray-green, densely pubescent; inflorescences numerous, panicle-like; mericarps 4–5 mm long . . . . . . . . . . . . . C. amabile 3b. Limb of corolla 2.5–4.5 mm wide; leaf blades green to gray-green, sparsely pubescent; inflorescences fewer, less crowded, raceme-like, paniculate; mericarps 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. microglochin 1. Cynoglossum amabile Stapf & Drummond E Chinese hound’s-tongue. MA, NH. Fields, roadsides, gardens, yards. 2. Cynoglossum microglochin Bentham E small-bristled hound’s-tongue. MA. Dumps, compost heaps. 3. Cynoglossum officinale L. E common hound’s-tongue. CT, MA, NH, RI, VT; also reported from ME by Johnston (1924), but specimens are unknown. Fields, pastures, waste areas, wool waste. Johnston’s report of this species in ME is without an exclamation point (i.e., he did not see a voucher specimen from ME). 4. Cynoglossum virginianum L.

Fig. 493  Schizocarp of Cynoglossum virginianum ssp. virginianum showing glochidiate prickles.

N C Fig. 493

wild hound’s-tongue.  4a. Cynoglossum boreale Fern.; C. virginianum L. var. boreale (Fern.) Cooperrider • CT, MA, ME, NH, VT. Deciduous and mixed evergreen-deciduous forests, sometimes associated with disturbances such as trails and old logging roads. 1a. Limb of corolla 5–8 mm wide in life, the lobes oblong and not overlapping; some stem leaves with a petiole; calyx 2–3 mm long at anthesis; mericarps 3.5–5 mm long; northern taxon, extending as far south as northern Connecticut . . . . . . . . . 4a. C. virginianum ssp. boreale (Fern.) A. Haines 1b. Limb of corolla 8–12 (–16) mm wide in life, the lobes suborbicular and overlapping; leaves broad at the base, clasping or sessile; calyx 3–4 mm long at anthesis; mericarps 6–8 mm long; southern taxon, reaching as far north as southwestern Connecticut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. C. virginianum var. virginianum Subspecies boreale is known from CT, MA, ME, NH, VT. Subspecies virginianum is known from CT.

Echium 1a. Stamens not or scarcely exserted from the corolla, not exceeding the longer corolla lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. plantagineum 1b. Stamens conspicuously exserted from the corolla, surpassing the lobes [Fig. 494] 2a. Pubescence of stem coarse, spreading, with an evident, pustulose base . . . E. pustulatum 2b. Pubescence of stem firm, bristle-like, but without a pustular base 3a. Lower branches of the inflorescence 2.5–6 cm long, bearing flowers from base to apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. vulgare 3b. Lower branches of the inflorescence 12–22 cm long, not bearing flowers in the basal portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. creticum

Bo r ag i n ac e a e   46 1

1. Echium creticum L. E Cretan viper’s-bugloss. Echium australe Lam. • MA, ME. Fields, roadsides, disturbed soil, dumps. 2. Echium plantagineum L. E plantain viper’s-bugloss. MA. Fields, roadsides, disturbed soil. 3. Echium pustulatum Sibthorp & Sm. E pustulose viper’s-bugloss. Echium vulgare L. var. pustulatum (Sibthorp & Sm.) Coincy • MA. Fields, roadsides, disturbed soil. 4. Echium vulgare L. E Fig. 494 common viper’s-bugloss. CT, MA, ME, NH, RI, VT. Fields, roadsides, disturbed soil, railroads, wool waste. Fig. 494  Flower of Echium vulgare.

Ellisia 1. Ellisia nyctelea (L.) L. E Aunt Lucy. Nyctelea nyctelea (L.) Britt. • CT, MA. Fields, roadsides, pastures.

Hackelia 1a. Mericarps with a single row of glochids on each margin, sometimes with 1–3 (–10) small glochids on the outer surface; intact schizocarp broad-pyramidal; basal leaf blades narrowovate to oblanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. deflexa 1b. Mericarps with 10–25 glochids on the outer surface that are of similar length to the marginal glochids [Fig. 495]; intact schizocarp subglobose; basal leaf blades broad-ovate to cordate-ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. virginiana 1. Hackelia deflexa (Wahlenb.) Opiz ssp. americana (Gray) A. & D. Löve

NC

nodding stickseed. Hackelia americana (Gray) Fern; H. deflexa (Wahlenb.) Opiz var. americana (Gray) Fern. & I.M. Johnston; Lappula americana (Gray) Rydb.; L. deflexa (Gray) Rydb. ssp. americana (Gray) Hultén; L. deflexa (Gray) Rydb. var. americana (Gray) Greene • ME, NH, VT. Rocky forests and cliff bases, in regions of high-pH bedrock. 2. Hackelia virginiana (L.) I.M. Johnson N Fig. 495 Virginia stickseed. Lappula virginiana (L.) Greene; Myosotis virginiana L. • CT, MA, NH, VT; also reported from RI by Magee and Ahles (1999), but specimens are unknown. Mesic, deciduous forests, talus, cliff bases, usually in regions of high-pH bedrock. Reports of this species in ME by Fernald (1950b) are erroneous and based on Hackelia deflexa.

Heliotropium 1a. Leaf blades linear to narrow-oblanceolate, 2–5 mm wide, glabrous [Fig. 496]; calyx and axis of inflorescence glabrous [Fig. 496] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. curassavicum 1b. Leaf blades lanceolate or oblanceolate to elliptic, ovate, or triangular-ovate, the principal ones 10–87 mm wide, pubescent; calyx and axis of inflorescence hirsute 2a. Schizocarp 4-lobed prior to separation, splitting into 4 mericarps at maturity; corolla white (less frequently blue) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. europaeum 2b. Schizocarp 2-lobed prior to separation, splitting into 2 segments, each segment consisting of a pair of cohering mericarps; corolla blue to purple 3a. Principal leaf blades lanceolate to oblanceolate, ca. 15 mm wide, sessile or shortpetioled; plants perennial; mericarps tuberculate; cymes usually solitary at the ends of branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. amplexicaule

Fig. 495  Schizocarp of Hackelia virginiana showing glochidiate prickles.

462 tricolpate s

3b. Principal leaf blades ovate to triangular-ovate, 25–87 mm wide, long-petioled; plants annual; mericarps longitudinally striate; cymes usually 2–5 at the summit of a common peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. indicum 1. Heliotropium amplexicaule Vahl E clasping heliotrope. Cochranea anchusifolia (Poir.) Guerke; Heliotropium anchusifolium Poir. • MA. Roadsides, waste areas, vacant lots, gardens.

2. Heliotropium curassavicum L. var. curassavicum E Fig. 496 seaside heliotrope. ME. Roadsides, waste areas, wool waste. 3. Heliotropium europaeum L. E European heliotrope. MA. Roadsides, waste areas, gardens, wool waste. 4. Heliotropium indicum L. E Indian heliotrope. Tiaridium indicum (L.) Lehm. • MA. Dumps, compost heaps, waste areas. Fig. 496  Inflorescence of Heliotropium curassavicum.

Hydrophyllum 1a. Principal stem leaf blades broad-ovate to broad-triagular in outline, pinnately lobed with deep sinuses that extend nearly to the midrib; calyx lobes ciliate with stiff hairs, not alternating with appendages in the sinuses; peduncles usually longer than the petioles of the subtending leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. virginianum 1b. Principal stem leaf blades ± orbicular in outline, palmately lobed with relatively shallow sinuses [Fig. 497]; calyx lobes usually glabrous, sometimes alternating with minute appendages in the sinuses; peduncles usually shorter than the petioles of the subtending leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. canadense 1. Hydrophyllum canadense L.

N C Fig. 497

blunt-leaved waterleaf. MA, VT. Rich, mesic florests, high-terrace floodplain forests, usually in regions of high-pH bedrock, often associated with cliff bases and streams. 2. Hydrophyllum virginianum L. N eastern waterleaf. Hydrophyllum virginianum L. var. atranthum (Alexander) Constance • CT, MA, NH, VT. Mesic, often rich, deciduous forests, riparian forests. Fig. 497  Leaf of Hydrophyllum canadense.

Lappula 1a. Mericarp bodies (i.e., excluding the prickles) 2–3 mm long, with a single row of prickles around the margin, the prickles with more or less confluent bases; calyx 3–3.5 mm long in fruit; corolla 1.5–2.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. occidentalis 1b. Mericarp bodies 3–4 mm long, with 2 (rarely 3) rows of prickles around the margin, the prickles distinct; calyx 2.5–3 mm long in fruit; corolla 2–4 mm wide . . . . . . . . . . . . L. squarrosa 1. Lappula occidentalis (S. Wats.) Greene var. occidentalis E flat-spined sheepburr. Lappula echinata Gilib. var. occidentalis (S. Wats.) Boivin; L. redowskii, auct. non (Hornem.) Greene; L. redowskii (Hornem.) Greene var. occidentalis (S. Wats.) Rydb. • MA. Wool waste, dumps. This species is over-reported and most collections from New England are in fact Lappula squarrosa. 2. Lappula squarrosa (Retz.) Dumort. E bristly sheepburr. Echinospermum lappula (L.) Lehm.; Lappula echinata Gilib.; L. lappula (L.) Karst.; L. squarrosa (Retz.) Dumort. var. erecta (A. Nels.) Dorn • CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas, farm yards, wool waste.

Bo r ag i n ac e a e   463

Lithospermum Lithospermum latifolium Michx. was reported from MA by Magee and Ahles (1999), but specimens are unknown. 1. Lithospermum officinale L. E European gromwell. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas.

Mertensia 1a. Erect or ascending, cultivated plants; flowers 18–25 mm long, not subtended by leafy bracts; leaf blades mostly 4–10 cm long, herbaceous, green; mericarps obtusely angled, dull, becoming wrinkled in drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. virginica 1b. Spreading to prostrate, native plants of Atlantic coast beaches; flowers 6–9 mm long, often subtended by leafy bracts; leaf blades mostly 2–6 cm long, fleshy, glaucous; mericarps acutely angled, shiny, smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. maritima 1. Mertensia maritima (L.) S.F. Gray var. maritima N seaside bluebells. Pneumaria maritima (L.) Hill • MA, ME, NH; rare outside of eastern ME. Atlantic coast beaches, often on cobble and gravel substrate. 2. Mertensia virginica (L.) Pers. ex Link E eastern bluebells. MA, ME, VT. Mesic forests, riparian forests, gardens.

Myosotis Reference: Grau and Merxmüller (1972). 1a. Calyx pubescent with erect to appressed hairs that are not hooked at the tip [Fig. 499]; plants commonly of wet-mesic to hydric soils, even growing in shallow water 2a. Limb of the corolla 2–5 mm wide; mericarps taller than the style; stems terete, not stoloniferous; calyx usually deciduous in fruit, with the lobes approximately as long as or longer than the basal connate portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. laxa 2b. Limb of the corolla 5–10 mm wide; mericarps shorter than the style; stems angled, often stoloniferous; calyx usually persistent in fruit, with the lobes shorter than the basal connate portion (rarely the lobes and connate portion approximately equal) . . . . . . . . . M. scorpioides 1b. Calyx pubescent, at least in part with spreading, uncinate hairs [Fig. 498]; plants of xeric to mesic soils, typically found in fields, disturbed lots, and along roads (though sometimes invading forest floors) 3a. Limb of corolla 1–2 mm wide in life; fruiting pedicels up to 3.5 mm long, shorter than the associated calyx 4a. Calyx 4–7 mm long, weakly zygomorphic, 2 or 3 of the lobes shorter than and separated by a gap from the other ones; corolla white . . . . . . . . . . . . . . . . . . . . . . . M. verna 4b. Calyx 3–5 mm long, ± actinomorphic; corolla usually blue or yellow changing to blue 5a. Inflorescence occupying much of the total height of the plant, the lower flowers scattered among the leaves; adaxial surface of leaf blades (at least along the midvein) with uncinate hairs; style surpassed by the mature mericarps; corolla blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. stricta 5b. Inflorescence occupying about half or less of the total height of the plant, the lower flowers not from the axils of leaves or sometimes the lowest 1 or 2 flowers subtended by leafy bracts; adaxial surface of the leaf blades lacking uncinate hairs (though other types of hairs may be present); style often surpassing the mature mericarps; corolla at first yellow, then usually changing to blue . . . . . . . . . . M. discolor

464 tricolpates

3b. Limb of corolla 2–8 mm wide in life; fruiting pedicels 3–9 mm long, equal in length or longer than the associated calyx 6a. Limb of the corolla 5–8 mm wide, horizontally spreading; plants short-lived perennials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. sylvatica 6b. Limb of the corolla 2–4 mm wide, spreading-ascending [Fig. 498]; plants annual or biennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. arvensis 1. Myosotis arvensis (L.) Hill E Fig. 498 field forget-me-not. Myosotis scorpioides L. var. arvensis L. • CT, MA, ME, NH, RI, VT. Fields, lawns, roadsides, waste areas.

Fig. 498  Flower of Myosotis arvensis showing calyx with uncinate hairs.

2. Myosotis discolor Pers. E yellow-and-blue forget-me-not. Myosotis versicolor (Pers.) Sm. • MA. Fields, roadsides, waste areas. 3. Myosotis laxa Lehm. N smaller forget-me-not. CT, MA, ME, NH, RI, VT. Stream banks and channels, pond shores, seepages. 4. Myosotis scorpioides L. E Fig. 499 water forget-me-not. Myosotis palustris (L.) Hill • CT, MA, ME, NH, RI, VT. Stream banks and channels, pond shores, seepages.

Fig. 499  Flower of Myosotis scorpioides showing calyx with hairs that are not hooked at the apex.

5. Myosotis stricta Link ex Roemer & J.A. Schultes E blue forget-me-not. Myosotis micrantha, auct. non Pallas ex Lehm. • CT, MA, ME, NH, VT. Fields, roadsides, waste areas. This species is responsible for early reports of Myosotis collina Bieb. in New England. 6. Myosotis sylvatica Ehrh. ex Hoffmann E woodland forget-me-not. MA, ME, NH, RI, VT. Fields, lawns, forest fragments, often in areas of habitation. 7. Myosotis verna Nutt. N spring forget-me-not. Myosotis virginica, auct. non (L.) B.S.P. • CT, MA, ME, NH, RI, VT. Ledges, pastures, woodlands, dry, open banks, waste areas.

Nonea 1. Nonea rosea (Bieb.) Link E rose monk’s-wort. ME. Yards, dumps, waste areas.

Onosmodium Onosmodium bejariense DC. ex A. DC. var. hispidissimum (Mackenzie) B.L. Turner was reported from ME and NH by Turner (1995). The report was certainly in error given that (1) Turner’s manuscript was internally inconsistent (e.g., the identification key gave one range, the discussion of the taxon gave another, the map provided gave yet another range) and (2) no Onosmodium has ever been collected north of MA in New England. 1a. Corollas (8–) 11–16 (–20) mm long, the lobes green or white, triangular to broad-triangular, and 1–2 (–3) times as long as wide; midstem leaf blades mostly 20–40 mm wide; mericarps 3.5–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. bejariense 1b. Corollas (7–) 8–10 (–14) mm long, the lobes yellow to yellow-orange, narrow-lanceolate to narrow-triangular, and (2–) 3–4 times as long as wide; midstem leaf blades mostly 10–20 mm wide; mericarps 2.5–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. virginianum 1. Onosmodium bejariense DC. ex A. DC. var. occidentale (Mackenzie) B.L. Turner E soft-haired false gromwell. Onosmodium molle Michx. ssp. occidentale (Mackenzie) Cochrane; O. molle Michx. var. occidentale (Mackenzie) I.M. Johnston; O. occidentale Mackenzie • CT. Waste areas, yards. New England’s only known collection of this species was found in the yard of a rubber reclaiming plant in New Haven County, CT.

Bo r ag i n ac e a e   46 5

2. Onosmodium virginianum (L.) DC. N C Fig. 500 eastern false gromwell. Onosmodium virginianum (L.) DC. var. hirsutum Mackenzie • CT, MA, RI. Sandy beaches, river banks, dry ledges, grasslands, sandy fields, woodlands, disturbed soil.

Phacelia Reference: Wilken et al. (1993). 1a. Lobes of the corolla fimbriately fringed on the margin [Fig. 501]; upper stem leaves conspicuously clasping the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. purshii 1b. Lobes of the corolla not fringed; upper stem leaves not clasping the stem 2a. Principal leaf blades linear to narrow-lanceolate, entire or with 1 or 2 pairs of linear lobes near the base; capsules 5–8 mm tall with 6–15 seeds . . . . . . . . . . . . . . . . . . . . P. linearis 2b. Principal leaf blades lanceolate or oblanceote to ovate or orbicular, toothed to bipinnately lobed (the stem leaves usually entire in P. egena); capsules 2–5 mm tall with 1–25 seeds or 7–13 mm tall with 30–80 seeds 3a. Plants perennial; stems pubescent with eglandular hairs only; principal stem leaf blades 10–25 cm long, usually entire; capsule with 1 or 2 seeds . . . . . . . . . . . . . . . P. egena 3b. Plants annual; stems pubescent with both glandular and eglandular hairs; principal stem leaf blades 1–10 (–15) cm long (or to 20 cm in P. tanacetifolia), usually toothed to lobed; capsule with 1–80 seeds 4a. Pedicels 2–15 (–20) mm long; corolla 8–40 mm long; capsules 7–13 mm tall with 30–80 seeds; leaf blades ovate to orbicular, simple with coarse and/or irregular teeth 5a. Corolla purple, with pairs of scale-like appendages near the base that alternate with and are adnate to the filaments; pedicels 10–15 (–20) mm long; stamens 15–35 mm long; leaf blade usually shorter than to ± equaling the length of the petiole . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. minor 5b. Corolla with a white or purple basal connate portion and a blue limb, lacking scale-like appendages near the base; pedicels 2–10 mm long; stamens 8–16 mm long; leaf blade usually longer than the petiole . . . . . . . . . . . . . . . . . . . . . . . P. viscida 4b. Pedicels up to 4 mm long; corolla 6–10 mm long; capsules 2–5 mm tall with 2–25 seeds; leaf blades elliptic to oblanceolate, lobed or compound 6a. Calyx lobes 4–6 mm long in flower, 6–8 mm long in fruit; style 11–15 mm long; capsules with 1 or 2 seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. tanacetifolia 6b. Calyx lobes 3–4 mm long in flower, 4–5 mm long in fruit; style 3–12 mm long; capsules with 2–25 seeds 7a. Corolla white to blue, with a pair of ovate appendages near the base between the filaments; styles 7–12 mm long, glabrous capsule 2–3 mm tall with 2–4 seeds 2–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. distans 7b. Corolla with a yellow basal, connate portion and a white to pink limb, without a pair of appendages between the filaments; style 3–4 mm long, pubescent; capsule 4–5 mm tall with 10–25 seeds ca. 0.5 mm long . . . . . . . . . . . P. brachyloba 1. Phacelia brachyloba (Benth.) Gray E yellow-throated scorpion-weed. MA. Wool waste, disturbed soil. 2. Phacelia distans Benth. E distant scorpion-weed. Phacelia cinerea Eastw. ex J.F. Macbr. • MA. Wool waste, fields. 3. Phacelia egena (Greene ex Brand) Greene ex J.T. Howell E rock scorpion-weed. Phacelia magellanica (Lam.) Coville, pro parte • MA. Wool waste, gardens.

Fig. 500  Inflorescence of Onosmodium virginianum.

466  tricolpate s

4. Phacelia linearis (Pursh) Holz. E thread-leaved scorpion-weed. Hydrophyllum linarea Pursh • CT, ME. Fields, yards, gardens, disturbed soil. 5. Phacelia minor (Harvey) Thellung ex F. Zimmerman E wild scorpion-weed. Phacelia whitlavia Gray • MA. Wool waste. 6. Phacelia purshii Buckl. E Fig. 501 purple scorpion-weed. Phacelia bicknellii Small • CT, RI. Fields, gardens, disturbed soil. 7. Phacelia tanacetifolia Benth. E lacy scorpion-weed. MA, ME, NH. Gardens, waste areas. 8. Phacelia viscida (Benth. ex Lindl.) Torr. E tacky scorpion-weed. CT. Fields, gardens, disturbed soil. Fig. 501  Flowers of Phacelia purshii.

Plagiobothrys Plagiobothrys scouleri var. scouleri was reported from ME by Kartesz (1999), but specimens are unknown. 1a. Attachment scar of mericarp and often basal portion of inner keel positioned within a shallow trough created by folds of the pericarp; outer suface of mericarp with cross-ribs that lack teeth or prickles; calyx 2–4 mm long; corolla 1.5–3.5 mm wide in life . . . . . . P. reticulatus 1b. Attachment scar of mericarp not situated within a trough; outer surface of mericarp with cross-ribs that are ± toothed (rarely the teeth well-developed and nearly forming prickles); calyx 4–6 mm long; corolla 1–2 mm wide in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. trachycarpus 1. Plagiobothrys reticulatus (Piper) I.M. Johnson E netted popcorn-flower. Allocarya californica (Fisch. & C.A. Mey.) Greene; A. reticulata Piper; Krynitzkia californica Gray; Myosotis californica Fisch. & C.A. Mey. • MA. Wool waste. This species was responsible for the report of Plagiobothrys scouleri (Hook. & Arn.) I.M. Johnston by Sorrie and Somers (1999)—North Chelmsford, Alcott s.n. (NEBC!). However, that species does not possess a mericarp attachment scar positioned within a fold (Chambers 2007). Mericarp maturity presents an obstacle for confirming the variety of this collection (var. reticulatus or var. rossianorum I.M. Johnston). The original determination, as Krynitzkia californica, places this collection with P. reticulatus var. rossianorum (both of these names are nomenclatural synonyms of Myosotis californica). 2. Plagiobothrys trachycarpus (Gray) I.M. Johnson E rough-fruited popcorn-flower. Allocarya trachycarpa (Gray) Greene • MA. Waste areas, wool waste.

Pulmonaria 1. Pulmonaria saccharata P. Mill. E Fig. 502 Bethlehem lungwort. CT, VT. Forest edges, lawns, roadsides.

Symphytum Reference: Gadella (1984). Fig. 502  Inflorescence of Pulmonaria saccharata.

1a. Leaves decurrent as wings on the stem and branches; mericarps smooth; filaments nearly as wide as the anthers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. officinale 1b. Leaves not decurrent; mericarps tuberculate-roughened; filaments narrower than the anthers 2a. Root thick, but neither tuberous nor constricted at intervals; corolla pink turning blue; stem usually branched, pubescent, in part, with stout, basally flattened hairs . . . . . S. asperum

Bo r ag i n ac e a e   467

2b. Root tuberous, enlarged and constricted at intervals; corolla pale yellow; stem usually simple, pubescent with subterete hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tuberosum 1. Symphytum asperum Lepechin E prickly comfrey. Symphytum asperrimum Donn ex Sims • CT, MA, ME, VT. Roadsides, dumps, fields. 1‌ × 2. Symphytum ×uplandicum Nyman is a very rare comfrey hybrid in New England known from CT, VT. It resembles S. asperum in that the leaves are not decurrent on the stem (or infrequently shortly decurrent for a distance of less than 10 mm). However, the hybrid differs in that it has a corolla 13–16 mm long, purple or pink flower buds, a calyx 5–7 mm long, and short, broad papillae on the margins of the fornices (vs. corolla 9–14 mm long, red flower buds, calyx 3–5 mm long, and long, narrow papillae on the margins of the fornices). From S. officinale it can additionally be distinguished by ascending corolla lobes and dull brown schizocarps (vs. recurved corolla lobes and lustrous black schizocarps). Two cytotypes of this hybrid are known, depending on which cytotype of S. officinale was involved in the cross (see discussion under S. officinale). Thus far, only the 2n=40 type of this hybrid has been collected in New England (with a 2n=48 S. officinale parent). This hybrid shows softer stem pubescence, slightly longer leaf base decurrence (on average), and blunter fornices than the 2n=36 type (which is known from North America). 2. Symphytum officinale L. E Fig. 503 common comfrey. Symphytum officinale L. ssp. uliginosum (Kern.) Nyman; S. tanaicense Steven; S. uliginosum Kern. • CT, MA, ME, NH, RI, VT. Roadsides, ditches, former homesteads, gardens. At present their appears to be two cytotypes of Symphytum officinale that may warrant recognition at some level. The 2n=24 or 48 type (diploid and tretraploid, respectively) appears to be the common form in New England. It shows hispid stems that are not harsh to the touch, leaves with prominent decurrent wings, marginal setae of the sepals distributed in an irregular pattern, and white (usually diploid) or purple (usually tetraploid) corollas. The 2n=40 type, known from North America but yet documented in New England, has tuberculate-based hairs on the stem that are harsh to the touch, leaves with shorter decurrent wings, marginal setae of the sepals distributed in a uniform manner, and purple corollas. This latter cytotype has been named ssp. uliginosum (or S. tanaicense at the rank of species). 3. Symphytum tuberosum L. E tuberous comfrey. CT, MA; also reported from ME by Fernald (1950b), but specimens are unknown. Roadsides, fields, disturbed soil.

Brassicaceae Matthiola incana (L.) Ait. was reported from VT by Atwood et al. (1973); however, the specimen was collected from a cultivated plant (Blanchard collection at HNH). Reference: Rollins (1993). 1a. Fruit a silicle (i.e., fruit up to 3 times as long as wide), elliptic or oblong to suborbicular or obtriangular in outline, sometimes with a notch or sinus at the base and/or apex [Figs. 506, 521, 531] 2a. Silicle indehiscent or tardily dehiscent, containing 1–4 (–5) seeds, inflated, corky, or spongy at least between the seeds (firm in Neslia) 3a. Plants pubescent in part with branched hairs; petals 1–2.5 mm long 4a. Silicle evidently reticulate-textured, glabrous, tipped by the slender, persistent, uncurved style; petals yellow, 2–2.5 mm long; leaves auriculate clasping . . . . . . Neslia 4b. Silicle not reticulate-textured, pubescent with both branched and unbranched, white hairs, with an apical beak that curves away from the inflorescence rachis; petals white, 1–1.3 mm long; leaves petiolate or sessile, but not clasping . . . . . . . . . Euclidium

Fig. 503  Branch of inflorescence of Symphytum officinale.

468  tricolpates

3b. Plants glabrous or pubescent entirely with unbranched hairs; petals 3–20 mm long 5a. Silicle 2.5–3 mm long; leaves auriculate-clasping; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lepidium 5b. Silicle 6–60 mm long; leaves not auriculate-clasping; plants annual 6a. Petals 15–20 mm long; silicle 30–60 mm long; plants from a fleshy, elongated taproot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Raphanus 6b. Petals 5–10 mm long, sometimes reduced in Cakile and bristle-like; silicle 6–25 mm long; plants not from a fleshy, enlarged taproot 7a. Petals pale purple to purple or white; leaf blades glabrous; upper segment of fruit lanceoloid to ovoid, without a terminal beak or with a stocky beak [Fig. 510] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cakile 7b. Petals yellow; leaf blades hirsute; upper segment of fruit subglobose, terminated by a slender beak 1–4 mm long . . . . . . . . . . . . . . . (in part) Rapistrum 2b. Silicle dehiscent, containing 2–many seeds, appearing inflated or not 8a. Silicle compressed at right angles to the septum, the septum therefore much narrower than the width of the fruit [Fig. 531] 9a. Leaves subulate, not flat, all basal [Fig. 530]; plants aquatic, frequently in submerged populations that become emergent as water levels decline . . . . . Subularia 9b. Leaves not subulate, definitely flat, not confined to the base of the plant except in Teesdalia; plants terrestrial weeds 10a. Silicle with 2 seeds 11a. Petals all of similar size (or absent) [Fig. 522]; silicle with either a short or obsolete style or with a truncate apex or both [Figs. 521, 522] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lepidium 11b. Outer two petals larger than the inner two; silicle with a long style projecting from the prominently retuse apex . . . . . . . . . . . . . . . . . . . . . . . . . . . Iberis 10b. Silicle with 4 or more seeds 12a. Outer two petal larger than the inner two; median filaments with a white scale at the base; plants scapose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Teesdalia 12b. Petals all of similar size; median filaments without a basal scale; plants with leafy stems 13a. Silicle oval to orbicular to outline, wing-margined, rounded to broadcuneate at the base, conspicuously retuse at the apex [Fig. 531]; plants glabrous or pubescent with unbranched hairs . . . . . . . . . . . . . . . . . . . . . Thlaspi 13b. Silicle obtriangular to obtriangular-cordate in outline, not wingmargined, cuneate at the base, convex to slightly concave at the apex; plants pubescent with branched hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Capsella 8b. Silicle either not compressed or compressed parallel to the septum, in either case the septum as wide as the fruit 14a. Silicle thin and flat, 20–35 mm wide, borne on a gynophore longer than 10 mm [Fig. 523]; stem leaf blades ovate to deltate; petals blue-purple . . . . . . . . . . . . . Lunaria 14b. Silicle compressed to plump, narrower than 6 mm, either sessile at the end of the pedicel or on a very short gynophore; stem leaf blades usually narrower or absent; petals white or yellow (sometimes tinged with purple in Lobularia) 15a. Petals with a deep, apical notch [Fig. 506]; fruiting pedicels erect to appressed (or widely ascending in B. mutabilis) [Fig. 506]; seeds winged . . . . . . . . . . . . Berteroa

Br a ss i c ac e a e   46 9

15b. Petals entire at the apex or slightly retuse (deeply notched in Draba verna, a small, scapose annual); fruiting pedicels spreading to ascending; seeds unwinged (winged in Aurinia) 16a. Silicle irregularly tuberculate, often asymmetrical (i.e., one locule larger than the other); plants frequently with sessile, tubercle-like glands on the leaf blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bunias 16b. Silicle not tuberculate, usually symmetrical; plants without tuberclelike glands 17a. Silicle conspicuously compressed; pubescence, at least in part, of branched hairs 18a. Petals yellow; seeds winged all around . . . . . . . . . . . . . . . . . . . . Aurinia 18b. Petals cream to white; seed unwinged 19a. Flower-bearing stems arising from a basal rosette of leaves; silicle lanceolate to narrow-oblong or oblong in outline, 4–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Draba 19b. Flower-bearing stems lacking a conspicuous basal rosette; silicle broad-ovate or broad-obovate to orbicular in outline, 2–4 mm long 20a. Plants pubescent with stellate-branched hairs; silicle with a thin, wing-like margin, with seeds aligned in 2 rows in each locule; fruiting pedicels 2–6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . Alyssum 20b. Plants pubescent with 2-pronged hairs; silicle lacking a thin margin, with seeds aligned in 1 row in each locule; fruiting pedicels 5–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lobularia 17b. Silicle with definite, convex valves, scarcely compressed if at all; pubescence, if present, of unbranched hairs (except Camelina with some branched hairs) 21a. Plants pubescent with both unbranched and branched hairs (sometimes subglabrous in C. sativa); leaves conspicuously auriculateclasping the stem; each valve of the silicle with a thin but visible and often sinuous midnerve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camelina 21b. Plants glabrous or pubescent with unbranched hairs; leaves narrowed to the base or auriculate-clasping; each valve of the silicle without a midnerve or with an obscure midnerve 22a. Petals white; leaf blades simple; seeds rarely maturing; silicle with a persistent style tip 0.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . Armoracia 22b. Petals yellow (white in 1 aquatic species); principal leaf blades pinnately lobed, divided, or dissected (simple in R. austriaca and R. amphibia); seeds usually maturing; silicle with a persistent style tip 0.4–3 mm long [Figs. 525, 526] . . . . . . . . . . . . . . . . . . . . . . (in part) Rorippa 1b. Fruit a silique (i.e., fruit greater than 3 times as long as wide), narrow-oblong to linear in outline, without a notch or sinus at the base or apex [Figs. 504, 507, 524] 23a. Silique indehiscent, separating at maturity into 1-seeded segments, prominently corky or spongy, at least between the seeds, or indurate and cross-septate in Chorispora [Figs. 510, 524] 24a. Stems and axis of inflorescence conspicuously stipitate-glandular; seeds embedded in cavities of the septum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chorispora 24b. Stems and axis of inflorescence lacking stipitate glands; seeds not embedded in cavities of the septum

470  tricolpates

25a. Petals 15–20 mm long; siliques 40–80 mm long, with 4–12 seeds [Fig. 524]; fruiting pedicels 10–25 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Raphanus 25b. Petals 5–10 mm long, sometimes reduced in Cakile and bristle-like; siliques 6–25 mm long, with 1–3 seeds; fruiting pedicels 1.5–8 mm long 26a. Petals pale purple to purple or white; leaf blades glabrous; upper segment of fruit lanceoloid to ovoid, without a prominent, slender beak . . . . . . (in part) Cakile 26b. Petals yellow; leaf blades hirsute; upper segment of fruit subglobose, terminated by a prominent, slender beak 1–4 mm long . . . . . . . (in part) Rapistrum 23b. Silique longitudinally dehiscent (with an apical, indehiscent beak in Brassica, Eruca, and Sinapis), the open valves revealing the seeds and septum, generally not corky or spongy, lacking cross-septa [Figs. 504, 509, 514] 27a. Leaf blades simple, entire or toothed, but lacking pronounced lobes (except for basal lobes in cordate- or sagittate-shaped leaf blades) 28a. Plants pubescent partly or entirely with branched hairs [Figs. 517, 518] 29a. Middle stem leaves sessile and sagittate- or auriculate-clasping 30a. Pubescence of basal leaves, when present, comprised entirely of unbranched hairs or, in part, of short-stalked, 2- to 4 (–7)-rayed hairs, the stalks mostly 0.01–0.06 mm long [Fig. 517]; siliques 1–2.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Boechera 30b. Pubescence of basal leaves comprised, in part, of long-stalked, 2- to 4-rayed hairs, the stalks mostly 0.12–0.6 mm long [Fig. 518]; siliques 0.8–1.3 mm wide 31a. Petals pale yellow; outer sepals not saccate at base; middle and upper stem leaves glabrous and glaucous; siliques terete . . . . . . . . . . . . . . . . . Turritis 31b. Petals white, sometimes tinged with pink; outer sepals somewhat saccate at base; middle and upper stem leaves pubescent, not glaucous; siliques compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Arabis 29b. Middle stem leaves narrowed to base, sometimes ± shortly petiolate [Fig. 507] 32a. Petals yellow; siliques pubescent with branched hairs; some of the hairs on the leaf blades malpighiaceous (i.e., hairs 2-pronged, less frequently 3-pronged, mesifixed and appressed to the surface) . . . . . . . . (in part) Erysimum 32b. Petals white or pink to purple; siliques glabrous or pubescent in some Draba; hairs of the leaf blades unbranched or branched, but none of them malpighiaceous (except in the rare introduction Arabis procurrens, a species with compressed siliques) 33a. Petals 18–25 mm long, white or pink to purple; compound hairs all forked (i.e., with a stalk and 2 ascending branches) . . . . . . . . . . . . . . . Hesperis 33b. Petals 2–10 mm long, usually white (infrequently pink or rosepurple); compound hairs with 2 or more ascending to spreading branches (i.e., compound hairs not uniformly forked with ascending branches) 34a. Siliques narrow-lanceolate or narrow-oblong to ovate or obovate in outline, 4–20 (–22) mm long, up to 5 times as long as wide; seeds aligned in 2 rows in each locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Draba 34b. Siliques linear in outline, 10–100 mm long, more than 5 times as long as wide; seeds aligned in 1 row in each locule 35a. Siliques often torulose; fruiting pedicels (1–) 1.5–6 (–8) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Braya 35b. Siliques scarcely, if at all, constricted between the seeds; fruiting pedicels 5–15 mm long

Br a ss i c ac e a e   471

36a. Plants 3–10 dm tall, usually with unbranched stems; siliques 2–3.5 mm wide; seeds winged . . . . . . . . . . . . . . . . . . (in part) Boechera 36b. Plants 0.8–4 dm tall, frequently branched at the base or with stolons, producing multiple upright stems; siliques 0.8–1.5 mm wide; seeds without wings (winged in Arabis procurrens) 37a. Primary leaf blades usually with some form of dentition or lobing; siliques 0.8–1 mm wide; petals 2–8 (–10) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Arabidopsis 37b. Primary leaf blades usually entire; siliques 1–1.5 mm wide; petals 8–10 mm long; . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Arabis 28b. Plants glabrous or pubescent entirely with unbranched hairs 38a. Stem leaves conspicuously cordate- or sagittate-clasping at the base 39a. Petals 6–25 (–30) mm long, narrowed to a distinct claw; siliques 2–5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Brassica 39b. Petals 3–12 mm long, narrowed to the base, but usually without a distinct claw; siliques 1–2.5 (–3.5) mm wide 40a. Plants annual, with entire leaf blades; petals yellow to yellow-white, 7–12 mm long; siliques quadrangular in cross-section; seeds without wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conringia 40b. Plants biennial or perennial, with at least the lower leaf blades toothed to pinnatifid (rarely all entire); petals 3–10 mm long, white (infrequently yellow-white or light purple); siliques compressed; seeds with a partly or completely encircling wing-margin usually wider than 0.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Boechera 38b. Stem leaves not clasping at the base [Fig. 507] 41a. At least the basal leaf blades triangular to suborbicular or reniform, broadrounded to cordate at the base 42a. Basal leaf blades entire or undulate-margined, the stem leaf blades entire, remotely toothed, or lobed; siliques compressed; septum bordered by a thin wing-like replum set perpendicular to the plane of the septum, therefore, the septum and replum combined are I-beam-shaped in crosssection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cardamine 42b. Basal and stem leaf blades prominently dentate; siliques somewhat 4-angled in cross-section; replum essentially the same thickness as the septum, the septum and replum combined flat in cross-section . . . . . Alliaria 41b. Leaf blades broad-linear or oblanceolate to elliptic, ovate, or obovate, narrowed to the base 43a. Sepals spreading to reflexed [Fig. 527]; siliques with a prominent, apical, indehiscent, conical beak 10–12 mm long . . . . . . . . . . . . . . . . . (in part) Sinapis 43b. Sepals erect to ascending; siliques without an apical, indehiscent beak or the beak present and only 1–5 (–6) mm long 44a. Siliques quadrangular, erect (i.e., ± appressed to the axis of the inflorescence); petals narrowed to a distinct claw . . . . . . (in part) Brassica 44b. Siliques terete to compressed, spreading to erect; petals narrowed to the base, with or without a distinct claw 45a. Petals yellow; seeds aligned in 2 rows in each locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Diplotaxis 45b. Petals white to pink or light purple; seeds aligned in 1 row in each locule

47 2 tricolpates

46a. Siliques 15–35 mm long; septum bordered by a thin wing-like replum set perpendicular to the plane of the septum, therefore, the septum and replum combined are I-beam-shaped in cross-section; dwarf alpine plants 2–10 cm tall . . . . . . . . . . . . . . (in part) Cardamine 46b. Siliques 40–100 mm long; replum essentially the same thickness as the septum, the septum and replum combined flat in cross-section; herbs of forests, cliffs, and talus, 10–90 cm tall 47a. Slender plants 10–40 cm tall, frequently branched at the base, producing multiple upright stems; siliques 0.8–1 mm wide; seeds without wings . . . . . . . . . . . . . . . . . . . . . (in part) Arabidopsis 47b. Stouter plants 30–100 cm tall, usually with unbranched stems; siliques 2–3.5 mm wide; seeds winged . . . . . . . . . (in part) Boechera 27b. At least the lower leaf blades lobed or divided 48a. Principal leaf blades palmately lobed or palmately divided into 3–5 segments; stem gradually narrowing at base to a fragile junction with a swollen, fleshy rhizome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cardamine 48b. Principal leaves pinnately lobed or pinnately divided; stems in most genera arising from taproots or fibrous roots, when arising from rhizomes the stem widest at the base and narrowing upward and/or continuous with the rhizome 49a. Plants pubescent, at least in part, with branched hairs 50a. Inflorescence leafy-bracteate, most or all of the pedicels subtended by bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Tropidocarpum 50b. Inflorescence not bracteate, only the lower pedicels, if any, subtended by bracts 51a. Leaf blades 1–3 times pinnately divided; branched hairs dendritic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Descurainia 51b. Leaf blades pinnately lobed; branched hairs malpighiaceous (i.e., hairs 2-pronged, less frequently 3-pronged, mesifixed and appressed to the surface) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Erysimum 49b. Plants glabrous or pubescent entirely with unbranched hairs 52a. Stem leaves sessile and auriculate-clasping (only the emergent leaves in Nastustium sometimes with auricles at the base of the petiole) [Figs. 505, 514] 53a. Petals white and 2–3 mm long or absent; septum bordered by a thin wing-like replum set perpendicular to the plane of the septum, therefore, the septum and replum combined are I-beam-shaped in cross-section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cardamine 53b. Petals yellow, or white in Nasturtium and some Boechera, but then 3–7 mm long; replum essentially the same thickness as the septum, the septum and replum combined flat in cross-section 54a. Leaves with pinnately compound blades, the leaflets often petiolulate; plants aquatic, with submersed or floating stems (or prostrate on mud as water levels decline), freely rooting from the nodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Nasturtium 54b. Middle and upper stem leaves with toothed to pinnately lobed, but not compound, leaf blades, the leaflets connected to each other by at least a wing of tissue along the leaf rachis; plants terrestrial or of wetlands, the stems upright or sometimes prostrate, not or only sparingly rooting from the lower nodes

Br a ss i c ac e a e   473

55a. Petals white to yellow-white; seeds with an encircling wing; siliques wide-spreading to recurved at maturity [Fig. 507] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Boechera 55b. Petals yellow; seeds without wings; siliques ascending to spreading 56a. Siliques terminated by an indehiscent beak 7–15 mm long; seeds subglobose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Brassica 56b. Siliques without an apical, indehiscent beak, tipped only by a persistent style up to 3 mm long if at all; seeds oblong to quadrate or cordiform (rarely maturing in some Rorippa) 57a. Siliques 3–15 (–20) mm long (rarely maturing in some Rorippa), with seeds aligned in 2 rows in each locule; valves of silique ± nervelss; stems terete . . . . . . . . . . . . . . . . (in part) Rorippa 57b. Siliques 10–80 mm long, with seeds aligned in 1 row in each locule; valves of silique with a prominent midnerve; stems angled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Barbarea 52b. Stem leaves petiolate or sessile, but not clasping the stem 58a. Siliques flat or strongly compressed 59a. Petals yellow in life; seeds aligned in 2 rows in each locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Diplotaxis 59b. Petals white to pink to purple; seeds aligned in 1 row in each locule 60a. Lower leaf blades pinnately divided [Figs. 514, 516]; septum bordered by a thin wing-like replum set perpendicular to the plane of the septum, therefore, the septum and replum combined are I-beamshaped in cross-section . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cardamine 60b. Lower leaf blades pinnately lobed to dentate; replum essentially the same thickness as the septum, the septum and replum combined flat in cross-section . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Arabidopsis 58b. Siliques terete or quadrangular in cross-section, each valve strongly convex 61a. Seeds aligned in 2 rows in each locule 62a. Silique terminated by a broad, flat, indehiscent beak 5–12 mm long that is ½ to fully as long as the valves; petals white to whiteyellow with red-purple veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eruca 62b. Silque without an apical beak or with a short beak 1–3 mm long that is much shorter than the valves; petals entirely white or yellow 63a. Petals white; leaves with pinnately compound blades, the leaflets often petiolulate; plants aquatic, with submersed or floating stems (or prostrate on mud as water levels decline), freely rooting from the nodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Nasturtium 63b. Petals yellow or absent; middle and upper stem leaves with toothed to pinnately lobed, but not compound, leaf blades, the leaflets connected to each other by at least a wing of tissue along the leaf rachis; plants terrestrial or of wetlands, the stems upright or sometimes prostrate, not or only sparingly rooting from the lower nodes 64a. Inflorescence leafy-bracteate, most or all of the pedicels subtended by bracts . . . . . . . . . . . . . . . . . (in part) Tropidocarpum 64b. Inflorescence not bracteate, only the lower pedicels, if any, subtended by bracts

474  tricolpates

65a. Siliques 3–15 (–20) mm long, the valves of silique ± nerveless; fruiting pedicels 0.5–14 mm long . . . (in part) Rorippa 65b. Siliques 20–45 mm long, the valves with a prominent midnerve; fruiting pedicels (6–) 10–30 (–35) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Diplotaxis 61b. Seeds aligned in 1 row in each locule 66a. Petals white; leaves with pinnately compound blades, the leaflets often petiolulate; plants aquatic, with submersed or floating stems (or prostrate on mud as water levels decline), freely rooting from the nodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Nasturtium 66b. Petals pale yellow to yellow in life (sometimes white-yellow in the cultivated Brassica oleracea); middle and upper stem leaves with toothed to pinnately lobed, but not compound, leaf blades, the leaflets connected to each other by at least a wing of tissue along the leaf rachis (those of Sisymbrium altissimum with linear to filiform segments that often lack connecting tissue along the rachis); plants primarily terrestrial weeds with upright stems that do not root from the nodes 67a. Seeds subglobose; silique tipped by an indehiscent beak 1–30 mm long 68a. Each valve of the silique with 1 prominent nerve; indehiscent beak of fruit terete or angled, without seeds (very rarely with 1 or 2 seeds); sepals erect to ascending . . . . . . . . . . . (in part) Brassica 68b. Each valve of the silique with 3 prominent nerves; indehiscent beak of fruit compressed or compressedquadrangular, often containing 1 (–3) seeds; sepals spreading to reflexed [Fig. 527] . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Sinapis 67b. Seeds obloid or ovoid; silique tipped by the persistent style up to 3 mm long 69a. Inflorescence not bracteate, only the lower pedicels, if any, subtended by bracts [Fig. 528]; valves of silique with 3 nerves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sisymbrium 69b. Inflorescence leafy-bracteate, most or all of the pedicels subtended by bracts; valves of silique with 1 nerve . . . . Erucastrum

Alliaria 1. Alliaria petiolata (Bieb.) Cavara & Grande E Fig. 504 garlic-mustard. Alliaria alliaria (L.) Britt.; A. officinalis Andrz. ex Bieb.; Erysimum alliaria L.; Sisymbrium alliaria (L.) Scop. • CT, MA, ME, NH, RI, VT. Roadsides, forests, riparian terraces, frequently in partially shaded plant communities.

Alyssum Fig. 504  Upper portion of inflorescence of Alliaria petiolata.

1. Alyssum alyssoides (L.) L. E pale alyssum. Alyssum calycinum L.; Clypeola alyssoides L. • CT, MA, ME, RI, VT. Fields, roadsides, disturbed soil.

Br a ss i c ac e a e   475

Arabidopsis 1a. Siliques (15–) 20–40 (–45) × 0.8–1 mm, flat; petals (4–) 6–8 (–10) mm long; seeds 0.8–1.4 mm long; native biennial or perennial plants of cliffs, outcrops, and rocky slopes . . . . . . . . . . . A. lyrata 1b. Siliques (8–) 10–15 (–18) × 0.5–0.8 mm, ± terete in cross-section; petals 2–3.5 (–4) mm long; seeds 0.3–0.5 mm long; non-native annuals of disturbed soil . . . . . . . . . . . . . . . . . . . A. thaliana 1. Arabidopsis lyrata (L.) O’Kane & Al-Shehbaz ssp. lyrata N lyre-leaved thale-cress. Arabis lyrata L.; A. lyrata L. var. glabra (DC.) M. Hopkins • CT, MA, VT; mainly in western New England. Cliffs, balds, rocky ridges, commonly in regions of high-pH bedrock. 2. Arabidopsis thaliana (L.) Heynh. E mouse-ear thale-cress. Arabis thaliana L.; Sisymbrium thalianum (L.) J. Gay & Monn. • CT, MA, ME, NH, RI, VT. Roadsides, cultivated fields, lawns, waste areas.

Arabis The traditionally defined Arabis has been shown to be a highly artificial genus that was composed of four different taxa in New England—Arabidopsis, Arabis s.s., Boechera, and Turritis (Al-Shehbaz 2003). It is now known that the morphological characters used to delimit Arabis s.l. evolved independently many times within the family, and features such as base chromosome number more accurately depict the evolutionary history of this group of genera (though particulars of fruit and trichome morphology do show trends between the genera). Arabis alpina L. was attributed to ME by Magee and Ahles (1999) based on a specimen taken from a private residence—Gould 17523 (NHA!). The label provided no evidence of naturalization, and this species is not accepted here as a part of the New England flora. 1a. Petals 8–10 mm long; fruiting pedicels ± spreading; siliques 12–35 mm long; leaves with entire blades (rarely with a single tooth on one or both margins), those along the stem not clasping at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. procurrens 1b. Petals 3–5 mm long; fruiting pedicels erect; siliques 30–50 mm long; leaves with entire to dentate blades, usually the lower with at some form of dentition, those along the stem auriculate-clasping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pycnocarpa 1. Arabis procurrens Waldst. & Kit. E running eared-rockcress. MA. Fields, gardens. 2. Arabis pycnocarpa M. Hopkins var. pycnocarpa N hairy eared-rockcress. Arabis hirsuta (L.) Scop. ssp. pycnocarpa (M. Hopkins) Hultén; A. hirsuta (L.) Scop. var. pycnocarpa (M. Hopkins) Rollins; Turritis oblongata Raf. • CT, MA, ME, NH, VT. Ledges, rock outcrops, rocky woodlands, in regions of high-pH bedrock. Arabais pycnocarpa, which is native to North America, is frequently treated as an infraspecific form of the European Arabis hirsuta. However, the two taxa differ in geographic range, ploidy level, fruit length and venation, style morphology, and seed number (i.e., ample justification exists for recognition of two species).

Armoracia 1. Armoracia rusticana P.G. Gaertn., B. Mey., & Scherb. E horse-radish. Armoracia armoracia (L.) Britt.; A. lapathifolia Gilib.; Cochlearia armoracia L.; Radicula armoracia (L.) B.L. Robins.; Rorippa armoracia (L.) A. Hitchc. • CT, MA, ME, NH, RI, VT. Fields, roadsides, areas of cultivation.

Aurinia 1. Aurinia saxatilis (L.) Desv. E rock goldentuft. Aethionema saxatile (L.) R. Br.; Alyssum saxatile L. • CT, MA, NH, VT. Roadsides, waste areas, ledges.

47 6 tricolpates

Barbarea Cilia near the apex of the sepals and auricles of upper leaves are an important identification characters for New England Barbarea. However, the cilia are sometimes very sparse and may number only 1 or a few on the margins of these organs. 1a. Basal leaf blades with 4–10 pairs of lateral lobes; siliques (40–) 53–70 (–80) mm long; fruiting pedicels 1.2–1.8 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. verna 1b. Basal leaf blades simple or with 1–4 pairs of lateral lobes; siliques 10–40 (–45) mm long; fruiting pedicels up to 1 mm thick 2a. Silique with a persistent style beak 2–3 (–3.5) mm long; petals 6–8 mm long; auricles of uppermost leaves eciliate [Fig. 505]; sepals glabrous . . . . . . . . . . . . . . . . . . . . . . . . . B. vulgaris 2b. Silique with a persistent style beak 0.5–1.6 (–2.3) mm long; petals 3–6 mm long; auricles of uppermost leaves ciliate; sepals usually pubescent near the apex (sometimes glabrous in B. stricta) 3a. Petals (1.5–) 2–3 mm wide; siliques (25–) 31–40 (–45) mm long; pedicels 3–7 mm long in fruit; upper leaf blades usually coarsely toothed to pinnately lobed . . . . . . . . B. orthoceras 3b. Petals 0.5–1 (–1.2) mm wide; siliques (12–) 18–28 (–30) mm long; pedicels 1–3 (–4) mm long in fruit; upper leaf blades usually toothed or with a few shallow sinuses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. stricta 1. Barbarea orthoceras Ledeb.

NC

American yellow-rocket. Barbarea americana Rydb.; B. orthoceras Ledeb. var. dolichocarpa Fern.; Campe orthoceras (Ledeb.) Heller • ME, NH. Ice-scoured river shores in high-pH bedrock and/or till regions, wet talus in subalpine setting. 2. Barbarea stricta Andrz. E upright yellow-rocket. CT, MA, ME, NH, RI, VT. Wet-mesic to hydric disturbed soil (e.g., river shores, low areas in cultivated fields). This species has been largely overlooked in New England and misidentified as Barbarea vulgaris. 3. Barbarea verna (P. Mill.) Aschers. E early yellow-rocket. Campe verna (P. Mill.) Heller; Erysimum vernum P. Mill. • CT, MA, ME, RI. Roadsides, fields, waste areas, dumps. 4. Barbarea vulgaris Ait. f. E Fig. 505 Fig. 505  Leaf of Barbarea vulgaris showing clasping base.

garden yellow-rocket. Barbarea arcuata (Opiz ex J. & K. Presl) Reichenb.; B. stricta, auct. non Andrz.; B. vulgaris Ait. f. var. arcuata (Opiz ex J. & K. Presl) Fries; Campe barbarea (L.) W.Wight ex Piper; C. stricta, auct. non (Andrz.) W. Wight ex Piper; Erysimum barbarea L. • CT, MA, ME, NH, RI, VT; nearly througout. Roadsides, fields, disturbed soil, river shores.

Berteroa 1a. Siliques 5–7 × 3–4 mm, inflated, with strongly convex valves, densely to moderately pubescent; fruiting pedicels erect to appressed [Fig. 506]; style 2–3 mm long; seeds with a narrow margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. incana 1b. Siliques 6–12 × 3.5–5 mm, compressed, ± glabrous or with some hairs near the margins of the valves; fruiting pedicels widely ascending; style 1–2 mm long; seeds with a winged margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. mutabilis 1. Berteroa incana (L.) DC. E Fig. 506 hoary false alyssum. Alyssum incanum L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, dry lawns. Fig. 506  Inflorescence of Berteroa incana.

2. Berteroa mutabilis (Vent.) DC. E roadside false alyssum. Alyssum mutabile Vent. • MA. Fields, roadsides, gardens.

Br a ss i c ac e a e   477

Boechera References: Mulligan (1995), Al-Shehbaz (2003). 1a. Fruiting pedicels erect; seeds aligned in 2 rows in each locule; petals 7–10 mm long, white to pink to rose-purple; lower stems usually with some mesifixed hairs or glabrous . . . . . . B. stricta 1b. Fruiting pedicels widely ascending to reflexed; seeds aligned in 1 row in each locule (rarely partially aligned in 2 irregular rows); petals 3–7 (–8) mm long, white to yellow-white (rarely light purple); lower stems glabrous or with simple or branched hairs, but none of the hairs mesifixed 2a. Stem leaves narrowed to the base, not clasping the stem [Fig. 507]; siliques 2.5–3.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. canadensis 2b. Stem leaves auriculate-clasping at the base; siliques 1–2.5 mm wide 3a. Basal leaf blades sparsely to densely pubescent with compound hairs bearing 2–4 (–7) rays; seeds aligned in 1 row or partially aligned in 2 irregular rows in each locule; sepals abaxially glabrous or pubescent; petals white or sometimes light purple; pollen subspherical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. grahamii 3b. Basal leaf blades glabrous to sparsely pubescent with simple hairs; seeds aligned in 1 row in each locule; sepals glabous; petals white to yellow-white; pollen ellipsoid 4a. Petals 3–5 mm long, equaling to slightly exceeding the length of the sepals (i.e., petals up to 1.3 times as long as the sepals); stems glaucous, with relatively fewer leaf-bearing nodes, averaging 13 internodes to the first flower; basal leaf blades serrate to subentire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. laevigata 4b. Petals 5–7 mm long, (1.3–) 1.4–2 times as long as the sepals; stems green, with relatively more leaf-bearing nodes, averaging 25 internodes to the first flower; basal leaf blades serrate to pinnately lobed [Fig. 508] . . . . . . . . . . . . . . . . . . B. missouriensis 1. Boechera canadensis (L.) Al-Shehbaz N Fig. 507 sicklepod rockcress. Arabis canadensis L.; A. falcata Michx. • CT, MA, NH, RI, VT. Rocky woodlands and forests. Reports of this species in ME by Fernald (1950b) and subsequent authors were based on a specimen of Hesperis matronalis. 2. Boechera grahamii (Lehmann) Windham & Al-Shehbaz NC Graham’s rockcress. Arabis brachycarpa (Torr. & Gray) Britt.; A. drummondii Gray var. brachycarpa (Torr. & Gray) Gray; A. holboellii Hornem. var. brachycarpa (Torr. & Gray) Welsh; Boechera brachycarpa (Torr. & Gray) Dorn; Turritis grahamii Lehmann • VT. Ledges, rocky banks and woodlands, all in regions of high-pH bedrock. This species has been usually referred to as Boechera brachycarpa (Torr. & Gray) Dorn or B. divaricarpa (A. Nels.) A. & D. Löve in New England literature (under one of the nomenclatural synonyms in the genus Arabis). The former name was published later than the epithet grahamii (1838 vs. 1831) and thus does not have priority. The latter name has been, in more recent literature, presumed to be a hybrid between Boechera holboellii (Hornem.) A. & D. Löve and B. stricta. However, B. holboellii is apparently restricted to Greenland, making it highly unlikely to be involved in species here in New England (Al-Shehbaz and Windham 2010). Further, recent taxonomic work reveals that B. divaricarpa is actually restricted to western North America. Our taxon (B. grahamii) likely originated through hybridization between B. collinsii (Fern.) A. & D. Löve and B. stricta (Windham and Al-Shehbaz 2007).

Fig. 507  Infructescence of Boechera canadensis.

3. Boechera laevigata (Muhl. ex Willd.) Al-Shehbaz N smooth rockcress. Arabis laevigata (Muhl. ex Willd.) Poir.; Turritis laevigata Muhl. ex Willd. • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Rocky woodlands and forests, cliffs, and talus in regions of high-pH bedrock. 4. Boechera missouriensis (Greene) Al-Shehbaz N C Fig. 508 green rockcress. Arabis laevigata (Muhl. ex Willd.) Poir. var. missouriensis (Greene) Ahles; A. missouriensis Greene; A. viridis Harger • CT, MA, ME, NH, RI, VT. Rocky woodlands and forests, cliffs, and rock balds in regions of moderate- to high-pH bedrock.

Fig. 508  Basal leaves of Boechera missouriensis.

47 8 tricolpate s

5. Boechera stricta (Graham) Al-Shehbaz N Canada rockcress. Arabis drummondii Gray; Boechera drummondii (Gray) A. & D. Löve; Turritis drummondii (Gray) Lunell; T. stricta Graham • CT, MA, ME, NH, RI, VT. Rocky woodlands and forests, cliffs, and talus slopes in regions of moderate to high-pH bedrock.

Brassica Reference: Heywood and Akeroyd (1993). 1a. Upper stem leaves either shortly petiolate or sessile and then narrowed to the base 2a. Siliques terete or subterete, 15–40 mm long, tipped by an indehiscent beak 5–10 mm long; fruiting pedicels ascending, mostly 10–15 mm long; plants usually glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. juncea 2b. Siliques quadrangular, 10–25 mm long, tipped by an indehiscent beak 1–3 mm long [Fig. 509]; fruiting pedicels erect to appressed, 2–5 mm long [Fig. 509]; plants usually hirsute-hispid in the lower portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. nigra 1b. Upper stem leaves sessile and auriculate-clasping the stem 3a. Petals (15–) 18–25 (–30) mm long; plants glabrous throughout; filaments all erect at base; sepals erect; indehiscent, apical beak of fruit (3–) 4–10 mm long, with (0–) 1 (–2) seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. oleracea 3b. Petals 6–14 mm long; plants sometimes sparsely pubescent near the base; filaments of lateral stamens curved at base; sepals ascending (rarely suberect); indehiscent, apical beak of fruit 7–15 mm long, with 0 (–1) seeds 4a. Plants green; petals 6–10 (–11) mm long, pale yellow; apical beak of silique (8–) 10–15 mm long; seeds 1–1.8 mm long; open flowers of raceme at the same level as or overtopping the flower buds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. rapa 4b. Plants glaucous; petals 10–14 mm long, deep yellow; apical beak of silique 7–10 (–11) mm long; seeds (1.2–) 1.5–2.5 (–3) mm long; open flowers of raceme usually lower than (rarely at the same level) the flower buds . . . . . . . . . . . . . . . . . . . . . . . . B. napus 1. Brassica juncea (L.) Czern. E Chinese mustard. Brassica juncea (L.) Czern. var. crispifolia Bailey; Sinapis juncea L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 2. Brassica napus L. E turnip. CT, MA, ME, NH, RI, VT. Cultivated fields, waste areas. 3. Brassica nigra (L.) W.D.J. Koch E Fig. 509 Fig. 509  Siliques of Brassica nigra.

black mustard. Sinapis nigra L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, river banks, waste areas, compost heaps. 4. Brassica oleracea L. E cabbage. Brassica oleracea L. var. acephala DC.; B. oleracea L. var. capitata L. • CT, MA, RI; also reported from VT by Warwick (2010a), but specimens are unknown. Cultivated fields, waste areas. This species has been reported from ME (e.g., Warwick 2010a); however, the only specimen I am aware of was taken from a cultivated plant—18 Jun 1955, Bicknell s.n. (MAINE!). Brassica oleracea is extensively cultivated and produces many different crop plants (e.g., broccoli, Brusell sprouts, cabbage, cauliflower, kale, kohlrabi). 5. Brassica rapa L. E rape. Brassica campestris L.; B. campestris L. var. rapa (L.) Hartman; B. rapa L. ssp. campestris (L.) Clapham; B. rapa L. var. campestris (L.) W.D.J. Koch; B. rapa L. ssp. olifera DC. • CT, MA, ME, NH, RI, VT. Fields, roadsides, gardens, waste areas, disturbed soil.

Br a ss i c ac e a e   479

Braya 1. Braya humilis (C.A. Mey.) B.L. Robins. ssp. humilis

NC

alpine northern-rockcress. Arabidopsis novae-angliae (Rydb.) Britt.; Braya humilis (C.A. Mey.) B.L. Robins. var. leiocarpa (Trautv.) Fern.; B. humilis (C.A. Mey.) B.L. Robins. var. novae-angliae (Rydb.) Fern.; B. novae-angliae (Rydb.) Sørensen; Neotorularia humilis (C.A. Mey.) Hedge & J. Léonard; Pilosella novae-angliae Rydb.; Sisymbrium humile C.A. Mey.; Torularia humilis (C.A. Mey.) O.E. Schulz • VT; northern portion of state. Cliffs and gravel slides in high-pH bedrock regions.

Bunias 1. Bunias orientalis L. E Turkish warty-cabbage. MA; also reported from VT by Hultén and Fries (1986), but specimens are unknown. Waste areas. The report for NH by Kartesz (1999) was in error.

Cakile 1. Cakile edentula (Bigelow) Hook. ssp. edentula var. edentula N Fig. 510 American sea-rocket. Bunias edentula Bigelow; Cakile americana Nutt.; C. lanceolata (Willd.) O.E. Schulz ssp. edentula (Bigelow) O.E. Schulz; C. maritima L. var. americana (Nutt.) Torr. • CT, MA, ME, NH, RI. Atlantic coast beaches on a variety of substrates.

Camelina 1a. Plants rough-pubescent with simple hairs 1–2.5 mm long that exceed the length of the branched hairs; silicle body (i.e., excluding the beak and the short, stipe-like base) (2.5–) 3.5– 5 (–6) × 2–4 (–5) mm; fruiting pedicels 4–14 (–17) mm long; seeds 0.8–1.4 (–1.5) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. microcarpa 1b. Plants often ± glabrate, when pubescent the simple hairs not exceeding the length of the branched hairs; silicle body (5.5–) 7–9 × (4.5–) 6–7 mm; fruiting pedicels 10–25 (–30) mm long; seeds (1.5–) 1.7–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sativa 1. Camelina microcarpa Andrz. ex DC. E small-seeded false flax. Camelina sativa (L.) Crantz ssp. microcarpa (Andrz. ex DC.) Schmid • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, gardens. 2. Camelina sativa (L.) Crantz ssp. sativa E large-seeded false flax. Myagrum sativum L. • CT, MA, ME, NH, RI, VT. Fields, roadsides.

Capsella 1. Capsella bursa-pastoris (L.) Medik. E shepherd’s-purse. Bursa bursa-pastoris (L.) Britt.; Thlaspi bursa-pastoris L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, waste areas, gardens.

Cardamine Some species of Cardamine are frequently treated as nothospecies (e.g., C. incisa, C. maxima), with the often-cited reasons of morphological intermediacy and failure to produce mature fruits in some populations. It is important to note that these plants are not exactly intermediate (i.e., they have character states that are novel or extreme compared with the putative parents; Sweeney and Price 2001) and often produce fertile siliques. Further, some of the populations occur great distances from where the parental taxa are sympatric. These observations suggest that these taxa, which are quite possibly derived through hybridization, are acting as species and are treated as such here. References: Sweeney and Price (2001), Al-Shehbaz et al. (2010).

Fig. 510  Inflorescence of Cakile edentula.

480   tricolpate s

1a. Principal leaf blades palmately lobed or palmately divided into 3–5 segments; stems gradually narrowing at base to a fragile junction with a swollen, fleshy rhizome; siliques with a persistent style beak 5–8 mm long 2a. Rhizome lacking tooth-like projections or these very obscure, formed of segments joined by fragile connectives, the rhizome readily breaking into segments [Fig. 511]; stem above the leaves usually pubescent with spreading hairs; stems usually with 3 leaves in a single who. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. concatenata 2b. Rhizome with tooth-like projections, either continuous [Fig. 513] or with alternately enlarged and constricted areas, but not readily breaking into segments; stem above the leaves usually glabrous; stems with usually 2 opposite leaves or with 3 alternate leaves 3a. Cilia on leaf margin mostly 0.2–0.3 mm long and spreading; rhizome alternately enlarged and constricted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. maxima 3b. Cilia on leaf margin mostly 0.1 mm long and ascending or appressed; rhizome continuous or alternately enlarged and constricted 4a. Leaflets sessile; rhizome alternately enlarged and constricted . . . . . . . . . C. incisa 4b. Leaflets usually petiolulate; rhizome continuous [Fig. 513] . . . . . . . . . . . C. diphylla 1b. Principal leaf blades simple or pinnately lobed or pinnately divided [Figs. 512, 514, 515]; stems arising from caudices, taproots, or rhizomes, when arising from rhizomes the stem widest at base and narrowing upward and/or continuous with rhizome; siliques with a persistent style beak 0.3–6 mm long 5a. Lower leaf blades, as well as usually the upper, simple 6a. Petals up to 5 mm long or absent; fruiting pedicels 1–10 mm long; stems arising from a vertical taproot and caudex or fibrous roots 7a. Petals absent or up to 1.2 mm long and concealed by the sepals; siliques 5–12 mm long, spreading to ascending [Fig. 512]; fruiting pedicels 1–3 mm long; wetland and aquatic plants, primarily of fresh-tidal river shores . . . . . . . . . . . . . . . . (in part) C. longii 7b. Petals 3–5 mm long, visible; siliques 15–35 mm long, erect; fruiting pedicels 4–10 mm long; dwarf alpine plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bellidifolia 6b. Petals 7–16 mm long; lower fruiting pedicels 20–30 mm long; stems arising from a stout, tuber-like rhizome 8a. Sepals green turning yellow; petals white (rarely pink); stem glabrous or pubescent with appressed hairs shorter than 0.15 mm; stems mostly 20–40 cm tall to the first pedicel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bulbosa 8b. Sepals purple turning brown; petals pink to purple (rarely white); stem pubescent with spreading hairs 0.2–0.8 mm long; stems mostly 8–20 cm tall to the first pedicel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. douglassii 5b. Lower leaf blades, as well as usually the upper, pinnately divided or pinnately lobed with sinuses extending nearly to the midrib 9a. Leaf blades with 2 or 3 (rarely up to 5) leaflets [Fig. 512]; siliques 5–12 mm long; fruiting pedicels 1–3 mm long [Fig. 512]; petals absent or up to 1.2 mm long and not exceeding the sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. longii 9b. Principal leaf blades with 5–17 leaflets [Figs. 514, 516]; siliques 10–30 mm long; lower fruiting pedicels (4–) 5–18 mm long; petals 1.5–4 mm long, visible (sometimes absent in C. impatiens) 10a. Stem leaves sagittate-auriculate, with sharply toothed to lacerate-margined leaflets that are usually acuminate at the apex [Fig. 514]; petals absent or present and 2–3 mm long, scarcely exceeding the sepals . . . . . . . . . . . . . . . . . . . . . . . . . C. impatiens 10b. Stem leaves shortly petiolate, not auriculate, with entire to crenate-dentate leaflets that are rounded to acute at the apex [Fig. 516]; petals present, 1.5–15 mm long

Br a ss i c ac e a e   48 1

11a. Petals 8–15 mm long; styles 3–6 mm long; lower fruiting pedicels 12–18 mm long; plants perennial 12a. Primary veins of terminal leaflet of basal leaves terminating in a ± triangular, callus tip that is prominently excurrent beyond the leaflet blade tissue [Fig. 516]; petals pink to white in life, drying pink to pink-purple; leaflets of upper stem leaves commonly sessile . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pratensis 12b. Primary veins of terminal leaflet of basal leaves terminating in a callus region, but the callus either not excurrent beyond the leaflet blade tissue or scarcely excurrent as a semicircular (i.e., dome-shaped) tip [Fig. 512]; petals white in life, drying white, cream, yellow-white, or brown-white; leaflets of upper stem leaves commonly petiolulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. dentata 11b. Petals 1.5–4 mm long; styles 0.3–1.5 mm long; lower fruiting pedicels (3–) 5–14 mm long; plants annual or biennial (sometimes short-lived perennial in C. flexuosa) 13a. Basal leaves numerous, persistent, with hirsute petioles; stems with 2–5 (–6) leaf-bearing nodes; flowers with 4 stamens; siliques usually erect and parallel or nearly so to the raceme axis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hirsuta 13b. Basal leaves few, sometimes absent at anthesis, with glabrous petioles (always with hirsute petioles in C. flexuosa, a very rare introduction); stems with 4–10 (–20) leaf-bearing nodes; flowers with 6 stamens (some individuals rarely with only 4 stamens); siliques usually erect-ascending to ascending and not parallel to the raceme axis (except sometimes in C. parviflora) 14a. Stems glabrous; stem leaves mostly 20–40 mm long, with linear to oblanceolate lateral leaflets or segments that are neither petiolulate nor decurrent on the leaf rachis; styles 0.5–1 mm long; seeds 0.7–0.9 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. parviflora 14b. Stems usually at least sparsely hispid near the base (except in inundated plants); stem leaves mostly 40–100 mm long, with oblong to suborbicular lateral leaflets or segments that are either petiolulate or decurrent on the leaf rachis; styles (0.3–) 0.5–2 mm long; seeds (0.9–) 1–1.5 mm long 15a. Lateral leaflets or segments of stem leaves decurrent on leaf rachis; petioles glabrous; stems erect, not flexuous; native plant of shorelines and wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pensylvanica 15b. Lateral leaflets or segments of stem leaves not decurrent on leaf rachis; petioles usually hirsute; stems flexuous, sometimes decumbent-based; introduced plants primarily of human-disturbed places . . . . . . . . C. flexuosa 1. Cardamine bellidifolia L. var. bellidifolia

NC

alpine bitter-cress. ME, NH. Cliffs, ravines, and wet shelves in alpine areas. The report of this plant occurring in Cumberland County, ME, by Furbish (1901) is considered to be a labeling error. The plants were, in all likelihood, collected by Fernald from Piscataquis County, ME, as is noted in handwriting on the voucher collection (10 Sep 1900, Furbish s.n. NEBC!). 2. Cardamine bulbosa (Schreb. ex Muhl.) B.S.P. N bulbous bitter-cress. Arabis bulbosa Schreb. ex Muhl.; A. rhomboidea Pers.; Cardamine rhomboidea (Pers.) DC. • CT, MA, NH, VT; also reported from ME by Al-Shehbaz et al. (2010) and from RI by George (1992), but specimens are unknown. Swamps, ditches, low areas in florests, streams shores. 3. Cardamine concatenata (Michx.) Sw. N Fig. 511 cut-leaved toothwort. Cardamine laciniata (Muhl. ex Willd.) Wood; Dentaria concatenata Michx.; D. laciniata Muhl. ex Willd. • CT, MA, ME, NH, VT. Rich, mesic, upland and riparian forests.

Fig. 511  Section of rhizome of Cardamine concatenata showing constrictions.

482   tricolpate s

4. Cardamine dentata Schult.

N C Fig. 512

white cuckoo bitter-cress. Cardamine palustris (Wimmer & Grab.) Peterm.; C. pratensis L. ssp. paludosa (Knaf) Čelak; C. pratensis L. ssp. palustris (Wimmer & Grab.) Clausen; C. pratensis L. var. palustris Wimmer & Grab. • CT, MA, VT; western New England. Swamps and fens, usually in regions of high-pH bedrock. 5. Cardamine diphylla (Michx.) Wood N Fig. 513 two-leaved toothwort. Dentaria diphylla Michx.; D. incisa Small • CT, MA, ME, NH, VT. Mesic, deciduous upland and riparian forests. Fig. 512  Basal leaf blade of Cardamine dentata.

6. Cardamine douglassii Britt.

NC

pink bitter-cress. Arabis douglassii Torr. in Torr. & Gray; Cardamine rhomboidea (Pers.) DC. forma purpurea (Torr.) O.E. Schulz • CT, MA; also reported from NH by Gleason and Cronquist (1991), but specimens are unknown. Swamps, low areas in forests, and forested riparian terraces in regions of high-pH bedrock. 7. Cardamine flexuosa With. E

Fig. 513  Section of rhizome of Cardamine diphylla showing uniform diameter.

wavy bitter-cress. Cardamine hirsuta L. ssp. flexuosa (With.) Forbes & Hemsl.; C. sylvatica Link • CT, MA, RI, VT. Gardens, nurseries, greenhouses, stream banks, wet spots, logging roads, disturbed soil. What has been called Cardamine flexuosa in North America is actually comprised of two separate entities—“European C. flexuosa” and “Asian C. flexuosa.” These two entities show strong divergence in cpDNA, possess different ploidy levels, and have slightly different ecologies (Lihová et al. 2006). Unfortunately, more work is needed to identify morphological characters that will allow discrimination of these genetically distinct entities. The RI occurrence is known to be the latter (Lihová et al. 2006). 8. Cardamine hirsuta L. E hairy bitter-cress. CT, MA, ME, NH, VT; also reported from RI by George (1997), but specimens are unknown. Gardens, nurseries, lawns, roadsides. 9. Cardamine impatiens L. E Fig. 514 narrow-leaved bitter-cress. CT, MA, ME, NH, VT. Lawns, gardens, forest edges, roadsides. 10. Cardamine incisa (Eames) Schumann

Fig. 514  Leaf blade of Cardamine impatiens.

NC

Eames’ toothwort. Dentaria incisa Eames; D. incisifolia Eames ex Britt. • CT; also reported from MA by Sorrie and Somers (1999), but specimens are unknown. Rich, mesic, upland and riparian forests. 11. Cardamine longii Fern.

N C Fig. 515

Long’s bitter-cress. CT, MA, ME, NH; also reported from RI by George (1992), but specimens are unknown. Fresh to brackish-tidal river shores, rarely inland in streams and about lake inlets and outlets. 12. Cardamine maxima (Nutt.) Wood N large toothwort. Dentaria maxima Nutt. • CT, MA, ME, NH, VT. Rich, mesic, upland and riparian forests. 13. Cardamine parviflora L. var. arenicola (Britt.) O.E. Schulz N small-flowered bitter-cress. Cardamine arenicola Britt.; C. flexuosa With. var. gracilis O.E. Schulz • CT, MA, ME, NH, RI, VT. Rocky ridges and balds, cliffs, woodlands. 14. Cardamine pensylvanica Muhl. ex Willd. N Pennsylvania bitter-cress. Cardamine flexuosa With. ssp. pensylvanica (Muhl. ex Willd.) O.E. Schulz; C. pensylvanica Muhl. ex Willd. var. brittoniana Farw.; Dracamine pensylvanica (Muhl. ex Willd.) Nieuwl. • CT, MA, ME, NH, RI, VT; throughout. Streams, muddy banks, lake shores, swamps. Forms along tidal river shores are sometimes transitional to Cardamine longii in that they possess fewer pairs of leaflets. 15. Cardamine pratensis L. E Fig. 516 Fig. 515  Leaves and infructescence of Cardamine longii.

pink cuckoo bitter-cress. Dracamine pratensis (L.) Nieuwl. • CT, MA, ME, NH, RI, VT. Wet meadows, lawns, pond shores, riparian forests, swamps.

Br a ss i c ac e a e   4 83

Chorispora 1. Chorispora tenella (Pallas) DC. E crossflower. Chorispermum tenellum (Pallas) R. Br.; Raphanus tenellus Pallas • MA. Roadsides, fields, waste areas, often in sandy soil.

Conringia 1. Conringia orientalis (L.) Dumort. E hare’s-ear-mustard. Brassica orientalis L. • CT, MA, ME, NH, VT; also reported from RI by Warwick (2010b), but specimens are unknown. Fields, roadsides, railroads, waste areas.

Descurainia

Fig. 516  Basal leaf blade of Cardamine pratensis.

1a. Siliques clavate or subclavate, 1–2 mm wide, rounded to obtuse at apex; seeds aligned in 2 rows in each locule; leaf blades, stems, and axis of inflorescence stipitate-glandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. pinnata 1b. Siliques narrow-linear, 0.5–1.3 (–1.5) mm wide, acute at the apex; seeds aligned in 1 row in each locule; plants usually eglandular 2a. Fruiting pedicels ascending to spreading, 8–15 mm long; siliques 10–20 (–30) mm long; seeds 10–20 in each locule; leaf blades usually twice or thrice pinnately divided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sophia 2b. Fruiting pedicels erect to ascending, 2–4 (–6) mm long; siliques 5–10 mm long; seeds 4–8 in each locule; leaf blades usually once pinnately divided, the leaflets entire to lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. incana 1. Descurainia incana (Bernh. ex Fisch. & C.A. Mey.) Dorn

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mountain tansy-mustard. Descurainia richardsonii (Sweet) O.E. Schulz; Sisymbrium incanum Bernh. ex Fisch. & C.A. Mey.; Sophia richardsonii (Sweet) Rydb. • CT, MA, ME, VT. Cliffs, talus, wool waste, disturbed soil, railroads, yards. This species is native to Aroostook and Piscataquis Counties, ME, it is introduced elsewhere. Though the report by Bean et al. (1961) for MA was based on Descurainia pinnata ssp. brachycarpa, D. incana is known from the state by a recently discovered specimen at DINH! that was originally determined incorrectly. 2. Descurainia pinnata (Walt.) Britt. ssp. brachycarpa (Richards.) Detling

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western tansy-mustard. Descurainia brachycarpa (Richards.) O.E. Schulz; D. pinnata (Walt.) Britt. var. brachycarpa (Richards.) Fern.; Sisymbrium brachycarpum Richards.; Sophia brachycarpa (Richards.) Rydb. • MA, ME, NH, VT. Cliffs, limestone headlands, wool waste. This species is native to NH and VT and is introduced elsewhere. 3. Descurainia sophia (L.) Webb ex Prantl E fine-leaved tansy-mustard. Sisymbrium sophia L.; Sophia sophia (L.) Britt. • CT, MA, ME, NH, RI, VT. Farm yards, fields, roadsides, waste areas, railroads.

Diplotaxis 1a. Plants perennial, with ± woody stem bases; leaves borne throughout the lower half of the stem; silique borne on a stipe 1–2 mm long; fruiting pedicels (15–) 20–30 mm long; sepals (4–) 5–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. tenuifolia 1b. Plants annual or biennial, herbaceous throughout; leaves borne chiefly at the base of the plant; silique not stipitate; fruiting pedicels (6–) 10–13 (–15) mm long; sepals 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. muralis 1. Diplotaxis muralis (L.) DC. E annual wall-rocket. Sisymbrium murale L.; Sinapis muralis (L.) R. Br. • CT, MA, ME. Fields, roadsides, wool waste, lawns, waste areas.

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2. Diplotaxis tenuifolia (L.) DC. E perennial wall-rocket. Sisymbrium tenuifolium L. • CT, MA, ME. Fields, roadsides, railroads, yards, waste areas.

Draba Micromorphological details of pubescence requiring 20× or greater magnification are important for confident determinations of some species. Reference: Mulligan (1976). 1a. Petals bilobed with a deep, apical sinus; stems scapose (i.e., all the leaves confined to a basal cluster) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. verna 1b. Petals entire or emarginate at apex; stems with at least 1 leaf-bearing node 2a. Plants annual; rachis of infructescence congested, 1–16 (–35) mm long; stem leaves all borne at base just above the basal rosette of leaves . . . . . . . . . . . . . . . . . . . . . . . . . D. reptans 2b. Plants perennial from a caudex; rachis of infructescence relatively elongate, (16–) 20–78 mm long; stem leaves more evenly distributed, usually some produced in the apical half of the stem 3a. Lowest 1–3 flowers each subtended by a leafy bract; siliques pubescent with stellate hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cana 3b. None of the flowers subtended by leafy bracts; siliques glabrous or sparsely pubescent with simple or forked hairs 4a. Stellate hairs of abaxial surface of basal leaves borne on short stalks, the radiating branches of the hairs elevated above the surface of the leaf blade [Fig. 518]; styles up to 0.3 (–0.4) mm long; stigmas often bilobed . . . . . . . . . . . . . . . . . . D. glabella 4b. Stellate hairs of abaxial surface of basal leaves sessile, the radiating branches of the hairs appressed to the leaf surface [Fig. 517]; styles (0.3–) 0.4–1 mm long; stigmas scarcely bilobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. arabisans 1. Draba arabisans Michx. N C Fig. 517 rock whitlow-mustard. Draba incana L. var. arabisans (Michx.) S. Wats. • ME, VT. Cliffs, talus, and headlands in high-pH bedrock regions, ascending to subalpine situations in northern VT. Fig. 517  Compound hairs on basal leaves of Draba arabisans showing nearly obsolete stalk.

Fig. 518  Compound hairs on basal leaves of Draba glabella showing evident stalk.

2. Draba cana Rydb. NC canescent whitlow-mustard. Draba breweri S. Wats. var. cana (Rydb.) Rollins; D. lanceolata, auct. non Royle • ME, NH, VT; northern portion of states. Cliffs and talus, usually in regions of high-pH bedrock, ascending to subalpine situations in northern VT. 3. Draba glabella Pursh N C Fig. 518 smooth whitlow-mustard. Draba arabisans Michx. var. canadensis (Burnet) Fern. & Knowlt.; D. canadensis Burnet; D. daurica DC.; D. glabella Pursh var. megasperma (Fern. & Knowlt.) Fern.; D. glabella Pursh var. orthocarpa (Fern. & Knowlt.) Fern.; D. megasperma Fern. & Knowlt. • ME, VT; northern portions of states. Cliffs and headlands in high-pH bedrock regions. 4. Draba reptans (Lam.) Fern. N Carolina whitlow-mustard. Arabis reptans Lam.; Draba caroliniana Walt.; D. micrantha Nutt. ex Torr. & Gray; D. reptans (Lam.) Fern. var. micrantha (Nutt. ex Torr. & Gray) Fern.; D. umbellata Muhl. • CT, MA, RI. Sandy and rocky fields, ledges, balds. 5. Draba verna L. E spring whitlow-mustard. Draba verna L. var. aestivalis Lej.; Draba verna L. var. boerhaavii van Hall; Erophila verna (L.) Bess.; E. verna (L.) Bess. ssp. praecox (Stev.) S.M. Walters • CT, MA, ME, NH, RI, VT. Lawns, fields, roadsides, gardens, waste areas.

Eruca 1. Eruca vesicaria (L.) Cav. ssp. sativa (P. Mill.) Thelung E garden-rocket. Brassica eruca L.; Eruca sativa P. Mill. • CT, MA, VT. Fields, roadsides, gardens.

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Erucastrum 1. Erucastrum gallicum (Willd.) O.E. Schulz E common dog-mustard. Brassica erucastrum L.; Sisymbrium gallicum Willd. • CT, MA, ME, NH, VT. Fields, roadsides, railroads, waste areas.

Erysimum Reference: Al-Shehbaz (2010a). 1a. Fruiting pedicels thick, ± the same diameter as the siliques, widely spreading at or nearly at right angles to the axis [Fig. 519]; rachis of inflorescence often flexuous . . . . . . E. repandum 1b. Fruiting pedicels slender, narrower than the siliques, erect to spreading-ascending; rachis of inflorescence straight 2a. Petals (7–) 8–25 (–30) mm long; sepals (4–) 5–14 mm long; siliques (20–) 40–70 mm long 3a. Petals 13–25 (–30) mm long, commonly orange to orange-yellow; sepals 7–14 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. capitatum 3b. Petals (7–) 8–18 mm long, commonly yellow; sepals (4–) 5–10 mm long 4a. Petals 12–18 mm long; sepals 6–10 mm long; anthers 2 mm long or longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. odoratum 4b. Petals (7–) 8–10 (–12) mm long; sepals (4–) 5–6 mm long; anthers 1–2.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. hieraciifolium 2b. Petals 3.5–7 (–8) mm long; sepals 2–5 (–6) mm long; siliques 15–50 mm long 5a. Anthers 0.4–0.6 mm long; sepals 2–3.5 mm long; petals bright yellow, 3.5–5 mm long; siliques glabrous to sparsely pubescent, 15–20 (–30) mm long, pubescent on the inner surface of the valves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. cheiranthoides 5b. Anthers 1.5–2.5 mm long; sepals 4–5 (–6) mm long; petals pale yellow, 5–7 (–8) mm long; siliques pubescent, 30–50 mm long, glabrous on the inner surface of the valves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. inconspicuum 1. Erysimum capitatum (Dougl. ex Hook.) Greene var. capitatum E sand-dune wallflower. Cheiranthus capitatus Dougl. ex Hook.; Erysimum arkansanum Nutt; E. asperum (Nutt.) DC. var. purshii Dur.; E. capitatum (Dougl. ex Hook.) Greene var. purshii (Dur.) Rollins • ME. Forests and forest edges, margins of trails. 2. Erysimum cheiranthoides L. E wormseed wallflower. Cheirinia cheiranthoides (L.) Link • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, waste areas. 3. Erysimum hieraciifolium L. E European wallflower. Erysimum strictum P.G. Gaertn., B. Mey., & Scherb. • MA, VT. Fields, roadsides, waste areas. 4. Erysimum inconspicuum (S. Wats.) MacM. var. inconspicuum E shy wallflower. Cheirinia inconspicua (S. Wats.) Britt.; Erysimum parviflorum Nutt. • MA, ME, NH. Fields, roadsides, railroads, waste areas. 5. Erysimum odoratum Ehrh. E fragrant wallflower. Erysimum pannonicum Crantz • MA. Wool waste. 6. Erysimum repandum L. E Fig. 519 spreading wallflower. Cheirinia repanda (L.) Link • MA. Fields, roadsides, waste areas.

Euclidium 1. Euclidium syriacum (L.) Ait. f. E Syrian-mustard. Anastatica syriaca L. • MA. Farm yards.

Fig. 519  Inflorescence of Erysimum repandum.

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Hesperis 1. Hesperis matronalis L. E dame’s-rocket. CT, MA, ME, NH, RI, VT. Roadsides, forest edges, gardens, riparian forests, waste areas.

Iberis 1a. Leaf blades usually with few and distant, irregular, large teeth (rarely entire); siliques 3–5 mm long; stems hirsute, at least on the lower portion; axis of the inflorescence elongating in fruit, the infructescence resembling a raceme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. amara 1b. Leaf blades entire or nearly so; siliques 5.5–7 (–10) mm long; stems glabrous; axis of the inflorescence elongating or not 2a. Plants perennial, with evergreen leaves; petals white (sometimes tinged with lilac); axis of the inflorescence elongating in fruit, the infructescence resembling a raceme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. sempervirens 2b. Plants annual, with deciduous leaves; petals pink to purple; axis of the inflorescence not or scarcely elongating in fruit, the infructescence resembling an umbel . . I. umbellata 1. Iberis amara L. E rocket candytuft. CT, MA, ME. Gardens, forest edges, waste areas, dumps, ledges near areas of habitation. The report of this species from VT by Atwood et al. (1973) is based on a collection of a cultivated specimen (Blanchard collection at HNH). 2. Iberis sempervirens L. E edging candytuft. NH, VT. Waste areas. 3. Iberis umbellata L. E globe candytuft. ME, NH. Gardens, railroads, waste areas.

Lepidium Lepidium ramosissimum A. Nels. was reported from ME by Rollins (1993), but specimens are unknown. Reference: Al-Shehbaz and Gaskin (2010). 1a. Silicles with a wrinkled texture due to raised veins or reticulum; racemes borne laterally from the axils of leaves 2a. Silicles with an evident basal and apical sinus, reticulate to rugose over the exterior surface [Fig. 521]; styles obsolete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. didymum 2b. Silicles lacking both basal and apical sinuses, coarsely veined and with protruding, wartlike processes over the exterior surface; styles present, ca. 0.5 mm long . . . . . . L. squamatum 1b. Silicles with a smooth surface; racemes chiefly terminal and at the tips of branches 3a. Silicles without a notch at the apex; plants perennial from rhizomes or stolons 4a. Leaf blades sagittate-clasping; silicles subglobose; flowering raceme typically flat or sometimes even concave-topped, the lower pedicels equaling the height of the inflorescence; plants 2–5 dm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. draba 4b. Leaf blades shortly petiolate or sessile, but not clasping the stem; silicles flattend perpendicular to the septum; flowering raceme typically round-topped, the lower pedicels shorter than the height of the inflorescence; plants (5–) 7–13 dm tall . . . . . . . . L. latifolium 3b. Silices with a notch or sinus at the apex; plants annual from taproots or fibrous roots 5a. Stem leaves either sagittate- or cordate-clasping [Fig. 520]; flowers with 6 stamens 6a. Stem leaves cordate-clasping, the basal lobes rounded; lower leaf blades 2 or 3 times divided into linear segments; petals yellow, 1–1.5 mm long; fruiting pedicels glabrous; silicles ca. 3.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. perfoliatum

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6b. Stem leaves auriculate-clasping, the basal lobes usually acute [Fig. 520]; lower leaf blades entire to toothed or sometimes once pinnately lobed; petals white, 1.8–2.8 mm long; fruting pedicels pubescent; silicles 5–6 mm long 7a. Style 0.2–0.5 (–0.7) mm long, not or only shortly exserted beyond the apical notch of the fruit; valves of silique papillose; plants annual, without a caudex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. campestre 7b. Style (0.6–) 1–1.5 mm long, prominently exserted beyond the apical notch of the fruit; valves of silique usually not papillose; plants perennial, with a caudex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. heterophyllum 5b. Stem leaves narrowed to the base, not surrounding the stem; flowers with 2 or 4 stamens except L. sativum with 6 stamens 8a. Flowers with 6 stamens; silicles 5–6 mm long, prominently winged; cotyledons 3-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sativum 8b. Flowers with 2 (rarely 4) stamens; silicles 2–4 mm long, without a wing or winged only in the apical portion; cotyledons entire 9a. Petals typically present, 1–2 times as long as the sepals [Fig. 521]; rachis of raceme, when pubescent, with curved, cylindrical hairs; cotyledons accumbent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. virginicum 9b. Petals obsolete or rudimentary and shorter than the sepals; rachis of raceme pubescent with straight, cyclindrical to subclavate hairs; cotyledons incumbent 10a. Racemes densely flowered, with 9–18 fruits per cm; silicles obovate to broad-obovate; plants inodorous; lower surface of fruiting pedicels smooth; seeds with an apical, narrow wing-margin . . . . . . . . . . . . . . . . . L. densiflorum 10b. Racemes relatively sparsely flowered, with 6–10 fruits per cm; silicles elliptic; plants foul-scented; lower surface of fruiting pedicels sparsely papillose; seeds lacking a wing margin . . . . . . . . . . . . . . . . . . . . . . . . L. ruderale 1. Lepidium campestre (L.) Ait. f. E Fig. 520 field pepperweed. Neolepia campestris (L.) W.A. Weber; Thlaspi campestre L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, waste areas. 2. Lepidium densiflorum Schrad. var. densiflorum E prairie pepperweed. Lepidium densiflorum Schrad. var. typicum Thellung; L. neglectum Thellung • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, waste areas.

Fig. 520  Inflorescence and upper leaves of Lepidium campestre.

3. Lepidium didymum L. E Fig. 521 lesser swine-cress. Carara didyma (L.) Britt. in Britt. & Brown; Coronopus didymus (L.) Sm.; Senebiera didyma (L.) Pers. • CT, MA, ME, NH, RI. Fields, roadsides, railroads, waste areas, lawns. 4. Lepidium draba L. E heart-podded pepperweed. Cardaria draba (L.) Desv. • CT, MA, ME, RI, VT. Fields, gardens, roadsides, waste areas. 5. Lepidium heterophyllum Benth. E variable-leaved pepperweed. MA, ME. Lawns, waste areas. 6. Lepidium latifolium L. E broad-leaved pepperweed. Cardaria latifolia (L.) Spach • CT, MA, NH. Fields, roadsides, waste areas, shorelines, sea beaches, coastal marshes. 7. Lepidium perfoliatum L. E clasping pepperweed. CT, MA, ME; also reported from RI by George (1992), but specimens are unknown. Fields, roadsides, railroads, waste areas. 8. Lepidium ruderale L. E stinking pepperweed. CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, waste areas.

Fig. 521  Silicles of Lepidium didymum.

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9. Lepidium sativum L. E garden pepperweed. CT, MA, ME, NH, RI, VT. Fields, roadsides, gardens, waste areas. 10. Lepidium squamatum Forsk. E greater swine-cress. Carara coronopus (L.) Medik.; Cochlearia coronopus L.; Coronopus squamatus (Forsk.) Aschers. • MA, RI; also reported from CT by Hultén and Fries (1986), but specimens are unknown. Fields, roadsides, waste areas. 11. Lepidium virginicum L. var. virginicum N Fig. 521 poor-man’s pepperweed. Lepidium virginicum L. ssp. eu-virginicum Thellung • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, gardens, waste areas.

Lobularia 1. Lobularia maritima (L.) Desv. E sweet-alyssum. Alyssum maritimum (L.) Lam.; Clypeola maritima L.; Koniga maritima (L.) R. Br. • CT, MA, ME, RI, VT. Gardens, waste areas. Fig. 522  Inflorescence and upper leaves of Lepidium virginicum.

Lunaria 1. Lunaria annua L. E Fig. 523 annual honesty. CT, MA, ME, RI, VT. Fields, roadsides, banks, waste areas.

Nasturtium The leaves in deep-water plants can have simple blades, contrary to the normal condition of pinnately compound blades. 1a. Seeds aligned in 1 row in each locule, with 75–150 (–175) polygonal depressions on each surface; silique 15–25 × 1–1.5 mm wide, terete in cross-section, with a slender, persistent style beak 0.5–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. microphyllum Fig. 523  Silicle of Lunaria annua.

1b. Seeds aligned in 2 rows in each locule, with 25–50 (–60) polygonal depressions on each surface; silique 10–15 × 1.5–3 mm, compressed somewhat perpendicular to the septum, with a short, thick, persistent style beak up to 0.5 (–1) mm long or the beak absent altogether . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. officinale 1. Nasturtium microphyllum Boenn. ex Reichenb. E one-rowed water-cress. Nasturtium officinale Ait. f. var. microphyllum (Boenn. ex Reichenb.) Thellung; Rorippa microphylla (Boenn. ex Reichenb.) Hyl. ex A. & D. Löve • CT, MA, ME, NH, RI, VT. Streams, ditches, outlets. 1‌ × 2. Nasturtium ×sterile (Airy-Shaw) Oefel. is a rare water-cress hybrid in New England known from CT, NH. It is recognized by its sterile siliques, which do not produce well-formed seeds. 2. Nasturtium officinale Ait. f. E two-rowed water-cress. Baeumerta nasturtium-aquaticum (L.) Hayek; Rorippa nasturtiumaquaticum (L.) Hayek; Sisymbrium nasturtium-aquaticum L. • CT, MA, ME, NH, RI, VT. Streams, ditches, outlets.

Neslia 1. Neslia paniculata (L.) Desv. E yellow ball-mustard. Myagrum paniculatum L. • CT, MA, ME, NH, VT. Fields, roadsides, railroads, waste areas.

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Raphanus Yang et al. (2002) have shown that species of Raphanus are derived from an ancestor that originated by hybridization between two different lineages of Brassica—the B. nigra clade and the B. rapa clade. Therefore, species of Raphanus are actually members of the genus Brassica. However, formal combinations under the genus Brassica have not yet been made. 1a. Siliques 3–6 mm thick, of uniform diameter through most of its length except for the beak and prominent constrictions between the seeds [Fig. 524], longitudinally striate, containing 4–12 seeds; petals usually yellow fading to white; taproot slender . . . . . . . . . R. raphanistrum 1b. Siliques 5–10 mm thick, widest below the middle and tapering toward the beak, not or scarcely constricted between the seeds, smooth or inconspicuously striate, containing 1–3 (–5) seeds; petals usually purple; taproot usually conspicuously thickened . . . . . R. sativus 1. Raphanus raphanistrum L. ssp. raphanistrum E Fig. 524 wild radish. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, Atlantic coast beaches. 2. Raphanus sativus L. E cultivated radish. Raphanus raphanistrum L. var. sativus (L.) G. Beck • CT, MA, ME, NH, RI, VT. Fields, roadsides, gardens.

Rapistrum 1. Rapistrum rugosum (L.) All. E annual bastard-cabbage. Myagrum rugosum L. • MA. Fields, roadsides, waste areas. Three subspecies of Rapistrum rugosum are recognized in Europe (where this species is native). In New England, most specimens are ± referable to ssp. orientale (L.) Arcang.; however, several collections at GH! and NEBC! approach ssp. rugosum in one or more characterisitcs. Given the variability of fruiting characters, I follow Rollins (1993) and Warwick (2010c) in not recognizing infraspecific taxa.

Rorippa Reference: Stuckey (1972). 1a. Petals white; submersed leaf blades highly dissected into narrow segments; septum incomplete, residual as narrow strips on the inner margins of the replum . . . . . . . . R. aquatica 1b. Petals yellow or absent; leaf blades subentire to pinnately lobed; septum complete 2a. Petals absent; fruiting pedicels 0.5–1.5 (–2) mm long . . . . . . . . . . . . . . . . . . R. sessiliflora 2b. Petals present; fruiting pedicels 2–15 mm long 3a. Petals 0.7–3.5 mm long, shorter than to approximately equaling the length of the sepals; plants usually annual or biennial (rarely short-lived perennial), from tap roots; anthers 0.2–0.5 mm long 4a. Silicles ovoid-obloid to broad-ovoid, (2–) 3–10 mm long; seeds papillose; petals 0.8–3.5 mm long; abaxial leaf blades and lower stem glabrous or pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. palustris 4b. Silicles spherical to subspherical, 2–3 (–3.5) mm long; seeds reticulate-patterned; petals 0.7–1.3 (–1.5) mm long; abaxial leaf blades and lower stem pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. globosa 3b. Petals (2–) 2.8–5.5 mm long, longer than the sepals; plants perennial from rhizomes or creeping stem bases; anthers 0.4–1.3 mm long 5a. Fertile siliques 10–20 mm long, linear in outline, those from unfertilized ovaries developing only 4–8 (–10) mm long; middle and upper stem leaves usually pinnately lobed to pinnately divided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. sylvestris

Fig. 524  Silique of Raphanus raphanistrum showing constrictions between the seeds.

490  tricolpates

5b. Siliques (2.5–) 3–5 (–6) mm long, ovoid-obloid to subspherical; middle and upper stem leaves entire to coarsely serrate 6a. Fruiting pedicels weakly sigmoid-curved downward to somewhat recurved [Fig. 525]; silicles ovoid-obloid [Fig. 525]; stem leaves clasping at the base or, more commonly, not clasping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. amphibia 6b. Fruiting pedicels straight, widely ascending [Fig. 526]; silicles spherical or subspherical [Fig. 526]; lower and middle stems leaves auriculate-clasping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. austriaca 1. Rorippa amphibia (L.) Bess. E Fig. 525 great yellow-cress. Armoracia amphibia (L.) Peterm.; Nasturtium amphibium (L.) Ait. f.; Radicula amphibia (L.) Druce; Sisymbrium amphibium L. • MA, ME. Hydric soils of streams, pond shores, and disturbed places. Previous reports of Rorippa amphibia from CT were based on specimens of R. × (fide Les Mehrhoff). ‌prostrata Fig. 525  Inflorescence of Rorippa amphibia.

‌1 × 7. Rorippa ×prostrata (Bergeret) Schinz & Thellung is a rare hybrid of two nonnative yellow-cress species. It is known from CT, MA. It has solid stems (hollow in R. amphibia), usually pinnately lobed lower leaf blades, sessile or subsessile upper leaves, obloid silicles mostly 5–6 mm long, and does not root from the lower, leafbearing nodes (as often does R. amphibia). Rorippa × is usually found on dry ‌prostrata banks, along roadsides, and in fields (whereas R. amphibia is a species of hydric soils). Records from MA previously thought to be R. austriaca were in fact this hybrid. Likewise, CT records previously thought to be R. amphibia were also this hybrid. 2. Rorippa aquatica (Eat.) Palmer & Steyermark

NC

lake yellow-cress. Armoracia aquatica (Eat.) Wieg.; A. lacustris (Gray) Al-Shehbaz & Bates; Nasturtium lacustre Gray; Neobeckia aquatica (Eat.) Greene; Rorippa americana (Gray) Britt. • MA, VT; western portion of states. Lakes and slow-moving rivers in high-pH bedrock regions. 3. Rorippa austriaca (Crantz) Bess. E Fig. 526 Austrian yellow-cress. Nasturtium austriacum Crantz • CT. Cultivated fields, roadsides, waste ‌prostrata areas. Previous reports of Rorippa austrica from MA were based on records of R. × (fide Les Mehrhoff). 4. Rorippa globosa (Turcz. ex Fisch. & C.A. Mey.) Hayek E globe yellow-cress. Nasturtium globosum Turcz. ex Fisch. & C.A. Mey. • MA. Dumps, waste areas. Fig. 526  Inflorescence of Rorippa austriaca.

5. Rorippa palustris (L.) Bess. N common yellow-cress. 5a. Nasturtium palustre (L.) DC. var. hispidum (Desv.) Gray; Radicula hispida (Desv.) Britt.; Rorippa hispida (Desv.) Britt.; R. islandica (Oeder ex Murr.) Borbás var. hispida (Desv.) Butters & Abbe; R. palustris ssp. hispida (Desv.) Jonsell; 5b. Radicula palustris (L.) Moench; Rorippa hispida (Desv.) Britt. var. glabrata Lunell; R. islandica (Oeder ex Murr.) Borbás ssp. fernaldiana (Butters & Abbe) Hultén; R. islandica (Oeder ex Murr.) Borbás var. fernaldiana Butters & Abbe; R. islandica (Oeder ex Murr.) Borbás var. glabra (O.E. Schulz) Welsh & Reveal; R. islandica (Oeder ex Murr.) Borbás var. glabrata (Lunell) Butters & Abbe; R. islandica (Oeder ex Murr.) Borbás var. microcarpa (Regel) Fern.; R. palustris (L.) Bess. ssp. fernaldiana (Butters & Abbe) Jonsell; R. palustris (L.) Bess. var. fernaldiana (Butters & Abbe) R. Stuckey; R. palustris (L.) Bess. ssp. glabra (Butters & Abbe) R. Stuckey; R. palustris (L.) Bess. var. glabra (O.E. Schulz) R. Stuckey; R. palustris (L.) Bess. var. glabrata (Lunell) R. Stuckey; R. palustris (L.) Bess. var. williamsii (Britt.) Hultén • CT, MA, ME, NH, RI, VT. Streams, lake shores, ditches, swamps, tidal and non-tidal marshes. 1a. Abaxial surface of leaf blades usually hirsute; stem normally hirsute up to the base of the terminal inflorescence, less frequently glabrate on the upper portion of the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5a. R. palustris var. hispida (Desv.) Rydb. 1b. Abaxial surface of leaf blades glabrous; stem glabrous or sparsely hirsute near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5b. R. palustris var. palustris

Br a ss i c ac e a e   49 1

Variety hispida is known from CT, MA, ME, NH, RI, VT. Variety palustris is known from CT, MA, ME, NH, RI, VT. Variety palustris has been further divided based primarily on plant size and leaf texture (among other characters, plants with taller and thicker stems and thicker leaves were recognized as var. fernaldiana). However, variation in these characters is continuous and overlapping. 6. Rorippa sessiliflora (Nutt. ex Torr. & Gray) A.S. Hitchc. E stalkless yellow-cress. Nasturtium sessiliflorum Nutt. ex Torr. & Gray; Radicula sessiliflora (Nutt. ex Torr. & Gray) Greene • MA. Gardens, rivers, streams, ditches. 7. Rorippa sylvestris (L.) Bess. E creeping yellow-cress. Nasturtium sylvestre (L.) R. Br.; Radicula sylvestris (L.) Druce; Sisymbrium sylvestre L. • CT, MA, ME, NH, RI, VT. Wet spots in cultivated fields, stream shores, disturbed, hydric areas.

Sinapis 1a. Silique glabrous or sparsely bristly, ca. 2 mm wide, with 5–12 seeds; indehiscent beak of silique compressed-quadrangular, 10–12 mm long, up to 1 times as long as the silique body; fruiting pedicels ascending, 3–5 mm long; leaf blades, especially the middle and upper ones, often merely toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. arvensis 1b. Silique densely bristly, ca. 4 mm wide, with 4–8 seeds; indehiscent beak of silique flat, 15–30 mm long, 1 or more times as long as the silique body; fruiting pedicels wide-spreading, 5–12 mm long; most of the leaf blades prominently pinnately lobed . . . . . . . . . . . . . . . . . . S. alba 1. Sinapis alba L. E white charlock. Brassica alba Rabenh.; B. hirta Moench • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, waste areas. 2. Sinapis arvensis L. n Fig. 527 corn charlock. Brassica arvensis Rabenh.; B. kaber (DC.) L.C. Wheeler; B. kaber (DC.) L.C. Wheeler var. pinnatifida (Stokes) L.C. Wheeler • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, waste areas. Ethnobotanical studies reveal this species was present in New England long before European visitors reached North America (Jacobson et al. 1988). However, most populations today are likely introductions from Europe.

Sisymbrium Reference: Al-Shehbaz (2010b). 1a. Fruiting pedicels erect to appressed, 2–3 mm long [Fig. 529]; siliques subulate in outline, (7–) 10–14 (–18) mm long [Fig. 529]; petals 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . S. officinale 1b. Fruiting pedicels spreading to nearly erect, 3–20 mm long [Fig. 528]; siliques linear-cylindric, (20–) 30–100 (–130) mm long [Fig. 528]; petals 6–10 mm long (only 2.5–4 mm in S. irio) 2a. Fruiting pedicels nearly as thick as the silique they support [Fig. 528]; mature siliques 30–100 mm long 3a. Upper stem leaf blades pinnately divided into linear to lanceolate segments; outer sepals corniculate; fruiting pedicels (4–) 6–10 mm long . . . . . . . . . . . . . . . . . S. altissimum 3b. Upper stem leaf blades simple or hastate (i.e., with 2 outward-pointing basal lobes); outer sepals not corniculate; fruiting pedicels 3–6 mm long . . . . . . . . . . . . . . . S. orientale 2b. Fruiting pedicels slender, thinner than the silique they support; mature siliques 20–40 mm long 4a. Immature siliques often overtopping the flowers in the apical portion of the inflorescence; petals 2.5–4 mm long; plants glabrous or sparsely pubescent; mature siliques 30–40 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. irio

Fig. 527  Flower of Sinapis arvensis.

492  tricolpates

4b. Immature siliques not overtopping the flowers in the apical portion of the inflorescence; petals 6–8 mm long; plants sparsely to densely pubescent below; mature siliques 20–35 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. loeselii 1. Sisymbrium altissimum L. E Fig. 528 tumbling hedge-mustard. Hesperis altissima (L.) Kuntze; Norta altissima (L.) Britt. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 2. Sisymbrium irio L. E London hedge-mustard. Descurainia irio (L.) Webb & Berth.; Erysimum irio (L.) Farw.; Norta irio (L.) Britt. • CT, MA. Fields, roadsides, waste areas. 3. Sisymbrium loeselii L. E tall hedge-mustard. Erysimum loeselii (L.) Farw.; Hesperis loeselii (L.) Kuntze; Norta loeselii (L.) Rydb.; Turritis loeselii (L.) R. Br. • CT, MA, NH, VT; also reported from ME by Kartesz (1999), but specimens are unknown. Fields, roadsides, waste areas. 4. Sisymbrium officinale (L.) Scop. E Fig. 529 Fig. 528  Inflorescence of Sisymbrium altissimum.

common hedge-mustard. Erysimum officinale L.; Sisymbrium officinale (L.) Scop. var. leiocarpum DC. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 5. Sisymbrium orientale L. E Indian hedge-mustard. Brassica kaber (DC.) L.C. Wheeler var. orientalis (L.) Scoggan; Sisymbrium columnae Jacq. • MA. Fields, roadsides, waste areas.

Subularia 1. Subularia aquatica L. ssp. americana Mulligan & Calder

N C Fig. 530

American water-awlwort. Subularia aquatica L. var. americana (Mulligan & Calder) Boivin • ME, NH, VT. Shallow water of lakes, usually on sand substrate.

Teesdalia 1. Teesdalia nudicaulis (L.) Ait. f. E common shepherd’s-cress. Iberis nudicaulis L. • CT, MA. Fields, roadsides, disturbed soil.

Fig. 529  Inflorescence of Sisymbrium officinale.

Thlaspi 1. Thlaspi arvense L. E Fig. 531 field penny-cress. Teruncius arvensis (L.) Lunnell • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas.

Tropidocarpum 1. Tropidocarpum gracile Hook. E Dobie-pod. Tropidocarpum dubium A. Davids.; T. gracile Hook. var. dubium (A. Davids.) Jepson • MA. Wool waste, waste areas.

Turritis 1. Turritis glabra L. N Fig. 530  Habit of Subularia aquatica.

tower-mustard. Arabis glabra (L.) Bernh. • CT, MA, ME, NH, RI, VT. Woodlands, dry fields, roadsides, railroads, disturbed soil.

B ux ac e a e   49 3

Buxaceae Pachysandra 1. Pachysandra terminalis Sieb. & Zucc. E Japanese mountain-spurge. CT, MA, NH. Forest borders, lawn edges, areas of habitation.

Cactaceae

Fig. 531  Silicles of Thlaspe arvense.

Opuntia 1. Opuntia humifusa (Raf.) Raf. n Fig. 532 eastern prickly-pear. Cactus humifusus Raf.; Opuntia ammophila Small; O. calcicola Wherry; O. compressa J.F. Macbr. • CT, MA, RI; mainly along the coastal plain. Dry fields, ledges, Atlantic coast dunes and beaches. Introduced to Middlesex County, MA.

Campanulaceae Fig. 532  Flower and portion of stem of Opuntia humifusa.

1a. Corolla zygomorphic [Figs. 539, 540]; stamens connate . . . . . . . . . . . . . . . . . . . . . . . . . Lobelia 1b. Corolla actinomorphic [Figs. 536, 537]; stamens distinct or connate only at base 2a. Flowers and fruits solitary, sessile or subsessile in the axils of leaves [Fig. 541]; capsules slender-cylindric; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triodanis 2b. Flowers and fruits solitary or borne in a spike, raceme, or glomerule, sessile or pedicellate [Figs. 534, 536, 537]; capsules globose or ovoid to obovoid or obconic; plants perennial 3a. Corolla slenderly tubular, cleft into 5 lobes; anthers connate at base; capsule dehiscing by 2 apical valves; inflorescence a dense, capitulum-like umbel that is subtended by a foliaceous involucre [Fig. 538] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jasione 3b. Corolla ± campanulate (narrow-campanulate in one species of Campanula); anthers distinct; capsule dehiscing by basal, lateral, or subapical pores; inflorescence open and not congested (congested in Campanula glomerata) 4a. Inflorescence a terminal panicle of primarily erect to ascending flowers [Fig. 537] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gadellia 4b. Inflorescece of solitary pedicellate flowers, or of sessile glomerulate flowers, or a spike or raceme, the flowers spreading to nodding or erect [Figs. 533, 534, 535] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campanula

494 tricolpates

Campanula Phylogenetic work using nrDNA suggests that the genus Campanula is not monophyletic (Eddie et al. 2003). Specifically, some bellflowers (e.g., C. aparinoides, C. carpatica, C. persicifolia, C. rotundifolia) may need to be segregated from the remainder of Campanula s.s. in order that recognized taxa be monophyletic. Campanula bononiensis L. was reported from ME by Magee and Ahles (1999), but the specimen was misidentified and is C. rapunculoides—Fernald 2139 (NEBC!). 1a. Style conspicuously exserted from the corolla; corolla globose-campanulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. divaricata 1b. Style included within the corolla to ± reaching the tips of the corolla lobes; corolla campanulate, tubular-campanulate, funnelform-campanulate, or rotate-campanulate 2a. Corolla 4–13 mm long; stems 3-angled, slender, weak, leaning on other vegetation [Fig. 533] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. aparinoides 2b. Corolla 10–55 mm long; stems terete or obscurely angled, erect to ascending 3a. Calyx with an appendage between the lobes 4a. Corolla broad-campanulate, yellow-white, unspotted . . . . . . . . . . . . . . . . C. lanata 4b. Corolla tubular-campanulate, usually white to light purple, usually spotted within . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. punctata 3b. Calyx without appendages between the lobes 5a. Flowers sessile in terminal and sometimes also axillary glomerules [Fig. 534] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. glomerata 5b. Flowers pedicellate, not arranged in compact clusters 6a. Calyx lobes linear; stem leaf blades linear to narrow-lanceolate 7a. Basal leaf blades obovate; capsules erect, dehiscing by lateral or subapical pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. patula 7b. Basal leaf blades broad-ovate to cordate-orbicular; capsules nodding, dehiscing by basal pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rotundifolia 6b. Calyx lobes lanceolate to triangular or oblong; stem leaf blades lanceolate to ovate, cordate-ovate, or triangular-ovate (narrower in C. persicifolia) 8a. Stem leaf blades linear to narrow-lanceolate . . . . . . . . . . . . . . C. persicifolia 8b. Stem leaf blades lanceolate to ovate, cordate-ovate, or triangular-ovate

Fig. 533  Flowers and upper stem of Campanula aparinoides.

9a. Corolla broadly rotate-campanulate; capsules dehiscing by subapical pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. carpatica 9b. Corolla campanulate, funnelform-campanulate, or tubular-campanulate; capsules dehiscing by basal pores 10a. Calyx lobes spreading to reflexed during anthesis [Fig. 536]; inflorescence an elongate, secund raceme [Fig. 536] . . . . C. rapunculoides 10b. Calyx lobes erect during anthesis [Fig. 535]; inflorescence of solitary, axillary flowers or of non-secund racemes of 1–4 flowers at the apex of the stem and branches [Fig. 535] 11a. Stem sharply angled; lower leaf blades evidently cordate at the base; inflorescenc a loose raceme of 1–4 flowers at the apex of the stem and branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. trachelium

Fig. 534  Inflorescence of Campanula glomerata.

11b. Stem bluntly angled; lower leaf blades rounded to cuneate at the base; inflorescence of solitary flowers in the axils of leaves, collectively appearing to be a terminal, leafy raceme [Fig. 535] . . . . . . . . C. latifolia

Ca m pa nu l ac e a e   49 5

1. Campanula aparinoides Pursh N Fig. 533 marsh bellflower. Campanula aparinoides Pursh var. grandiflora Holz.; C. aparinoides Pursh var. uliginosa (Rydb.) Gleason; C. uliginosa Rydb. • CT, MA, ME, NH, RI, VT. Marshes, shorelines, stream banks, wet meadows. 2. Campanula carpatica Jacq. E tussock bellflower. CT. Fields, roadsides, gardens. 3. Campanula divaricata Michx. E southern bellflower. Campanula flexuosa Michx. • CT, NH. Fields, roadsides, railroads, gardens. 4. Campanula glomerata L. E Fig. 534 clustered bellflower. MA, ME, NH, VT. Fields, roadsides, gardens. 5. Campanula lanata Friv. E woolly bellflower. MA. Fields, roadsides, gardens. 6. Campanula latifolia L. E Fig. 535 giant bellflower. ME, VT; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, waste areas.

Fig. 535  Flowers of Campanula latifolia.

7. Campanula patula L. E spreading bellflower. CT, MA, NH. Fields, roadsides, dumps, gardens. 8. Campanula persicifolia L. E peach-leaved bellflower. Campanula persicifolia L. var. alba Horton • CT, MA, VT. Yards, gardens, roadsides. 9. Campanula punctata Lam. E spotted bellflower. CT, NH. Gardens, waste areas, roadsides. 10. Campanula rapunculoides L. E Fig. 536 creeping bellflower. CT, MA, ME, NH, RI, VT. Roadsides, waste areas, gardens. 11. Campanula rotundifolia L. N Scotch bellflower. CT, MA, ME, NH, VT. Cliffs, ledges, river shore outcrops, dry fields. 12. Campanula trachelium L. E nettle-leaved bellflower. MA, VT; also reported from ME by Campbell et al. (1995), but specimens are unknown. Fields, roadsides, dumps.

Fig. 536  Flowers of Campanula rapunculoides.

Gadellia 1. Gadellia lactiflora (Bieb.) Schulkina E Fig. 537 milky-bellflower. Campanula lactiflora Bieb. • NH. Gardens, roadsides, waste areas.

Jasione 1. Jasione montana L. E Fig. 538 sheepbit. CT, MA, RI. Roadsides, sandy banks and fields.

Fig. 537  Inflorescence of Gadellia lactiflora.

496  tricolpate s

Lobelia Reference: McVaugh (1936). 1a. Flowers 20–45 mm long; corolla fenestrate 2a. Corolla red (rarely pink or white); flowers 30–45 mm long; filament tube 24–33 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. cardinalis 2b. Corolla blue (rarely white); flowers 20–30 mm long; filament tube 12–15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. siphilitica 1b. Flowers 7–18 mm long; corolla lacking open slits Fig. 538  Inflorescence of Jasione montana.

3a. Leaves all basal, with 2 hollow tubes extending the entire length; all the anthers pubescent at the apex; plants aquatic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. dortmanna 3b. Leaves borne on a stem, flat, lacking interior hollow tubes; only the 2 smaller anthers pubescent at the apex; plants terrestrial 4a. Stem leaf blades linear to narrow-lanceolate, (0.5–) 1–4 (–5) mm wide; lower lip of the corolla glabrous on the adaxial (i.e., inside) surface . . . . . . . . . . . . . . . . . . . . . . L. kalmii 4b. Stem leaf blades lanceolate or oblanceolate to ovate or obovate, the principal blades 10–35 (–50) mm wide; lower lip of the corolla pubescent on the adaxial surface near the base 5a. Corolla 6–8 mm long; hypanthium nearly as long as the distinct portion of the calyx, evidently inflated in fruit [Fig. 539]; middle stem leaf blades ± ovate-oblong; stems long-pubescent, at least below; plants annual . . . . . . . . . . . . . . . . . . . . . L. inflata 5b. Corolla 7–11 mm long; hypanthium shorter than the distinct portion of the calyx, scarcely inflated in fruit [Fig. 540]; middle stem leaf blades ± oblanceolate; stems short-pubescent; plants biennial or perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . L. spicata 1. Lobelia cardinalis L. N red lobelia. CT, MA, ME, NH, RI, VT. Stream and river shores, lake shores, marshes. 2. Lobelia dortmanna L. N water lobelia. CT, MA, ME, NH, RI, VT. Shallow water of lakes and ponds. Usually occurring as a submerged rosette of leaves with a stem that elevates the flowers above the water level.

Fig. 539  Flower of Lobelia inflata.

3. Lobelia inflata L. N Fig. 539 bladder-pod lobelia. CT, MA, ME, NH, RI, VT. Fields, roadsides, forest clearings, trail edges, ledges, shorelines. 4. Lobelia kalmii L. N brook lobelia. Lobelia kalmii L. var. strictiflora Rydb.; L. strictiflora (Rydb.) Lunell • CT, MA, ME, NH, VT; restricted to the western portion of the southern New England states. Fens, stream shores, seepy river shore outcrops, and disturbed soil in regions of high-pH bedrock. 5. Lobelia siphilitica L. var. siphilitica n great lobelia. CT, MA, ME, NH, VT. Marshes, shorelines, wet meadows, open swamps. Considered introduced in eastern MA, but native to Berkshire County. 6. Lobelia spicata Lam. N Fig. 540 pale-spiked lobelia.  6a. Lobelia hirtella (Gray) Greene; 6c. Lobelia bracteata Small; L. spicata Lam. var. originalis McVaugh • CT, MA, ME, NH, RI, VT. Fields, openings.

Fig. 540  Flower of Lobelia spicata.

1a. Stems, bracts, and calyx lobes bristly pubescent with abundant, short, stiff hairs; leaves often confined to lower half of stem . . . . . . . . . . . . . . . . . . . . . . 6a. L. spicata var. hirtella Gray 1b. Stems, bracts, and calyx lobes glabrous or pubescent with sparser and/or softer hairs; leaves usually distributed throughout stem

C a m pa nu l ac e a e   497

2a. Anthers white; calyx terete in cross-section during anthesis; corolla dark purple-blue; racemes with 10–30 flowers; capsules globose . 6b. L. spicata var. campanulata McVaugh 2b. Anthers blue; calyx compressed during anthesis; corolla light blue; racemes with 30–60 (–100) flowers (infrequently fewer); capsules short-hemispherical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6c. L. spicata var. spicata Variety hirtella is known from ME, NH, VT. Variety campanulata is known from CT, MA, ME, NH; also reported from RI by Kartesz (1999), but specimens are unknown. Variety spicata is known from CT, MA, ME, NH, RI, VT.

Triodanis The genus Triodanis (along with many other genera) is nested within Campanula as it is currently circumscribed (Roquet et al. 2008). Further work is needed to determine if Campanula should be dismantled (in which case Triodanis will stand as a genus) or maintained as a large, inclusive genus (in which case Triodanis and many other genera will need to be subsumed under Campanula). 1. Triodanis perfoliata (L.) Nieuwl. N Fig. 541 clasping-leaved Venus’-looking-glass. Legousia perfoliata (L.) Britt.; Specularia perfoliata (L.) A. DC. • CT, MA, ME, NH, RI, VT. Deciduous forests, cliff bases, rocky slopes, disturbed sites, lawns, sometimes weedy. The lower flowers of this species are cleistogamous and have a 3- or 4-merous calyx and show reduced corollas and shorter capsules compared with the chasmogamous flowers.

Cannabaceae Reference: Mitchell (1988). 1a. Plants tall shrubs or small trees; leaf blades simple, unlobed; fruit a drupe . . . . . . . . . Celtis 1b. Plants herbs or vines; leaf blades palmately lobed or palmately divided; fruit an achene 2a. Leaf blades palmately lobed; stems vining, retrorsely spinulose; petioles with forked hairs; fruits concealed by large, accrescent bracteoles, collectively the infructescence appearing as “hops” [Fig. 542] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Humulus 2b. Leaf blades palmately divided; stems erect, not spinulose; petioles without forked hairs; fruits subtended, but not concealed, by accrescent bracteoles, the infructescence not appearing as “hops” . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cannabis

Cannabis 1. Cannabis sativa L. E hemp. Cannabis sativa L. ssp. indica (Lam.) E. Small & Cronq. • CT, MA, ME, NH, RI, VT. Roadsides, barn yards, waste areas, frequently planted in forest clearings.

Celtis 1. Celtis occidentalis L. N common hackberry. Celtis occidentalis L. var. canina (Raf.) Sarg.; C. occidentalis L. var. pumila (Pursh) Gray • CT, MA, NH, RI, VT. Forests, riparian terraces, regenerating fields, commonly on high-pH soils.

Fig. 541  Inflorescence of Triodanis perfoliata.

498  tricolpates

Humulus Reference: Small (1978). 1a. Plants annual; larger leaf blades commonly 5- to 9-lobed, with very narrow sinuses, without glandular dots; carpellate spikes with spinulose-ciliate bracts that are narrow and do not completely conceal the fruits; anthers without glands . . . . . . . . . . . . . . . . . . . . . H. japonicus 1b. Plants perennial; larger leaf blades commonly 3- to 5-lobed, with broad and open sinuses, gland-dotted on the abaxial surface; carpellate spikes with entire-margined bracts that are broad and conceal the fruits [Fig. 542]; anthers glandular . . . . . . . . . . . . . . . . . . . . . . . H. lupulus 1. Humulus japonicus Sieb. & Zucc. E Japanese hop. CT, MA, ME, NH, RI, VT. Roadsides, waste areas. 2. Humulus lupulus L. n Fig. 542 common hop.  2a. Humulus americanus Nutt.; H. lupulus L. var. lupuloides E. Small • CT, MA, ME, NH, RI, VT. Riparian forests and shrub thickets, abandoned homesteads, roadsides,

waste areas. 1a. Abaxial leaf midrib with 20–100 hairs per cm; abaxial leaf surface with more than 25 glands per 10 mm² . . . . . . . . . . . . . . . . . . 2a. H. lupulus ssp. americanus (Nutt.) A. & D. Löve Fig. 542  Carpellate spikes of Humulus lupulus.

1b. Abaxial leaf midrib with fewer than 20 hairs per cm; abaxial leaf surface with fewer than 25 glands per 10 mm² . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. H. lupulus ssp. lupulus Supspecies americanus is known from CT, MA, ME, NH, RI, VT, and is native to New England, usually occurring in riparian habitats. Subspecies lupulus is known from CT, MA, ME, NH, VT and is non-native, often occurring in human-disturbed habitats.

Caprifoliaceae 1a. Plants herbaceous or suffrutescent; flowers with 3 or 4 stamens (with 5 stamens in Triosteum) 2a. Flowers and fruits sessile in the axils of leaves [Fig. 551]; stamens 5 per flower; fruit an orange-red or orange-yellow drupe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triosteum 2b. Flowers and fruits borne on a peduncle [Figs. 544, 545, 546]; stamens 3 or 4 per flower; fruit an achene or cypsela (sometimes with sterile locules and therefore the fruits asymmetrical) 3a. Flowers borne in a capitulum-like cyme [Figs. 543, 544, 549]; corolla 4-lobed (5-lobed in Scabiosa); ovary and fruit partially enclosed in a gamophyllous, tubular involucel that may be expanded apically 4a. Stem, and often the involucral bracts and abaxial leaf midrib, with prickles [Fig. 543]; receptacular bracts awn-tipped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dipsacus 4b. Stem, bracts, and leaves without prickles; receptacular scales absent or present and pointed at the apex, but without awn-tips 5a. Calyx-teeth or -setae (6–) 8–16 (–24); receptacle pubescent, without scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Knautia 5b. Calyx-teeth or -setae 4 or 5 or absent altogether; receptacle glabrous, with scales 6a. Involucral bracts ± ovate, rigid, in more than 3 series; marginal flowers of cyme enlarged and falsely radiate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephalaria

Ca p r i fo l i ac e a e   49 9

6b. Involucral bracts narrow-lanceolate to lanceolate, herbaceous, in 1–2 (–3) series; marginal flowers of cyme enlarged or not 7a. Corolla 5-lobed; marginal flowers of cyme enlarged and falsely radiate; stem leaves once or twice pinnately lobed with deep sinuses (i.e., appearing pinnately compound) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scabiosa 7b. Corolla 4-lobed; marginal flowers of cyme not notably enlarged [Fig. 549]; stem leaves simple 8a. Calyx-setae 4 or 5; involucel 4-angled and pubescent . . . . . . . Succisa 8b. Calyx-setae absent; involucel 8-angled and glabrous . . . . . Succisella 3b. Flowers borne separately in an open inflorescence [Figs. 545, 552]; corolla 5-lobed; ovary and fruit not enclosed in an involucel 9a. Stems trailing [Fig. 545]; corolla 10–15 mm long; androecium with 4 stamens; flowers nodding, paired [Fig. 545] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Linnaea 9b. Stems upright; corolla 1.5–7 mm long; androecium with 3 stamens; flowers neither nodding nor paired [Fig. 545] 10a. Stem leaves pinnately divided [Fig. 552]; calyx composed of 5–20 narrow segments, these inconspicuous at anthesis but unrolling and forming a plumose, pappus-like crown in fruit; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . Valeriana 10b. Stem leaves simple; calyx obsolete or minute; plants annual . . . Valerianella 1b. Plants definitely woody; flowers with 5 stamens (with 4 stamens in Kolkwitzia) 11a. Stamens 4 per flower; pedicels and ovary densely bristly pubescent; fruit a bristly achene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kolkwitzia 11b. Stamens 5 per flower; pedicels and ovary glabrous or pubescent, but not bristly pubescent; fruit a capsule, berry, or drupe 12a. Leaf blades regularly serrate; fruit a capsule 13a. Corolla 12–20 mm long, zygomorphic, yellow, turning red in age; capsules glabrous; seeds not winged; petioles glabrous to sparsely pubescent . . . . . . Diervilla 13b. Corolla 25–30 mm long, nearly actinomorphic, dark red when open (sometimes white or pink); capsules pubescent; seeds winged; petioles ± densely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Weigela 12b. Leaf blades entire or with a few, irregular, tooth-like lobes; fruit fleshy 14a. Corolla 10–50 mm long; ovary with 2 or 3 locules, each locule bearing several ovules; fruit red, blue, or black; leaf blades strictly entire; leaf scars with 3 vascular bundles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lonicera 14b. Corolla 2–8 mm long; ovary 4-locular, 2 of the locules with several abortive ovules, the other 2 locules with a solitary, fertile ovule; fruit white (red in one rare introduction); leaves entire or with irregular, tooth-like lobes; leaf scars with 1 vascular bundle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symphoricarpos

Cephalaria 1. Cephalaria gigantea (Ledeb.) Bobrov E yellow giant-scabious. Scabiosa gigantea Ledeb. • MA. Waste areas, railroads.

5 00   tricolpate s

Diervilla 1. Diervilla lonicera P. Mill. N bush-honeysuckle. Diervilla diervilla (L.) MacM.; D. lonicera P. Mill. var. hypomalaca Fern. • CT, MA, ME, NH, RI, VT. Woodlands, forest borders, sand plains, roadsides, talus slopes, open rights-of-way, subalpine meadows.

Dipsacus 1a. Stem leaf blades crenate-serrate to entire; involucral bracts linear . . . . . . . . . . . D. fullonum 1b. Stem leaf blades irregularly laciniate-pinnatifid; involucral bracts narrow-subulate [Fig. 543] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. laciniatus Fig. 543  Inflorescence and upper peduncle of Dipsacus laciniatus.

1. Dipsacus fullonum L. E fuller’s teasel. Dipsacus fullonum L. ssp. sylvestris (Huds.) Clapham; D. sylvestris Huds. • CT, MA, nh, RI, VT. Fields, pastures, roadsides, dumps, waste areas.

2. Dipsacus laciniatus L. E Fig. 543 cut-leaved teasel. MA, VT. Fields, roadsides, waste areas.

Knautia 1. Knautia arvensis (L.) Coult. E Fig. 544 bluebuttons. Scabiosa arvensis L. • CT, MA, ME, NH, VT. Fields, roadsides, lawns, waste areas.

Kolkwitzia 1. Kolkwitzia amabilis Graebn. E beautybush. MA, VT. Forest borders, roadsides, areas of habitation. Fig. 544  Inflorescence of Knautia arvensis.

Linnaea 1. Linnaea borealis L. ssp. americana (Forbes) Hultén ex Clausen N Fig. 545 American twinflower. Linnaea americana Forbes; L. borealis L. var. americana (Forbes) Rehd. • CT, MA, ME, NH, RI, VT; becoming rare in southern New England. Mesic to wet-mesic, evergreen and mixed evergreen-deciduous forests, ascending to boreal situations.

Lonicera 1a. Flowers paired, borne on axillary peduncles [Figs. 546, 548]; plants upright (twining or climbing in L. japonica); leaves all distinct 2a. Plants twining or climbing; leaves persistent; corolla 30–50 mm long; fruit black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. japonica 2b. Plants upright; leaves deciduous; corolla 10–22 mm long; fruit red or blue Fig. 545  Habit of Linnaea borealis.

3a. Style glabrous; corolla ± actinomorphic [Fig. 546] 4a. Ovaries definitely united; fruit blue; corolla gibbous at the base on one side; bractlets subtending the flowers mostly 4–6 mm long; winter buds with 2 scales; plants mostly of hydric soils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. villosa 4b. Ovaries separate [Fig. 546]; fruit red; corolla with a short spur at the base; bractlets subtending the flowers mostly 1.2–3 mm long; winter buds with more than 2 scales; plants mostly of mesic soils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. canadensis

Ca p r i fo l i ac e a e   5 0 1

3b. Style pubescent; corolla weakly to strongly zygomorphic [Fig. 548] 5a. Branchlets with solid, white pith; bractlets subtending the flowers obsolete; plants native, of circumneutral fens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. oblongifolia 5b. Branchlets with hollow, darker pith; bractlets subtending the flowers 2–8 mm long; plants introduced, of fields, roadsides, and disturbed areas 6a. Peduncles shorter than 5 mm; leaf blades abruptly tapering to an acuminate apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. maackii 6b. Peduncles 5–25 mm long [Fig. 548]; leaf blades obtuse to acute at the apex 7a. Upper lip of the corolla with shallow sinuses, the sinuses extending ca. half the distance to the base of the lip; filaments pubescent; bractlets and ovary glandular; winter buds fusiform, the scales conspicuously ciliate . . . L. xylosteum 7b. Upper lip of the corolla with deep sinuses, the sinuses extending nearly to the lip’s base [Fig. 548]; filaments glabrous or pubescent only near base; bractlets and ovary eglandular; winter buds conical or ovoid, ciliate or not 8a. Leaf blades pubescent on the abaxial surface; peduncles 5–15 mm long; corolla white (turning yellow), pubescent on the abaxial (i.e., outside) surface, saccate at the base; branchlets pubescent [Fig. 548] . . . . . . . . . . . . L. morrowi 8b. Leaf blades glabrous on the abaxial surface; peduncles 15–25 mm long; corolla usually pink to red (fading yellow), glabrous on the abaxial surface, not saccate at the base; branchlets glabrous . . . . . . . . . . . . . . . . . . . L. tatarica 1b. Flowers in 3-flowered cymules, sessile in the axils of leaves [Fig. 547]; plants twining or climbing; upper 1 or 2 pairs of leaves gamophyllous (all distinct in L. periclymenum) 9a. Leaf blades pubescent on abaxial surface, green on adaxial surface . . . . . . . . L. hirsuta 9b. Leaf blades glabrous on abaxial surface, thinly to conspicuously glaucous on adaxial surface 10a. All the leaves distinct, with petioles; basal connate tube of the corolla glandularpubescent; distal end of the branchlets usually glandular-pubescent . . . L. periclymenum 10b. Upper 1–4 pairs of leaves gamophyllous and lacking petioles [Fig. 547]; basal connate tube of the corolla eglandular; branchlets usually glabrous 11a. Corolla 30–50 mm long, weakly zygomorphic, the 5 lobes nearly equal; seeds 4–4.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sempervirens 11b. Corolla 15–30 mm long, strongly zygomorphic, with an upper and lower lip; seeds either 3–3.5 mm long or 4.5–5 mm long 12a. Uppermost gamophyllous leaf blades green adaxially, bluntly pointed to pointed at the apex, collectively forming a rhombic-elliptic to double-ovate disk [Fig. 547]; seeds 3–3.5 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. dioica

Fig. 546  Flowers of Lonicera canadensis.

12b. Uppermost gamophyllous leaf blades glaucous adaxially, rounded to retuse at the apex, collectively forming a suborbicular to orbicular disk; seeds 4.5–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. reticulata 1. Lonicera canadensis Bartr. ex Marsh. N Fig. 546 American honeysuckle. Xylosteum ciliatum Pursh • CT, MA, ME, NH, VT. Mesic to wet-mesic forests and forest borders. 2. Lonicera dioica L. N Fig. 547 wild honeysuckle. Lonicera dioica L. var. glaucescens (Rydb.) Butters; L. glaucescens (Rydb.) Rydb. • CT, MA, ME, NH, RI, VT. Forests, forest borders, woodlands, sometimes associated with rocky and/or ledgy areas. Forests and forest borders, rocky woodlands, thickets, roadsides.

Fig. 547  Inflorescence and gamophyllous leaves of Lonicera dioica.

5 02  tricolpates

3. Lonicera hirsuta Eat. hairy honeysuckle. Lonicera hirsuta Eat. var. interior Gleason • MA, VT. Rocky forests and woodlands, ledges, usually in high-pH bedrock regions.

NC

4. Lonicera japonica Thunb. E Japanese honeysuckle. Nintooa japonica (Thunb.) Sweet • CT, MA, ME, NH, RI. Forests, forest fragments, forest edges, roadsides, clearings. 5. Lonicera maackii (Rupr.) Herder E Amur honeysuckle. Ct, mA, VT. Forest borders, roadsides, areas of habitation. 6. Lonicera morrowii Gray E Fig. 548 Morrow’s honeysuckle. CT, MA, ME, NH, RI, VT. Upland and riparian forests, edges of swamps, field edges, fence rows, roadsides, areas of habitation. 6 ‌ × 11. Lonicera ×bella Zabel is a frequent honeysuckle hybrid known from CT, MA, ME, NH, RI, VT. It is variable but usually displays a pink (fading yellow) corolla that is scarcely saccate at the base, sparsely pubescent branchlets and leaf blades, and peduncles 5–15 mm long (compare character states with those presented in identification key, couplet 8).

Fig. 548  Flowers of Lonicera morrowii.

7. Lonicera oblongifolia (Goldie) Hook. N swamp honeysuckle. Lonicera oblongifolia (Goldie) Hook. var. altissima (Jennings) Rehd. • ME, VT. Fens, evergreen swamps dominated by Thuja occidentalis. 8. Lonicera periclymenum L. E European honeysuckle. ME. Fence rows, abandoned homesteads, yards. 9. Lonicera reticulata Raf. E grape honeysuckle. Lonicera prolifera (Kirchn.) Booth ex Rehd.; L. prolifera (Kirchn.) Booth ex Rehd. var. glabra Gleason; L. sullivantii Gray • MA. Forest borders, fence rows, areas of habitation. 10. Lonicera sempervirens L. var. sempervirens nC trumpet honeysuckle. Phenianthus sempervirens (L.) Raf. • CT, MA, ME, NH, RI, VT. Forest borders, woodlands, fence rows, roadsides, ledges, open rights-of-way. Most collections of this plant are clearly introduced (intentionally or unintentionally) and many that away from homes and farms are located on tracts of land surrounded by urban and suburban development. Some collections from CT, MA, and RI (a minority of them) do appear to be growing as native plants and probably should be conserved as such (though it is possible that all populations in New England are introduced). 11. Lonicera tatarica L. E Tatarian honeysuckle. CT, MA, ME, NH, RI, VT. Riparian forests, field edges, waste areas, roadsides, areas of habitation. Though frequently included on state lists of invasive plants, this species is not nearly as common as Lonicera morrowii (with which it has been confused by some collectors) and is infrequently encountered. 12. Lonicera villosa (Michx.) J.A. Schultes N mountain honeysuckle. Lonicera caerulea L. var. villosa (Michx.) Torr. & Gray; L. villosa (Michx.) J.A. Schultes var. calvescens (Fern. & Wieg.) Fern.; L. villosa (Michx.) J.A. Schultes var. fulleri Fern. L. villosa (Michx.) J.A. Schultes var. solonis (Eat.) Fern.; L. villosa (Michx.) J.A. Schultes var. tonsa Fern. • CT, MA, ME, NH, RI, VT. Mesic to, more commonly, wet-mesic and hydric forests, fens, bogs, wet meadows, low and open rights-of-way, boggy outlets, ascending to open, alpine areas in some ranges (e.g., Presidentials of NH). 13. Lonicera xylosteum L. E fly honeysuckle. CT, MA, ME, NH, RI, VT. Forest borders, roadsides, clearings, areas of habitation.

C a p r i fo l i ac e a e   5 03

Scabiosa Some species of Scabiosa (e.g., S. ochroleuca) are considered to be a complex of intergrading microspecies by European botanists (Jasiewicz 1976). Given that the characters that separate these races are trivial and intermediate morphologies abound, the species of Scabiosa are here treated in the broad sense. Scabiosa atropurpurea L. was reported from MA by Sorrie and Somers (1999) based on the following specimen: 20 Aug 1880, Manning s.n. (NEBC!). It was incorrectly determined. This species has also been reported from ME by Campbell et al. (1995), but specimens are unknown. 1a. Corolla yellow to yellow-white; capitula 15–25 mm in diameter . . . . . . . . . . . . S. ochroleuca 1b. Corolla purple to red-purple or blue-purple; capitula 20–40 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. columbaria 1. Scabiosa columbaria L.

E

dove pincushions. MA. Fields, roadsides, waste areas. 2. Scabiosa ochroleuca L. E cream pincushions. MA. Fields, roadsides, waste areas, borrow pits.

Succisa 1. Succisa pratensis Moench. E Devil’s-bit. Scabiosa succisa L. • MA, RI. Fields, roadsides, river banks, waste areas.

Succisella 1. Succisella inflexa (Kluk) G. Beck E Fig. 549 southern succisella. Scabiosa australis Wulfen; Succisa australis (Wulfen) Reichenb. • CT, MA. Swamps, ditches, meadows.

Fig. 549  Inflorescence of Succisella inflexa.

Symphoricarpos 1a. Fruit purple-red; corolla 2–4 mm long; style villous; pith solid . . . . . . . . . . . . . S. orbiculatus 1b. Fruit white; corolla 5–8 mm long; style glabrous; pith hollow . . . . . . . . . . . . . . . . . . . . S. albus 1. Symphoricarpos albus (L.) Blake n Fig. 550 common snowberry.  1a. Symphoricarpos albus (L.) Blake var. laevigatus (Fern.) Blake;  1b. Symphoricarpos racemosus Michx. • CT, MA, ME, NH, RI, VT. Roadsides, forest borders, abandoned homesteads, forest fragments, areas of habitation, woodlands, rocky forests and woodlands, cliffs, lake shore outcrops. 1a. Branchlets glabrous; leaf blades glabrous abaxially; drupe mostly (10–) 12–15 mm in diameter; cultivated shrubs mostly 1–2 m tall . . . . . 1a. S. albus ssp. laevigatus (Fern.) Hultén 1b. Branchlets usually pubescent [Fig. 550]; leaf blades usually pubescent abaxially; drupe 6–10 (–12) mm in diameter; native shrubs of western New England up to 1 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. S. albus ssp. albus Subspecies laevigatus is known from CT, MA, ME, NH, RI,VT. It is non-native and usually found in areas of human disturbance and habitation. Subspecies albus is known from MA, VT; western portion of states. It is found on thin soils and ledges in regions of high-pH bedrock. 2. Symphoricarpos orbiculatus Moench. E red snowberry. Symphoricarpos symphoricarpos (L.) MacM. • CT, MA, RI, VT. Roadsides, forest borders, yards, areas of habitation.

Fig. 550  Fruits and branchlet with leaves of Symphoricarpos albus ssp. albus.

5 04  tricolpates

Triosteum Reference: Wiegand (1923). 1a. Sepals 9–12 mm long, conspicuously hispid-ciliate, the abaxial surface glabrous or shortpubescent; stem long-pubescent, the longer hairs 1.5–3 mm long; leaf blades 2–6 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. angustifolium 1b. Sepals 10–18 mm long, the margins and abaxial surface pubescent with hairs of uniform length; stem pubescent with hairs up to 1.5 mm long; leaf blades 4–15 cm wide 2a. Leaf blades of the middle nodes broadly connate at base (the lower ones and the upper ones sometimes sessile or petiolate); stem short-pubescent, the eglandular hairs up to 0.5 mm long (a few scattered hairs may range to 1.5 mm); sepals 0.9–2 mm wide (mean=1.4 mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. perfoliatum 2b. Leaf blades not basally connate (sometimes the middle nodes with slightly gamophyllous leaves); stem with longer pubescence, the eglandular hairs 0.5–1.5 mm long; sepals 1.5–2.8 mm wide (mean=2 mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. aurantiacum 1. Triosteum angustifolium L.

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lesser horse-gentian. CT; western portion of state. Rocky forests and woodlands, ledges, railroads. 1‌ × 2. Triosteum ×eamesii (Wieg.) A. Haines is a rare horse-gentian hybrid that is known from CT and was formerly treated as Triosteum angustifolium var. eamesii Wieg. It most resembles T. angustifolium due to its short sepals and long-pubescent stems. It has leaf blades mostly 2.2–3.2 times a long as wide that are moderately puberulent abaxially and dried stems (3–) 4–7 mm thick at the midpoint (vs. leaf blades mostly 3.5–7 times as long as wide that are sparsely pubescent abaxially and dried stems 2–2.7 mm thick at the midpoint in T. angustifolium). Recognition of this hybrid shows T. angustifolium to be far more rare than previously believed because many collections of T. angustifolium were of the var. eamesii (i.e., they were hybrid plants). 2. Triosteum aurantiacum Bickn. var. aurantiacum N Fig. 551 Fig. 551  Fruits of Triosteum aurantiacum.

orange-fruited horse-gentian. Triosteum perfoliatum L. var. aurantiacum (Bickn.) Wieg. • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic forests, woodlands, and forest borders. 3. Triosteum perfoliatum L.

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perfoliate-leaved horse-gentian. CT, MA, RI. Forests, woodlands, fields, pastures, often on dry-mesic, sandy substrate.

Valeriana 1a. Basal leaf blades simple or sometimes with a single pair of basal lobes; stem leaf blades with 3–13 leaflets, the rachis and abaxial surface ± glabrous . . . . . . . . . . . . . . . . . . . . V. uliginosa 1b. Basal leaf blades pinnately divided; stem leaf blades with 11–21 leaflets, the rachis and abaxial surface sparsely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. officinalis 1. Valeriana officinalis L. E Fig. 552 common valerian. CT, MA, ME, NH, VT. Fields, roadsides, open banks, yards, waste areas. 2. Valeriana uliginosa (Torr. & Gray) Rydb. Fig. 552  Inflorescence of Valeriana officinalis.

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marsh valerian. Valeriana sitchensis Bong. ssp. uliginosa (Torr. & Gray) F.G. Mey.; V. sitchensis Bong. var. uliginosa (Torr. & Gray) Boivin • ME, NH, VT. Evergreen swamps dominated by Thuja occidentalis, fen woodlands.

Valerianella Valerianella have unusual, asymmetrical fruits due to the development of only one locule (of three) of the fruit. Many taxa have been named based solely on differences in fruit morphology.

C a p r i fo l i ac e a e   5 0 5

However, Ware (1983) determined that many of these sometimes striking fruit differences are the result of a single gene (i.e., a single species may produce 2 or 3 types of fruits within a given population, each type of fruit specific to a given plant). Therefore, the number of species recognized here is fewer than some previous treatments. Reference: Dyal (1938). 1a. Fertile locule of fruit conspicuously thickened by a corky mass on the abaxial (i.e., back) surface [Fig. 553]; cymules with spatulate outer bractlets and spinulose-ciliate inner bractlets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. locusta 1b. Fertile locule of fruit lacking a corky mass on the abaxial surface [Fig. 554]; cymules with lanceolate or oblanceolate outer bractlets and usually eciliate inner bractlets (these sometimes with cilia near apex or rarely ciliate throughout) 2a. Corolla 3–5 mm long, with lobes 1–2 mm long; outer bracts of cymule usually eciliate (except sometimes near apex) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. umbilicata 2b. Corolla 1.5–2 mm long, with lobes 0.4–0.8 mm long; outer bracts of cymule spinuloseciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. radiata 1. Valerianella locusta (L.) Lat. E Fig. 553 European corn-salad. Valerianella olitoria (L.) Pollich • CT, me; also reported from MA and VT by Hultén and Fries (1986), but specimens are unknown. Fields, pastures, river banks, and other open, often disturbed sites. 2. Valerianella radiata (L.) Dufr.

Fig. 553  Cross-section of fruit of Valerianella locusta showing corky mass on abaxial side of fertile locule.

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beaked corn-salad. Valerianella radiata (L.) Dufr. var. fernaldii Dyal; V. radiata (L.) Dufr. var. missouriensis Dyal; V. stenocarpa (Engelm. ex Gray) Krok var. parviflora Dyal • CT. Open banks and terraces of rivers, often associated with graminoid vegetation. 3. Valerianella umbilicata (Sullivant) Wood N Fig. 554 navel corn-salad. Valerianella intermedia Dyal; V. patellaria (Sullivant ex Gray) Wood; V. radiata (L.) Dufr. var. intermedia (Dyal) Gleason • CT, MA. Hydric fields, low pastures, and other open, often disturbed, sites.

Weigela 1. Weigela floribunda (Sieb. & Zucc.) C.A. Mey. E crimson weigela. Diervilla floribunda Sieb. & Zucc. • CT, MA. Forest borders, old homesteads, ledgy woodlands, areas of habitation.

Caryophyllaceae Reference: Mitchell (1993). 1a. Sepals distinct or essentially so (appearing connate in Scleranthus, but the tube actually a hypanthium) [Figs. 564, 565, 578]; ovary sessile; petals without a prominent, narrow, basal portion (i.e., lacking a claw or the claw obscure) 2a. Leaves with scarious or hyaline stipules [Figs. 561, 577] 3a. Leaves alternate (rarely some subopposite); gynoecium with 3 sessile stigmas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Corrigiola 3b. Leaves opposite or whorled [Fig. 561]; gynoecium with 2–5 stigmas (with 3 stigmas in Polycarpon and Spergularia but these elevated on styles) 4a. Flowers without petals [Fig. 565]; gynoecium with a single ovule; ovary maturing as a utricle

Fig. 554  Cross-section of fruit of Valerianella umbilicata, which lacks a corky mass on abaxial side of fertile locule.

5 06   tricolpate s

5a. Sepals white, spongy, each with a prominent awn . . . . . . . . . . . . . . . . Illecebrum 5b. Sepals ± green (but white-scarious at the margin in Paronychia, the actual color may be partly obscured by white or gray hairs on the abaxial surface), not spongy, with or without awns 6a. Stipules with entire margins, (0.5–) 1–8 mm long; calyx 1–4 mm long, the lobes white-scarious at the margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paronychia 6b. Stipules with fringed margins [Fig. 561], 0.5–1 (–1.5) mm long; calyx 0.5–1.5 mm long, the lobes green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Herniaria 4b. Flowers with white or pink to red-purple petals (but these sometimes caducous in Polycarpon); gynoecium with many ovules; ovary maturing as a capsule 7a. Flowers with a single style that is obscurely lobed near the apex; leaf blades elliptic to ovate or obovate; valves of capsules twisted after dehiscence and forming a tube . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polycarpon 7b. Flowers with 3–5 styles; leaf blades linear to broad-linear; valves of capsules spreading to recurved but not twisted into a tube 8a. Leaves clustered at the nodes in 2 sets of 6–8, appearing whorled, with small stipules 0.5–1 (–1.5) mm long; flowers usually with 5 styles; capsules with usually 5 valves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spergula 8b. Leaves opposite (with axillary fascicles in S. rubra), with stipules 1–5 mm long; flowers usually with 3 styles; capsules with usually 3 valves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spergularia 2b. Leaves without stipules 9a. Flowers perigynous (i.e., with a cupuliform hypanthium) [Fig. 568]; fruit a 1-seeded utricle; perianth sepaloid, composed of only sepals that have a narrow, scarious border . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scleranthus 9b. Flowers hypogynous or essentially so; fruit a many-seeded capsule; perianth in part petaloid, composed of both sepals and petals (petals absent and, therefore, the perianth sepaloid in some Sagina and Stellaria) 10a. Perianth monochlamydeous, only the sepals present 11a. Flowers with 3 styles; capsule dehiscing by 6 valves; leaves linear-oblong to elliptic, 1.5–6 (–10) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Stellaria 11b. Flowers with 4 or 5 styles; capsule dehiscing by 4 or 5 valves; leaves linearsubulate, up to 1 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Sagina 10b. Perianth dichlamydeous, both the sepals and the petals present [Figs. 556, 557] 12a. Petals deeply notched, in some Stellaria so deeply as to appear as 2 distinct, but proximate, petals [Figs. 557, 579] 13a. Gynoecium with 3 styles; capsules dehiscing by 6 valves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Stellaria 13b. Gynoecium with 4 or 5 styles [Fig. 557]; capsules dehiscing by 5 valves or by 8 or 10 apical teeth 14a. Capsules cylindric, dehiscing by 8 or 10 apical teeth; vegetative leaf blades 1–15 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cerastium 14b. Capsules ovoid, dehiscing by 5 valves, each valve notched or shortly bifid at the apex; vegetative leaf blades 10–40 mm wide . . . . . . . . . Myosoton 12b. Petals entire, at most the apex retuse or erose [Figs. 562, 566] 15a. Plants fleshy; seeds 3–5 mm long; petals and stamens inserted on a conspicuous, 10-lobed disk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Honckenya

C a ryo p hyl l ac e a e   5 07

15b. Plants not fleshy; seeds 0.4–1.4 mm long; staminal disk inconspicuous 16a. Flowers with 4 or 5 styles [Fig. 567]; capsules dehiscing by 4 or 5 valves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Sagina 16b. Flowers with 3 styles; capsules dehiscing by 3 or 6 valves 17a. Inflorescence an umbel with 3–15 flowers; petals minutely erose at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holosteum 17b. Inflorescence a cyme or a solitary flower; petals not erose at apex 18a. Seeds 1–1.6 mm long, strophiolate; plants rhizomatous perennials, the stems not tufted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Moehringia 18b. Seeds 0.4–0.8 mm long, not strophiolate; plants annuals or perennials, the stems tufted 19a. Capsules dehiscing by 6 valves; leaf blades lanceolate to ovate, 3–5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arenaria 19b. Capsules dehiscing by 3 valves; leaf blades linear to narrowoblanceolate, 0.3–0.8 mm wide . . . . . . . . . . . . . . . . . . . . . . . . Minuartia 1b. Sepals connate in the basal portion [Figs. 570, 572]; ovary stipitate; petals with a prominent, narrow, basal portion (i.e., possessing a claw) 20a. Flowers with 5 (rarely 4) styles or 1 style in staminate individuals of dioecious Silene; capsules dehiscing by 5 or 10 valves or teeth (or dehiscing by 8 valves in 4-styled flowers) 21a. Style conspicuously pubescent; calyx lobes 20–40 mm long, much longer than the connate portion; petals lacking both auricles and appendages at the junction of the claw and blade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agrostemma 21b. Style glabrous; calyx lobes 2.5–9 mm long, much shorter than the connate portion; petals commonly with auricles and/or appendages at the junction of the claw and blade 22a. Plants dioecious (i.e., with unisexual flowers), the staminate flowers with a single, undivided style; capsules dehiscing by 5 valves that eventually split again . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Silene 22b. Plants synoecious (i.e., with bisexual flowers); capsules dehiscing by 5 valves or 5 bifid teeth 23a. Plants annual; calyx 15–28 mm long; carpophores (5–) 7–12 mm long; leaf blades linear to narrow-lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eudianthe 23b. Plants perennial; calyx 6–17 mm long; carpophores 0–6 mm long; leaf blades narrow-lanceolate to narrow-oblong or ovate 24a. Style bases persistent in fruit and reflexed; capsule unilocular, septicidally dehiscent, each valve lacking a distinct midline; petals 2- or 4-lobed, the lobes varying from short and separated by a shallow sinus to elongate and separated by a deep sinus [Fig. 563] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lychnis 24b. Style bases not persistent in fruit; capsule 5-locular in the basal portion, loculicidally dehiscent, each valve with a distinct midline, sometimes further splitting on this line; petals ± entire at the apex . . . . . . . . . . . . . . . . . . . . . Viscaria 20b. Flowers with 2 or 3 styles; capsules dehiscing by 4 or 6 valves 25a. Calyx subtended by 1–3 pairs of scarious or foliaceous bracts [Figs. 558, 559]; seeds disciform 26a. Calyx with 5 principal nerves (with an additional 10 fine nerves in 1 species), prominently scarious between the 5 principal nerves . . . . . . . . . . . . . . . . . Petrorhagia 26b. Calyx 20- to 40-nerved [Fig. 559], without scarious regions between the nerves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dianthus

5 08   tricolpate s

25b. Calyx not subtended by bracts, 5- to 30-nerved; seeds orbicular to reniform 27a. Flowers with 3 styles; capsules dehiscing by 6 valves 28a. Capsules borne on a carpophore 7–8 mm long [Fig. 555]; pedicels 1–5 mm long; stem and calyx glarous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atocion 28b. Capsules borne on a carpophore as long as 6 mm (5–8 mm long in S. caroliniana); pedicels (2–) 5–50 mm long; commonly either the stem and/or the calyx pubescent, but both glabrous in a few species . . . . . . . . . . (in part) Silene 27b. Flowers with usually 2 styles; capsules dehiscing by 4 valves 29a. Adaxial surface of petals with a subulate appendage at the junction of the claw and blade; calyx with 20, often inconspicuous, nerves . . . . . . . . . . . Saponaria 29b. Petals without an appendage; calyx with 5 nerves 30a. Petals 18–22 mm long; seeds globose; calyx nerved with stout ribs, these becoming wing angles in fruit, not at all scarious . . . . . . . . . . . . . . . . . . . Vaccaria 30b. Petals 2–15 mm long; seeds subreniform; calyx merely nerved, without prominent wing angles in fruit, evidently scarious between its green nerves [Fig. 560] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gypsophila

Agrostemma 1. Agrostemma githago L. var. githago E common corncockle. Lychnis githago (L.) Scop. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, areas of habitation and cultivation.

Arenaria 1. Arenaria serpyllifolia L. E thyme-leaved sandwort. 1a. Arenaria leptoclados (Reichenb.) Guss.; A. serpyllifolia L. var. tenuior Mert. & Koch • CT, MA, ME, NH, RI, VT. Fields, roadsides, dry lawns, waste areas, gardens, railroads, thin soil of high-pH ledges. 1a. Fruiting calyx 2–3 mm long; capsules slender-ovoid to subcylindric (i.e., not or scarcely swollen at the base), with fragile valves; seeds 0.4–0.5 mm long; leaf blades lanceolate to narrow-ovate . . . . . . . . . . . . . . . . . . 1a. A. serpyllifolia ssp. leptoclados (Reichenb.) Nyman 1b. Fruiting calyx 3–4 mm long; capsules ovoid to ovoid-conic (i.e., swollen at the base), with firm, resistant valves; seeds 0.5–0.6 mm long; leaf blades ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. A. serpyllifolia ssp. serpyllifolia Subspecies leptoclados is known from MA, ME, VT. Subspecies serpyllifolia is known from CT, MA, ME, NH, RI, VT.

Atocion 1. Atocion armeria (L.) Raf. E Fig. 555 sweet-William-catchfly. Silene armeria L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, gardens.

Cerastium Fig. 555  Flower of Atocion armeria showing carpophore above pedicel.

Cerastium velutinum was reported from New England by Morton (2005a) based on a collection from Franklin County, MA—29 May 1892, Churchill s.n. (MO!). However, the specimen, though vegetatively robust, has anthers, sepals, and petals too short for C. velutinum. The specimen is our native C. strictum. Reference: Morton (2005a).

C a ryo p hy l l ac e a e  5 0 9

1a. Plants perennial, with sprawling stems that root at the nodes or with rhizomes and axillary shoots that produce overwintering leaves 2a. Stems and leaf blades densely white-tomentose, the surfaces ± concealed by tomentum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. tomentosum 2b. Stems and leaf blades pubescent with villous hairs (rarely the stems subglabrous), the surfaces visible 3a. Petals 5–7 mm long, ± equal in length to the sepals; axillary shoots usually lacking; eglandular or occasionally with stipitate glands confined to the inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. fontanum 3b. Petals 7.5–15 mm long, ca. 2 times as long as the sepals [Fig. 557]; axillary clusters of leaves or short shoots produced on lower stem; plants stipitate-glandular in the inflorescence and often also on the upper portion of the stem 4a. Petals 7.5–9 (–9.5) mm long; sepals 3.5–6 (–7) mm long; anthers 0.8–0.9 (–1.1) mm long; reproductive stems 5–20 (–30) cm tall; plants growing in pristine as well as human-disturbed habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. strictum 4b. Petals 10–15 mm long; sepals 5–8 mm long; anthers 0.9–1.2 (–1.5) mm long; reproductive stems (15–) 25–35 (–40) cm tall; plants mainly from human-disturbed habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. arvense 1b. Plants annual, usually from taproots, lacking stolons, rhizomes, and axillary shoots 5a. Flowers usually 4-merous throughout (very rarely 5-merous); capsules usually dehiscing by 8 apical teeth (by 10 teeth in 5-merous flowers) . . . . . . . . . . . . . . . . C. diffusum 5b. Flowers 5-merous, with 5 or 10 stamens; capsules dehiscing by 10 apical teeth 6a. Sepals and often the bracts apically with long, forward-pointing, white, eglandular hairs that potrude beyond the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. glomeratum 6b. Sepals and bracts lacking elongate, eglandular hairs that protrude beyond the apex 7a. Bracts of inflorescence herbaceous throughout, lacking a scarious margin; leaf blades (10–) 15–50 (–60) mm long; androecium with usually 10 stamens (often only 5 in apetalous forms) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nutans 7b. Bracts of inflorescence with a translucent, scarious, distal margin and tip; leaf blades 5–18 mm long; androecium with usually 5 stamens 8a. Veins of petals unbranched; petals shorter than the sepals, only slightly notched, the apical sinus up to 0.5 (–1) mm deep [Fig. 556]; bracts with a broad scarious margin and apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. semidecandrum 8b. Veins of petals branched; petals equaling or shortly surpassing the sepals, prominently notched, the apical sinus 1–1.5 mm deep; bracts with a narrow scarious margin and apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pumilum 1. Cerastium arvense L. E field chickweed. CT, MA, ME, VT. Fields, roadsides, lawns, cemeteries. Cerastium arvense was introduced from Europe and tends to occur in human-disturbed habitats. Native members of this complex (e.g., C. strictum) commonly (but not always) occur in undisturbed habitats, including high-pH and serpentine communities. 2. Cerastium diffusum Pers. ssp. diffusum E four-stamened chickweed. Cerastium tetrandrum W. Curtis • MA. Fields, roadsides, waste areas. 3. Cerastium fontanum Baumg. ssp. vulgare (Hartman) Greuter & Burdet E mouse-ear chickweed. Cerastium vulgatum, sensu L. 1762 • CT, MA, ME, NH, RI, VT. Fields, lawns, roadsides, yards, ledges, waste areas.

5 10  tricolpates

4. Cerastium glomeratum Thuill. E sticky chickweed. Cerastium glomeratum Thuill. var. apetalum (Dumort.) Fenzl; C. viscosum, auct. non L. • MA, RI; also reported from CT by Morton (2005a), but specimens are unknown. Roadsides, waste areas. 5. Cerastium nutans Raf. ssp. nutans

NC

nodding chickweed. Cerastium longepedunculatum Muhl. ex Britt. • CT, MA, VT; primarily western New England. Rocky woodlands and forests, balds, cliffs, streams, sandy banks, borrow pits. 6. Cerastium pumilum W. Curtis ssp. pumilum E European chickweed. CT, MA. Fields, roadsides, waste areas. 7. Cerastium semidecandrum L. E Fig. 556 Fig. 556  Flower of Cerastium semidecandrum.

five-stamened chickweed. CT, MA, RI. Fields, roadsides, gardens, disturbed soil. 8. Cerastium strictum L. N Fig. 557 American field chickweed. Cerastium angustatum Greene; C. arvense L. var. angustifolium Fenzl; C. arvense L. var. latifolium Fenzl; C. arvense L. ssp. strictum (L.) Ugborogho; C. campestre Greene; C. pubescens Goldie; C. vestitum Greene • CT, MA, ME, NH, RI, VT. Cliffs, talus slopes, quarries, fields, rocky beaches, coastal headlands and points. This species most closely resembles Cerastium arvense and with it forms a difficult complex with nearly worldwide distribution. Cerastium velutinum Raf., one of the elements of this complex native to North America, was recognized as distinct from C. arvense by Morton (2005a) on the grounds of morphological and cytological differences. The same arguments can be used for the recognition of C. strictum (rather than recognizing it as a variety or subspecies of C. arvense). In fact, to be consistent with the use of data, both C. strictum and C. velutinum need to be recognized at the same rank.

Fig. 557  Flower of Cerastium strictum.

9. Cerastium tomentosum L. E tomentose chickweed. MA, ME, VT; also reported as “status undetermined” for RI by Gould et al. (1998), but specimens are unknown. Fields, waste areas, gardens. The very similar Cerastium biebersteinii DC. was reported from New England (e.g., Kartesz 1999). Herbarium surveys performed by Morton (2005a) did not locate vouchers of this species in North America (marked by its flat capsule teeth and absence of rhizomes and stolons, as opposed to revolute margined capsule teeth and possession of rhizomes and stolons in C. tomentosum).

Corrigiola 1. Corrigiola litoralis L. ssp. litoralis E strapwort. MA. Gardens, disturbed soil.

Dianthus 1a. Plants with scattered, solitary flowers borne on slender pedicels 10–40 mm long [Fig. 559]; bracts subtending the flowers up to ½ as long as calyx [Fig. 559] 2a. Basal leaf blades oblanceolate, 15–30 mm long; at least the lower internodes of the stem puberulent; blade of petals 5–10 mm long, the apex toothed [Fig. 559]; calyx equaling the length of the fruit; bracts ca. 50% as long as the calyx . . . . . . . . . . . . . . . . . . D. deltoides 2b. Basal leaf blades linear, 20–80 mm long; stems glabrous; blade of petals 12–18 mm long, fringed-cleft to near the middle; calyx shorter than the length of the fruit; bracts 25–36% as long as the calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. plumarius 1b. Plants with closely crowded, sessile or short-pedicellate flowers borne in terminal cymes [Fig. 558]; bracts subtending flowers nearly equaling to exceeding the length of the calyx [Fig. 558]

Ca ryo phy l l ac e a e   5 1 1

3a. Calyx, including the basal connate portion, pubescent [Fig. 558]; plants annual or biennial; stems usually strigose below the nodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. armeria 3b. Calyx completely glabrous or only the lobes pubescent [Fig. 558]; plants perennial; stems glabrous (sometimes scabrous) 4a. Stem leaf blades lanceolate to oblanceolate, (10–) 15–25 (–30) mm wide, sheathing the stem for less than 3 times the stem diameter; calyx lobes glabrous . . . . D. barbatus 4b. Stem leaf blades linear, up to 5 mm wide, sheathing the stem more than 3 times the stem diameter; calyx lobes pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. carthusianorum 1. Dianthus armeria L. E Fig. 558 Deptford pink. CT, MA, ME, NH, RI, VT. Fields, roadsides, and other sandy or gravelly, open areas. 2. Dianthus barbatus L. E sweet William pink. CT, MA, ME, NH, RI, VT. Fields, roadsides, yards, waste areas. 3. Dianthus carthusianorum L. E cluster-headed pink. Dianthus atrorubens All. • NH. Fields, roadsides, gardens. 4. Dianthus deltoides L. E Fig. 559

Fig. 558  Inflorescence of Dianthus armeria showing crowded inflorescence and pubescent calyx.

maiden pink. CT, MA, ME, NH, VT. Fields, roadsides, waste areas, ballast. 5. Dianthus plumarius L. E garden pink. CT, MA, ME, NH, VT. Fields, roadsides, and other dry, often grassy, open areas.

Eudianthe 1. Eudianthe coeli-rosa (L.) Reichenb. E rose catchfly. Agrostemma coeli-rosa L.; Lychnis coeli-rosa (L.) Desr.; Silene coeli-rosa (L.) Godr. • MA. Fields, roadsides, gardens.

Gypsophila Reference: Pringle (2005). 1a. Pedicels and sepals glandular-puberulent; leaf blades clasping at the base; apical portion of stem glandular-puberulent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. scorzonerifolia 1b. Pedicels and sepals not glandular-puberulent [Fig. 560]; leaf blades not clasping (basal leaves clasping in G. elegans); apical portion of stems glabrous (sometimes the stems pubescent with eglandular or glandular hairs, but then the hairs usually confined to the basal half) 2a. Slender annuals with green stems 0.4–3 (–4) dm tall; leaf blades linear, 0.2–2 (–3) mm wide; capsule oblong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. muralis 2b. Stout annuals or perennials with glaucous stems 2–10 dm tall; leaves lanceolate, (1–) 2–16 mm wide; capsule spherical 3a. Plants perennial; petals 1–4 mm long; calyx 1–3 mm long, slightly shorter to nearly as long as the petals; longer pedicels 0.5–12 mm long . . . . . . . . . . . . . . . . . . . . G. paniculata 3b. Plants usually annual; petals 6–15 mm long; calyx 2.5–5 mm long, ca. 36–50% as long as the petals; longer pedicels 10–35 mm long . . . . . . . . . . . . . . . . . . . . . . . . G. elegans 1. Gypsophila elegans Bieb. E showy baby’s-breath. CT, MA, ME, NH, RI. Fields, roadsides, waste areas, gardens. Reports of this species from VT (e.g., Seymour 1982, Kartesz 1999) were based on collections from cultivated material (i.e., this species is not known to be naturalized in VT; specimens at VT).

Fig. 559  Inflorescence of Dianthus deltoides showing open inflorescence and glabrous calyx.

5 12   tricolpate s

2. Gypsophila muralis L. E Fig. 560 low baby’s-breath. Psammophiliella muralis (L.) Ikonnikov • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 3. Gypsophila paniculata L. E tall baby’s-breath. CT, MA, ME, NH, VT. Fields, roadsides, waste areas. 4. Gypsophila scorzonerifolia Ser. E glandular baby’s-breath. CT, MA. Fields, roadsides, waste areas, edges of saline marshes.

Herniaria Fig. 560  Flower of Gypsophila muralis.

1a. Sepals glabrous or short-ciliate; leaf blades glabrous . . . . . . . . . . . . . . . . . . . . . . . . . H. glabra 1b. Sepals densely pubescent; leaf blades pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. hirsuta 1. Herniaria glabra L. E Fig. 561 smooth rupturewort. ME. Roadsides, waste areas. 2. Herniaria hirsuta L. ssp. cinerea (DC.) Coutinho E hairy rupturewort. Herniaria cinerea DC. • MA. Roadsides, gardens, waste areas.

Holosteum 1. Holosteum umbellatum L. ssp. umbellatum E jagged-chickweed. CT, MA, RI, VT. Fields, gardens, nurseries, waste areas. Holosteum umbellatum has an extremely brief life cycle and senesces by May (or soon afterwards). Therefore, the plant may be more common than collections indicate. Fig. 561  Leaves of Herniaria glabra showing stipules with fringed margins.

Honckenya 1. Honckenya peploides (L.) Ehrh. ssp. robusta (Fern.) Hultén N Fig. 562 seaside-sandwort. Arenaria peploides L. var. robusta Fern. Honckenya peploides (L.) Ehrh. var. robusta (Fern.) House • CT, MA, ME, NH, RI; becoming rare in southern New England. Atlantic coast beaches, commonly on sand and gravel substrate.

Illecebrum 1. Illecebrum verticillatum L. E coral necklace. Paronychia verticillata (L.) Lam. • MA. Gardens, nursery lots, waste areas.

Fig. 562  Flower and upper leaves of Honckenya peploides.

Lychnis 1a. Plants densely tomentose; calyx lobes twisted; inflorescence with mostly 3–7 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. coronaria 1b. Plants glabrous to pubescent but not tomentose; calyx lobes planar, not twisted; inflorescence with mostly 5–26 flowers 2a. Stems pubescent with long hairs, some hairs longer than 1.5 mm, with mostly 10–20 pairs of lanceolate to ovate leaf blades 20–60 mm wide; petals with 2 apical lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. chalcedonica 2b. Stems glabrous or pubescent with short hairs, the hairs shorter than 1 mm, with mostly 2–5 pairs of linear to lanceolate leaf blades 4–15 (–30) mm wide; petals cleft into 4 lobes [Fig. 563] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. flos‑cuculi

C a ryo phyl l ac e a e   5 13

1. Lychnis chalcedonica L. E scarlet lychnis. Silene chalcedonica (L.) Krause in Sturm • CT, MA, ME, NH, VT; also reported from RI by George (1997), but specimens are unknown. Fields, roadsides, gardens, areas of habitation. 2. Lychnis coronaria (L.) Murr. E mullein lychnis. Agrostemma coronaria L.; Silene coronaria (L.) Clairville • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 3. Lychnis flos-cuculi L. ssp. flos-cuculi E Fig. 563 ragged robin lychnis. Coronaria flos-cuculi (L.) A. Braun; Silene flos-cuculi (L.) Clairville • CT, MA, ME, NH, RI, VT. Low fields and pastures, ditches, riverside meadows.

Minuartia References: Weaver (1970), Rabeler et al. (2005). Fig. 563  Flower of Lychnis floscuculi.

1a. Sepals nerveless or obscurely nerved 2a. Leaf blades rigid, triangular in cross-section; stem usually glandular-pubescent above; sepals ovate to broad-ovate; petals entire at apex . . . . . . . . . . . . . . . . . . . . . . . M. caroliniana 2b. Leaf blades soft, flat; stem glabrous; sepals oblong to oblong-ovate; petals slightly retuse at apex 3a. Plants perennial, forming small mats, with often dense, sterile, leafy shoots at the base of the plant, found usually in alpine settings at or above 1000 m elevation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. groenlandica 3b. Plants annual, not forming mats, lacking sterile, leafy shoots at the base of the plant, found usually on acidic ridges, balds, and outcrops well below 1000 m elevation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. glabra 1b. Sepals with 3 or 5 strong, longitudinal ribs 4a. Sepals herbaceous, rounded at apex [Fig. 564]; lower portion of branches densely clothed by coriaceous, marcescent leaves; capsules 6–10 mm long, with valves that are emarginate at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. marcescens 4b. Sepals rigid, acute at apex; leaves deciduous or marcescent, but not densely clothing the lower portion of branches; capsules 3–5 mm long, with valves that are entire at apex 5a. Tufts of leaves produced in the axils of the principal leaves; leaf blades with the midvein more prominent than the lateral veins; petals 5–8 mm long, longer than the sepals; seeds 0.8–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. michauxii 5b. Axillary leaves not produced; leaf blades with 3 ± equally prominent veins; petals 2–4.2 mm long, usually shorter than the sepals; seeds 0.4–0.7 mm long . . . . . M. rubella 1. Minuartia caroliniana (Walt.) Mattf.

N C

pine barren sandplant. Arenaria caroliniana Walt.; Minuopsis caroliniana (Walt.) W.A. Weber; Sabulina caroliniana (Walt.) Small • RI. Sandy flats and back dunes near the Atlantic shore. Residential development and, possibly, a hurricane (Rick Enser, personal communication), have claimed the only known mainland population of this species in New England. The Block Island population is also known historically, not known to have been seen since the late 1800s. 2. Minuartia glabra (Michx.) Mattf.

NC

Appalachian sandplant. Arenaria glabra Michx.; A. groenlandica (Retz.) Spreng. var. glabra (Michx.) Fern.; Minuartia groenlandica (Retz.) Ostenf. ssp. glabra (Michx.) A. & D. Löve; Porsildia groenlandica (Retz.) A. & D. Löve ssp. glabra (Michx.) A. & D. Löve; Sabulina glabra (Michx.) Small • CT, ME, NH, RI. Ledges, balds, rock slabs, usually on relatively low summits and ridges, occasionally near sea level on sandy gravel and slabs, typically on granite or soils derived from

5 14   tricolpate s

granite. Minuartia glabra and M. groenlandica appear to be distinct taxa in the southeastern United States (Weaver 1970). However, critical examination of herbarium specimens reveals a different situation in New England. The vast majority of morphological characters used to separate these taxa by previous authors (e.g., plant height, length of leaves, pedicels, sepals, and petals, flower number, anther color) show significant overlap and do not discriminate taxa. The only reliable characters appear to be habit and presence/absence of dense, basal growth (perhaps not enough differentiation to warrant species status). Plants of M. groenlandica are generally found in high elevation, alpine habitats, whereas M. glabra is found on low summits and even bedrock slabs near the coast. However, M. groenlandica can be found on the low, coastal hills of Mt. Desert Island, me, blurring the ecological distinction. 3. Minuartia groenlandica (Retz.) Ostenf.

NC

mountain sandplant. Arenaria groenlandica (Retz.) Spreng.; Porsildia groenlandica (Retz.) A. & D. Löve; Sabulina groenlandica (Retz.) Small • ME, NH, VT. High, open, alpine summits on sandy gravel and in the cracks of rock, infrequently on low lying coastal hills. Rare apetalous forms have been collected on Mount Desert Island, Hancock County, ME. 4. Minuartia marcescens (Fern.) House

N C Fig. 564

serpentine sandplant. Arenaria laricifolia (L.) B.L. Robins. var. marcescens (Fern.) Boivin; A. marcescens Fern. • VT. Serpentine cliffs; northern portion of state. Known from only one site in New England at ca. 970 meters in Orleans County, VT. 5. Minuartia michauxii (Fenzl) Farw. var. michauxii N Fig. 564  Flower of Minuartia marcescens showing sepals that are rounded at the apex.

Michaux’s sandplant. Arenaria michauxii (Fenzl) Hook. f.; A. stricta Michx.; Minuopsis michauxii (Fenzl) W.A. Weber; Sabulina stricta (Michx.) Small • CT, MA, ME, NH, RI, VT. Dry ledges, frequently on moderately high-pH substrate (e.g., limestone, trap rock), infrequently on disturbed soil such as railroads and other dry, gravelly areas. 6. Minuartia rubella (Wahlenb.) Hiern

NC

boreal sandplant. Arenaria rubella (Wahlenb.) Sm.; A. verna L. var. rubella (Wahlenb.) S. Wats.; Tryphane rubella (Wahlenb.) Reichenb. • ME, VT. Cliffs, slides, and gullies in boreal and subalpine settings, associated with high-pH bedrock.

Moehringia 1a. Sepals 3–6 mm long, acute to acuminate at the apex; leaf blades glabrous, acute to acuminate (rarely obtuse) at the apex; seeds 1.3–1.6 mm long . . . . . . . . . . . . . . . M. macrophylla 1b. Sepals 2–3 mm long, obtuse to subacute at the apex; leaf blades ciliate, obtuse to subacute at the apex; seeds 1–1.3 (–1.4) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. lateriflora 1. Moehringia lateriflora (L.) Fenzl N blunt-leaved grove-sandwort. Arenaria lateriflora L. • CT, MA, ME, NH, RI, VT. Forests, woodlands, river terraces, borders of fields, shorelines. 2. Moehringia macrophylla (Hook.) Fenzl

NC

large-leaved grove-sandwort. Arenaria macrophylla Hook. • CT, MA, VT. On cliffs, talus, and thin soil over ledges and balds, usually associated with serpentine bedrock, but infrequently on limestone and trap rock.

Myosoton 1. Myosoton aquaticum (L.) Moench E giant-chickweed. Alsine aquatica (L.) Britt.; Cerastium aquaticum L.; Stellaria aquatica (L.) Scop. • CT, MA, NH, VT; also reported from RI by George (1997), but specimens are unknown. River banks, riparian forests.

C a ryo p hy l l ac e a e   5 15

Paronychia Reference: Core (1941). 1a. Stipules 2.5–8 mm long; calyx 3.5–5 mm long, each lobe tipped by a prominent awn 0.9–2 mm long; flowers concealed by conspicuous white bracts; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. argyrocoma 1b. Stipules 0.5–3 mm long; calyx 1–3 mm long, each lobe with a tiny apiculus or short awn up to 0.3 mm long; flowers not concealed by bracts; plants annual 2a. Stems glabrous; calyx 1–1.5 mm long, the lobes terminated by an inconspicuous apiculus; mature fruit noticeably exceeding the calyx . . . . . . . . . . . . . . . . . . . . . P. canadensis 2b. Stems puberulent; calyx 2–3 mm long, the lobes terminated by a minute awn; mature fruit not or only shortly exceeding the calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. fastigiata 1. Paronychia argyrocoma (Michx.) Nutt.

NC

silvery whitlow-wort. Paronychia argyrocoma (Michx.) Nutt. ssp. albimontana (Fern.) Maguire; P. argyrocoma (Michx.) Nutt. var. albimontana Fern. • MA, ME, NH, VT. Thin, sandy or gravelly soils of ledges, balds, and slides, often on low lying hills and ridges of low-pH bedrock, but ascending to boreal and (rarely) subalpine settings, rarely along sand and gravel river shores (e.g., Saco River) and river shore ledges (e.g., Merrimac River). The VT record is based on a specimen collected by Ridler s.n. (NEBC!). However, the specimen lacks any additional information (e.g., date, township, location) and is believed to represent a label mix-up by some authors. 2. Paronychia canadensis (L.) Wood N smooth forked whitlow-wort. Anychia canadensis (L.) B.S.P.; A. dichotoma Michx.; Paronychia dichotoma (Michx.) A. Nels. • CT, MA, NH, RI, VT; becoming rare in northern New England. Woodlands, rocky forests, balds, trail edges. 3. Paronychia fastigiata (Raf.) Fern.

n C Fig. 565

hairy forked whitlow-wort.  3a. Anychia polygonoides Raf.; 3b. Anychia nuttallii Small • CT, MA. Woodlands, grasslands, roadsides, fields, railroads, dumps. 1a. Sepal awns 0.05–0.15 mm long . . . . . . . . . . . . . . . . . . . . . . . . 3a. P. fastigiata var. fastigiata 1b. Sepal awns 0.2–0.3 mm long [Fig. 565] . . . . . . . 3b. P. fastigiata var. nuttallii (Small) Fern. Variety fastigiata is known from CT, MA. It is of conservation concern. This variety was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated this taxon had questionable occurrence in RI and was unaware of any collections. Variety nuttallii is known from ME and is likely introduced given its high degree of disjunction (the variety is mainly of the mid-Atlantic states).

Petrorhagia 1a. Inflorescence a dense, head-like cyme of (1–) 3–7 flowers at the stem summit; bracts brown-scarious, broad-ovate, ± enclosing the 10–13 mm long calyx . . . . . . . . . . . . . . P. prolifera 1b. Inflorescence a solitary flower (rarely up to 3 flowers) at the tips of branches; bracts of inflorescence herbaceous, narrow-ovate, enclosing up to 50% of the 3–6 mm long calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. saxifraga 1. Petrorhagia prolifera (L.) P.W. Ball & Heywood E proliferous childing-pink. Dianthus prolifer L.; Tunica prolifera (L.) Scop. • MA. Fields, roadsides, waste areas. 2. Petrorhagia saxifraga (L.) Link var. saxifraga E saxifrage childing-pink. Dianthus saxifragus L.; Tunica saxifraga (L.) Scop. • MA, ME, NH. Fields, roadsides, lawns, cliffs, waste areas.

Fig. 565  Flower of Paronychia fastigiata var. nuttallii.

5 16 tricolpates

Polycarpon 1. Polycarpon tetraphyllum (L.) L. ssp. tetraphyllum E four-leaved manyseed. Mollugo tetraphylla L. • MA. Wool waste.

Sagina Reference: Crow (1978). 1a. Petals 3–4.5 mm long, ca. 2 times as long as the sepals [Fig. 566]; upper leaf axils of stem usually bearing fascicles of minute, succulent leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nodosa 1b. Petals 0.75–2.5 (–3) mm long, shorter than to shortly exceeding the sepals [Fig. 567], or the petals absent; upper stem lacking axillary fascicles of leaves 2a. Sepals 4 per flower (rarely some flowers with 5) [Fig. 567], spreading in fruit; plants perennial, with basal rosettes of narrow leaves (at least on younger plants), completely glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. procumbens 2b. Sepals 5 per flower (rarely some flowers with 4), erect in fruit; plants annual (rarely short-lived perennial in S. maxima), usually without basal rosettes of leaves, at least the sepals and apical portion of pedicels stipitate-glandular (infrequently glabrous throughout in S. decumbens) 3a. Leaf blades herbaceous; seeds light brown, obliquely triangular in outline, with an abaxial groove; capsules broad-ellipsoid prior to dehiscence, with relatively thin valves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. decumbens 3b. Leaf blades somewhat succulent; seeds red-brown or dark brown, reniform to globose, plump, lacking an abaxial groove; capsules globose prior to dehiscence, with relatively thick valves 4a. Seeds dark brown, conspicuously tuberculate with magnification; sepals 2–2.5 mm long; petals 1–2 mm long; capsules 2.5–3 mm long . . . . . . . . . . . S. japonica 4b. Seeds red-brown, smooth or inconspicuously tuberculate; sepals (2–) 2.5–3.5 mm long; petals 2–2.5 (–3) mm long; capsules (3–) 3.5–4.5 mm long . . . . . . . . . . S. maxima 1. Sagina decumbens (Ell.) Torr. & Gray ssp. decumbens

NC

trailing pearlwort. Sagina decumbens (Ell.) Torr. & Gray var. smithii (Gray) S. Wats. • CT, MA, VT. Roadsides, trail edges, dry fields. 2. Sagina japonica (Sw.) Ohwi E Japanese pearlwort. CT, MA, RI, VT. Roadsides, waste areas, disturbed soil near buildings. 3. Sagina maxima Gray ssp. maxima E sticky-stemmed pearlwort. Sagina crassicaulis S. Wats. var. littoralis (Hultén) Hultén; S. littoralis Hultén; S. maxima Gray var. littorea (Mackenzie) Hara • MA. Fields, roadsides, waste areas. 4. Sagina nodosa (L.) Fenzl

n C Fig. 566

knotted pearlwort.  4a. Sagina nodosa (L.) Fenzl var. borealis (Crow) Cronq.; 4b. Sagina nodosa (L.) Fenzl var. glandulosa (Bess.) Aschers.; S. nodosa (L.) Fenzl var. pubescens (Bess.) Mert. & Koch; Spergella nodosa (L.) Reichenb.; Spergula nodosa L. • MA, ME, NH. Open, coastal fields, headlands, turfs, and upper beaches, often on thin soils over ledge, in rock crevices, or among stones. 1a. Stems and leaf blades glabrous; pedicels and sepals often glabrous (sometimes stipitateglandular) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a. S. nodosa ssp. borealis Crow Fig. 566  Flowers and upper stem of Sagina nodosa ssp. nodosa.

1b. Stems and usually the leaf blades stipitate-glandular [Fig. 566]; pedicels and sepals stipitate-glandular [Fig. 566] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. S. nodosa ssp. nodosa

C a ryo phy l l ac e a e   5 17

Subspecies borealis is known from ME, NH. It is native and of conservation concern. Subspecies nodosa is known from MA, ME, NH. It is non-native. The two subspecies sometimes co-occur in populations (e.g., Little Moose Island, Hancock County, ME, Isles of Shoals, Rockingham County, NH). 5. Sagina procumbens L. N Fig. 567 bird’s-eye pearlwort. Sagina procumbens L. var. compacta Lange • CT, MA, ME, NH, RI, VT. Mesic, or more commonly, wet-mesic to hydric soils of river banks, ledges, lawns, roadsides, cracks in pavement, and shorelines.

Saponaria 1a. Plants cespitose; stems 1-flowered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pumilio 1b. Plants not cespitose; stems usually many-flowered 2a. Plants upright, 30–90 cm tall; calyx 15–25 mm long; capsule 10–20 mm . . S. officinalis 2b. Plants trailing, procumbent, or ascending, 5–25 cm tall; calyx 7–12 mm long; capsule 6–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. ocymoides

Fig. 567  Flowers and upper stem of Sagina procumbens.

1. Saponaria ocymoides L. E rock soapwort. MA. Fields, roadsides, waste areas. 2. Saponaria officinalis L. E common soapwort. Lychnis saponaria Jessen • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, railroads, waste areas. 3. Saponaria pumilio (L.) Fenz ex A. Braun E pygmy soapwort. Cucubalus pumilio L.; Silene pumilio (L.) Wulfen • NH. Alpine ridges and plateaus. This species was once collected on the Amanoosuc Ravine Trail of Mount Washington, Coos County, NH (Harris 1965). This colony is believed to have been intentionally planted.

Scleranthus 1a. Sepals acute at the apex [Fig. 568], spreading to erect in fruit, separate, with a narrow, white-scarious margin up to 0.1 mm wide [Fig. 568]; floral bracts equaling or exceeding than flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. annuus 1b. Sepals obtuse to rounded at apex, usually introrse in fruit and overlapping, with a broader white-scarious margin 0.3–0.5 mm wide; floral bracts mostly shorter than flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. perennis 1. Scleranthus annuus L. E Fig. 568 annual knawel. CT, MA, ME, NH, RI, VT. Dry, open, sandy areas such as roadsides, yards, and dry fields. 2. Scleranthus perennis L. E perennial knawel. CT, MA. Dry, open, sandy, areas such as roadsides, yards, and dry fields.

Silene The genus Silene has seen numerous revisions to its circumscription. Definitions have varied, from a very inclusive Silene (e.g., Morton 2005b) to a narrowly defined Silene (e.g., Oxelman et al. 2000). The latter is followed here as it was based on morphology and DNA sequence data interpreted in a phylogenetic fashion, and the entities (i.e., genera) they proposed are very distinguishable (Atocion, Eudianthe, Lychnis, Silene, and Viscaria in the New England flora). Reference: Morton (2005b).

Fig. 568  Flowers of Scleranthus annuus.

5 18 tricolpates

1a. Plants dioecious, the carpellate flowers with 5 styles and the staminate flowers with 1 style; capsules dehiscing by 5 valves, each valve usually splitting again 2a. Flowers with white petals, fragrant, opening in the evening; leaf blades oblonglanceolate to elliptic, 6–30 mm wide; carpophore 1–2 mm long; capsules ovoid, the valves erect or slightly recurved after dehiscence; seeds coarsely tuberculate . . . . . . . . S. latifolia 2b. Flowers with red petals, inodorous, opening during the day; leaf blades ovate to elliptic, 10–50 mm wide; carpophore absent; capsules broad-ovoid to subglobose, the valves recurved after dehiscence; seeds densely and uniformly papillose . . . . . . . . . . . . . . . S. dioica 1b. Plants synoecious, the flowers with 3 (rarely 4) styles; capsules dehiscing by 3 valves that eventually split again (rarely 4 valves that split again) 3a. Plants matted, pulvinate, alpine, 3–6 cm tall [Fig. 569]; flowers solitary at the tips of branches; petals purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. acaulis 3b. Plants upright, simple or branched, but not matted, of low-elevation habitats, 8–150 cm tall; flowers solitary in the axils of leaves or in terminal cymes; petals white to pink to red (rarely purple) 4a. Flowers solitary on peduncles from the axils of the upper leaves . . . . . . . . . . S. nivea 4b. Flowers in a terminal cyme 5a. Inflorescence a monochasial cyme, appearing as a secund raceme [Fig. 571] 6a. Axis of the inflorescence commonly forking; petals lobed to below the middle; stems coarse, 20–100 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. dichotoma 6b. Axis of the inflorescence usually simple; petals retuse or lobed nearly to the middle; stems shorter, 15–45 cm tall 7a. Calyx 7–10 mm long in fruit, pubescent with both short, glandular hairs and longer eglandular hairs, the lobes acute at the apex; petals entire to retuse at the apex; flowers erect to ascending; stems erect . . . . . . . . . . . . . . . . . . S. gallica 7b. Calyx 12–18 mm long in fruit, pubescent with only short, glandular hairs, the lobes obtuse at the apex; petals lobed nearly to the middle; flowers divergent or nodding; stems procumbent to ascending . . . . . . . . . . . . . . . . . . . . . S. pendula 5b. Inflorescence a dichasial cyme (note: the cyme sometimes panicle-like) [Fig. 573] 8a. Calyx glabrous (or sometimes minutely pubescent near the base) 9a. Stems glutinous below the nodes; calyx fitting rather tightly over the capsule, 10-nerved; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. antirrhina 9b. Stems not glutinous; calyx inflated, fitting loosely over the body of the capsule, 20-nerved; plants biennial or perennial (rarely annual in S. csereii) 10a. Calyx with 20 obscure veins of equal length connected by many, smaller, anastomosing veins, 12–18 mm long in fruit; filaments pale for much of their length, sometimes purple near the apex; seeds ± black, 1–1.5 mm long, finely tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. vulgaris 10b. Calyx with 10 long veins and 10 short veins, with few or no anastomosing veins, 9–13 mm long in fruit; filaments dark purple; seeds 0.6–1 mm long, gray-brown, with concentric rings of papillae . . . . . S. csereii 8b. Calyx pubescent, especially on the nerves [Fig. 572] 11a. Stem leaves mostly in whorls of 4 (the lower and upper nodes may have opposite leaves); petals fimbriately lobed into 8–12 apical segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. stellata 11b. Stem leaves opposite throughout; petals entire, retuse, or notched at the apex

C a ryo phyl l ac e a e   5 19

12a. Petals entire or merely retuse at the apex; calyx lobes rounded to obtuse at apex; plants 8–20 (–25) cm tall; seeds 1.3–1.5 mm long, papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. caroliniana 12b. Petals with a prominent notch, creating a 2-lobed apex; calyx lobes acute to attenuate at apex; plants (15–) 20–80 cm tall; seeds 0.6–1 mm long, tuberculate 13a. Calyx with ca. 30 longitudinal nerves [Fig. 570], umbilicate at the base in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. conica

Fig. 569  Habit of Silene acaulis.

13b. Calyx with 10 nerves, not umbilicate at base 14a. Calyx 15–24 mm long in flower, becoming 25–30 (–40) mm long in fruit; petals pink on the adaxial surface, yellow on the abaxial surface, opening at night, with auricles 1–1.5 mm long; leaf blades lanceolate to narrow-ovate, 20–40 mm wide; plants not clumped, annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. noctiflora 14b. Calyx 7–10 mm long; petals white or pink, open during the day, with inconspicuous auricles; leaf blades narrow-lanceolate to oblanceolate, 5–15 mm wide; plants subcespitose, perennial . . . . . . . . . . . . . . . S. nutans 1. Silene acaulis (L.) Jacq.

N C Fig. 569

moss campion. Silene acaulis (L.) Jacq. var. exscapa (All.) DC.; S. acaulis (L.) Jacq. ssp. arctica A. & D. Löve • ME, NH. Alpine summits and plateaus. 2. Silene antirrhina L. N sleepy campion. Silene antirrhina L. forma apetala Farw.; S. antirrhina L. var. confines Fern.; S. antirrhina L. forma deameana Fern.; S. antirrhina L. var. divaricata B.L. Robins.; S. antirrhina L. var. laevigata Engelm. & Gray • CT, MA, ME, NH, RI, VT. Fields, roadsides, railroads, dry ledges, open woodlands.

Fig. 570  Flowers of Silene conica.

3. Silene caroliniana Walt. ssp. pensylvanica (Michx.) Clausen N wild campion. Silene caroliniana Walt. var. pensylvanica (Michx.) Fern.; S. pensylvanica Michx. • CT, MA, NH, RI. Deciduous woodlands, roadsides, grassy banks, clearings. 4. Silene conica L. ssp. conica E Fig. 570 striate campion. Pleconax conica (L.) Sourkova • MA. Dry and/or sandy fields, roadsides, waste areas. 5. Silene csereii Baumg. E Balkan campion. MA, ME, NH, VT. Fields, roadsides, waste areas. 6. Silene dichotoma Ehrh. E forked campion. CT, MA, ME, NH, VT. Fields, roadsides, waste areas. 7. Silene dioica (L.) Clairville E

Fig. 571  Inflorescence of Silene gallica.

red campion. Lychnis dioica L.; Melandrium dioicum (L.) Coss. & Germ.; M. dioicum (L.) Coss. & Germ. ssp. rubrum (Wieg.) D. Löve • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. ‌7 × 9. Silene ×hampeana Meusel & K. Werener is an infrequent hybrid between two dioecious species known from CT, MA, ME, NH, VT. It usually has pale pink petals. 8. Silene gallica L. E Fig. 571 windmill campion. Silene anglica L. • CT, MA, ME, NH. Fields, roadsides, waste areas. 9. Silene latifolia Poir. ssp. alba (P. Mill.) Greuter & Burdet E Fig. 572 white campion. Lychnis alba P. Mill.; Melandrium album (P. Mill.) Garcke; Silene alba (P. Mill.) Krause; S. pratensis (Raf.) Godr. & Gren. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, areas of cultivation. Fig. 572  Flower of Silene latifolia.

5 20   tricolpate s

10. Silene nivea (Nutt.) Muhl. ex Otth E snowy campion. ME, VT. River banks, riparian forests, field edges. Growing as if native in some locations (e.g., Sandy River, Franklin County, ME) but highly disjunct from its main range. 11. Silene noctiflora L. E night-flowering campion. Melandrium noctiflorum (L.) Fries • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, areas of cultivation. 12. Silene nutans L. E Eurasian campion. ME; also reported from VT by Kartesz (1999), but specimens are unknown. Fields, roadsides, gardens. 13. Silene pendula L. E nodding campion. ME. Fields, roadsides, waste areas. 14. Silene stellata (L.) Ait. f.

NC

starry campion. Cucubalis stellatus L.; Silene stellata (L.) Ait. f. var. scabrella Palmer & Steyermark • CT, RI, VT; also reported from MA by Fernald (1950b), but specimens are unknown. Deciduous woodlands and rocky forests, river banks, roadsides. 15. Silene vulgaris (Moench) Garcke ssp. vulgaris E Fig. 573 bladder campion. Behen vulgaris Moench; Silene cucubalus Wibel; S. inflata Sm. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads, open shorelines.

Spergula Reference: Hartman and Rabeler (2005a). Fig. 573  Inflorescence of Silene vulgaris.

1a. Seeds subglobose, sometimes with an equatorial keel or wing up to 0.1 mm wide; flowers with usually 10 stamens; leaf blades revolute, appearing terete in cross-section at low magnification, 15–30 (–50) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. arvensis 1b. Seeds compressed, with a proment equatorial wing 0.2–0.6 mm wide; flowers with 5 or 10 stamens; leaf blades flat or weakly revolute, 3–15 (–20) mm long 2a. Petals lanceolate, ± acute at the apex, generally not contiguous in intact flowers; flowers with usually 5 stamens; equatorial wing of seed white to light brown, (0.3–) 0.4–0.6 mm wide, approximately as wide as the seed body . . . . . . . . . . . S. pentandra 2b. Petals ovate, narrowly rounded to obtuse at the apex, generally contiguous in intact flowers; flowers with usually 10 stamens; equatorial wing of seed light brown to brownblack (rarely white), 0.2–0.3 (–0.5) mm wide, narrower than the seed body [Fig. 574] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. morisonii 1. Spergula arvensis L. E corn spurry. Spergula arvensis L. var. sativa (Boenn.) Reichenb.; S. sativum Boenn. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, areas of cultivation. 2. Spergula morisonii Boreau E Fig. 570 Morison’s spurry. MA. Fields, roadsides, waste areas.

Fig. 574  Seed of Spergula morisonii.

3. Spergula pentandra L. E five-stamened spurry. CT. Fields, roadsides, waste areas.

Spergularia Rossbach (1940) ambiguously reported Spergularia diandra (Guss.) Boiss. from MA based on the following collection—Cushman 8512 (NEBC!). The same specimen bears a 1937 annotation by Rossbach indicating the collection may be S. purpurea (Pers.) D. Don (i.e., the annotation bears

Ca ryo p hy l l ac e a e   5 2 1

a question mark indicating uncertainty in the determination). Examination of the specimen revealed it is likely neither of these species. It was originally determined as S. atheniensis Heldr. & Sart. ex Nyman (a taxonomic synonym of S. bocconii (Scheele) Asch. & Graeb.). It is very close to that taxon but has very dark seeds. Unfortunately, the specimen’s identity remains a question. Spergularia media (L.) C. Presl has been reported from New England by several sources; however, the specimen (at NEBC!) is Stellaria media and has been determined as such since 1898. References: Rossbach (1940), Hartman and Rabeler (2005b). 1a. Seeds 0.4–0.6 mm long, without an equatorial wing; flowers with 6–10 stamens [Fig. 577]; plants with evident fascicles of leaves in the axils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rubra 1b. Seeds 0.6–1.4 mm long, with or without an equatorial wing; flowers with (1–) 2–4 (–5) stamens; plants without fascicles of leaves in the axils or these sparse and poorly developed 2a. Leaf blades obtuse to acute at the apex, but without a minute mucro; seeds 0.8–1.4 mm long, shiny, smooth or with irregular reticulate thickenings on the faces, usually with a ± white, erose, equatorial wing 0.2–0.3 mm wide [Fig. 575]; stipules 1–2.8 mm long; pedicels and sepals usually glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. canadensis 2b. Leaf blades minutely mucronate at the apex; seeds 0.6–0.8 mm long, dull, smooth or minutely glandular-papillose on the faces, usually unwinged [Fig. 576]; stipules 2–4 mm long; pedicels and sepals usually stipitate-glandular . . . . . . . . . . . . . . . . . . . . . . . . . . S. marina

Fig. 575  Seed of Spergularia canadensis.

1. Spergularia canadensis (Pers.) G. Don ssp. canadensis N Fig. 575 Canada sand-spurry. Arenaria canadensis Pers.; Tissa canadensis (Pers.) Britt. • CT, MA, ME, NH, RI; coastal counties. Saline to brackish marshes, coastal beaches and strands. 2. Spergularia marina (L.) Griseb. n Fig. 576 saltmarsh sand-spurry. Spergularia leiosperma (Kindb.) F. Schmidt; S. marina (L.) Griseb. var. leiosperma (Kindb.) Guerke; S. salina J. & K. Presl; Tissa marina (L.) Britt. • CT, MA, ME, NH, RI, VT; primarily coastal counties. Saline and brackish marshes, coastal beaches and strands, salted roadsides and edges of parking lots. This species has been collected from some interior counties (e.g., Berkshire County, MA, Washington County, VT), where it should be considered non-native. Spergularia marina has been called S. salina in recent literature, but the epithet marina has priority over salina at the rank of species (published in 1785 vs. 1819). However, application of epithet marina has been inconsistent, and some favor its rejection as an ambiguous name.

Fig. 576  Seed of Spergularia marina.

3. Spergularia rubra (L.) J. & K. Presl E Fig. 577 red sand-spurry. Arenaria rubra L.; Spergularia rubra (L.) J. & K. Presl var. perennans (Kindb.) B.L. Robins.; Tissa rubra (L.) Britt. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, yards, dry lawns.

Stellaria Stellaria longipes Goldie was reported from ME by Kartesz (1999), and was taken up by Magee and Ahles (1999). The record was based on maps presented in Chinnappa and Morton (1976), where dots could be interpreted as being very close to or within the state of ME. However, the authors did not explicitly attribute this species to New England, nor did the second author consider this species part of the New England flora in his later work on the genus (see reference that follows). Therefore, Stellaria longipes should not be considered part of the New England flora. Stellaria palustris (Murr.) Retz. was reported from ME by Campbell et al. (1995), but specimens are unknown. Reference: Morton (2005c). 1a. At least the lower leaves with evident petioles; stems pubescent in 1 or 2 lines [Fig. 578] 2a. Sepals (3.5–) 4.5–5 (–6) mm long; leaf blades 3–25 mm long; capsules shortly exceeding the persistent sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. media

Fig. 577  Flower and upper stem of Spergularia rubra showing axillary fascicles of leaves.

5 22   tricolpate s

2b. Sepals (7.5–) 8–11 mm long; leaf blades 10–100 mm long; capsules shorter than the persistent sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. corei 1b. All the leaves sessile; stems glabrous or minutely scabrous (sometimes apically hispidulous in S. holostea) 3a. Bracts subtending the pedicels herbaceous and green throughout [Fig. 578]; inflorescence either of flowers in the axils of normal foliage leaves or at branches in the stem or the inflorescence a terminal cyme 4a. Sepals 6–8 mm long; petals 8–12 mm long, conspicuously exceeding the sepals (the petals rarely absent); seeds 1.5–2 mm long, coarsely papillose . . . . . . S. holostea 4b. Sepals 2–5 mm long; petals 1–6 mm long, shorter than or barely exceeding the sepals (petals sometimes absent) [Fig. 578]; seeds 0.7–1 mm long, smooth or obscurely marked or roughened 5a. Leaf blades 4–15 × 1–5 mm, fleshy, many of the axils bearing sterile tufts or branchlets; capsules slightly shorter than to equaling the length of the persistent sepals; plants halophytic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. humifusa 5b. Leaf blades 7–60 × 2–8 mm, not fleshy, usually without sterile tufts or branchlets; capsules much longer than the persistent sepals; plants not halophytic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. borealis 3b. Bracts subtending the flowers either wholly scarious or with only a central green strip [Fig. 579]; inflorescence usually a terminal cyme [Fig. 579] 6a. Cymes produced in the axils of leaves; sepals 2.5–3.5 mm long; petals shorter than the sepals (or the petals absent); seeds 0.3–0.7 mm long . . . . . . . . . . . . . S. alsine 6b. Cymes produced at the apex of the stem; sepals 2–7 mm long; petals nearly as long as or longer than the sepals; seeds 0.7–1.2 mm long 7a. Stems smooth; sepals conspicuously 3-veined, 3–7 mm long; leaf blades narrow-lanceolate to elliptic-lanceolate, usually widest below the middle; seeds coarsely rugose-tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. graminea 7b. Stems papillate-scabrous on the angles; sepals obscurely 3-veined, 2–4 mm long; leaf blades linear to oblanceolate or narrow-elliptic, usually widest at or above the middle; seeds obscurely sculpted, nearly smooth . . . . . . . . S. longifolia Fig. 578  Portion of inflorescence of Stellaria borealis showing entirely green bracts.

1. Stellaria alsine Grimm N bog stitchwort. Alsine uliginosa (Murr.) Britt.; Stellaria uliginosa Murr. • CT, MA, ME, NH, VT. Stream sides, springs, swamps, pool edges, wet ditches. 2. Stellaria borealis Bigelow ssp. borealis N Fig. 578 boreal stitchwort. Alsine borealis (Bigelow) Britt.; Stellaria borealis Bigelow var. floribunda Fern.; S. borealis Bigelow var. isophylla Fern.; S. calycantha (Ledeb.) Bong. var. floribunda (Fern.) Fern.; S. calycantha (Ledeb.) Bong. var. isophylla (Fern.) Fern.; S. calycantha (Ledeb.) Bong. var. laurentiana Fern. • CT, MA, ME, NH, ri, VT. Stream sides, springs, low fields, swamps, ascending high into the mountains and occasionally found in gullies and other wet-mesic to hydric areas above treeline. 3. Stellaria corei Shinners E Tennessee stitchwort. Alsine tennesseensis (C. Mohr) Small; Stellaria pubera Michx. var. sylvatica (Beguinot) Weatherby; S. sylvatica (Beguinot) Maguire; S. tennesseensis (C. Mohr) Strausbaugh & Core • CT. Yards, waste areas.

Fig. 579  Portion of inflorescence of Stellaria graminea showing bracts with hyaline margins.

4. Stellaria graminea L. E Fig. 579 grass-leaved stitchwort. Alsine graminea (L.) Britt.; Stellaria graminea (Ledeb.) Bong. var. latifolia Peterm. • CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings.

C a ryo p hyl l ac e a e   5 23

5. Stellaria holostea L. E greater stitchwort. Alsine holostea (L.) Britt. • CT, MA. Woodlands, field edges. Reports of this species in ME (e.g., Kartesz 1999) are based on a collection taken from a cultivated plant— Cape Elizabeth, Lowe s.n. (MAINE!). 6. Stellaria humifusa Rottb.

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saltmarsh stitchwort. Alsine humifusa (Rottb.) Britt. • ME; eastern coastal portion of state. Saline to brackish marshes. 7. Stellaria longifolia Muhl. ex Willd. var. longifolia N long-leaved stitchwort. Alsine longifolia (Muhl. ex Willd.) Britt.; Stellaria longifolia Muhl. ex Willd. var. atrata J.W. Moore; S. longifolia Muhl. ex Willd. var. eciliata (Boivin) Boivin • CT, MA, ME, NH, RI, vT. Low fields and meadows, graminoid marshes, open riverbanks, wetland edges. 8. Stellaria media (L.) Vill. E Fig. 580 common stitchwort. Alsine media L.; Stellaria apetala Ucria ex Roemer; S. media (L.) Vill. var. procera Klatt & Richter • CT, MA, ME, NH, RI, VT. Fields, lawns, gardens, trail edges, about buildings.

Vaccaria 1. Vaccaria hispanica (P. Mill.) Rauschert E cowcockle. Saponaria vaccaria L.; Vaccaria pyramidata Medik.; V. vaccaria (L.) Britt.; V. vulgaris Host • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, river banks, railroads, areas of cultivation.

Viscaria 1. Viscaria vulgaris Bernh. E clammy-campion. Lychnis viscaria L.; Silene viscaria (L.) Jessen • CT, MA, ME, NH. Gardens, lawn edges, areas of habitation.

Celastraceae 1a. Leaves alternate; perianth 5-merous; fruit an orange or orange-yellow capsule; some of the flowers unisexual; plants lianas by means of twining stems with terete branchlets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Celastrus 1b. Leaves opposite [Fig. 582]; perianth 4-merous; fruit a pink or purple capsule; flowers bisexual; plants upright shrubs (liana usually by means of aerial roots in 1 species) often with quadrangular or 4-winged branchlets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euonymus

Celastrus New England genera of this family have, until recently, been appropriately treated as grammatically feminine (their classical gender). However, most authors have usually treated the genera (incorrectly) as masculine, as did Linnaeus. This now appears to be remedied as the names in Celastrus are recommended to be considered masculine (Brummitt 2005), and some authors are applying these results to Euonymus as well. Therefore, specific epithets will generally show an “-us” termination rather than an “-a” termination. 1a. Inflorescence an axillary cyme with usually 2 or 3 flowers; leaf blades broad-ovate to suborbicular, crenate, 1.2–1.7 times as long as wide; pollen white; branchlets with low but prominent and long ridges formed by lines of decurrence extending down from the leaf scars;

Fig. 580  Upper stem of Stellaria media with detail showing stem pubescence arranged in lines.

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expanding leaves weakly conduplicate, the inner leaves progressively enfolded by the outer leaves, the margins not or scarcely involute; capsules with (1–) 4–7 (–8) seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. orbiculatus 1b. Inflorescence a terminal panicle with 6 or more flowers [Fig. 581]; leaf blades elliptic to oblong or ovate, serrate, 1.8–2.6 times as long as wide; pollen yellow; branchlets lacking ridges formed by lines of decurrence or these obscure; expanding leaves neither conduplicate nor obviously enfolding successively inner leaves, the margins initially tightly involute; capsules with 1–4 (–8) seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. scandens 1. Celastrus orbiculatus Thunb. E Asian bittersweet. CT, Ma, Me, RI, VT. Forests and forest edges, roadsides, clearings, open rightsof-way. 1 × 2. This rare bittersweet hybrid is known from MA. It is best identified by its blending of characters (e.g., intermediate capsule and/or anther color) or by its combination of characters (e.g., leaf blades of one species and inflorescences of the other species). The outer and inner valves of the capsule often display an intermediate color between the parental species—yellow to orange-yellow in Celastrus orbiculatus, deep orange in C. scandens, and bright orange in the hybrid. 2. Celastrus scandens L. N Fig. 581 American bittersweet. CT, MA, ME, NH, RI, VT. Forests and forest edges, rocky slopes, river banks.

Euonymus Fig. 581  Terminal inflorescence of Celastrus scandens.

1a. Petioles 1–3 (–5) mm long; capsules divided to near base into 4 almost separate sections (often 1 or more of these sections abortive); branchlets with (2–) 4 conspicuous corky wings [Fig. 582]; leaf blades turning bright red in autumn prior to dropping from plant . . . . . E. alatus 1b. Petioles 5–20 mm long; capsules 4-lobed; branchlets without corky wings, though sometimes with low ridges on the angles (branchlets occasionally with corky wings in E. phellomana); leaf blades either remaining green or turning yellow-green to yellow or dull red-purple in autumn prior to dropping from plant 2a. Plants usually climbing by means of aerial roots (rare forms merely spreading and lacking aerial roots); leaves evergreen; branchlets subterete; capsule pink to nearly white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. fortunei 2b. Plants upright shrubs; leaves deciduous; branchlets 4-angled or 4-winged; capsule purple or red to pink 3a. Petals brown-purple; anthers purple; cymes with usually 7–15 flowers; capsule purple; leaf blades pubescent on the abaxial surface with erect hairs up to ca. 0.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. atropurpureus 3b. Petals green-white; anthers yellow; cymes with usually 2–7 flowers; capsule red to pink; leaf blades glabrous or essentially so on the abaxial surface 4a. Seeds white with an orange aril; cyme usually with 2–5 flowers, borne on a peduncle 20–35 mm long; branchlets lacking prominent corky wings . . . . E. europaeus 4b. Seeds dark brown with a red aril; cyme usually with 7 flowers, borne on a peduncle 5–10 mm long; branchlets sometimes with prominent corky wings [Fig. 582] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. phellomanus 1. Euonymus alatus (Thunb.) Sieb. E Fig. 582

Fig. 582  Internodes of branchlet of Euonymus alatus with corky wings.

winged spindle-tree. Celastrus alatus Thunb. • CT, MA, ME, NH, RI, VT. Roadsides, forest fragments, field edges, areas of habitation. This species has a purple capsule and an orange to orange-red aril surrounding the brown seed. These are additional characters that aid in separation from Euonymus phellomana, another species that can have prominently winged branches.

Ce l a st r ac e a e   5 25

2. Euonymus atropurpureus Jacq. var. atropurpureus E eastern spindle-tree. CT, MA, ME, NH, RI. Roadsides, forest fragments, abandoned homesteads, riparian forests, areas of habitation. This species has brown seeds with a red aril. 3. Euonymus europaeus L. E European spindle-tree. CT, MA, ME, NH, RI, VT. Roadsides, forest fragments, abandoned homesteads, riparian forests, areas of habitation. 4. Euonymus fortunei (Turcz.) Hand.-Mazz E climbing spindle-tree. Euonymus fortunei (Turcz.) Hand.-Mazz var. radicans (Sieb. ex Miq.) Rehd.; E. fortunei (Turcz.) Hand.-Mazz var. vegetus (Rehd.) Rehd.; E. radicans Sieb. ex Miq.; E. radicans Sieb. ex Miq. var. vegetus Rehd. • CT, MA, ME, NH. Roadsides, forest fragments and edges, areas of habitation. The common form of Euonymus fortunei that is found naturalized in New England is a liana that climbs by means of aerial roots and has prominent, lightcolored veins on the adaxial surface (especially the midrib). Non-climbing forms are known from at least MA and have been reported as var. vegetus by Sorrie and Somers (1999), which is a cultivar of this species. 5. Euonymus phellomanus Loes. ex Diels E cork spindle-tree. CT, MA. Roadsides, forest fragments, areas of habitation.

Cercidiphyllaceae Cercidiphyllum 1. Cercidiphyllum japonicum Sieb. & Zucc. ex J. Hoffman & H. Schult. E Katsura-tree. CT, MA. Abandoned homesteads, forest fragments, roadsides.

Cistaceae 1a. Petaliferous flowers with 3 inconspicuous, red petals; petals flat in bud; plants overwintering by a basal rosette of crowded, leafy branches . . . . . . . . . . . . . . . . . . . . . . . Lechea 1b. Petaliferous flowers with 5 conspicuous, yellow petals; petals wrinkled in bud; plants lacking a basal rosette of crowded branches 2a. Leaf blades narrow-oblong to elliptic or elliptic-oblanceolate [Figs. 583, 584], 20–30 (–40) mm long, widest near or above the middle, not imbricate; plants erect to ascending (with horizontal branches in 1 species), not forming mats, with stellate pubescence on the stem; style very short, not over 1 mm long; flowers of 2 types— chasmogamous flowers with large, conspicuous petals and cleistogamous flowers with reduced petals or lacking petals altogether . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crocanthemum 2b. Leaf blades linear-subulate or narrow-ovate to triangular, 1–6 mm long, widest below the middle, densely imbricate; plants diffusely branched, forming low mats and mounds, with simple pubescence on the stem; style ca. 2 mm long; flowers of 1 type—all chasmogamous, with conspicuous petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hudsonia

5 26   tricolpate s

Crocanthemum Our frostweeds have traditionally been treated as part of the genus Helianthemum; however, recent phylogenetic evidence, as well as morphology and distribution, supports separating New World (Crocanthemum) from Old World (Helianthemum) species (Arrington and Kubitzki 2003). Species of Crocanthemum produce two types of flowers—chasmogamous flowers with showy yellow petals and cleistogamous flowers that lack petals. The chasmogamous flowers also have larger sepals and larger capsules that contain more seeds than the cleistogamous ones. Reference: Daoud and Wilbur (1965). 1a. Chasmogamous (i.e., petaliferous) flowers solitary (rarely paired) at the summit of the stem during anthesis, later overtopped by lateral branches [Fig. 584]; capsules of cleistogamous flowers with 5–14 seeds; seeds papillose; pedicels and calyx sometimes with red, glandular hairs (in addition to stellate and simple hairs); adaxial surface of leaf blades, especially the young ones, with both short, stellate hairs and longer simple hairs 0.5–1 mm long 2a. Mature stems with widely divergent branches; cleistogamous capsules 3–4.5 mm wide, with 8–14 seeds; seeds with low, broad papillae; adaxial surface of mid-stem leaf blades typically with dense stellate pubescence, the hairs so abundant as to have overlapping branches that obscure view of the epidermis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. dumosum 2b. Mature stems with ascending branches [Fig. 584]; cleistogamous capsules 2–3 mm wide, with 5–10 seeds; seeds with long, slender papillae; adaxial surface of mid-stem leaf blades typically with moderate stellate pubescence, the hairs usually spaced so that the branches are separate and not overlapping and the epidermis is visible . . . . . . . . . . . . . . C. canadense 1b. Chasmogamous flowers in corymbs of 2–20 flowers at the summit of the stem during anthesis, only sometimes overtopped later in season by lateral branches [Fig. 583]; capsules of cleistogamous flowers with 1 or 2 (–3) seeds; seeds indistinctly reticulate; pedicels and calyx lacking glandular hairs, with only stellate and simple, eglandular hairs; adaxial surface of the leaf blades with only stellate hairs, lacking simple hairs

Fig. 583  Inflorescence of Crocanthemum bicknellii showing corymbose chasmogamous flowers.

3a. Stems scattered along a creeping rhizome; outer sepals (i.e., the narrower ones) of cleistogamous flowers apically distinct for a distance of 0.2–0.5 mm, the distinct tip 1–2 times as long as wide; capsules of cleistogamous flowers somewhat rounded in crosssection; leaf blades narrow-cuneate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. propinquum 3b. Stems ± clustered at the tips of caudex branches; outer sepals of cleistogamous flowers apically distinct for a distance of (0.3–) 0.6–1.2 (–1.8) mm, the distinct tip mostly 3–5 times as long as wide; capsules of cleistogamous flowers sharply 3-angled in cross-section; leaf blades cuneate at base [Fig. 583] . . . . . . . . . . . . . . . . . . . . . . C. bicknellii 1. Crocanthemum bicknellii (Fern.) Barnh. N Fig. 583 hoary frostweed. Crocanthemum majus, sensu Britt.; Helianthemum bicknellii Fern. • CT, MA, ME, NH, RI, VT; largely absent from extreme northern New England. Open, sandy soils of woodlands, roadsides, clearings, dry fields, and sandplains. Flowering ca. 10–15 days later than Crocanthemum canadense when both are present at the same site. 2. Crocanthemum canadense (L.) Britt. N Fig. 584 Canada frostweed. Helianthemum canadense (L.) Michx.; H. canadense (L.) Michx. var. sabulonum Fern. • CT, MA, ME, NH, RI, VT; largely absent from extreme northern New England. Open, sandy soils of woodlands, roadsides, clearings, dry fields, and sandplains. Typically flowering ealier than the other species of Crocanthemum in New England, save for C. dumosum. 3. Crocanthemum dumosum Bickn.

Fig. 584  Inflorescence of Crocanthemum canadense showing solitary chasmogamous flower.

bushy frostweed. Helianthemum dumosum (Bickn.) Fern. • CT, MA, RI; southeastern New England. Open, sandy soils of woodlands, roadsides, clearings, dry fields, and sandplains. Usually the first flowering species of Crocanthemum in New England.

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C i stac e a e   5 27

4. Crocanthemum propinquum (Bickn.) Bickn. N low frostweed. Helianthemum propinquum Bickn. • CT, MA, RI; also reported from NH by Gleason and Cronquist (1991), but specimens are unknown. Open, sandy soils of woodlands, roadsides, clearings, dry fields, and sandplains. Usually flowering after Crocanthemum canadense but prior to C. bicknellii in New England where these species are sympatric.

Hudsonia Reference: Morse (1979). 1a. Leaf blades narrow-ovate to triangular, closely appressed, densely tomentose, 1–3 mm long; ovary and capsule glabrous; pedicels absent or up to 1 (–3) mm long . . . . . H. tomentosa 1b. Leaf blades linear-subulate, erect to spreading, sparsely pubescent, 3–4.5 (–6) mm long; ovary and capsule pubescent; pedicels 5–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . H. ericoides 1. Hudsonia ericoides L. N pine-barren false heather. Hudsonia ericoides L. ssp. andersonii Nickerson & Skog • CT, MA, ME, NH, RI, VT; present mostly in coastal plain counties but also scattered in some interior New England counties (e.g., Carroll County, NH, Rutland County, VT). Woodlands, sandy roadsides, grasslands, sandy river beaches, summit balds. 1 × 2. This rare hybrid is known from MA, ME, NH. It is found typically where the parental taxa are in close association, such as coastal dunes adjacent to upland woodlands and on disturbed roadsides of Cape Cod, MA. It is marked by pedicels 3–6 mm long and broad-lanceolate to narrow-triangular leaf blades that are 2–4 mm long and usually ‌spectabilis by Morse (1979); however, appressed. This hybrid was named Hudsonia × the name was never effectively published. Some of the plants formerly referred to as H. tomentosa var. intermedia are actually this hybrid. 2. Hudsonia tomentosa Nutt. N sand false heather. Hudsonia ericoides L. ssp. intermedia (Peck) Nickerson & Skog; H. intermedia (Peck) Erskine; H. tomentosa Nutt. var. intermedia Peck • CT, MA, ME, NH, RI, VT; present mostly in coastal plain counties, but also scattered in some interior New England counties (e.g., Carroll County, NH, Chittenden County, VT). Dunes, sandplains, sandy river beaches, lake shores. Longpedicelled forms of Hudsonia tomentosa have been referred to by the epithet intermedia. Though these plants can generally be separated from short-pedicelled forms (the common form) in New England, Morse (1979) found many intermediate plants throughout the Midwest, demonstrating the impracticality of recognizing two taxa within H. tomentosa.

Lechea The basal leaf blades (i.e., those on basal shoots) are formed in the late summer when the plants begin to fruit (typically in August), overwinter on the plant, and then senesce when the flowering shoots are produced in the spring and summer. These leaves, which are important for idenfitication, are, unfortunately, absent for part of the growing season when botanical inventories are performed. I have little faith in many of the infraspecific taxa proposed by Hodgdon (1938) as the names appear to represent endpoints in clinal morphological variation (i.e., subtle morphological differences do seem to exist between northern vs. southern or coastal plain vs. inland populations of some Lechea species, but morphological gaps are not apparent). References: Hodgdon (1938), Wilbur and Daoud (1961), Barringer (2004). 1a. Outer 2 sepals as long as or longer than the inner 3 sepals [Fig. 586] 2a. Pubescence of stem spreading; inner (i.e., wider) sepals with a prominent keel, glabrous except along the keel; seeds lustrous, bright brown to clear yellow, with nearly transparent endosperm, therefore, the embryo clearly visible . . . . . . . . . . . . . . . . . . . . . . . . . L. mucronata

5 28   tricolpate s

2b. Pubescence of stem appressed to ascending; inner sepals with a midrib but without a prominent keel, pubescent [Fig. 586]; seeds dull, varying from light to dark brown, opaque and without a visible embryo or somewhat translucent with a visible, but not conspicuous, embryo in L. tenuifolia 3a. Leaf blades mostly less than 9 times as long as wide, those of the basal shoots broad-elliptic to ovate, 2–4 (–6) mm wide; capsules equaling or slightly exceeding the sepals, the summit of the fruit visible . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. minor 3b. Leaf blades mostly more than 10 times as long as wide, those of the basal shoots narrow-linear to linear, up to 0.5–1 (–1.5) mm wide; capsules usually slightly shorter than and enclosed by the sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. tenuifolia 1b. Outer 2 sepals up to 60% as long as the inner 3 sepals [Fig. 585] 4a. Leaf blades pubescent across the abaxial surface, those of the basal shoots thick, dull, 1.5–3.5 mm wide; seeds light brown to brown, the endosperm somewhat translucent and the embryo visible . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. maritima 4b. Leaf blades usually pubescent abaxially on the midrib margins only, sometimes nearly glabrous, those of the basal shoots thinner, bright green, 0.7–2 (–2.5) mm wide; seeds light brown to dark brown, opaque, the embryo not visible 5a. Seeds 1–3 (–4) per capsule, compressed-ovoid, brown to dark brown, without an investing membrane; fruiting calyx obovoid, acutely tapering at the base; stem leaf blades with a hardened, yellow to brown, conical apiculus at the apex . . . . . L. pulchella 5b. Seeds 4–6 per capsule, with 3 faces, the outer face convex (i.e., the seeds shaped like sections of an orange), light brown to brown, irregularly covered by a thin, gray membrane that imparts a reticulate surface to seed; fruiting calyx subglobose, rounded at the base; stem leaf blades merely acute at the apex, without an apiculus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. intermedia 1. Lechea intermedia Leggett ex Britt. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N Fig. 585 round-fruited pinweed.  1b. Lechea juniperina Bickn. • CT, MA, ME, NH, RI, VT. Sandy fields, roadsides, woodlands, dry clearings, gravel lots. Fig. 585  Calyx of Lechea intermedia showing outer (i.e., narrower) sepals shorter than inner sepals.

1a. Inner sepals broad-ovate, obtuse at the apex, with 3 or 5 veins; capsule depressed globose, equal to or slightly shorter than the persistent sepals; basal leaf blades oblong, dark green and often purple tinged . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. L. intermedia var. intermedia 1b. Inner sepals narrow-ovate, acute to subacute at the apex, with 3 veins; capsule globose, equal to or slightly exceeding the persistent sepals; basal leaf blades narrow-elliptic, bright green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. L. intermedia var. juniperina (Bickn.) B.L. Robins. Variety intermedia is known from CT, MA, ME, NH, RI, VT. It has a much wider regional distribution than var. juniperina, being found throughout much of New England. Variety juniperina is known from MA, ME, NH. It is more restricted than the previous variety, found commonly along the coastal plain and inland in ME and NH. 2. Lechea maritima Leggett ex B.S.P. var. maritima N beach pinweed. Lechea minor L. var. maritima (Leggett ex B.S.P.) Gray • CT, MA, ME, NH, RI, VT; most abundant in coastal counties but known from many inland counties in most states (but very limited and disjunct in VT). Atlantic coast beaches and dunes, sandy openings and waste areas. 3. Lechea minor L.

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thyme-leaved pinweed. CT, MA, RI, VT; also reported from NH by Wilbur and Daoud (1961), but specimens are unknown. Woodlands, ledges, roadsides, sandy rights-of-way, coastal plain pond shores, dry fields and clearings. 4. Lechea mucronata Raf. N hairy pinweed. Lechea minor L. var. villosa (Ell.) Boivin; L. villosa Ell. • CT, MA, NH, RI, VT. Fields, roadsides, waste areas, woodlands, clearings.

C i stac e a e   5 2 9

5. Lechea pulchella Raf. N Leggett’s pinweed. Lechea leggettii Britt. & Hollick; L. leggettii Britt. & Hollick var. moniliformis (Bickn.) Hodgdon; L. leggettii Britt. & Hollick var. ramosissima Hodgdon; L. moniliformis Bickn.; L. pulchella Raf. var. moniliformis (Bickn.) Seymour • CT, MA, RI. Woodlands, rocky forests, dry clearings, sandy fields and pond shores. I follow Wilbur (1966) in not recognizing infraspecific variation in Lechea pulchella. Review of the characters used by Hodgdon (1938) to distinguish infraspecific taxa shows them to overlap substantially and likely to be, at least in large part, environmentally controlled. The names L. moniliformis and L. ramosissima were used for plants with remote fruits, but this morphology is not distinct because numerous intermediates can be observed in collections. Barringer (2004) also discusses these infraspecific taxa and the nomenclatural status of the names (all of Hodgdon’s varietal names under L. leggettii are illegitimate). 6. Lechea tenuifolia Michx. N Fig. 586 narrow-leaved pinweed. CT, MA, ME, NH, RI; also reported from VT by Magee and Ahles (1999), but specimens are unknown; primarily southern New England, very rare in the northern states. Woodlands, rocky forests, open ridges, dry clearings, sandy fields.

Cleomaceae 1a. Petals retuse at the apex (sometimes merely truncate); flowers with (7–) 10–27 stamens; fruit elevated on a gynophore 0–2 (–6) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polanisia 1b. Petals entire at the apex; flowers with 6 stamens; fruit elevated on a gynophore 1–80 (–152) mm long 2a. Principal leaves with 5–7 leaflets and a pair of short spines at the base of the petioles; stem viscid-pubescent; petals 20–30 (–45) mm long, all oriented toward one side of the flower; gynophores 45–80 (–152) mm long in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tarenaya 2b. Leaves with 3 leaflets and unarmed petioles; stem glabrous (sparsely villous when young); petals 7–12 mm long, nearly evenly distributed around the flower; gynophores 1–15 (–20) mm long in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Peritoma

Peritoma 1. Peritoma serrulata (Pursh) DC. E stinking bee-plant. Cleome serrulata Pursh; C. serrulata Pursh var. angusta (M.E. Jones) Tidestrom • CT, MA, ME. Fields, roadsides, waste areas.

Polanisia 1. Polanisia dodecandra (L.) DC. n red-whiskered clammyweed. 1a. Polanisia graveolens Raf.; 1b. Polanisia dodecandra (L.) DC. var. trachysperma (Torr. & Gray) Iltis; P. trachysperma Torr. & Gray • CT, MA, NH, VT; also reported from ME by Tucker (2010), but specimens are unknown. Sandy and stony lake shores, dry banks, railroads. 1a. Petals 3.5–6.5 (–8) mm long; longest stamens 4–10 (–14) mm long, scarcely if at all exceeding the petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. P. dodecandra ssp. dodecandra 1b. Petals (7–) 8–13 (–16) mm long; longest stamens (9–) 12–30 mm long, much exceeding the petals . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. P. dodecandra ssp. trachysperma (Torr. & Gray) Iltis Subspecies dodecandra is known from CT, MA, NH, VT. It is native to CT and VT and introduced in MA and NH. Subspecies trachysperma is known from CT, MA. It is non-native to New England.

Fig. 586  Calyx of Lechea tenuifolia showing outer (i.e., narrower) sepals as long as inner sepals.

5 30   tricolpate s

Tarenaya 1. Tarenaya hassleriana (Chod.) Iltis E giant spider-flower. Cleome hassleriana Chod.; C. spinosa, auct. non Jacq. • CT, MA, RI. Fields, roadsides, waste areas.

Clethraceae Clethra 1. Clethra alnifolia L. N coastal sweet-pepperbush. Clethra alnifolia L. var. tomentosa (Lam.) Michx.; Clethra tomentosa Lam. • CT, MA, ME, NH, RI. Swamps, lake shores, rarely in woodlands and ridgetops.

Comandraceae 1a. Flowers borne in axillary cymes, lacking a hypanthium, usually in groups of 2–4, the lateral flowers functionally staminate and deciduous, the central flower bisexual; tepals green-purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geocaulon 1b. Flowers borne in terminal, panicle-like cymes, with a hypanthium, all bisexual; tepals white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Comandra

Comandra 1. Comandra umbellata (L.) Nutt. ssp. umbellata N bastard-toadflax. Comandra richardsiana Fern. • CT, MA, ME, NH, RI, VT. Woodlands, dry fields, balds, clearings, open rights-of-way.

Geocaulon 1. Geocaulon lividum (Richards.) Fern.

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false toadflax. Comandra lividum Richards. • ME, NH, VT. Subalpine heaths, coastal peatlands, mossy openings in boreal and alpine, evergreen forests. Found mainly in northern and/or high elevation areas of New England, Geocaulon lividum occurs also in coastal peatlands of downeast ME.

Convolvulaceae 1a. Plants parasitic, with haustoria, lacking chlorophyll; stems pink-yellow to orange, with scale-like leaves; corolla 1.5–5 mm long [Figs. 589, 590, 591] . . . . . . . . . . . . . . . . . . . . . . . Cuscuta 1b. Plants autotrophic, without haustoria, with chlorophyll; stems green or tinged with red, with foliaceous leaves; corolla 15–80 mm long [Figs. 587, 592]

Co nv o lv ul ac e a e   5 3 1

2a. Stigma 1, capitate unlobed or with 2 or 3 small lobes; ovary and capsule 2- or 3-locular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ipomoea 2b. Stigmas 2, filiform or obloid, unlobed; ovary and capsule 1-, 2-, or 4-locular 3a. Pedicels subtended by 2 foliaceous bracts that are positioned near and partly conceal the calyx [Figs. 587, 588]; corolla 20–70 mm long; stigmas obloid . . . . . . . . . . . . . Calystegia 3b. Pedicels subtended by small bracts that are positioned well below the calyx and do not conceal it; corolla 15–25 mm long; stigmas filiform . . . . . . . . . . . . . . . . . . Convolvulus

Calystegia Calystegia hederacea Wallich has been reported from most states in New England by various authors. However, these reports actually refer to C. pubescens (see discussion under that species). Reference: Brummitt (1980). 1a. Stems erect, only the tip twining; leaf blades broad-cuneate to subcordate at the base; leaves subtending the flowers with petioles less than 50% of the length of the leaf midvein, those of the basal portion of the plant conspicuously reduced; bracts subtending the calyx cuneate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. spithamaea 1b. Stems twining or trailing; leaf blades sagittate to hastate at the base; leaves subtending the flowers with petioles more than 50% of the length of the leaf midvein, those of the basal portion of the plant only slightly reduced; bracts subtending the calyx rounded to cordate at the base 2a. Sinus of leaf blade usually rectangular, the leaf tissue not beginning on the interior of the basal lobes for 2–5 (–10) mm from the petiole attachment [Fig. 588]; bracteoles rounded to obtuse at apex, saccate at the base [Fig. 588], overlapping; peduncles frequently paired at some nodes on the plant, not exceeding the leaves; corolla usually white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. silvatica 2b. Sinus of leaf blade usually V-shaped or U-shaped, the leaf tissue beginning on the interior of the basal lobes 0–2 (–3) mm from the petiole attachment [Fig. 587]; bracteoles long-acute to obtuse at the apex, not or slightly saccate at the base [Fig. 587], not or scarcely overlapping; peduncles usually solitary, often equaling or exceeding the leaves; corolla white or pink 3a. Flowers usually with stamens modified into a second cycle of petals; stems pubescent; petioles of proximal leaves 1–6 cm long . . . . . . . . . . . . . . . . . . . . C. pubescens 3b. Flowers not modified, with a single series of connate petals; stems glabrous or pubescent in ssp. americana; petioles of proximal leaves (1.5–) 5–10 cm long . . . C. sepium 1. Calystegia pubescens Lindl. E hairy false bindweed. Calystegia japonica Choisy; C. pellitus Ledeb. forma anestius Fern.; Convolvulus pubescens (Lindl.) Thellung • CT, MA, ME, NH, VT. Fields, roadsides, waste areas, fence rows. This species, which is introduced in New England and usually shows a double corolla (i.e., flore pleno), has a long history of names being misapplied to it. Fernald (1950b) used the name Convolvulus pellitus Ledeb., but that is a separate Asian species and is not the one introduced in New England. Gleason and Cronquist (1991) used the name Calystegia hederacea Wallich inclusively, but that is also a separate species with smaller floral parts and “single” corollas. Calystegia pubescens usually shows a pink corolla, but white forms are also known. 2. Calystegia sepium (L.) R. Br. n Fig. 587 hedge false bindweed.  2a. Calystegia sepium (L.) R. Br. var. americana (Sims) Matsuda; Convolvulus americanus (Sims) Greene; C. sepium L. var. americanus Sims; 2b. Convolvulus sepium L.; 2c. Calystegia sepium (L. ) R. Br. var. angulata (Brummitt) N. Holmgren; C. sepium (L.) R. Br. var. repens (L.) Gray; Convolvulus repens L.; C. sepium L. var. repens (L.) Gray • CT, MA, ME, NH, RI, VT. Fields, roadsides, fence rows, shorelines, margins of tidal marshes, forest clearings, about buildings.

Fig. 587  Stem, leaf, and flower of Calystegia sepium ssp. americana showing pubescent internodes, petioles, and peduncles.

5 32   tricolpate s

1a. Plants pubescent on the distal stems, petioles, peduncles, and abaxial blade surface of new leaves [Fig. 587]; sinus of leaf blade commonly V-shaped [Fig. 587]; corolla pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a. C. sepium ssp. americana (Sims) Brummitt 1b. Plants glabrous or with a few hairs on the distal portion of the petiole (rarely the peduncle bases also pubescent); sinus of leaf blade U-shaped or V-shaped; corolla white to pink 2a. Sinus of leaf blade commonly V-shaped [Fig. 587]; corolla white or rarely pale pink, usually 30–50 mm long; stamens (15–) 17–23 (–25) mm long . . . . 2b. C. sepium ssp. sepium 2b. Sinus of leaf blade commonly U-shaped; corolla white or pink, usually 45–64 mm long; stamens (19–) 23–30 (–32) mm long 3a. Corolla usually white; bracteoles (6–) 10–18 mm wide, usually approximate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2c. C. sepium ssp. angulata Brummitt 3b. Corolla usually pink; bracteoles (12–) 14–26 (–28) mm wide, usually overlapping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2d. C. sepium ssp. appalachiana Brummitt Subspecies americana is native and known from CT, MA, ME, NH, RI, VT. It is most frequent in, but not confined to, coastal counties and along Lake Champlain, VT. It is frequently misinterpreted as the correct name for all of our native subspecies of Calystegia sepium (e.g., Tucker 1995, Sorrie and Somers 1999). Subspecies angulata is native and known from CT, MA, ME, NH, RI, VT. Subspecies appalachiana is native and known from CT, MA, ME, NH, VT. Subspecies sepium is nonnative and known from MA, NH, RI, VT. 3. Calystegia silvatica (Kit.) Griseb. ssp. fraterniflora (Mackenzie & Bush) Brummitt

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N C Fig. 588 short-stalked false bindweed. Calystegia fraterniflora (Mackenzie & Bush) Brummitt; C. sepium (L.) R. Br. var. fraterniflora (Mackenzie & Bush) Shinners; Convolvulus fraterniflorus (Mackenzie & Bush) Mackenzie & Bush; C. sepium L. var. fraterniflorus Mackenzie & Bush • CT, MA, RI, VT. Fields, roadsides, clearings. Rare in New England and usually overlooked as Calystegia sepium. 4. Calystegia spithamaea (L.) Pursh ssp. spithamaea Fig. 588  Stem, leaf, and paired flowers of Calystegia silvatica showing rectangular leaf blade sinus.

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upright false bindweed. Convolvulus spithamaeus L. • CT, MA, ME, NH, VT. Sandy fields, roadsides, and clearings, railroads, woodlands, sand plain grasslands. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it could occur in RI and was unaware of any collections

Convolvulus Convolvulus tricolor L. was reported from MA by Magee and Ahles (1999), but specimens are unknown. 1. Convolvulus arvensis L.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E field bindweed. Strophocaulos arvensis (L.) Small • CT, MA, ME, NH, RI, VT. Fields, waste areas.

Cuscuta Herbarium specimens of Cuscuta are much more difficult to interpret than living populations as many structures dry and become less obvious. Careful notes on corolla lobe orientation, numbers of perianth parts, etc., will make specimen review easier. C. epilinum Weihe was reported from MA by Knowlton and Deane (1923b), but specimens are unknown. Reports of C. epilinum from VT were based on a collection of C. epithymum. References: Yuncker (1932), Costea et al. (2006a, 2006b, 2006c). 1a. Stigmas filiform, elongate-tapering, much taller than wide [Fig. 589]; fruit a pyxis, circumscissile near the base 2a. Perianth mostly 4-merous; lobes of the calyx obtuse at the apex; infrastaminal scales bifid and lacking a fringe; seeds ca. 1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . C. europaea

Co nv o lv u l ac e a e  5 3 3

2b. Perianth mostly 5-merous [Fig. 589]; lobes of the calyx acute at the apex [Fig. 589]; infrastaminal scales simple and fringed; seeds up to 1.3 mm long 3a. Connate portion of the calyx gold-yellow, lustrous, and with a prominent cellularreticulate pattern; calyx lobes terminated by a short, fleshy, cylindrical appendage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. approximata 3b. Connate portion of the calyx not gold-yellow, dull, without a prominent cellularreticulate pattern; calyx lobes not terminated by a cylindral appendage . . . C. epithymum 1b. Stigmas capitate, dilated near the apex, not or only slightly taller than wide; fruit a utricle or an irregularly rupturing capsule 4a. Sepals distinct; flowers subtended by 3–5 imbricate bracts; seeds (1.9–) 2–2.6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. compacta 4b. Sepals connate at the base; flowers not individually provided with bracts; seeds 0.9–2.2 mm long 5a. All or many flowers with a 4-merous corolla [Fig. 591] 6a. Corolla lobes obtuse or rounded at the apex; tips of the calyx lobes not reaching the sinuses between the corolla lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cephalanthi 6b. Corolla lobes acute to acuminate at the apex; tips of the calyx lobes reaching or surpassing the sinuses between the corolla lobes 7a. Calyx lobes obtuse at the apex; corolla lobes relatively smooth, not granulatetextured, the tip of lobes erect or ascending; pedicels absent or shorter than the flowers; marcescent corolla persistent around the base of the fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. polygonorum 7b. Calyx lobes acute at the apex; corolla lobes appearing granulate-textured due to thickened outer cells, the tip of lobes inflexed [Fig. 590]; pedicels usually as long or longer than the flowers; marcescent corolla forming a cap over the fruit (though sometimes falling off early) 8a. Flowers 2–5.3 mm long; infrastaminal scales adnate to the connate portion of the corolla for ⅓ to ½ (rarely ¾) of their length, subspathulate to spathulate in outline, rarely 2- or 3-lobed, rounded to truncate at the apex, with (6–) 20–35 (–50) fimbria; dried capsules brown-yellow to light brown, somewhat translucent, 0.8–1.5 times as long as wide, with scarcely depressed suture lines beteen the 2 carpels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. indecora 8b. Flowers 1.7–2.6 (–3) mm long; infrastaminal scales united with the connate portion of the corolla for most of their length, oblong, bifid with 1–3 fimbria on each side of the filament attachment (rarely truncate and with 3–6 fimbria); dried capsules brown, opaque, 1.6–2.4 times as long as wide, with depressed suture lines between the carpels that form a groove on each side of the fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. coryli 5b. Flowers with a 5-merous corolla [Fig. 590] 9a. Corolla lobes obtuse at apex, not inflexed at the tip [Fig. 590]; tips of the calyx lobes not reaching the sinuses between the corolla lobes (except in var. latiflora); fruit globose to ovoid to conical; stylopodium present . . . . . . . . . . . . . . . . . . C. gronovii 9b. Corolla lobes acute at apex, usually inflexed at the tip [Fig. 591]; tips of the calyx lobes ± reaching or slightly surpassing the sinuses between the corolla lobes (usually not reaching the sinuses in C. indecora); fruit usually globose to depressed-globose; stylopodium absent 10a. Calyx lobes acute to narrow-obtuse at apex; corolla lobes erect to ascending; flowers with a faint sweet smell in life; marcescent corolla calyptrate (i.e., forming a cap over the fruit, eventually detaching at base and pushed off like a lid) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. indecora

5 34   tricolpate s

10b. Calyx lobes broad-obtuse at apex; corolla lobes spreading to reflexed; flowers inodorous in life; marcescent corolla persistent around the base of the fruit 11a. Calyx lobes ovate-triangular, not or scarcely overlapping at base (except in some newly opened flowers), thus the calyx ± circular when viewed from above; flowers (1.9–) 2.1–3.6 mm long; anthers (0.3–) 0.4–0.5 mm long; seeds 1.1–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. campestris 11b. Calyx lobes broad-ovate to rhombic, broadly overlapping at base, protruding to form angles (note: this most visible on mature flowers, it is often obscured in fruit), thus the calyx 5-angled when viewed from above; flowers 1.4–2.1 (–2.5) mm long; anthers 0.25–0.3 (–0.35) mm long; seeds 0.9–1.1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pentagona 1. Cuscuta approximata Bab. E alfalfa dodder. Cuscuta approximata Bab. var. urceolata (Kunze) Yuncker; C. epithymum L. ssp. approximata (Bab.) Rouy; C. planiflora Ten. • MA. Parasitic on tracheophytes, commonly species of Medicago. 2. Cuscuta campestris Yuncker E field dodder. Cuscuta arvensis Beyrich ex Hook. var. calycina Engelm.; C. pentagona Engelm. var. calycina Engelm.; Grammica campestris (Yuncker) Hadac & Chrtek • MA. Parasitic on a large number of herbaceous tracheophytes, in New England observed on Persicaria. 3. Cuscuta cephalanthi Engelm. N buttonbush dodder. Epithymum cephalanthi (Engelm.) Nieuwl. & Lunnell; Grammica cephalanthi (Engelm.) Hadac & Chrtek • CT, MA, nh, RI; also reported from ME by several sources (e.g., Campbell et al. 1995, Kartesz 1999), but specimens are unknown. Parasitic on a large number of tracheophytes, commonly on coarse herbeaceous and shrub types (e.g., Cephalanthus, Decodon, Impatiens). Number of perianth parts is known to vary in this species in some regions of the United States (Musselman 1986). Specimens with 5-merous perianths would key to Cuscuta gronovii but are separable based on the absence of a stylopodium and globose to depressed-globse fruit (a stylopodium is present in C. gronovii, which has a globose to ovoid or conical fruit). 4. Cuscuta compacta Juss. ex Choisy var. compacta N compact dodder. Cuscuata compacta Juss. ex Choisy var. adpressa Engelm.; Cuscuta glomerata Choisy var. adpressa (Engelm.) Choisy in DC.; Lepidanche adpressa Engelm. • CT, MA, RI; also reported from NH by Kartesz (1999), but specimens are unknown. Parasitic on many tracheophytes, commonly shrub species. Fig. 589  Flowers of Cuscuta epithymum showing elongate stigmas.

5. Cuscuta coryli Engelm.

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hazel dodder. Epithymum coryli (Engelm.) Nieuwl. & Lunnell; Grammica coryli (Engelm.) Hadac & Chrtek • CT, MA, RI. Parasitic on a diversity of tracheophytes, especially on members of the Asteraceae (e.g., Euthamia, Solidago, Symphyotrichum), usually found on the coastal plain. 6. Cuscuta epithymum L. E Fig. 589 clover dodder. CT, MA, ME, RI, VT. Parasitic on members of the Fabaceae, especially Medicago and Trifolium, as well as other herbaceous tracheophytes. 7. Cuscuta europaea L. E greater dodder. ME; western portion of state. Parasitic on Diervilla, Doellingeria, Solidago, and Symphyotrichum. 8. Cuscuta gronovii Willd. ex J.A. Schultes N Fig. 590

Fig. 590  Flowers of Cuscuta gronovii showing 5-merous perianth.

common dodder.  8a. Cuscuta saururi Engelm.; 8b. Cuscuta vulgivaga Engelm; Grammica gronovii (Willd. ex J.A. Schultes) Hadac & Chrtek • CT, MA, ME, NH, RI, VT. Parasitic on a large number of tracheophytes, including herbaceous and shrub types.

Con v o lvul ac e a e   5 3 5

1a. Calyx on most or all flowers ± as long as the basal, connate portion of the corolla (i.e., the tips of the sepals reaching the sinuses between the petals), with oblong to ovate lobes that barely overlap one another; basal, connate portion of corolla 1–1.5 mm long; corolla lobes ca. ½ to fully as long as long as the basal, connate portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8a. C. gronovii var. latiflora Engelm. 1b. Calyx ca. ½ as long as the basal, connate portion of corolla (i.e., the tips of the sepals not reaching the sinuses between the petals), with ovate to suborbicular lobes that overlap one another; basal, connate portion of corolla 1.5–2.5 mm long; corolla lobes ca. ⅓ (rarely only ¼) as long as the basal, connate portion . . . . . . . . . . . . . . . . . . . . . . . . 8b. C. gronovii var. gronovii Variety laterifolora is known from CT, MA, ME, NH, RI, VT. It is less common than the typical variety and is rare in some states (e.g., ME). Variety gronovii is known from CT, MA, ME, NH, RI, VT. It is the most common taxon of Cuscuta in New England. 9. Cuscuta indecora Choisy var. indecora

N C Fig. 591

collared dodder. Cuscuta indecora Choisy var. neuropetala (Engelm.) A.S. Hitchc.; C. neuropetala Engelm.; C. pulcherrima Scheele; Grammica indecora (Choisy) W.A. Weber ssp. neuropetala (Engelm.) W.A. Weber • CT, MA, RI. Parasitic on a large number of tracheophytes, including herbaceous and shrub types, especially on members of the Asteraceae (e.g., Coreopsis, Euthamia, Solidago, Symphyotrichum), found usually on the coastal plain. Cuscuata indecora normally displays a 5-merous perianth. However, recent collections from Rhode Island have been confusing in that they have mainly 4-merous perianths. These plants can be separated from C. coryli by features presented in the key. 10. Cuscuta pentagona Engelm.

NC

bush-clover dodder. Cuscuta arvensis Beyrich ex Hook.; Epithymum arvense (Beyrich ex Engelm.) Nieuwl. & Lunnell; Grammica pentagona (Engelm.) W.A. Weber • CT, MA, RI; also reported from NH by Seymour (1982), but specimens are unknown. The report for NH (Ossipee) by Seymour (1982) was based on a specimen of Cuscuta gronovii. Parasitic on a large number of tracheophytes, especially members of the Asteraceae (e.g., Ambrosia, Centaurea, Liatris, Symphyotrichum), mainly on or near the coastal plain. 11. Cuscuta polygonorum Engelm.

NC

smartweed dodder. Cuscuta chlorocarpa Engelm. • CT, MA; also reported from ME by Campbell et al. (1995) and RI by Seymour (1982), but specimens are unknown. Parasitic on various tracheophytes, especially coarse herbaceous types.

Ipomoea 1a. Corolla usually red (but white and yellow forms are known), ± salverform with an elongate, narrow, tube-like, basal connate portion; stamens and style exserted from the corolla 2a. Leaf blade pinnately divided into 11–31 linear segments; sepals obtuse to shortapiculate, lacking prolonged awns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. quamoclit 2b. Leaf blade entire to lobed with 3–7 lanceolate to ovate lobes, but not divided; sepals with evident, elongate, terminal or subterminal awns 3a. Calyx (5–) 6–8 (–9) mm long; fruits reflexed . . . . . . . . . . . . . . . . . . . . . . . . . . . I. coccinea 3b. Calyx 4–4.5 mm long; fruits erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. hederifolia 1b. Corolla blue to purple or pink to white, funnelform to nearly campanulate [Fig. 592], the basal connate portion expanding from below the middle; stamens and style not exserted from corolla [Fig. 592] 4a. Stigmas 3-lobed; ovary 3-locular; sepals conspicuously hirsute on the basal half 5a. Leaf blades usually entire; sepals 10–15 mm long, acute to acuminate at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. purpurea

Fig. 591  Flowers of Cuscuta indecora showing 4-merous perianth and incurved petal apices.

5 36   tricolpate s

5b. Leaf blades usually 3-lobed [Fig. 592]; sepals 15–25 mm long, with a linear, spreading to recurved tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. hederacea 4b. Stigmas 2-lobed or entire; ovary 2-locular; sepals glabrous or sparsely villous on the basal half 6a. Corolla abaxially white below, the limb usually blue to blue-purple (fading to red-purple) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. tricolor 6b. Corolla abaxially white throughout (rarely pink to pale purple) 7a. Corolla white (rarely pink to pale purple), 10–23 mm long; sepals 10–15 mm long, acuminate at the apex, often abrubtly so, hirsute-ciliate along the margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. lacunosa 7b. Corolla white with a red to purple center, 50–80 mm long; sepals 13–20 mm long, obtuse or rounded at the apex, eciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. pandurata 1. Ipomoea coccinea L. E red morning-glory. Quamoclit coccinea (L.) Moench • CT, MA, RI, VT. Roadsides, waste areas, railroads, gardens. 2. Ipomoea hederacea Jacq. E Fig. 592 ivy-leaved morning-glory. Ipomoea barbigera Sweet; I. hederacea Jacq. var. integriuscula Gray; Pharbitis barbigera (Sweet) G. Don; P. hederacea (Jacq.) Choisy • CT, MA, ME, NH, VT. Dumps, yards, waste areas, roadsides, railroads. 3. Ipomoea hederifolia L. E scarlet morning-glory. Ipomoea coccinea L. var. hederifolia (L.) Gray • MA, VT. Roadsides, waste areas, dumps. Fig. 592  Flower and leaf of Ipomoea hederacea.

4. Ipomoea lacunosa L. E white morning-glory. MA. Roadsides, waste areas, dumps, railroads. 5. Ipomoea pandurata (L.) G.F.W. Mey. n wild sweet potato morning-glory. Convolvulus panduratus L. • CT, MA; also reported from RI by George (1997), but specimens are unknown. Fields, roadsides, woodlands, forest clearings, waste areas. This species is thought to be native to CT but is introduced in MA. 6. Ipomoea purpurea (L.) Roth E common morning-glory. Convolvulus purpureus L.; Pharbitis purpurea (L.) Voigt • CT, MA, ME, NH, RI, VT. Roadsides, waste areas, gardens, abandoned homesteads, dumps. 7. Ipomoea quamoclit L. E Cypress-vine morning-glory. Quamoclit quamoclit (L.) Britt.; Quamoclit vulgaris Choisy • VT. Roadsides, waste areas, dumps. 8. Ipomoea tricolor Cav. E granny morning-glory. MA. Roadsides, waste areas, dumps.

Cornaceae The Cornaceae have seen a number of changes, including the inclusion of the Nyssaceae and the dismantling of the long familiar genus Cornus (accepted by some authors). Multiple studies have shown that Cornus can be confidently split into four (or more) groups worldwide that are morphologically and phylogenetically distinct (Fan and Xiang 2001, Xiang et al. 2006). Further supporting the recognition of these groups are patterns of hybridization (interspecific

Co rn ac e a e   5 37

hybrids are restricted to within the genera recognized here). Three of the four genera are represented in New England. Cornus s.s. (Cornelian cherry) does not occur in the region. 1a. Flowers with 5–8 petals, of 2 types—staminate with 8–15 stamens and functionally carpellate with 5–10 stamens that have nonfunctional anthers; pith diaphragmed; leaves alternate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nyssa 1b. Flowers with 4 petals, of 1 type—bisexual with 4 stamens; pith of woody species solid; leaves opposite, whorled, or alternate 2a. Aerial stems herbaceous, 0.1–0.2 m tall, with whorled leaves (sometimes opposite at some nodes); petals terminated by awns; leaves strongly dimorphic, those of the lower nodes many times smaller than those of the upper node (+/- monomorphic in rare putative hybrid individuals); inflorescence pseudanthial, a corymb-like cyme subtended by 4 white, petaloid bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chamaepericlymenum 2b. Aerial stems woody, 1–10 m tall, with opposite or whorled leaves; petals not awned at apex; leaves essentially monomorphic; inflorescence pseudanthial or not, with or without showy, petaloid bracts 3a. Inflorescence pseudoanthial, with 4, white to yellow-white or pink, petaloid bracts [Fig. 593]; flowers sessile; petals green-white to yellow; fruits separate or fused together into a multiple, red; terminal flower bud large, globose or discoid . . . . . . . . Benthamidia 3b. Inflorescence not pseudanthial, with minute and usually caducous bracts [Fig. 595]; flowers pedicellate [Fig. 595]; petals ± white; fruits separate, white to blue, black, or green; terminal flower bud smaller, ovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Swida

Benthamidia Benthamia Spach is the oldest name for this genus; however, it is a later homonym of Benthamia A. Rich. The next oldest genus is Benthamidia Spach (and the one used here). Benthamidia is here treated conservatively as including Dendrobenthamia (the multiple-fruit species). 1a. Bracts of inflorescence truncate to emarginate at apex [Fig. 593]; fruits separate; leaf blades with 6 or 7 pairs of veins; petioles 5–15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . B. florida 1b. Bracts of inflorescence acuminate at apex; fruits fused together into a multiple; leaf blades with (3–) 4 or 5 pairs of veins; petioles 4–6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . B. japonica 1. Benthamidia florida (L.) Spach N Fig. 593 flowering big-bracted-dogwood. Benthamia florida (L.) Spach; Cornus florida L.; Cynoxylon floridum (L.) Raf. ex B.D. Jackson • CT, MA, ME, NH, RI, VT; mainly restricted to the southern portion of the northern New England states (though occurring in some northern counties in NH). Dry-mesic to mesic forests, forest edges, and woodlands. 2. Benthamidia japonica (Siebold & Zucc.) Hara E kousa big-bracted-dogwood. Benthamia japonica Siebold & Zucc.; B. kousa (Buerger ex Miq.) Nakai; Cornus kousa Buerger ex Miq.; Dendrobenthamia japonica (Siebold & Zucc.) W.P. Fang • CT. Fields, roadsides, waste areas.

Chamaepericlymenum Rare individuals of this genus in New England show multiple nodes with opposite leaves and/ or purple-tipped petals. These plants are hypothesized by Murrell (1994) to be the product of hybridization between Chamaepericlymenum canadense and C. suecica (L.) Aschers. & Graebn. backcrossed to C. canadense. Reference: Murrell (1994). 1. Chamaepericlymenum canadense (L.) Aschers. & Graebn. N Canada dwarf-dogwood. Cornella canadensis (L.) Rydb.; Cornus canadensis L. • CT, MA, ME, NH, RI, VT. Forests, forest clearings, swamps.

Fig. 593  Inflorescence of Benthamidia florida.

5 38   tricolpate s

Nyssa 1. Nyssa sylvatica Marsh. N black tupelo. Nyssa sylvatica Marsh. var. caroliniana (Poir.) Fern.; N. sylvatica Marsh. var. dilatata Fern.; N. sylvatica Marsh. var. typica Fern. • CT, MA, ME, NH, RI, VT. Swamps, pond shores, stream-side fens, occasionally one or more individuals occur along field edges, in forests, and on hillsides.

Swida Swida is treated conservatively here as including Bothrocaryum (the alternate-leaved species). 1a. Leaves alternate, crowded at the apex of the twig [Fig. 594]; petioles (8–) 20–50 mm long; endocarp pitted at apex; branchlets sometimes yellow with minute red-brown to black dots due to infection of Cryptodioporthe corni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. alternifolia 1b. Leaves opposite, ± evenly spaced along the twig; petioles 3–15 (–25) mm long; endocarp not pitted; branchlets without above-mentioned fungal infection 2a. Drupes black or green at maturity; style conspicuously widened in the apical portion; infructescence with red branches and pedicels . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sanguinea 2b. Drupes white to blue at maturity; style of uniform diameter throughout (apically widened in S. amomum); infructescence with yellow or gray-brown to brown branches and pedicels (red in S. racemosa) 3a. Pith of branchlets brown; style conspicuously widened in the apical portion [Fig. 595]; sepals 1–2 mm long; drupes blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. amomum 3b. Pith white (infrequently light brown in S. racemosa); style of nearly uniform diameter throughout; sepals shorter than 1 mm; drupes white to blue 4a. Winter branchlets gray-brown; inflorescence convex or pyramidal, nearly as tall as wide or taller, with red branches and pedicels when in fruit; leaf blades with 3 or 4 (rarely 5) pairs of lateral veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. racemosa 4b. Winter branchlets not uniformly gray-brown (see descriptions in couplet 5); inflorescence flat-topped, often wider than tall, with branches yellow or gray-brown to brown; leaf blades with 5–8 pairs of lateral veins 5a. Branchlets red to red-purple; drupe white (rarely light blue); leaf blades lanceolate to ovate; seeds brown, with 7–9 yellow stripes; buds without scales, densely pubescent throughout with red-brown and some white hairs . . S. sericea

Fig. 594  Alternate leaves crowded near apex of branchlet of Swida alternifolia.

5b. Branchlets green to yellow-green, often mottled with purple or red; drupe light blue (rarely white); leaf blades ovate to suborbicular; seeds light brown, without stripes; buds covered about ⅔ of their length by sparsely pubescent scales, only the tips of the densely pubescent leaf primordia visible . . . . . . . . . . . . . . . S. rugosa 1. Swida alternifolia (L. f.) Small N Fig. 594 alternate-leaved dogwood. Bothrocaryum alternifolia (L. f.) Pojark.; Cornus alternifolia L. f. • CT, MA, ME, NH, RI, VT; nearly throughout. Forests, forest edges, thickets.

2. Swida amomum (P. Mill.) Small N Fig. 595 silky dogwood.  2a. Cornus amomum P. Mill.; 2b. Cornus amomum P. Mill. ssp. obliqua (Raf.) J.S. Wilson; C. amomum P. Mill. var. schuetzeana (C.A. Mey.) Rickett; C. obliqua Raf.; C. purpusii Koehne; C. sericea L. var. schuetzeana C.A. Mey.; Swida obliqua (Raf.) Moldenke • CT, MA, ME, NH, RI, VT. Swamps, shorelines, low fields, wetland margins.

Fig. 595  Inflorescence of Swida amomum.

1a. Leaf blades mostly 6–12 cm long, ovate to broad-elliptic, 1–2.2 times as long as wide, subtruncate to rounded at the base, with 4–6 pairs of lateral veins, green and usually nonpapillose on the abaxial surface, with white or, more commonly, red-brown pubescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a. S. amomum var. amomum

Co rnac e a e   5 3 9

1b. Leaf blades mostly 6–9 cm long, lanceolate to narrow-ovate-oblong, 2.2–4 times as long as wide, cuneate at the base, with 3–5 pairs of lateral veins, usually white and papillose on the abaxial surface, with white or, only rarely, red-brown pubescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. S. amomum var. schuetzeana (Raf.) A. Haines Variety amomum is known from CT, MA, ME, NH, RI, VT. Variety schuetzeana is known from CT, MA, ME, NH, VT. It is less common than the preceding variety. This species is sometimes confused with Swida sericea because it frequently produces a red stem. ‌2b × 3. Swida ×arnoldiana (Rehd.) Soják is a very rare dogwood hybrid known from the Arnold Arboretum in MA, where it originated spontaneously. It was described as being rather similar to Swida racemosa but with red-purple winter branchlets (instead of gray). 3. Swida racemosa (Lam.) Moldenke N gray dogwood. Cornus foemina P. Mill. ssp. racemosa (Lam.) J.S. Wilson; C. paniculata L’Hér.; C. racemosa Lam. • CT, MA, ME, NH, RI, VT. Succeeding fields, riparian forests, roadsides, forests. 4. Swida rugosa (Lam.) Rydb. N round-leaved dogwood. Cornus rugosa Lam. • CT, MA, ME, NH, RI, VT. Forests, riparian forests, forest edges, ledges, talus slopes. ‌4 × 6. Swida ×slavinii (Rehd.) Holub is a very rare dogwood hybrid known from ME (though it is likely overlooked and more common than collections indicate). It resembles S. sericea in habit but is more upright, and the winter branchlets are red-purple with sparse, dark, longitudinal lines. The leaf blade shape is more similar to S. rugosa, but they are glaucous abaxially, as well as villous abaxially on flowering stems (the hairs are more appressed in S. sericea). The drupes are usually tinged with blue (as in S. rugosa), but the seeds tend to have yellow stripes (as in S. sericea). The phenology is intermediate between the putative parents—S. sericea flowers first, then the hybrid, then S. rugosa. 5. Swida sanguinea (L.) Opiz E blood-twig dogwood. Cornus sanguinea L. • MA. Fields, roadsides, areas of habitation. 6. Swida sericea (L.) Holub N red-osier dogwood. Cornus alba L. ssp. stolonifera (Michx.) Wangerin; C. sericea L.; C. sericea L. ssp. stolonifera (Michx.) Fosberg; C. stolonifera Michx.; Swida stolonifera (Michx.) Rydb. • CT, MA, ME, NH, RI, VT. River shores, including fresh-tidal systems, hydric fields and pastures, swamps, fens.

Crassulaceae 1a. Plants annual, aquatic or amphibious; flowers solitary in the axils of the leaves; perianth 3- or 4-merous; stamens 3 or 4 per flower (i.e., as many as the petals); leaves connate around the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crassula 1b. Plants perennial, terrestrial; flowers usually borne in cymes; perianth 4- to 16-merous; stamens usually 8–32 per flower (i.e., twice as many as the petals); leaves not connate around the stem 2a. Perianth mostly 12- to 16-merous; stamens mostly 24–32 per flower; leaves in crowded, basal rosettes; plants monocarpic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sempervivum 2b. Perianth mostly 4- or 5-merous; stamens mostly 8 or 10 per flower; leaves borne on a stem; plants polycarpic

5 40  tricolpates

3a. Leaf blades entire, often very thick (i.e., elliptic, semi-circular, or terete in crosssection) [Fig. 599]; leaves alternate, opposite, or whorled; plants forming mats by means of creeping stems (not mat-forming in S. hispanicum) . . . . . . . . . . . . . . . . . . . . . . . . . Sedum 3b. Leaf blades toothed (sometimes inconspicuously so), broad and flat [Figs. 596, 597, 598]; leaves alternate (opposite in Phedimus spurius); plants with upright stems, sometimes basally decumbent in age (mat-forming by creeping stems in Phedimus spurius) 4a. Flowers 4-merous (rarely 5-merous), unisexual; reproductive stems produced from the axils of brown, scale-like leaves on a partly subterranean rootstock . . . . Rhodiola 4b. Flowers 5-merous [Fig. 596], bisexual (sometimes with defective and missing parts in Hylotelephium erythrostictum); reproductive stems produced from underground, tuberous roots or arising from creeping, horizontal stems 5a. Plants forming mats by creeping, horizontal stems; roots not tuberous; leaves opposite 6a. Petals yellow, spreading from base [Fig. 596] . . . . . . . . . . . . (in part) Aizopsis 6b. Petals pink to pink-purple, erect to ascending at the base and spreading above [Fig. 598] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phedimus 5b. Plants not forming mats; roots tuberous, carrot-shaped; leaves alternate, opposite, whorled, or sometimes varied on a single plant 7a. Ovaries and follicles narrowly tapering at base, stipitate, lacking an adaxial outgrowth; petals red-purple to purple or white . . . . . . . . . . . . . . . Hylotelephium 7b. Ovaries and follicles not tapering at base, sessile, with an adaxial outgrowith; petals yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Aizopsis

Aizopsis Aizopsis has been treated as part of Phedimus. However, base chromosome number (x=16 vs. x=14), petal color (yellow vs. white or pink), petal orientation (spreading from base vs. erect at base), and distribution (primarily Asian vs. primarily European) justify Aizopsis being treated as distinct from Phedimus. Reference: Gontcharova et al. (2006). 1a. Roots tuberous, carrot-shaped; leaf blades narrow-lanceolate to elliptic-lanceolate or oblong-lanceolate (i.e., usually widest at or below the middle); stems erect, 20–50 cm tall; sepals linear . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. aizoon 1b. Roots not tuberous, relatively slender; leaf blades oblanceolate to obovate or spatulate (i.e., widest above the middle) [Fig. 596]; stems decumbent, 15–40 cm tall; sepals lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. kamtschatica 1. Aizopsis aizoon (L.) V. Grulich E aizoön stonecrop. Phedimus aizoon (L.) ’t Hart; Sedum aizoon L. • MA. Gardens, areas of habitation, lawns. 2. Aizopsis kamtschatica (Fisch. & C.A. Mey.) Grulich E Fig. 596 orange stonecrop. Aizopsis ellacombeana (Praeger) P.V. Heath; Phedimus ellacombeanus (Fisch. & C.A. Mey.) ’t Hart; Sedum ellacombeanum Praeger; S. kamtschaticum Fisch. & C.A. Mey.; S. kamtschaticum Fisch. & C.A. Mey. ssp. ellacombeanum (Praeger) Clausen, nom. inval. • ME, NH. Gardens, rock walls, roadsides, lawns. Aizopsis kamtschatica is recognized here in the broad sense because there is no detailed treatment available that discusses its distinctions from A. ellacombeana. Further study is needed to clarify the taxonomy of these two species. Fig. 596  Inflorescence and leaves of Aizopsis kamtschatica.

Cr a ssu l ac e a e   5 41

Crassula Crassula saginoides (Maxim.) Bywater & Wickens was reported from MA by Kartesz (1999), but specimens are unknown. 1. Crassula aquatica (L.) Schoenl. N pygmy-weed. Bulliarda aquatica (L.) DC.; Hydrophila aquatica (L.) House; Tillaea angustifolia Nutt.; T. aquatica L.; Tillaeastrum aquaticum (L.) Britt. • CT, MA, ME, NH, VT. Open, often muddy, shorelines, these typically brackish-tidal rivers and large, non-tidal rivers, but rarely of pond shores as well. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it could be in RI and was unaware of any collections.

Hylotelephium Hylotelephium forms a well-supported clade with Orostachys (Gontcharova et al. 2006). Unfortunately, the name Orostachys has priority as a generic epithet. Therefore, all our species of Hylotelephium need to be transferred to Orostachys. Hylotelephium telephioides (Michx.) H. Ohba was reported from CT by Magee and Ahles (1999), but specimens are unknown. Reference: Fu and Ohba (2001). 1a. Leaves tending to be alternate (but subopposite, opposite, and whorled leaves may also be present on a plant) [Fig. 597]; anthers yellow; cymes rounded at the apex . . . . . . . H. telephium 1b. Leaves tending to be opposite (but alternate and whorled leaves may also be present on a plant); anthers violet to purple; cymes relatively flat-topped 2a. Flowers with white to green-white petals, 0–10 stamens, and 0–5 carpels; stamens slightly shorter than to ± equaling the petals in length; leaf blades serrate to crenateserrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. erythrostictum 2b. Flowers with pink to purple petals, 10 stamens, and 5 carpels; stamens longer than the petals; leaf blades entire to undulate-dentatate (infrequently crenate-serrate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. spectabile 1. Hylotelephium erythrostictum (Miq.) H. Ohba E garden orpine. Sedum alboroseum Baker; S. erythrostictum Miq. • MA, ME, VT. Roadsides, waste areas, gardens, lawns. This plant was thought by Clausen (1949) to be the hybrid product of Hylotelephium spectabile (Boreau) H. Ohba and H. viridescens (Nakai) H. Ohba. Though frequently lacking some or all of the stamens or carpels in a given flower, in its native range (Asia), this species also produces normal flowers. 2. Hylotelephium spectabile (Boreau) H. Ohba E showy orpine. Sedum spectabile Boreau • CT, MA. Gardens, yards, roadsides, fields. 3. Hylotelephium telephium (L.) H. Ohba E Fig. 597 purple orpine. 3a. Sedum fabaria W.D.J. Koch; S. telephium L. ssp. fabaria (W.D.J. Koch) Kirschl.; 3b. Sedum purpureum (L.) J.A. Schultes; S. purpurascens W.D.J. Koch; S. telephium L.; • CT, MA, ME, NH, RI, VT. Roadsides, gardens, riparian forests, fields, areas of habitation. Though typically producing alternate leaves, this species can produce opposite leaves on shoots from axillary buds. 1a. Upper leaf blades cuneate at base, often shortly petiolate; follicles not grooved on abaxial (i.e., outer) surface . . . . . . . . . . . . . . . . . . . . 3a. H. telephium ssp. fabaria (W.D.J. Koch) H. Ohba 1b. Upper leaf blades shortly truncate, sessile; follicles grooved on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3b. H. telephium ssp. telephium Subspecies fabaria is known from CT. Subspecies telephium is known from CT, MA, ME, NH, RI, VT. It is our most common Hylotelephium in New England.

Fig. 597  Inflorescence and leaves of Hylotelephium telephium.

5 42   tricolpate s

Phedimus 1a. Leaf blades crenate-serrate on the distal portion [Fig. 598], ciliate with stout, white hairs along the margin; petals erect at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. spurius 1b. Leaf blades entire to obscurely crenate near apex, not or only sparingly ciliate short, blunt, white hairs; petals ascending at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. stoloniferus 1. Phedimus spurius (Bieb.) ’t Hart E Fig. 598 false stonecrop. Sedum spurium Bieb.; Spathulata spuria (Bieb.) A. & D. Löve • CT, MA, ME, NH, VT. Ledges, gardens, stone steps, dumps, road cuts. 2. Phedimus stoloniferus (Gmel.) ’t Hart E running false stonecrop. Sedum stoloniferum Gmel. • ME, NH. Gardens, yards. Fig. 598  Inflorescence and leaves of Phedimus spurius.

Rhodiola 1. Rhodiola rosea L. n roseroot. Rhodiola roanensis Britt.; Sedum roanense Britt.; S. roseum (L.) Scop.; S. roseum (L.) Scop. var. roanense (Britt.) Berger • CT, ME, VT; native populations along eastern half of ME coastline and southern VT. Rocks, ledges, and turf near the Atlantic ocean, inland cliffs in high-pH bedrock regions. This species in native to ME and VT and introduced in CT.

Sedum Sedum has been shown to be highly artificial (i.e., non-monophyletic) and has been split into smaller, more homogeneous genera that better reflect the evolutionary history of the family (’t Hart 1995, Ohba 2009). Recognized here apart from the broadly defined Sedum are Aizopsis, Hylotelphium, Phedimus, and Rhodiola. Reference: Ohba (2009). 1a. Upper stem and axis of inflorescence stipitate-glandular; petals white and usually some with a pink or green midstripe; flowers with both 6- to 9-merous corollas and horizontally spreading mature carpels; plants annual or biennial (rarely short-lived perennial) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hispanicum 1b. Upper stem and axis of inflorescence glabrous (though stipitate-glandular near base of plant in S. album); petals yellow to yellow-white or entirely white; flowers not combining the stated characters, with either 4- or 5-merous corollas, or erect follicles, or both; plants perennial 2a. At least the leaves of the sterile shoots and basal portion of reproductive shoots whorled (i.e., 3 at each node) or less frequently opposite [Fig. 599], the blades relatively flat 3a. Petals yellow; flowers mostly 5-merous; leaf blades elliptic-oblanceolate to oblonglanceolate or narrow-ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sarmentosum 3b. Petals white; flowers mostly 4-merous; leaf blades obovate (except the upper ones of reproductive shoots, which are oblanceolate to ± linear). . . . . . . . . . . . . . . . S. ternatum 2b. Leaves alternate, sometimes tightly imbricate and the arrangement difficult to discern, the blades strongly biconvex to subterete in cross-section 4a. Flowers (5–) 6- or 7 (–9)-merous; leaf blades pointed or apiculate at the apex; follicles erect 5a. Sepals 5–6 mm long, lanceolate to broad-lanceolate, acute to acuminate at the apex, slightly concave on the abaxial surface; petals cream to green-white or whiteyellow, 8–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. ochroleucum 5b. Sepals 1.5–3 mm long, ovate, acute or obtuse at the apex, not concave on the abaxial surface; petals bright yellow, 6–7 mm long . . . . . . . . . . . . . . . . . . . . . S. reflexum 4b. Flowers usually 5-merous; leaf blades blunt at the apex; follicles divergent (erect in S. album)

C r a ssu l ac e a e   5 43

6a. Petals white; follicles erect; leaf blades 4–20 mm long . . . . . . . . . . . . . . . . S. album 6b. Petals yellow; follicles divergent; leaf blades 2–6 mm long 7a. Leaf blades ovoid, not arranged in consistent vertical ranks; petals 6–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. acre 7b. Leaf blades narrow-cylindric, arranged in 6 vertical ranks; petals 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sexangulare 1. Sedum acre L. E moss stonecrop. CT, MA, ME, NH, RI, VT. Rocks, ledges, roadsides, dry soils, lawns, often near dwellings. 2. Sedum album L. E white stonecrop. Oreosedum album (L.) Grulich • ME, VT. Ledges and rocks, often near dwellings. 3. Sedum hispanicum L. E Spanish stonecrop. MA, ME, VT. Gardens, fields, rock outcrops. 4. Sedum ochroleucum Chaix E European stonecrop. Sedum anopetalum DC. • ME. Roadsides. Early reports (e.g., Chamberlain 1912) of Sedum anophyllum DC. refer to this species (based on the description provided). Kartesz (1999) erroneously reported this species for MA. 5. Sedum reflexum L. E rock stonecrop. Petrosedum reflexum (L.) Grulich; Sedum rupestre L. • MA; also reported from ME by Kartesz (1999), but specimens are unknown. Roadsides, waste areas. 6. Sedum sarmentosum Bunge E Fig. 599 stringy stonecrop. CT, MA, ME, NH, RI, VT. Gardens, roadsides, forest fragments, waste areas, dumps. 7. Sedum sexangulare L. E six-angled stonecrop. CT, MA, NH, VT. Gardens, cemeteries, roadsides. 8. Sedum ternatum Michx. E woodland stonecrop. Clausenellia ternata (Michx.) A. & D. Löve • CT, MA, ME, VT. Roadsides, yards, forest fragments, about dwellings. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI and was unaware of any collections.

Sempervivum 1. Sempervivum tectorum L. E hens-and-chickens. CT, MA, ME, VT. Rocks, ledges, dry lawns, usually near dwellings.

Cucurbitaceae 1a. Ovary and fruit evidently prickly with narrow prickles [Figs. 601, 602]; petals shorter than 1 cm; fruit 1.3–6 cm long 2a. Petals yellow; fruit fleshy, with many seeds; leaf blades with deep, rounded sinuses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cucumis 2b. Petals white to green-white or green; fruit not fleshy, with 1 or 4 seeds; leaf blades with shallow sinuses or with moderately deep, V-shaped sinuses 3a. Fruit inflated, 4-seeded, 3–5 cm long, solitary [Fig. 601], dehiscent by 2 apical pores; staminate corolla 6-merous; herbage glabrous or nearly so . . . . . . . . . . . . . . Echinocystis

Fig. 599  Inflorescence and leaves of Sedum sarmentosum.

5 44   tricolpate s

3b. Fruit not inflated, 1-seeded, 1.3–1.5 cm long, in clusters of 3–10 [Fig. 602], indehiscent; staminate corolla 5-merous; herbage conspicuously pubescent . . . . Sicyos 1b. Ovary and fruit smooth, verrucose, or with small prickles in longitudinal stripes; petals 1 cm long or longer; fruit 4–60 (–90) cm long 4a. Staminate pedicels with a reniform to orbicular, foliaceous bract; fruits bluntly verrucose, splitting at maturity to reveal seeds with bright red arils . . . . . . . . . . Momordica 4b. Staminate pedicels ebracteate; fruits smooth or with small prickles in longitudinal strips, not dehiscent, the seeds without red arils 5a. Petals white; corolla rotate; fruit woody when mature . . . . . . . . . . . . . . . . . . Lagenaria 5b. Petals yellow to orange-yellow; corolla campanulate, open-campanulate, or rotate; fruit fleshy when mature 6a. Leaf blades pinnately lobed, the segments ± round at the apex; fruit exocarp green with irregular white or green-white stripes, the mesocarp red . . . . . . . Citrullus 6b. Leaf blades simple or palmately lobed and then with angular or pointed segments; fruit exocarp green, blue, or orange, the mesocarp white, yellow, orange, tan, or green 7a. Petals 5–10 cm long; anthers connivent; tendrils branched one or more times . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cucurbita 7b. Petals 1–3 cm long; anthers separate; tendrils simple 8a. Corolla campanulate, the free portion of the petals connate ca. ½ their length; ovary and fruit short-pubescent; leaf blade irregularly dentate, but not lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thladiantha 8b. Corolla open-campanulate, the free portion of the petals connate less than ½ their length; ovary and fruit spiny or roughened in longitudinal bands, but not pubescent (except in C. melo); leaf blades prominently to obscurely lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Cucumis

Citrullus 1. Citrullus lanatus (Thunb.) Matsumura & Nakai E watermelon. Citrullus citrullus (L.) Karst.; C. vulgaris Schrad.; Cucubertia citrullus L.; Momordica lanata Thunb. • CT, MA, ME, RI, VT. Gardens, fields, dumps, pond shores.

Cucumis 1a. Leaf blades prominently lobed, with deep, rounded sinuses; petals 4–8 mm long 2a. Pepo obloid to ellipsoidal, 3–6 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. anguria 2b. Pepo ± globose, 2–2.5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. myriocarpus 1b. Leaf blades simple or lobed with shallow, often angular, sinuses; petals 20–30 mm long 3a. Fruit usually cylindrical, the exocarp prickly with sharp elevations, the mesocarp not sweet, ± white; stems rough-pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sativus 3b. Fruit usually globose to subglobose, the exocarp without prickly elevations, though sometimes rough or warty, usually pubescent initially (often becoming glabrous), the mesocarp sweet tasting and orange, yellow, or green; stems soft-pubescent . . . . . . C. melo 1. Cucumis anguria L. E West India gherkin. MA. Yards, waste areas.

Cu c u r b i tac e a e   5 45

2. Cucumis melo L. E cantaloupe. CT, MA, NH, RI. Gardens, dumps, waste areas. 3. Cucumis myriocarpus Naud. E paddy melon. MA. Wool waste, dumps. 4. Cucumis sativus L. E cucumber. MA. Waste areas, dumps, fields.

Cucurbita 1a. Plants rough-pubescent with translucent setae; leaf blades palmately 3- to 7-lobed; peduncle strongly angled or cornered and expanded at attachment with pepo . . . . . . C. pepo 1b. Plants pilose, but not rough-pubescent; leaf blades not or only inconspicuously lobed; peduncle nearly cylindrical, not expanded at junction with fruit . . . . . . . . . . . . . . . . . C. maxima 1. Cucurbita maxima Duchesne E winter squash. MA, ME, VT. Dumps, fields, waste areas. This species includes several familiar cultivated squashes (e.g., acorn, hubbard). 2. Cucurbita pepo L. E Fig. 600 pumpkin. Pepo pepo (L.) Britt. ex Small • CT, MA, NH, VT. Fields, dumps, waste areas, river shores. This species includes several other cultivated plants, including most summer squashes (e.g., pattypan, spaghetti, summer crookneck, zucchini).

Fig. 600  Flower of Cucurbita pepo.

Echinocystis 1. Echinocystis lobata (Michx.) Torr. & Gray N Fig. 601 wild cucumber. Micrampelis lobata (Michx.) Greene; Sicyos lobata Michx. • CT, MA, ME, NH, RI, VT. Riparian forests, farmlands, roadsides and thickets in rich, mesic soil.

Lagenaria 1. Lagenaria siceraria (Molina) Standl. E bottle gourd. Cucurbita lagenaria L.; C. siceraria Molina; Lagenaria leucantha Rusby; L. vulgaris Ser. • MA. Gardens, beaches.

Fig. 601  Fruit of Echinocystis lobata.

Momordica 1. Momordica charantia L. E bitter-melon. CT. Dumps, waste areas.

Sicyos 1. Sicyos angulatus L. N Fig. 602 one-seeded burr-cucumber. CT, MA, ME, NH, RI, VT. Riparian forests, river banks, fallow fields.

Thladiantha 1. Thladiantha dubia Bunge E Manchu tuber gourd. MA, NH. Abandoned homesteads, waste areas.

Fig. 602  Fruits of Sicyos angulatus.

5 46  tricolpates

Diapensiaceae 1a. Leaves opposite, with entire blades 0.5–1.5 cm long; flowers lacking staminodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diapensia 1b. Leaves all basal, with toothed blades 2–10 cm long; flowers with 5 staminodes 2a. Flowers numerous, borne in a raceme; corolla 3–5 mm long; stamens monadelphous; anthers 1-locular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Galax 2b. Flowers solitary, at the summit of a scape [Fig. 603]; corolla 15–25 mm long; stamens distinct; anthers 2-locular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Shortia

Diapensia 1. Diapensia lapponica L. ssp. lapponica

N C Fig. 603

cushion-plant. ME, NH, VT; northern portion of states. Alpine plateaus and ridges.

Galax Fig. 603  Habit of Diapensia lapponica.

1. Galax urceolata (Poir.) Brummitt E beetleweed. Galax aphylla, auct. non L. • MA, NH. Thickets, fields, abandoned homesteads.

Shortia 1. Shortia galacifolia Torr. & Gray var. galacifolia E Oconee-bells. CT. Forest fragments, areas of habitation.

Droseraceae Drosera 1a. Leaf blades filiform, scarcely differentiated from the petiole; plants bulbous-based; petals purple, 8–15 × 5–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. filiformis 1b. Leaf blades broad-linear to tranverse-broad-elliptic, clearly differentiated from the petiole; plants not bulbous-based; petals white to pink, 3–6 × 2–4 mm 2a. Leaf blades suborbicular to orbicular [Fig. 604], sometimes even wider than long; petioles glandular-pubescent [Fig. 604]; seeds light brown, longitudinally striate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. rotundifolia 2b. Leaf blades obovate to broad-linear, definitely longer than wide; petioles glabrous (sometimes sparsely glandular-pubescent in D. anglica); seeds red-brown or black, papillose, pitted, or striate-alveolate 3a. Stipules distinct from the petiole most of their length; seeds red-brown, papillose; leaf blades (5–) 8–20 (–25) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. intermedia 3b. Stipules connate to the petiole most of their length; seeds black, striate-alveolate or pitted; leaf blades (10–) 15–50 mm long 4a. Leaf blades broad-linear, 1.5–3 mm wide; seeds 0.5–1 mm long, finely pitted, the testa close and not extending beyond the ends of the ovule . . . . . . . . . . . . D. linearis

Drose r ac e a e   5 47

4b. Leaf blades narrow-obovate to elongate-spatulate, 3–4 mm wide; seeds 1–1.5 mm long, striate-alveolate, the testa loose and extending beyond the ovule at each end . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. anglica 1. Drosera anglica Huds.

NC

English sundew. Drosera longifolia L. • ME; northern portion of state. Circumneutral fens. 2. Drosera filiformis Raf. var. filiformis N thread-leaved sundew. CT, MA, RI; coastal plain. Coastal plain pond shores, fens. 3. Drosera intermedia Hayne N spatulate-leaved sundew. CT, MA, ME, NH, RI, VT. Bogs, fens, lake shores, river shore seeps. 4. Drosera linearis Goldie

NC

slender-leaved sundew. ME; northern portion of state. Circumneutral fens, seepy river shores. 5. Drosera rotundifolia L. N Fig. 604 round-leaved sundew. Drosera rotundifolia L. var. comosa Fern. • CT, MA, ME, NH, RI, VT. Bogs, fens, lake shores, river shore seeps. Drosera rotundifolia var. comosa represents a set of plants with varying degrees of proliferation in the flowers. On a given plant, some flowers are normal, whereas others have the carpels (and sometimes other parts of the flower) modified into tiny, stipitate-glandular leaves. There is great variation regarding these features within a given population (i.e., plants range from normal to highly modified). Further, plants with proliferations are not known to be genetically different from plants without them. For these reasons, this taxon is included within the typical form of the species.

Ebenaceae Diospyros 1. Diospyros virginiana L.

nC

common persimmon. Diospyros mosieri Small; D. virginiana L. var. mosieri (Small) Sarg.; D. virginiana L. var. platycarpa Sarg.; D. virginiana L. var. pubescens (Pursh) Dippel • CT, MA; also reported from RI by George (1997), but specimens are unknown. Old pastures, sandy beaches. This tree is native to CT and introduced in MA.

Elaeagnaceae 1a. Leaves alternate; thorns commonly present; flowers with 4 stamens; plants synoecious or polygamous (i.e., some of the flowers staminate, the others bisexual) . . . . . . . . . . . . Elaeagnus 1b. Leaves opposite; thorns absent; staminate flowers with 8 stamens; plants dioecious . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Shepherdia

Elaeagnus 1a. Leaves persistent; plants flowering in late summer and fall . . . . . . . . . . . . . . . . . . E. pungens 1b. Leaves deciduous; plants flowering in spring

Fig. 604  Leaf blade of Drosera rotundifolia.

5 48   tricolpate s

2a. Leaf blades oblong-lanceolate to narrow-lanceolate, 3–8 times as long as wide; branchlets and leaf blades with silver scales only; calyx lobes yellow on the adaxial surface, about as long as the basal connate portion of the calyx; fruit yellow or silver . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. angustifolia 2b. Leaf blades elliptic to ovate-oblong, up to 3 times as long as wide; branchlets and leaf blades abaxially often with some brown scales in addition to the silver scales; calyx lobes yellow-white on the adaxial surface, about ½ as long as the basal connate portion of the calyx; fruit red with some silver scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. umbellata 1. Elaeagnus angustifolia L. E Russian-olive. CT, MA, RI, VT. Roadsides, fields, forest edges, waste areas. More work is needed to refine the range of this species in New England. Many specimens identified as Elaeagnus angustifolia in regional herbaria are in fact E. umbellata. 2. Elaeagnus pungens Thunb. E thorny-olive. MA. Roadsides, fields, forest edges. 3. Elaeagnus umbellata Thunb. var. parvifolia (Royle) Schneid. E autumn-olive. Elaeagnus parvifolia Royle • CT, MA, ME, NH, RI, VT. Roadsides, fields, forest edges, waste areas. An invasive species in our region that is rapidly increasing in abundance along open road shoulders and in fields.

Shepherdia 1. Shepherdia canadensis (L.) Nutt. N Fig. 605 Canada buffalo-berry. Elaeagnus canadensis (L.) A. Nels.; Hippophae canadensis L.; Lepargyrea canadensis (L.) Greene • ME, VT. River shore and lake shore outcrops and cliffs in regions of high-pH bedrock. Fig. 605  Flowers and expanding leaves of Shepherdia canadensis.

Elatinaceae Elatine Reference: Fernald (1941). 1a. Placentation basal-axile, therefore, the ovules/seeds all set at approximately the same level in the ovary/fruit; seeds marked with transversely elliptic, round-ended areoles, the ends of the areoles more or less meeting the other ends (i.e., not dovetailed with them), therefore, the longitudinal ridges of the seed appearing straight and distinct [Fig. 607]; leaf blades 0.7–5 mm long; perianth 2-merous; flowers with 2 stamens . . . . . . . . . . . . . . . . . . . . . . . E. minima 1b. Placentation axile, therefore, the ovules/seeds set at different levels in the ovary/fruit; seeds marked with transversely elongate, 6-sided, angular-ended areoles, the ends of the areoles alternating and dovetailed with areole ends from the adjacent rows, therefore, the longitudinal ridges of the seed broken and less conspicuous [Fig. 606]; leaf blades 2.8– 8 (–15) mm long; perianth (2–) 3-merous; flowers with (2–) 3 stamens 2a. Leaf blades obovate to broad-spatulate, usually rounded at the apex, the larger 1.5–5 mm wide; seeds borne from the lower half of the ovary’s central axis, ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. americana 2b. Leaf blades linear to lanceolate or narrow-spatulate, truncate or retuse at the apex, 0.5–3 mm wide; seeds borne throughout the length of the ovary’s central axis, divergent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. triandra

E l at i n ac e a e   5 49

1. Elatine americana (Pursh) Arn. N Fig. 606 American waterwort. Elatine triandra Schkuhr var. americana (Pursh) Fassett • CT, MA, ME, NH, RI, VT. Lake shores, river shores, including tidal reaches, typically on muddy substrate, frequently submerged in shallow water. 2. Elatine minima (Nutt.) Fisch. & C.A. Mey. N Fig. 607 small waterwort. CT, MA, ME, NH, RI, VT. Lake shores, typically on muddy substrate, frequently submerged in shallow water. 3. Elatine triandra Schkuhr E Eurasian waterwort. MA, ME. Lake shores, typically on muddy substrate, frequently submerged in shallow water. Fig. 606  Seed of Elatine americana.

Ericaceae The anthers of the Ericaceae are sometimes provided with specialized features that can aid in identification; however, the naming of these features has been confused in some manuals where they use different names interchangeably. Slender appendages that are found on the filament near or at the junction of the anther are here referred to as spurs. Similar appendages that are borne on the anthers are here referred to as awns. These names are used regardless of the orientation of the appendages. 1a. Corolla apopetalous, or absent in Corema [Figs. 618, 621] 2a. Plants lacking chlorophyll, the stems and capsules white, pink to red, or yellow to light brown; leaves reduced, scale-like, without green pigment 3a. Inflorescence a solitary flower; stems glabrous, usually white (rarely pink); sepals similar to the subtending bracts; anthers dehiscing by 2 clefts across the top; stigmas glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monotropa 3b. Inflorescence a raceme with 2–16 flowers; stems pubescent, yellow to light brown or pink to red; sepals not similar to the subtending bracts; anthers dehiscing by a single cleft; stigmas sparsely to densely villous around the margin . . . . . . . . . . . . . . . Hypopitys 2b. Plants with chlorophyll, the stems and capsules variously colored; leaves evident, though sometimes scale-like, photosynthetic 4a. Herbaceous plants with leaves all basal 5a. Flower solitary; petals widely spreading, forming a rotate corolla . . . . . . . Moneses 5b. Flowers borne in racemes; petals inwardly arching, forming a umbraculiform or subglobose corolla 6a. Flowers borne in a secund raceme, with an actinomorphic corolla; petals bituberculate at the base; style straight, exserted beyond the petals, with a 10-lobed hypogynous disk at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orthilia 6b. Flowers borne on a spirally arranged raceme, with a weakly zygomorphic corolla (actinomorphic in P. minor); petals without tubercles; style either strongly curved and exserted beyond the petals or straight and about equaling the petals, lacking a 10-lobed hypogynous disk at the base . . . . . . . . . . . . . . . . . . . . . . . . . Pyrola 4b. Woody or suffrutescent plants with alternate or opposite leaves 7a. Leaf blades linear to narrow-elliptic, ca. 1 mm wide; perianth inconspicuous; stamens, when present, 2 or 4 per flower

Fig. 607  Seed of Elatine minima.

5 50   tricolpate s

8a. Flowers on pedicels from the axils of leaves, with 3 sepals, 3 petals, usually 4 stamens, and usually 6–9 stigmas; fruit a fleshy drupe [Fig. 610]; leaf blades entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Empetrum 8b. Flowers in small, terminal clusters, with 3 or 4 sepals, 0 petals, 2 stamens (when present), and 2–5 stigmas (when present); fruit a dry drupe; leaf blades minutely denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Corema 7b. Leaf blades lanceolate, oblanceolate, elliptic-oblong, or elliptic-ovate to ovate, wider than 1 mm; perianth conspicuous; stamens 5–10 per flower 9a. Leaves opposite, sometimes verticillate, serrate; stamens 10 per flower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chimaphila 9b. Leaves alternate, entire; stamens 5–8 per flower 10a. Corolla 4-merous, pink to red; leaf blades 8–18 mm long, glaborous or sparsely pubescent; flowers with 8 stamens; fruit a red berry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Vaccinium 10b. Corolla 5-merous, white; leaf blades 20–50 mm long, densely villoustomentose on the abaxial surface; flowers with 5–7 stamens; fruit a septicidal capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Rhododendron 1b. Corolla gamopetalous [Figs. 608, 611, 619] 11a. Ovary inferior 12a. Perianth 4-merous; plants low, commonly prostrate or trailing 13a. Leaves glabrous; ovary wholly inferior; fruit a red or dark blue to black berry; plants without wintergreen odor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Vaccinium 13b. Leaves pubescent; ovary partly inferior; fruit a white berry; plants with wintergreen odor in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Gaultheria 12b. Perianth 5-merous; plants ascending (depressed in extreme conditions) 14a. Ovary 4- or 5-locular, with numerous ovules; leaves lacking resin dots and stipitate glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Vaccinium 14b. Ovary 10-locular, with 10 ovules; leaves resin-dotted or stipitate-glandular [Fig. 612] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gaylussacia 11b. Ovary superior 15a. Corolla campanulate to salverform or rotate, not constricted in the apical portion [Figs. 613, 614, 619] 16a. Corolla rotate, 10-saccate, the anthers fitting in the corolla sacs and later springing forward [Fig. 614] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Kalmia 16b. Corolla campanulate to salverform, not saccate 17a. Perianth 4-merous; fruit a 4-locular capsule or berry 18a. Leaves alternate; corolla deciduous; fruit a white berry; flowers with 4 stamens; subshrubs with prostrate, trailing stems, with wintergreen odor in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Gaultheria 18b. Leaves opposite or in whorls of 3 or 4; corolla persistent; fruit a capsule; flowers with 8 stamens; shrubs with straggling to upright stems, without wintergreen odor 19a. Leaves in whorls of 3 or 4 [Fig. 611], without basal auricles; calyx shorter than the corolla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Erica 19b. Leaves opposite, the larger with 2 basal auricles; calyx much longer than the corolla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calluna 17b. Perianth 5-merous; fruit a 2-, 3-, or 5-locular capsule

E r i c ac e a e   5 5 1

20a. Leaf blades 2–4 mm long, bristle-like [Fig. 613]; stamens with spreading or deflexed spurs; plants with moss-like aspect . . . . . . . . . . . . . . . . . . . Harrimanella 20b. Leaf blades 3–200 mm long, broader, not resembling bristles; stamens lacking spurs; plants not resembling mosses 21a. Leaf blades 3–8 mm long, opposite; corolla 3–5 mm long; fruit a capsule with 2 or 3 locules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Kalmia 21b. Leaf blades 10–200 mm long, alternate; corolla 14–45 mm long; fruit a capsule (berry-like in Epigaea) with 5 locules 22a. Calyx ebracteate; corolla rotate to campanulate or broad-funnelform [Figs. 619, 620]; anthers opening by 2 pores; fruit a septicidal capsule; upright shrubs (depressed in the alpine R. lapponicum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Rhododendron 22b. Calyx subtended by 2 bracts; corolla ± salverform; anthers opening by longitudinal slits; fruit a berry-like capsule; prostrate and trailing shrubs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epigaea 15b. Corolla globose, ovoid, urceolate, or tubular, slightly to strongly constricted in the apical portion [Figs. 608, 611, 616] 23a. Leaves in whorls of 3 or 4; perianth 4-merous . . . . . . . . . . . . . . . . . . . (in part) Erica 23b. Leaves alternate; perianth 5-merous 24a. Non-photosynthetic plants with pink to red or red-brown (eventually brown), stipitate-glandular stems bearing scale-like leaves and a terminal raceme of flowers [Fig. 616] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterospora 24b. Photosynthetic plants with green leaves, with or, more commonly, without stipitate glands, bearing various types of inflorescence 25a. Calyx closely subtended by 2 bracteoles (i.e., the bracteoles overlapping the calyx much of their length); inflorescence a solitary flower from the axils of normal foliage leaves or from the axils of reduced, bract-like leaves 26a. Calyx and abaxial surface of leaf blades lepidote-scaly; plants 30– 150 cm tall; stamens without awns; plants without wintergreen odor; fruit a loculicidal capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chamaedaphne 26b. Calyx and leaf blades without lepidote scales; plants up to 20 cm tall; each pollen sac of an anther with 2 awns; plants with wintergreen odor in life; fruit a red, berry-like capsule . . . . . . . . . . . . . . . . . . . . . (in part) Gaultheria 25b. Calyx not subtended by 2 bracteoles, though sometimes the associated pedicel may possess 2 bracteoles (these near the apex of the pedicel in Eubotrys); inflorescence composed of 2 or more flowers (sometimes a solitary flower in Phyllodoce) 27a. Corolla purple; leaf blades 4–10 × 1–1.3 mm; pedicels stipitate-glandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllodoce 27b. Corolla white to pink; leaf blades 10–150 × 2–50 mm; pedicels without stipitate glands 28a. Inflorescence produced from the axils of leaves, laterally from the apex of the previous season’s branch, or distributed along leafless branches; anthers essentially lacking apical awns or each with 4 awns 29a. Anthers with basal spurs; flowers borne in umbel-like clusters from the axils of leaves or these distributed along leafless branches forming a compound, raceme-like or panicle-like inflorescence; sutures of capsule prior to dehiscence a broad, pale strip ca. 0.5 mm wide [Fig. 615]; winter buds with 2 scales . . . . . . . . . . . . . . . . . . . . . . . . . . Lyonia

5 52   tricolpate s

29b. Anthers lacking spurs; flowers borne in axillary racemes or laterally from the apex of the previous seasons branch; sutures of capsule prior to dehiscence a thin strip ca. 0.1 mm wide; winter buds with 3 or more scales 30a. Leaf blades evergreen, coriaceous, lustrous; each pedicel with 2 bracteoles borne near the base . . . . . . . . . . . . . . . . . . . . . Leucothoe 30b. Leaf blades deciduous, membranaceous, dull; each pedicel with 2 bracteoles borne near the apex just below the calyx . . . . Eubotrys 28b. Inflorescence terminal; anthers each with 2 apical awns (without apical awns in Pieris) 31a. Corolla 5-saccate at base, the sacs alternating with the calyx lobes; inflorescence a panicle; leaf blades 10–25 mm wide . . . . . . Pieris 31b. Corolla not 5-saccate at base; inflorescence a few-flowered raceme or umbel-like raceme; leaf blades 2–12 (–18) mm wide 32a. Leaves pubescent on the abaxial surface, conspicuously revolute; sepals valvate in bud; fruit a loculicidal capsule . . . . . . . . Andromeda 32b. Leaves glabrous, with flat margins; sepals imbricate in bud; fruit a fleshy drupe 33a. Leaf blades entire, persistent and living throughout the season, not conspicuously veiny, with unwinged petioles; branchlets pubescent; inflorescence with 5–12 flowers; drupes red; pyrenes partly or entirely concrescent . . . . . Arctostaphylos 33b. Leaf blades serrate, marcescent, conspicuously rugosereticulate-veined, with winged petioles [Fig. 608]; branchlets glabrous; inflorescence with 2 or 3 flowers [Fig. 608]; drupes purple to purple-black; pyrenes distinct . . . . . . . . . . . . . . . . Arctous

Andromeda 1. Andromeda polifolia L. var. glaucophylla (Link) DC. N bog-rosemary. Andromeda glaucophylla Link; A. glaucophylla Link var. iodandra Fern.; A. polifolia L. ssp. glaucophylla (Link) Hultén • CT, MA, ME, NH, RI, VT. Open bogs and fens.

Arctostaphylos 1. Arctostaphylos uva-ursi (L.) Spreng. N red bearberry. Arctostaphylos adenotricha (Fern. & J.F. Macbr.) A. & D. Löve & Kapoor; A. uva-ursi (L.) Spreng. ssp. adenotricha (Fern. & J.F. Macbr.) Calder & Taylor; A. uva-ursi (L.) Spreng. var. adenotricha Fern. & J.F. Macbr.; A. uva-ursi (L.) Spreng. ssp. coactilis (Fern. & J.F. Macbr.) A. & D. Löve & Kapoor; A. uva-ursi (L.) Spreng. var. coactilis Fern. & J.F. Macbr.; Uva-ursi uva-ursi (L.) Britt. • CT, MA, ME, NH, RI, VT. Dry summits of hills and ridges, sand plains, dry, open, often sandy, clearings and rights-of-way. Some plants of Cape Cod and Nantucket Island, MA, are unusual in producing late-season flowers with red to purple corollas (in addition to the usual spring flowers with white to pink corollas).

Arctous 1. Arctous alpina (L.) Nied. Fig. 608  Flowers and leaves of Arctous alpina.

N C Fig. 608

alpine-bearberry. Arbutus alpina L.; Arctostaphylos alpina (L.) Spreng.; Mairania alpina (L.) Desv. • ME, NH; northern half of states. Alpine plateaus and ridges.

Er i c ac e a e   5 5 3

Calluna 1. Calluna vulgaris (L.) Hull E heather. CT, MA, ME, NH, RI, VT. Sandy fields, roadsides.

Chamaedaphne 1. Chamaedaphne calyculata (L.) Moench N leatherleaf. Cassandra calyculata (L.) D. Don var. angustifolia (Ait.) Seymour; C. calyculata (L.) D. Don var. latifolia (Ait.) Seymour; Chamaedaphne calyculata (L.) Moench var. angustifolia (Ait.) Rehd.; C. calyculata (L.) Moench var. latifolia (Ait.) Fern. • CT, MA, ME, NH, RI, VT. Bogs, swamps, mixed graminoid-shrub marshes, low floodplains of streams, pond shores.

Chimaphila 1a. Leaf blades ± oblanceolate, acute to obtuse at the apex, tapering to the base, not marked with white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. umbellata 1b. Leaf blades ± lanceolate, acute to acuminate at the apex, tapering to rounded at the base, marked with white along the midvein and primary veins [Fig. 609] . . . . . . . . . . . . . C. maculata 1. Chimaphila maculata (L.) Pursh N Fig. 609 spotted prince’s-pine. Pyrola maculata L. • CT, MA, ME, NH, RI, VT. Dry-mesic, deciduous or evergreen-deciduous woodlands and forests, infrequently on rich, mesic soils (particularly in the northern portion of its range). 2. Chimaphila umbellata (L.) W. Bart. ssp. cisatlantica (Blake) Hultén N noble prince’s-pine. Chimaphila corymbosa Pursh; C. umbellata (L.) W. Bart var. cisatlantica Blake • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic forests and shaded banks.

Corema

Fig. 609  Leaves and flowers of Chimaphila maculata.

1. Corema conradii (Torr.) Torr. ex Loud. N broom-crowberry. Empetrum conradii Torr. • MA, ME. Woodlands, heathlands, roadsides, and open sandy or ledgy areas along the coastal plain, often associated with Pinus rigida and various shrubs of the Ericaceae (e.g., Vaccinium angustifolium, Gaylussacia baccata). A specimen of this species stated to have been collected from the White Mountains of NH— Oakes s.n. (NY; image seen!)—is very likely a labeling error.

Empetrum Reference: Murray et al. (2009). 1a. Branchlets densely white-villous or white-tomentose [Fig. 610]; drupe purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. atropurpureum 1b. Branchlets stipitate-glandular; drupe black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. nigrum 1. Empetrum atropurpureum Fern. & Wieg. N Fig. 610 red crowberry. Empetrum eamesii Fern. & Wieg. ssp. atropurpureum (Fern. & Wieg.) D. Löve; E. nigrum L. var. atropurpureum (Fern. & Wieg.) Boivin; E. nigrum L. var. purpureum (Raf.) A. DC.; E. rubrum Vahl ex Willd. var. atropurpureum (Fern. & Wieg.) R. Good • ME, NH, VT. Open ledges and plateaus, usually at high elevation in subalpine and alpine situations. This species is frequently confused with Empetrum nigrum, especially when collections are vegetative.

Fig. 610  Fruit and branchlet of Empetrum atropurpureum showing white-tomentose internodes.

5 54   tricolpate s

2. Empetrum nigrum L. N black crowberry. Empetrum eamesii Fern. & Wieg. ssp. hermaphroditum (Lange ex Hagerup) D. Löve; E. hermaphroditum Lange ex Hagerup; E. nigrum L. ssp. hermaphroditum (Lange ex Hargerup) Böcher • ME, NH, VT. Open ledges, plateaus, headlands, and summits, along the ME coast and ascending to high elevations in the mountains of northern New England. It has long been considered that plants in New England were synoecious (i.e., with bisexual flowers) and tetraploid (ssp. hermaphroditum) as opposed to dioecious (i.e., with unisexual flowers) and diploid (ssp. nigrum, which are the type plants from Europe). It is now known that some dioecious, diploid plants are known from New England (ME; Murray et al. 2009) and some authors note that the correlation between ploidy level and reproductive biology is not perfect (Webb 1972). For these reasons, Empetrum nigrum is treated broadly (i.e., without infraspecific taxa).

Epigaea 1. Epigaea repens L. N trailing-arbutus. Epigaea repens L. var. glabrifolia Fern. • CT, MA, ME, NH, RI, VT. Forests, clearings, trail edges, usually found growing on acidic soils.

Erica Reference: Webb and Rix (1972). 1a. Corolla 2.5–3.5 (–4) mm long, cupuliform (i.e., not urceolate); anthers exserted from the corolla, lacking spurs; sepals broad-ovate; inflorescence a 2-flowered axillary raceme; petioles 0.5–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. vagans 1b. Corolla 5–9 mm long, urceolate [Fig. 611]; anthers included within the corolla [Fig. 611], with spurs; sepals linear to lanceolate; inflorescence a terminal raceme; petioles 0.1–0.3 mm long 2a. Leaf blades closely pubescent with short hairs and glandular-ciliate on the margin [Fig. 611], variably revolute but some portion of the abaxial surface visible; calyx glandularciliate [Fig. 611] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. tetralix 2b. Leaf blades glabrous and eciliate, the margin strongly revolute and contiguous, concealing the entire abaxial surface; calyx eciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . E. cinerea 1. Erica cinerea L. E Scotch heath. MA. Woodland fragments, heathlands, fields. 2. Erica tetralix L. E Fig. 611 cross-leaved heath. CT, MA, ME, NH. Fields, roadsides. 3. Erica vagans L. E

Fig. 611  Flowers and leaves of Erica tetralix.

Cornish heath. MA. Fields, roadsides. The leaf blades of Erica vagans are similar to those of E. cinerea in that the margins are strongly revolute, contiguous, and completely conceal the abaxial surface.

Eubotrys 1. Eubotrys racemosa (L.) Nutt. N swamp deciduous dog-laurel. Eubotrys elongata (Small) Small; E. racemosa (L.) Nutt. var. elongata (Small) Fern.; Leucothoe elongata Small; L. racemosa (L.) Gray; L. racemosa (L.) Gray var. projecta Fern. • CT, MA, RI; mainly along the coastal plain. Swamps, shorelines, stream banks.

Gaultheria 1a. Stems erect; leaf blades 20–50 mm long, glabrous; flowers with 5 petals and superior ovary; fruit a red, berry-like capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. procumbens

E r i c ac e a e   5 5 5

1b. Stems prostrate or trailing; leaves 5–10 mm long, bristly on the abaxial surface; flowers with 4 petals and a partly inferior ovary; fruit a white berry . . . . . . . . . . . . . . . . . . . . . G. hispidula 1. Gaultheria hispidula (L.) Muhl. ex Bigelow N creeping spicy-wintergreen. Chiogenes hispidula (L.) Torr. & Gray; Vaccinium hispidulum L. • CT, MA, ME, NH, RI, VT. Evergreen and mixed evergreen-deciduous forests in north-temperate

and boreal areas, evergreen swamps dominated by Chamaecyparis thyoides. 2. Gaultheria procumbens L. N eastern spicy-wintergreen. CT, MA, ME, NH, RI, VT; nearly throughout. Dry-mesic to mesic forests, woodlands, clearings, and fields.

Gaylussacia Reference: Sorrie and Weakley (2007). 1a. Leaf blades, bracts, sepals, and pedicels with stipitate glands [Fig. 612]; plants 2–5 dm tall; fruit stipitate-glandular; leaf blades lustrous on the adaxial surface . . . . . . . . . . G. bigeloviana 1b. Leaf blades, bracts, sepals, and pedicels with resin dots; plants 3–20 dm tall; fruit glabrous; leaf blades relatively dull on the adaxial surface 2a. Leaf blades with resin dots on both surfaces; inflorescence a short, compact, somewhat secund raceme usually not exceeding the leaves; pedicels commonly shorter than 7 mm; berry black, not glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. baccata 2b. Leaf blades with resin dots on only the abaxial surface; inflorescence a lax, open raceme commonly exceeding the leaves; pedicels 10–25 mm long; berry dark blue, thinly glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. frondosa 1. Gaylussacia baccata (Wangenh.) K. Koch N black huckleberry. Andromeda baccata Wangenh.; Decachaena baccata (Wangenh.) Small • CT, MA, ME, NH, RI, VT. Dry-mesic woodlands, forests, and fields. Plants with blue, glaucous

fruits and plants with white to pink fruits are known. 2. Gaylussacia bigeloviana (Fern.) Sorrie & Weakley N Fig. 612 dwarf huckleberry. Gaylussacia dumosa (Andr.) Torr. & Gray var. bigeloviana Fern.; Lasiococcus dumosus (Andr.) Small var. bigelovianus (Fern.) Fern. • CT, MA, ME, NH, RI. Bogs, acidic fens, heathlands 3. Gaylussacia frondosa (L.) Torr. & Gray ex Torr. N blue huckleberry. Decachaena frondosa (L.) Torr. & Gray; Vaccinium frondosum L. • CT, MA, NH, RI. Dry-mesic woodlands, forests, and clearings, sometimes along swamp edges.

Fig. 612  Flowers and leaves of Gaylussacia bigeloviana showing stipitate-glandular surfaces.

Harrimanella 1. Harrimanella hypnoides (L.) Coville

N C Fig. 613

moss-plant. Cassiope hypnoides (L.) D. Don • ME, NH. Gullies, ravines, and open slopes at high elevation.

Hypopitys Stem color, a critical character for identification, is lost in drying. Therefore, it is very important to note on herbarium labels (less than 1% of collections in regional herbaria note stem color in life). 1a. Stems pink to red in life; plants flowering during the latter half of August and during September; stigma usually densely retrorsely pubescent . . . . . . . . . . . . . . . . . . . . . H. lanuginosa

Fig. 613  Flower and leaves of Harrimanella hypnoides.

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1b. Stems light brown to yellow in life; plants flowering during June, July, and the early half of August; stigma usually sparsely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. monotropa 1. Hypopitys lanuginosa (Michx). Nutt. N hairy pine-sap. Monotropa lanuginosa Michx. • CT, MA, RI. Forests, woodlands, frequently associated with Quercus. 2. Hypopitys monotropa Crantz N yellow pine-sap. Hypopitys americana (DC.) Small; Monotropa hypopithys L. • CT, MA, ME, NH, RI, VT; nearly throughout. Forests, woodlands, frequently associated with Pinus and/or Quercus.

Kalmia Reference: Liu et al. (2009). 1a. Corolla 3–5 mm long, campanulate, lacking sacs, connate ca. ½ its length; leaf blades 0.3–0.8 cm long; androecium with 5 stamens; capsules 2- or 3-locular; dwarf, mat-forming shrubs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. procumbens 1b. Corolla 7.5–25 mm long, rotate, 10-saccate, the anthers fitting in the corolla sacs and later springing forward, connate most of its length [Fig. 614]; leaf blades 1.5–12 cm long; androecium with 10 stamens; capsules 5-locular; upright shrubs, sometimes colonial and forming thickets, but not mat-forming 2a. Leaves whorled, 3 at each node [Fig. 614]; inflorescence lateral [Fig. 614]; corolla 6–12 mm wide; capsules 3–5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. angustifolia 2b. Leaves opposite or alternate; inflorescence terminal; corolla 10–25 mm wide; capsules 6–8 mm wide 3a. Leaves mostly alternate, the blades 5–10 cm long, lanceolate to elliptic, borne on petioles 1–2 cm long, glabrous; corolla 20–25 mm wide . . . . . . . . . . . . . . . . . . . . K. latifolia 3b. Leaves opposite, the blades 1–4 cm long, linear to lanceolate, subsessile, pubescent on the abaxial surface; corolla 10–16 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. polifolia 1. Kalmia angustifolia L. ssp. angustifolia N Fig. 614 sheep American-laurel. CT, MA, ME, NH, RI, VT; throughout. Dry-mesic to wet-mesic forests and woodlands on acidic soils, bogs, lake shores, high-elevation slopes and ridges on folist soils. 2. Kalmia latifolia L. N mountain American-laurel. CT, MA, ME, NH, RI, VT; mainly in the southern half of the northern New England states. Forests, woodlands, field edges, evergreen swamps dominated by Chamaecyparis thyoides. Fig. 614  Flowers and leaves of Kalmia angustifolia.

3. Kalmia polifolia Wangenh. N bog American-laurel. CT, MA, ME, NH, RI, VT. Bogs, high-elevation slopes and ridges on folist soils. 4. Kalmia procumbens (L.) Gift, Kron, & P.F. Stevens ex Galasso, Banfi, & F. Conti

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alpine-azalea. Azalea procumbens L.; Chamaecistus procumbens (L.) Kuntze; Loiseleuria procumbens (L.) Desv. • ME, NH. Alpine ridges, plateaus, and gullies. This species was first combined under Kalmia without reference to the basionym in 2002 (as Kalmia procumbens (L.) Gift, Kron, & P.F. Stevens). It was later corrected by Gift and Kron in 2008 (as K. procumbens (L.) Gift & Kron), but only after this error was already remedied by Galasso et al. in 2005 (as K. procumbens (L.) Gift, Kron, & P.F. Stevens ex Galasso, Banfi, & F. Conti). Therefore, astute students may notice three different naming authorities ascribed to K. procumbens in various works.

Leucothoe 1. Leucothoe fontanesiana (Steud.) Sleumer E highland dog-laurel. Leucothoe axillaris (Lam.) D. Don var. editorum (Fern. & Schub.) Ahles; L. editorum Fern. & Schub. • MA. Roadsides, forest edges.

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Lyonia 1a. Corolla cylindric-ovoid, 8–13 mm long; sepals 3–9.5 mm long; leaf blades entire; inflorescence an umbel-like fascicle; capsules ovoid to ovoid-pyramidal, 4–6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. mariana 1b. Corolla globose-urceolate to ovoid-urceolate, 3–5 mm long; sepals 0.5–1.5 mm long; leaf blades often minutely serrulate; inflorescence a panicle; capsules globose to subglobse, 2–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. ligustrina 1. Lyonia ligustrina (L.) DC. var. ligustrina N Fig. 615 maleberry. Arsenococcus ligustrinus (L.) Small; Xolisma ligustrina (L.) Britt. • CT, MA, ME, NH, RI, VT. Mesic to hydric, deciduous to mixed evergreen-deciduous forests, edges of swamps, often associated with acidic soils. 2. Lyonia mariana (L.) D. Don

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piedmont staggerbush. Neopieris mariana (L.) Britt.; Xolisma mariana (L.) Rehd. • CT, RI. Deciduous to mixed evergreen-deciduous woodlands.

Fig. 615  Capsules with a thick, pale stripe on sutures of Lyonia ligustrina.

Moneses 1. Moneses uniflora (L.) Gray N one-flowered-shinleaf. Pyrola uniflora L. • CT, MA, ME, NH, RI, VT. Dry-mesic to hydric, deciduous to evergreen forests and swamps.

Monotropa 1. Monotropa uniflora L. N one-flowered Indian-pipe. CT, MA, ME, NH, RI, VT. Deciduous to mixed evergreen-deciduous forests.

Orthilia See Freudenstein (1999) for segregation of this species from Pyrola. 1. Orthilia secunda (L.) House N one-sided-shinleaf. Orthilia secunda (L.) House ssp. obtusata (Turcz.) Böcher; O. secunda (L.) House var. obtusata (Turcz.) House; Pyrola secunda L.; P. secunda L. ssp. obtusata (Turcz.) Hultén; P. secunda L. var. obtusata Turcz.; Ramischia secunda (L.) Garcke • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic, deciduous to evergreen forests, evergreen swamps dominated by Thuja occidentalis.

Phyllodoce 1. Phyllodoce caerulea (L.) Bab.

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purple mountain-heath. Andromeda caerulea L.; Bryanthus taxifolius Gray • ME, NH. Alpine gullies, ravines, and snow bank communities. Reports of this species in VT are considered erroneous (Zika 1992).

Pieris 1. Pieris floribunda (Pursh) Benth. & Hook. f. E mountain fetterbush. Andromeda floribunda Pursh • VT. Fields, roadsides.

Pterospora 1. Pterospora andromedea Nutt. pine-drops. NH, VT. Deciduous to mixed evergreen-deciduous forests.

N C Fig. 616

Fig. 616  Inflorescence and upper stem of Pterospora andromedea.

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Pyrola Reference: Freeman (2009). 1a. Style short and straight [Fig. 618], 0.5–1.5 mm long; corollas actinomorphic; stigmas broadly peltate, 5-lobed, without a subterminal collar; anthers connivent around the style, 0.8–1.4 mm long; petals 3–5 mm long; scape 5–15 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . P. minor 1b. Style elongate and strongly down-curved [Fig. 617], (4–) 6–10 mm long; corollas weakly zygomorphic; stigmas capitate, with a subterminal collar; anthers not connivent around the style, (1.7–) 2–4.1 mm long; petals 4–10 mm long; scape 10–30 cm tall 2a. Leaf blades 10–30 (–40) mm long, usually shorter than the petiole; anther tubes (0.6–) 0.7–1.1 mm long; petals green-white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. chlorantha 2b. Leaf blades 25–70 mm long, ± equaling to exceeding the length of the petiole; anther tubes 0.1–0.4 (–0.6) mm long; petals white or pink to pale purple 3a. Leaf blades elliptic or oblong to obovate, usually longer than the petiole; sepals as wide as or wider than long, ca. 25% as long as the petals; bracts below the flowers numbering 0–2 (–3), narrow-lanceolate, not clasping the scape . . . . . . . . . . . . P. elliptica 3b. Leaf blades broad-elliptic to reniform, usually ± equaling the length of the petiole; sepals longer than wide, ca. 35–60% as long as the petals; bracts below the flowers numbering 1–5, broad-lanceolate, clasping the scape 4a. Petals white; sepals oblong or ovate-oblong, not overlapping at the base, firm, 3- to 5-nerved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. americana 4b. Petals pink to pale purple; sepals triangular, overlapping at the base [Fig. 617], thin, nerveless or obscurely nerved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. asarifolia 1. Pyrola americana Sweet N American shinleaf. Pyrola asarifolia Michx. ssp. americana (Sweet) Krísa; P. rotundifolia L. ssp. americana (Sweet) Clausen; P. rotundifolia L. var. americana (Sweet) Fern. • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic forests. 2. Pyrola asarifolia Michx. ssp. asarifolia N Fig. 617 Fig. 617  Flower of Pyrola asarifolia.

pink shinleaf. Pyrola asarifolia Michx. var. incarnata (DC.) Fern.; P. asarifolia Michx. var. ovata Farw.; P. asarifolia Michx. var. purpurea (Bunge) Fern.; P. rotundifolia L. ssp. asarifolia (Michx.) A. & D. Löve; P. uliginosa Torr. & Gray ex Torr. • MA, ME, NH, VT; also reported from RI by Kartesz (1999), but specimens are unknown. Evergreen swamps dominated by Thuja occidentalis, riparian terraces, mesic forests. 3. Pyrola chlorantha Sw. N green-flowered shinleaf. Pyrola chlorantha Sw. var. convoluta (W. Bart.) Fern.; P. chlorantha Sw. var. paucifolia Fern.; P. convoluta W. Bart.; P. virens Schreb.; P. virens Schreb. var. convoluta (W. Bart.) Fern. • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic forests and woodlands. 4. Pyrola elliptica Nutt. N elliptic-leaved shinleaf. CT, MA, ME, NH, RI, VT; nearly throughout. Dry-mesic to mesic, deciduous to evergreen forests. 5. Pyrola minor L. N Fig. 618 little shinleaf. Braxilia minor (L.) House; Erxlebenia minor (L.) Rydb.; Pyrola minor L. var. parviflora Boivin • ME, NH, VT; mainly in the northern half of the states. Boreal to subalpine forests and evergreen swamps, often ascending to considerable elevation.

Rhododendron Fig. 618  Flowers of Pyrola minor.

Reference: Judd and Kron (2009). 1a. Corolla apopetalous, white; capsules opening from base to apex; leaf blades densely white-tomentose on the abaxial surface, turning rusty-tomentose . . . . . . . . R. groenlandicum

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1b. Corolla gamopetalous, white to purple, often spotted or striped [Figs. 619, 620]; capsules opening from apex to base; leaf blades variously scaly, scurfy, or pubescent on the abaxial surface, but not white- or rusty-tomentose 2a. Leaf blades lepidote, especially on the abaxial surface; ovary lepidote 3a. Leaf blades (6–) 10–15 (–20) mm long; corolla 10–20 mm wide; capsules 4–8 mm long; plants depressed shrubs 1–3 dm tall. . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. lapponicum 3b. Leaf blades (40–) 50–80 (–120) mm long; corolla 30–40 mm wide; capsules 8–10 mm long; plants upright shrubs to 20 dm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. carolinianum 2b. Leaf blades not lepidote; ovary glabrous or pubescent with eglandular and/or glandular hairs, but not lepidote 4a. Leaves coriaceous, persistent and living throughout the year, the blade (8–) 10– 20 cm long, eciliate; sepals 2–6 mm long (0.5–1.7 mm long in R. catawbiense) 5a. Leaf blades cuneate at base, acute at apex, mostly 3–5 times as long as wide; sepals 2–6 mm long; corolla usually largely white or pale pink . . . . . . . . . R. maximum 5b. Leaf blades rounded at base (infrequently cuneate or subcordate), obtuse at the apex, mostly 1.5–2.5 times as long as wide; sepals 0.5–1.7 mm long; corolla usually largely pink to purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. catawbiense 4b. Leaves membranaceous, deciduous, the blade 2–10 (–12) cm long, ciliate along the margin (sometimes eciliate in R. vaseyi); sepals 0.1–1.1 mm long 6a. Corolla strongly zygomorphic, divided into 3 portions—the upper portion 3-lobed, the 2 lower portions narrow and unlobed [Fig. 619]; flowers with 10 stamens; styles 1.5–2 cm long; capsules puberulent, sparsely, if at all, setose . . . . . . . . . R. canadense 6b. Corolla weakly zygomorphic [Fig. 620]; flowers with 5–7 stamens; styles longer than 2 cm; capsules glabrous, evidently strigose-setose, or stipitate-glandular 7a. Corolla yellow to orange-red to dark red, often with a prominent yellow blotch on the uppermost petal 8a. Corolla with a funnel-shaped basal, connate portion, eglandular on the abaxial (i.e., outside) surface, the yellow blotch divided into smaller dots by veins; branchlets glabrous (infrequently sparsely pubescent) . . . . R. japonicum 8b. Corolla with a nearly cylindrical basal, connate portion, stipitate-glandular on the abaxial surface, the yellow blotch (when present) not divided into smaller dots; branchlets pubescent . . . . . . . . . . . . . . . . . . . . . . R. calendulaceum 7b. Corolla white to pink, unmarked or with red, brown, or green spots in R. vaseyi 9a. Basal corolla tube 2–5 mm long, up to 25% as long as the lobes, glabrous on the abaxial (i.e., outside) surface; flowers with (5–) 7 stamens; fruit glabrous or with a few stipitate glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. vaseyi 9b. Basal corolla tube 13–30 mm long, as long as or longer than the lobes, pubescent or stipitate-glandular on the abaxial surface; flowers with 5 stamens; fruit setose or stipitate-glandular or both 10a. Flowers appearing after the expansion of the leaves, present June through July; corolla white (rarely pink); mature leaf blades (1.5–) 2–6 cm long, glabrous on the abaxial surface except for appressed strigae along the midrib and sometimes also the principal veins; pedicels stipitate-glandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. viscosum 10b. Flowers appearing before or with the expansion of the leaves, present May through June; corolla light to bright pink (rarely white); mature leaf blades (3–) 5–8 (–10) cm long, either softly pubescent across the surface or with strigae confined the major veins; pedicels with or without stipitate glands

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11a. Ovary, pedicels, and capsules stipitate-glandular [Fig. 620]; outer scales of vegetative winter buds pubescent on the abaxial surface, especially near the midvein; leaf blades permanently soft-pubescent on the abaxial surface, often also with strigae along the midrib . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. prinophyllum 11b. Ovary, pedicels, and capsules strigose with eglandular hairs; outer scales of vegetative winter buds ± glabrous (though ciliate on the margin); leaf blades glabrous on the abaxial surface except for appressed strigae along the midrib and sometimes also the principal veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. periclymenoides 1. Rhododendron calendulaceum (Michx.) Torr. E flame azalea. Azalea calendulacea Michx. • CT. Roadsides, areas of habitation. 2. Rhododendron canadense (L.) Torr. N Fig. 619 rhodora. Azalea canadensis (L.) Kuntze; Rhodora canadensis L. • CT, MA, ME, NH, RI, VT. Bogs, acidic fens, peaty shorelines, swamps, low roadsides, pastures. 3. Rhododendron carolinianum Rehd. E Carolina rhododendron. CT, MA. Roadsides, forest borders and fragments, areas of habitation. Fig. 619  Flower of Rhododendron canadense.

4. Rhododendron catawbiense Michx. E Catawba rosebay. MA. Roadsides, forest borders and fragments, areas of habitation. 5. Rhododendron groenlandicum (Oeder) Kron & Judd N Labrador-tea. Ledum groenlandicum Oeder; L. palustre L. ssp. groenlandicum (Oeder) Hultén • CT, MA, ME, NH, VT. Bogs, swamps, peaty shorelines, ascending to high elevation on open slopes and ridges. 6. Rhododendron japonicum (Gray) Sur. E Japanese azalea. Azalea japonica Gray • CT. Roadsides, fields, areas of habitation. This species is similar to Rhododendron vaseyi in that the corolla has a very short, basal tube that is much shorter than the lobes. 7. Rhododendron lapponicum (L.) Wahlenb. Lapland rosebay. Azalea lapponica L. • ME, NH. Alpine ridges and plateaus.

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8. Rhododendron maximum L. N great rosebay. CT, MA, ME, NH, RI, VT; unusual distribution with several well-known disjunct populations in the northern New England states. Evergreen to deciduous swamps, lake shores, forests. 9. Rhododendron periclymenoides (Michx.) Shinners N pink azalea. Azalea nudiflora L.; Rhododendron nudiflorum (L.) Torr.; R. nudiflorum (L.) Torr. var. glandiferum (Porter) Rehd.; R. periclymenoides (Michx.) Shinners var. eglandulosum Seymour • CT, MA, NH, RI, VT. Swamps, mesic forests. Rare plants of this species show stipitate-glands on the abaxial surface of the corolla. Early reports of Rhododendron canescens (Michx.) G. Don from New England (e.g., Knowlton and Deane 1922) refer to this species (specimens at A! and NEBC!). 10. Rhododendron prinophyllum (Small) Millais N Fig. 620 early azalea. Azalea prinophylla Small; Rhododendron nudiflorum (L.) Torr. var. roseum (Loisel.) Wieg.; R. roseum (Loisel.) Rehd. • CT, MA, ME, NH, RI, VT. Woodlands, often rocky or ledgy, talus slopes, swamps, shorelines. Fig. 620  Flowers of Rhododendron prinophyllum.

11. Rhododendron vaseyi Gray E pink-shell azalea. Biltia vaseyi (Gray) Small • MA; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Forest fragments, nurseries. 12. Rhododendron viscosum (L.) Torr. N clammy azalea. Azalea viscosa L.; Rhododendron viscosum (L.) Torr. var. glaucum (Michx.) Gray • CT, MA, ME, NH, RI, VT; rare in the northern New England states. Swamps, lake shores.

E r i c ac e a e   5 6 1

Vaccinium There are herbarium specimens from various locations in New England that have been referred to as hybrids. These include, for example, collections of Vaccinium angustifolium with entire leaves (considered to be V. angustifolium × V. myrtilloides; specimens at VT!) and V. angustifolium with glaucous leaves (considered to be V. angustifolium × V. uliginosum; specimens at VT!). Some of these determinations are doubtful, others are difficult to confirm. More work, including field and herbarium research, is needed to verify the authenticity of these putative hybrids. Reference: Vander Kloet (1988). 1a. Leaf blades coriaceous, lustrous, evergreen; shrubs prostrate or trailing, up to 2 dm tall; corolla with 4 lobes [Fig. 621]; fruit usually red or dark red, tart at maturity 2a. Corolla divided to near the middle; leaf blades with erect, black glands on the abaxial surface; pedicels less than 5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. vitis‑idaea 2b. Corolla divided to below the middle [Fig. 621]; leaf blades without glands on the abaxial surface; pedicels 10–50 mm long 3a. Leaf blades elliptic-oblong, rounded at the apex, with flat or slightly revolute margins; pedicels with green, leaf-like bracteoles 2–4 × 1–2 mm [Fig. 621]; styles 5–7 mm long; berry 10–15 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. macrocarpon 3b. Leaf blades ovate to triangular, acute at the apex, with strongly revolute margins; pedicels with red, scale-like bracteoles 0.5–2 × less than 1 mm; styles 3–4 mm long; berry 6–12 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. oxycoccos 1b. Leaf blades herbaceous, not lustrous, deciduous; shrubs prostrate or upright, never trailing, 0.5–30 dm tall; corolla with 4 or, more commonly, 5 lobes [Figs. 622, 623]; fruit blue to blue-black, sweet at maturity 4a. Branchlets not papillose; anthers with a pair of upward-projecting awns in addition to the terminal tubules 5a. Corolla scutelliform to open-campanulate [Fig. 623], open in bud; stamens exserted; berry (7–) 10–14 (–18) mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. stamineum 5b. Corolla urceolate [Fig. 622], closed in bud; stamens included within the corolla; berry 5–9 mm in diameter 6a. Flowers 5-merous; winter buds with 2 scales; leaf blades finely serrulate; calyx continuous with the pedicel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. cespitosum 6b. Flowers mostly 4-merous; winter buds with more than 2 scales; leaf blades entire; calyx articulated with the pedicel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. uliginosum 4b. Branchlets papillose; anthers lacking awns 7a. Crown-forming, single- to several-stemmed shrubs 1–3 (–4) m high; mature leaf blades (30–) 38–80 mm long; corolla (4–) 5–10 mm long 8a. Leaf blades darkening in drying, moderately to densely pubescent abaxially with brown-gray to brown hairs; berry black, not glaucous; winter bud scales brown-green to black; branchlets pubescent throughout circumference; flowers precocious . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. fuscatum 8b. Leaf blades not darkening in drying, glabrous or sparsely pubescent abaxially with white to gray-white hairs; berry blue to dark blue, glaucous; winter bud scales pink to red; branchlets glabrous, pubescent in 2 lines, or infrequently pubescent throughout circumference; flowers coetaneous 9a. Leaf blades 30–50 mm long, 15–20 mm wide, eciliate, glaucous abaxially, entire; corolla 4–6 mm long; berry 5–8 mm in diameter . . . . . . . . . . V. caesariense 9b. Leaf blades (30–) 38–80 mm long, 20–30 mm wide, ciliate, green or glaucous abaxially, entire or serrulate; corolla 5–10 mm long; berry 6–12 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. corymbosum

5 62  tricolpates

7b. Rhizomatous, colonial, many-stemmed shrubs 0.01–1 (–1.15) m tall; leaf blades 8–45 (–50) mm long; corolla 3–8 mm long 10a. Leaf blades entire or nearly so [Fig. 622] 11a. Branchlets and leaf blades evidently pubescent; winter bud scale apex acute; leaf blades not glaucous, elliptic to narrow-elliptic . . . . . . . . . . . . . . . V. myrtilloides 11b. Branchlets and leaves sparsely pubescent at most; winter bud scale apex rounded; leaf blades glaucous, ovate to broad-elliptic . . . . . . . (in part) V. pallidum 10b. Leaf blades finely serrulate 12a. Leaf blades 8–21 × 2–6 mm, narrow-elliptic (rarely elliptic); plants rarely reaching 9 cm tall; corolla 3–4 mm long; berries 3–5 mm in diameter . . . . . . . . . . . . . . V. boreale 12b. Leaf blades (10–) 15–50 × 6–22 mm, narrow-elliptic to broad-elliptic or ovate; plants 9–51 cm tall; corolla 4–8 mm long; berries (3–) 4–9 (–12) mm in diameter 13a. Leaf blades regularly serrulate, glaucous or, more commonly, not glaucous, (10–) 15–32 (–41) mm long, ca. ⅓ as wide, elliptic to, less commonly, narrowelliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. angustifolium 13b. Leaf blades often irregularly serrulate, glaucous, (15–) 30–50 mm long, ca. ½ as wide, broad-elliptic to ovate [Fig. 622] . . . . . . . . . . . (in part) V. pallidum 1. Vaccinium angustifolium Ait. N common lowbush blueberry. Vaccinium angustifolium Ait. var. hypolasium Fern.; V. angustifolium Ait. var. laevifolium House; V. angustifolium Ait. var. nigrum (Wood) Dole; V. nigrum (Wood) Britt. • CT, MA, ME, NH, RI, VT; nearly throughout. Dry-mesic to mesic fields, open slopes, woodlands, clearings, ridges, balds, headlands, and mountain summits. This species displays variation in fruit and leaf blade characteristics, and several different forms are known to occur within New England. Some of these forms, including that with glaucous leaf blades and black, non-glaucous fruits, appear very distinct in the field but are not worthy of recognition (e.g., I have observed ramets of this form growing within the same colony of plants as the typical form with non-glaucous leaf blades and blue, glaucous fruits). Forms with white fruits, narrow-leaf blades, and pubescent leaf blades are also known (the last may represent hybrids with Vaccinium myrtilloides). 1‌ × 5. Vaccinium ×atlanticum Bickn. is a rare blueberry hybrid known from MA, ME, NH. It most resembles plants of V. corymbosum, given its non-colonial habit, but typically is shorter than 1 m at maturity. It is further characterized by leaf blades (25–) 35–65 mm long and corollas (6.5–) 7–9.5 mm long (vs. leaf blades (10–) 15–32 (–41) mm long and corollas 4–6 mm long in V. angustifolium and leaf blades (30–) 38–80 mm long and corollas 5–10 mm long in V. corymbosum). Though F₁ plants do show intermediacy between the two parental taxa, later generation plants tend to become more similar to V. corymbosum (Sam Vander Kloet, personal communication). 2. Vaccinium boreale Hall & Aalders N northern blueberry. Vaccinium pensylvanicum Lam. var. angustifolium (Ait.) Gray; V. pensylvanicum Lam. var. alpinum Wood • ME, NH, VT. Found mainly at high elevation on alpine ridges and plateaus, also growing on low elevation balds and coastal peatlands in eastern ME. Reports by Seymour (1982) of Vaccinium angustifolium var. angustifolium mostly refer to this species. Vaccinium boreale most closely resembles V. angustifolium, and forms of the latter sometimes have small, narrow leaf blades that approximate the dimensions of V. boreale. However, Vaccinium boreale is a shorter, more compactly branched plant that flowers 10–20 earlier than V. angustifolium (regardless of leaf dimensions). 3. Vaccinium caesariense Mackenzie N New Jersey highbush blueberry. MA, ME, NH, RI; mainly along the coastal plain. Swamps, shorelines, bogs, fens. 4. Vaccinium cespitosum Michx. N dwarf blueberry. Vaccinium arbuscula (Gray) Merriam; V. cespitosum Michx. var. arbuscula Gray • ME, NH, VT. Alpine and subalpine ridges, plateaus, and summits, river shore outcrops, infrequently in fields.

E r i c ac e a e   5 63

5. Vaccinium corymbosum L. N highbush blueberry. Cyanococcus corymbosus (L.) Rydb.; Vaccinium corymbosum L. var. albiflorum (Hook.) Fern.; V. corymbosum L. var. glabrum Gray • CT, MA, ME, NH, RI, VT. Swamps, fens, bogs, shorelines, locally found in upland forests and woodlands. Vaccinium corybosum is a variable taxon, which has undoubtedly led some researchers to consider most of the highbush blueberries as a single species. However, field studies in various parts of the eastern United States consistently show that several morphologies can be distinguished. Therefore, additional highbush blueberry species are here recognized (V. caesariense, V. fuscatum). 6. Vaccinium fuscatum Ait. N black highbush blueberry. Cyanococcus fuscatus (Ait.) Small; Vaccinium atrococcum (Gray) Heller; V. corymbosum L. var. atrococcum Gray; V. corymbosum L. var. fuscatum (Ait.) Hook. • CT, MA, ME, NH, RI, VT. Swamps, fens, bogs, shorelines. Flowering 7–10 days earlier than other highbush blueberries (Vaccinium caesariense, V. corymbosum) when found growing in close proximity. This species has entire leaf blades. 7. Vaccinium macrocarpon Ait. N Fig. 621 large cranberry. Oxycoccus macrocarpus (Ait.) Pursh • CT, MA, ME, NH, RI, VT. Bogs, fens, marshes, peaty lake shores, abandoned borrow pits, sandy roadsides.

Fig. 621  Flower and leaves of Vaccinium macrocarpon.

8. Vaccinium myrtilloides Michx. N velvet-leaved blueberry. Cyanococcus canadensis (Kalm ex A. Rich.) Rydb.; Vaccinium angustifolium Ait. var. myrtilloides (Michx.) House; V. canadense Kalm ex A. Rich. • CT, MA, ME, NH, VT; becoming rare in southern New England. Forests, woodlands, clearings, fields, peatlands, ledges, balds, high mountain summits. 9. Vaccinium oxycoccos L. N small cranberry. Oxycoccus microcarpos Turcz. ex Rupr.; O. oxycoccos (L.) MacM.; O. palustris Pers.; O. quadripetalus Gilib.; Vaccinium microcarpos (Turcz. ex Rupr.) Schmalh.; V. oxycoccos L. var. intermedium Gray • CT, MA, ME, NH, RI, VT. Fens, bogs, marshes, peaty lake shores. 10. Vaccinium pallidum Ait. N Fig. 622 hillside blueberry. Cyanococcus pallidus (Ait.) Small; C. vacillans (Kalm ex Torr.) Rydb.; Vaccinium corymbosum L. var. pallidum (Ait.) Gray; V. vacillans Kalm ex Torr.; V. vacillans Kalm ex Torr. var. crinitum Fern. • CT, MA, ME, NH, RI, VT. Woodlands, ridges, balds, dry fields, ledges. This species is sometimes confused with forms of Vaccinium angustifolium that have glaucous leaf blades. However, those forms show regularly serrulate leaf blades that are relatively narrow (the leaf blades of V. pallidum are entire or irregularly serrulate and relatively broad). Similar to several other species of blueberry, this species is polymorphic for fruit color and bloom—glaucous, blue berries and non-glaucous, black berries are known.

Fig. 622  Flowers and leaves of Vaccinium pallidum.

11. Vaccinium stamineum L. N Fig. 623 deerberry. Polycodium neglectum Small; P. stamineum (L.) Greene; Vaccinium caesium Greene; V. neglectum (Small) Fern.; V. stamineum L. var. neglectum (Small) Deam • CT, MA, RI, VT; mainly southern New England, barely extending into southern VT. Rocky forests, woodlands, ridges, balds. 12. Vaccinium uliginosum L. N alpine blueberry. Vaccinium uliginosum L. ssp. alpinum (Bigelow) Hultén; V. uliginosum L. var. alpinum Bigelow • ME, NH, VT. Boreal, subalpine, and alpine summits, ridges, and plateaus. 13. Vaccinium vitis-idaea L. ssp. minus (Lodd.) Hultén N mountain cranberry. Vaccinium vitis-idaea L. var. minus Lodd. • CT, MA, ME, NH, VT; very rare in southern New England. Open ridges, ledges, balds, and summits at middle to high elevations, occurring at lower elevations in coastal peatlands and lake shores in boreal regions, also historically known from a dry pasture and an evergreen forest in northeastern MA.

Fig. 623  Flowers and leaves of Vaccinium stamineum.

5 64   tricolpate s

Euphorbiaceae 1a. Leaf blades palmately lobed with peltate petioles; stamens with repeatedly branched filaments; capsules covered with soft prickles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ricinus 1b. Leaf blades entire or toothed, but not lobed, with basifixed petioles; stamens with simple filaments; capsules without prickles (though sometimes pubescent) 2a. Plants with a milky latex; inflorescence a cyathium; flowers without a calyx, subtended by a cup-like involucre with glands (sometimes with petaloid appendages) at the margin, housing within a single, peduncled, carpellate flower and 1 to several staminate flowers, collectively resembling a single bisexual flower [Figs. 626, 627, 628] . . . . . . . . . . . Euphorbia 2b. Plants without latex, the sap watery; inflorescence an axillary cyme; flowers with a calyx, clearly unisexual [Figs. 624, 625] 3a. Plants with stellate hairs (the branches of the hairs basally fused in Crotonopsis to form a flat scale on some surfaces); anthers inflexed in bud 4a. Abaxial leaf surface with stellate hairs; staminate flowers with 7–14 stamens; fruit dehiscent, 3-locular; seeds with a caruncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Croton 4b. Abaxial leaf surface with stellate hairs that have the branches basally united to form a flat scale; staminate flowers with 5 stamens; fruit indehiscent, 1-locular; seeds without a caruncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crotonopsis 3b. Plants glabrous or with simple hairs; anthers erect in bud 5a. Leaves alternate; staminate calyx 4-merous; gynoecium with 3 carpels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acalypha 5b. Leaves opposite; staminate calyx 3-merous; gynoecium with 2 carpels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mercurialis

Acalypha Reference: Levin (1999). 1a. Leaf blades linear to narrow-oblong; petioles 2–14 mm long, 9–25 (–30)% as long as the leaf blade; carpellate bracts sparsely to densely beset with sessile, red glands, toothed with triangular to ovate teeth 8–28% as long as the bract . . . . . . . . . . . . . . . . . . . . . . . . . . A. gracilens 1b. Leaf blades broad-lanceolate to ovate; petioles 7–70 mm long, (23–) 33–89% as long as the leaf blade; carpellate bracts lacking sessile, red glands (though usually with stipitate glands), toothed with linear to lanceolate teeth 21–75% as long as the bract 2a. Carpellate bracts with 5–9 lobes, stipitate-glandular, without long, eglandular hairs [Fig. 624]; stems usually without long, eglandular hairs; petioles 34–89% as long as the leaf blades, the petioles of the larger leaves commonly more than ½ as long as the blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. rhomboidea 2b. Carpellate bracts with 9–15 lobes, hirsute with long, eglandular hairs, short stipitate glands may also be present; stems usually hirsute with long, eglandular hairs; petioles 23–66% as long as the leaf blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. virginica 1. Acalypha gracilens Gray N slender three-seeded-Mercury. Acalypha gracilens Gray var. delzii L. Mill.; A. gracilens Gray var. fraseri (Muell.-Arg.) Weatherby • CT, MA, NH, RI, VT; also reported from ME by Gleason (1963), but specimens are unknown. Fields, lawns, roadsides, waste areas, forest clearings. Fig. 624  Carpellate flower of Acalypha rhomboidea showing bracteole.

2. Acalypha rhomboidea Raf. N Fig. 624 common three-seeded-Mercury. Acalypha virginica L. var. rhomboidea (Raf.) Cooperrider • CT, MA, ME, NH, RI, VT. Fields, roadsides, riparian forests, river banks, gardens, waste areas.

Eu ph o r b i ac e a e   5 6 5

Levin (1999) noted that about 5% of plants he examined from North America showed hirsute pubescence on the stems, a character state normally associated with Acalypha virginica. 3. Acalypha virginica L. N Virginia three-seeded-Mercury. Acalypha digyneia Raf. • CT, MA, ME, NH, RI. Fields, woodlands, shorelines, disturbed soil. Reports of this species in VT were based on collections of Acalypha rhomboidea (specimens at NEBC! and VT!). Levin (1999) noted that about 10% of plants he examined from North America lacked the hirsute pubescence on the stem that is typical of A. virginica.

Croton Croton glandulosus L. var. septentrionalis Muell.-Arg. was reported from CT by Magee and Ahles (1999), but specimens are unknown. 1a. Each of the 3 styles divided nearly to the base into 4 or more branches; staminate flowers without petals; plants dioecious . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. texensis 1b. Each of the 3 styles dichotomously branched 2 or 3 times; staminate flowers with 5 petals; plants monoecious . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. capitatus 1. Croton capitatus Michx. var. capitatus E Fig. 625 woolly croton. MA. Fields, roadsides, waste areas. 2. Croton texensis (Klotzsch) Muell.-Arg. E Texas croton. MA. Disturbed soil, waste areas. Fig. 625  Flower of Croton capitatus.

Crotonopsis 1. Crotonopsis elliptica Willd.

NC

elliptic-leaved rushfoil. Croton willdenowii G.L. Webster • CT. Fields, sandy waste areas.

Euphorbia Chamaesyce, a distinctive group of usually ± prostrate plants, is here subsumed under the large genus Euphorbia. Though Chamaesyce has been shown to be a monophyletic group, its recognition creates an artificial and paraphyletic Euphorbia (Steinmann and Porter 2002). The phylogeny of Euphorbia is complex on a worldwide scale, and its dismantling into smaller, more homogeneous genera appears to be premature based on the present knowledge. Euphorbia agraria Bieb. was reported from MA by Kartesz (1999), but the report was erroneous. Euphorbia pubentissima Michx. was reported from NH by Pease (1964; as E. corollata var. mollis Millsp.), but the collection was misidentified and is E. corollata—Pease 36700 (NEBC!). Also, Euphorbia humistrata Engelm. ex Gray was reported from NH and VT by Magee and Ahles (1999), but specimens are unknown. Reference: Wheeler (1941). 1a. Leaves opposite throughout, the blades with inequilateral bases [Figs. 628, 629]; stipules present [Fig. 629]; involucre with 4 glands that possess petaloid appendages; plants annual, with prostrate to wide-spreading stems (upright in E. nutans) 2a. Leaf blades with entire margins [Fig. 629]; seeds 2–2.6 mm long (mostly 0.9-1 mm long in C. serpens), with ± smooth faces; glands with very small or obsolete appendages 3a. Leaf blades broad-oblong to suborbicular, 2–7 mm long; stipules united into a white, scale-like structure that is usually fringed or lobed near apex; capsules 1–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. serpens 3b. Leaf blades narrow-oblong to lanceolate-oblong [Fig. 629], 8–15 mm long; stipules distinct, usually colored similar to the stem, particularly in the basal ½ of the stipule; capsules 3–3.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. polygonifolia

5 66   tricolpate s

2b. Leaf blades with serrate or partially serrate margins; seeds 0.9–1.6 mm long, with finely wrinkled or ridged faces; glands with small, but evident, appendages 4a. Leaves and stem glabrous; faces of the seed with 2–4 prominent, transverse ridges (smooth to obscurely marked in E. serpyllifolia) 5a. Faces of the seed with 2–4 prominent, transverse ridges; each cyathium with 1–5 stamens; involucres 0.6–0.9 mm in diameter . . . . . . . . . . . . . . . . . . . . . E. glyptosperma 5b. Faces of seed smooth or obscurely roughened or pitted; each cyathium with 5–12 (–18) stamens; involucres 0.8–1.2 mm in diameter . . . . . . . . . . . . . . E. serpyllifolia 4b. Leaves (at least when young) and stems pubescent [Fig. 628]; faces of the seed merely wrinkled or with 3 or 4 prominent, transverse ridges in E. prostrata 6a. Ovary and capsule pubescent [Fig. 628] 7a. Styles 0.3–0.4 mm long, divided in the apical ⅓ to ¼; capsules appressedpuberulent; seeds obtusely angled, usually finely wrinked with inconspicuous transverse ridges on each face . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. maculata 7b. Styles 0.2–0.3 mm long, divided nearly to the base; capsules spreading-villous, at least on the angles; seeds acutely angled, with 3 or 4 prominent transverse ridges on each face . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. prostrata 6b. Ovary and capsule glabrous 8a. Stems prostrate to wide-spreading, usually branched from the base, rather uniformly spreading-hirsute with hairs 0.5–1.5 mm long . . . . . . . . . . E. vermiculata 8b. Stems ascending to suberect, usually unbranched in the lower portion, puberulent on the younger portions with incurved hairs up to 0.3 mm long, the hairs sometimes confined to a longitudinal strip . . . . . . . . . . . . . . . . . . . . . . E. nutans 1b. Leaves of the stem alternate (opposite in E. dentata and E. lathyris), the blades with equilateral bases, those subtending the cyathiescence whorled (regularly opposite in E. dentata and sometimes opposite in E. marginata); stipules absent (present in E. marginata); involucre with 1–4 glands that lack petaloid appendages or with 5 glands that possess petaloid appendages; plants annual or perennial, with ascending to erect stems 9a. Leaves of the stem all or mostly all opposite and dentate; cyathiescence irregularly clustered near the apex of the stem, without evident, umbel-like branches; involucre with a solitary gland (rarely more) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. dentata 9b. Leaves of the stem alternate and entire in most species, never both opposite and toothed; cyathiescence with umbel-like branches; involucre with 4 or 5 glands 10a. Involucre with 5 (rarely 4) glands, the glands with evident, white (rarely green), petaloid appendages (the petal appendages small in E. ipecacuanhae) [Fig. 626] 11a. Petaloid appendages 0.05–0.6 mm long, shorter than wide, usually green; stems 1.5–3 dm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. ipecacuanhae 11b. Petaloid appendages (0.5–) 1–4.4 mm long, as long as wide or longer, usually white [Fig. 626]; stems 3–10 dm tall 12a. Leaves of the cyathiescence white on the margin or entirely white, those of the stem with broad-ovate to elliptic or obovate-oblong blades that are acute at the apex; stems and involucres pubescent; plants annual, with stipules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. marginata 12b. Leaves of the cyathiescence green, those of the stem with linear to elliptic blades that are obtuse at the apex; stems and involucres glabrous or pubesenct; plants perennial, without stipules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. corollata 10b. Involucre with 4 glands, these without petaloid appendages

Eu p h o r bi ac e a e   5 67

13a. Leaves of the stem opposite; seeds 4–6 mm long . . . . . . . . . . . . . . . . . . E. lathyris 13b. Leaves of the stem alternate [Fig. 627]; seeds 1.2–2.5 mm long 14a. Leaf blades serrulate; glands of the involucre orbicular to elliptic 15a. Leaf blades blunt or retuse at the apex, cuneate at the base; casules smooth on the outer surface; seeds conspicuously aereolate-patterned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. helioscopia 15b. Leaf blades ± acute at the apex, the upper ones broad-based; capsules verrucose on the outer surface; seeds smooth or very obscurely reticulatepatterned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. platyphyllos 14b. Leaf blades entire; glands of the involucre crescent-shaped, the concave surface facing outwards, often narrowed to horn-like ends 16a. Umbel-like cyathiescence with 3–5 primary branches; plants annual; involucre 1–2 mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. peplus 16b. Umbel-like cyathiescence with usually 7–17 primary branches; plants perennial from rhizomes; involucre 2.5–3 mm tall 17a. Leaf blades of the stem 1–3 × 0.1–0.3 cm [Fig. 627]; seeds 1.5–2 mm long; plants 15–40 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. cyparissias 17b. Leaf blades of the stem 3–8 × 0.4–0.8 cm; seeds 2–2.5 mm long; plants 30–70 (–80) cm tall 18a. Leaf blades oblanceolate to obovate or obovate-elliptic, 7–15 (–22) mm wide, 3–5 (–6) times as long as wide, usually widest above the middle and tapering to the base, obtuse to emarginate at the apex, of relatively thinner texture; terminal cyathiescence with usually 8–17 primary branches; axillary cyathiescences usually numbering 8–20 . . . . . . E. esula 18b. Leaf blades linear to lanceolate, 3–10 mm wide, 6–15 times as long as wide, widest below the middle and sometimes widened and rounded at the base, acute to subacute at the apex, of relatively firmer texture; terminal cyathiescence with usually 5–9 primary branches; axillary cyathiescences usually numbering 2–12 . . . . . . . . . . . . . . . . . . . . . E. virgata 1. Euphorbia corollata L. E Fig. 626 flowering spurge. Tithymalopsis corollata (L.) Klotzsch • CT, MA, ME, NH, RI, VT. Fields, open woodlands.

Fig. 626  Cyathia of Euphorbia corollata.

2. Euphorbia cyparissias L. E Fig. 627 cypress spurge. Galarhoeus cyparissias (L.) Small ex Rydb.; Tithymalus cyparissias (L.) Hill • CT, MA, ME, NH, RI, VT. Fields, lawns, roadsides, cemeteries, waste areas.

‌2 × 4. Euphorbia ×pseudo-esula Schur is a very rare spurge hybrid in New England, known from VT. It shows leaf blade outline most similar to E. cyparissias (i.e., narrowlinear), with many blades approaching or reaching 35 mm in length and 3 mm in width. The arrangement of cyathia shows similarity to E. esula in that flowers are produced in the axils of the upper leaves (not just in a terminal cyathiescence). 3. Euphorbia dentata Michx. E toothed spurge. Anisophyllum dentatum (Michx.) Haw.; Euphorbia dentata Michx. var. linearis Engelm. ex Boiss.; Poinsettia dentata (Michx.) Klotzsch & Garcke • VT; northern portion of state. Urban and suburban streets, yards, and waste areas. 4. Euphorbia esula L. E leafy spurge. Euphorbia intercedens Podp.; Galarhoeus esula (L.) Rydb.; Tithymalus esula (L.) Hill • CT, MA, ME, NH, RI, VT. Streets, roadsides, fields, forest edges, lawns, waste areas. See Croizat (1945) for discussion of the taxonomy of this species.

Fig. 627  Cyathiescence and upper leaves of Euphorbia cyparissius.

5 68   tricolpate s

5. Euphorbia glyptosperma Engelm.

NC

rib-seeded sandmat. Chamaesyce glyptosperma (Engelm.) Small • MA, ME, NH, VT. Railroads, roadsides, fields, open, sandy areas. 6. Euphorbia helioscopia L. E sun spurge. Galarhoeus helioscopius (L.) Haw.; Tithymalus helioscopius (L.) Hill • MA, ME, NH, VT; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Roadsides, fields, waste areas. 7. Euphorbia ipecacuanhae L.

NC

ipecac spurge. Tithymalopsis ipecacuanhae (L.) Small • CT. Woodlands, open, sandy areas. 8. Euphorbia lathyris L. E gopher spurge. Galarhoeus lathyris (L.) Haw.; Tithymalus lathyris (L.) Hill • CT, MA. Roadsides, fields, waste areas. 9. Euphorbia maculata L. N Fig. 628 spotted sandmat. Chamaesyce maculata (L.) Small; C. supina (Raf.) Moldenke; Euphorbia supina Raf. • CT, MA, ME, NH, RI, VT. Roadsides, railroads, waste areas, yards. 10. Euphorbia marginata Pursh E Fig. 628  Flowers and leaves of Euphorbia maculata.

mountain snow spurge. Agaloma marginata (Pursh) A. & D. Löve; Dichrophyllum marginatum (Pursh) Klotzsch & Garcke; Lepadena marginata (Pursh) Nieuwl. • CT, ma, nh, RI, VT. Fields, roadsides, waste areas. 11. Euphorbia nutans Lag.

NC

eyebane sandmat. Chamaesyce nutans (Lag.) Small; C. preslii (Guss.) Arthur; Euphorbia preslii Guss. • CT, MA, NH, RI, VT. Fields, roadsides, railroads, waste areas. The name Euphorbia maculata has been misapplied to this species (e.g., Seymour 1982). 12. Euphorbia peplus L. E petty spurge. Galarhoeus peplus (L.) Rydb.; Tithymalus peplus (L.) Hill • CT, MA, ME, NH, VT. Gardens, lawns, waste areas, about dwellings. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI and was unaware of any collections. 13. Euphorbia platyphyllos L. E broad-leaved spurge. Galarhoeus platyphyllos (L.) Small; Tithymalus platyphyllos (L.) Hill • MA, VT. Dumps, roadsides, waste areas.

14. Euphorbia polygonifolia L. N Fig. 629 seaside sandmat. Chamaesyce polygonifolia (L.) Small • CT, MA, ME, NH, RI; coastal sites. Sea beaches, dunes, sandy areas near the Atlantic coast. 15. Euphorbia prostrata Ait. E Fig. 629  Flowers and leaves of Euphorbia polygonifolia.

prostrate sandmat. Chamaesyce prostrata (Ait.) Small; Euphorbia chamaesyce, auct. non L. • MA. Cotton waste. 16. Euphorbia serpens Kunth E matted sandmat. Anisophyllum serpens (Kunth) Klotzsch & Garcke; Chamaesyce serpens (Kunth) Small • NH. Fields, roadsides, waste areas. 17. Euphorbia serpyllifolia Pers. var. serpyllifolia E thyme-leaved sandmat. Chamaesyce serpyllifolia (Pers.) Small • NH. Pastures. 18. Euphorbia vermiculata Raf. N hairy sandmat. Chamaesyce rafinesquei (Greene) Arthur; C. vermiculata (Raf.) House • CT, MA, ME, NH, RI, VT. Roadsides, railroads, sandy fields, waste areas.

Eu p h o r bi ac e a e   5 6 9

19. Euphorbia virgata Waldst. & Kit. E slender leafy spurge. Euphorbia esula L. ssp. tommasiniana (Bertol.) Nyman; E. esula L. var. orientalis (Boiss. in DC.) Kuzmanov; E. esula L. var. uralensis (Fisch. ex Link) Dorn; E. uralensis Fisch. ex Link; E. virgata Waldst. & Kit. var. orientalis Boiss. in DC. • CT, MA. Streets, roadsides, fields, forest edges, lawns, waste areas. See Geltman (1998) for rationale of treating this species as distinct from Euphorbia esula.

Mercurialis 1. Mercurialis annua L. E annual mercury. MA, ME. Yards, roadsides, ballast, waste areas.

Ricinus 1. Ricinus communis L. E castor-bean. CT, MA, NH. Dumps, railroads, waste areas.

Fabaceae Reference: Isely (1998). 1a. Leaf blades simple or reduced to a tendril 2a. Plants armed by means of reduced, spine-tipped phyllodes; legume mostly enclosed in the persistent calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulex 2b. Vegetative portions of plants unarmed; fruit visible most of its length 3a. Leaf blade reduced to a tendril; flowers solitary (rarely paired) on slender peduncles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lathyrus 3b. Leaf blade broad, not tendril-like; flowers arranged in racemes, fascicles, or umbels 4a. Leaf blade obovate to narrow-obovate, gradually tapering toward the base into the petiole; legume spirally coiled, armed with thin prickles along the outer surface of the spiral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpiurus 4b. Leaf blades linear to lanceolate or elliptic, or elliptic to broad-cordate-ovate, either sessile or rather abruptly transitioning from blade to petiole; fruit not spirally coiled (but curved in Coronilla), without prickles 5a. Plants herbaceous, annual; fruits plump, ± terete in cross-section 6a. Stipules adnate to the stem forming conspicuous, decurrent wings [Fig. 634]; corolla 9–11 mm long; fruit a legume, very inflated, straight . . . . . . . . . Crotalaria 6b. Stipules not adnate to the stem (though connate to each other), not forming decurrent wings; corolla 4–8 mm long; fruit a schizocarp, not bladderlike, curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Coronilla 5b. Plants woody, perennial; legumes compressed 7a. Leaves without petioles, the blades elliptic to lanceolate; flowers serotinous, with a yellow corolla; legume 1.5–3 cm long . . . . . . . . . . . . . . . . . . . . . . . . . Genista 7b. Leaves with petioles, the blades broad-cordate-ovate; flowers precocious, with a deep pink corolla; legume 6–10 cm long . . . . . . . . . . . . . . . . . . . . . . . Cercis

570  tricolpate s

1b. Leaf blades compound (sometimes the upper 1-foliate in Cytisus) 8a. Leaves pinnately compound with more than 3 leaflets [Figs. 630, 632, 649] 9a. Plants definitely woody 10a. Leaf blade surface with minute, glandular dots; corolla with 1 petal—the banner petal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amorpha 10b. Leaf blade surface without glandular dots; corolla with 5 petals (note: the 2 lower petals fused to form the keel and thus the corolla appearing as 4 petals in Caragana, Cladrastris, Colutea, Robinia, and Wisteria) 11a. Plants lianas; petals blue, blue-purple, or purple (rarely white or pink) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wisteria 11b. Plants trees or shrubs; petals largely or wholly white, yellow, or pink to red-purple 12a. Leaves bipinnately compound with strongly asymmetrical leafules, the midvein of the leafule at or near one margin [Fig. 630]; each flower with numerous stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Albizia 12b. Leaves pinnately or bipinnately compound, when bipinnately compound with ± symmetrical leafules, the midvein of the leafules aligned along the center of the blade; each flower with 5–10 stamens 13a. Perianth actinomorphic; stamens distinct, with versatile anthers; legume (6–) 8–40 (–50) cm long; leaves pinnately or bipinnately compound 14a. Leaves all bipinnate, with petioles 20–40 mm long and entire leafules; hypanthium 6–10 mm long; legume (6–) 8–15 (–25) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnocladus 14b. Leaves pinnate on spurs and bipinnate on new, terminal growth, with petioles up to 20 mm and obscurely crenate leaflets or leafules (depending on division of blade); hypanthium 3–5 mm long; legume 15–40 (–50) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gleditsia 13b. Perianth zygomorphic; stamens diadelphous (distinct in Cladrastis), with basifixed anthers; legume 4–10 cm long; leaves pinnately compound 15a. Leaflets alternately arranged along the rachis, (4–) 6–15 (–20) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cladrastris 15b. Leaflets oppositely or suboppositely arranged along the rachis [Fig. 632], 1–6 cm long 16a. Leaves paripinnate [Fig. 632]; inflorescence a fascicle with 2–4 flowers; winter bud not concealed by petiole base . . . . . . . . . Caragana 16b. Leaves imparipinnate [Fig. 649]; inflorescence a raceme with 2–20 (or more) flowers; winter bud concealed within or surrounded by base of petiole 17a. Corolla yellow; legume very inflated; calyx with subequal lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colutea 17b. Corolla white or pink to pink-purple; legume flat; calyx ± bilabiate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Robinia 9b. Plants herbaceous 18a. Some or all of the filaments distally widened; inflorescence a dense umbel (a solitary, peduncled flower in Lotus unifoliatus) [Fig. 650]

Fa bac e a e   571

19a. Inflorescence subtended by palmately lobed bracts; terminal leaflet usually considerably longer than the adjacent lateral leaflets; fruit enclosed in the calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anthyllis 19b. Inflorescence without palmately lobed bracts; terminal leaflet of ± similar length to the adjacent lateral leaflets; fruit much longer than the calyx and visible most or all of its length 20a. Corolla yellow (later becoming orange marked with red); leaf blades appearing to have 5 leaflets, the basal 2, which are actually stipules, separated from the apical 3; fruit a legume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lotus 20b. Corolla pink and/or white; leaf blades with 11–25 evenly spaced leaflets; fruit a schizocarp 21a. Schizocarp terete or angled, not flat, 20–50 mm long, weakly constricted between the seeds; calyx zygomorphic; corolla 10–15 mm long, usually pink and white; plants perennial . . . . . . . . . . . . . . . . . . . . . . Securigera 21b. Schizocarp ± flat, 10–25 mm long, strongly constricted between the seeds; calyx nearly actinomorphic; corolla 6–8 mm long, usually pink with purple lines; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithopus 18b. All of the filaments slender, not dilated near the apex; inflorescence a raceme, spike, or a solitary flower 22a. Leaves paripinnate (i.e., the terminal leaflet absent or modified into a tendril) [Figs. 640, 641, 656] 23a. Leaves with 30–60 leaflets (the upper leaves with fewer leaflets); fruit 10–20 cm long, transversely septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sesbania 23b. Leaves with 2–22 leaflets; fruit 0.6–9 cm long (5–20 (–30) cm in Vicia faba, a rare escape), not transversely septate 24a. Androecium monadelphous; anthers dimorphic—alternately obloid and globose; fruit developing underground, indehiscent, with a fibrous-reticulate pericarp; tendrils lacking . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arachis 24b. Androecium diadelphous; anthers monomorphic; fruit developing above ground, dehiscent, the pericarp not fibrous-reticulate; tendrils present (absent in Vicia faba) 25a. Stipules foliaceous, ± as large as or larger than the leaflets; style longitudinally folded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pisum 25b. Stipules small to prominent, usually smaller than the leaflets [Figs. 640, 641]; style not longitudinally folded 26a. Calyx lobes all 2–4 times as long as the basal, connate portion; legume 10–14 × 5–9 mm, with 1 or 2 seeds . . . . . . . . . . . . . . . . . . . . . Lens 26b. Calyx lobes (or at least some of them) less than 2 times as long as the basal, connate portion; legume 6–300 mm long, when less than 14 mm long only 3–5 mm wide, with 2–15 or more seeds 27a. Styles slightly widened and flattened in the apical portion, pubescent along the distal, inner surface; wing petals essentially free from the keel petals; leaves with 2–12 leaflets 15–80 mm long; stems winged in some species . . . . . . . . . . . . . . . . . . (in part) Lathyrus 27b. Styles filiform-terete, pubescent in a ring around the apex, or on the outer surface, or glabrous; wing petals cohering to the keel petals; leaves with 4–22 leaflets 5–35 mm long (40–100 mm in V. faba); stems angled or ridged, but not winged . . . . . . . . . . . . . Vicia

572  tricolpates

22b. Leaves imparipinnate (i.e., the terminal leaflet present) [Figs. 631, 649] 28a. Sepals distinct; stamens distinct or shortly connate near base [Fig. 633]; uppermost petal internal to the adjacent lateral petals in bud; petioles (or rachises in 1 species) with 1 or more prominent glands on the adaxial surface 29a. Stipules longitudinally striate; pedicels subtended by two bracteoles; stamens with well-formed anthers; legume elastically dehiscent, not transversely septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chamaecrista 29b. Stipules without longitudinal striae; pedicels ebracteate; upper 3 stamens without well-formed anthers; legume indehiscent or inertly dehiscent, with transverse septa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Senna 28b. Sepals connate; stamens monadelphous or diadelphous; uppermost petal external to the adjacent lateral petals in bud; petiole lacking prominent, adaxial glands 30a. Leaf blades with minute glandular scales or dots (at least on one surface) 31a. Wing petals tiny, much shorter than keel; legume 1-seeded, woody, prominently rugose with raised veins, the lower margin toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Onobrychis 31b. Wing petals larger, at least ½ as long as the keel; legume with 1 or more seeds, membranaceous to coriaceous, not prominently rugose, lacking teeth on the lower margin (but covered in uncinate prickles in Glycyrrhiza) 32a. Inflorescence a raceme with pale yellow flowers; wing and keel petals attached to the receptacle; stamens concealed in intact flowers; fruit 12–15 mm long, armed with uncinate prickles . . . . . . . . Glycyrrhiza 32b. Inflorescence a dense, obloid to cylindrical spike with white to blue-purple flowers; wing and keel petals attached laterally or apically to the basal, connate portion of the stamens; stamens shortly exserted and visible; fruit 2.5–3 mm long, unarmed . . . . . . . . . . . . . . . . . . . . . Dalea 30b. Leaf blades without glandular scales or dots 33a. Leaves with 5–7 leaflets; corolla purple-brown; climbing or trailing vines with tuberous rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apios 33b. Leaves with 7–31 leaflets; corolla purple or white; upright to decumbent herbs, but not trailing, without tuberous rhizomes 34a. Leaflets coarsely serrate near apex; fruit stipitate-glandular . . . Cicer 34b. Leaflets entire; fruit not stipitate-glandular 35a. Racemes terminal or opposite the leaves; corolla distinctly bicolored—the banner petal yellow to ochroleucous, the wing and keel petals pink to pale purple; style barbellate . . . . . . . . . Tephrosia 35b. Racemes axillary; corolla not bicolored, purple with varying degress of pink, red, or blue tinting, or white to ochroleucous; style not barbellate 36a. Stipules semisagittate to hastate (i.e., with 1 or 2 basal lobes); fruit 30–50 mm long, longitudinally striate . . . . . . Galega 36b. Stipules without basal lobes; fruit 5–30 (–35) mm long, without longitudinal striae 37a. Fruit a schizocarp with 2–5 mericarps; raceme secund in flower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hedysarum 37b. Fruit a legume; raceme spirally arranged in flower

Fa b ac e a e   573

38a. Keel obtusely to acutely pointed; plants with short, but evident, stems; leaflets 1.3–3.5 times as long as wide (4–5 times as long as wide in A. eucosmus) . . . . . . . . Astragalus 38b. Keel abruptly short-pointed; plants acaulescent or nearly so; leaflets 3.5–6 times as long as wide . . . Oxytropis 8b. Leaves palmately compound or with 3 or fewer leaflets [Figs. 635, 639, 653] 39a. Plants definitely woody 40a. Branchlets 4- or 5-angled; inflorescence composed of solitary or paired flowers in the axils of the upper leaves; legume pubescent only on the margin . . . . . . Cytisus 40b. Branchlets terete; inflorescence a pendulous raceme 15–25 cm long; legume pubescent over the surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Laburnum 39b. Plants herbaceous or sub-woody 41a. Leaves palmately compound with 7–17 leaflets; androecium with monadelphous filaments and dimorphic anthers—one type of anther subglobose and versatile, the other linear and basifixed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lupinus 41b. Leaves pinnately or palmately compound with 3 or fewer leaflets; androecium with distinct or diadelphous filaments (monadelphous in Hylodesmum and submonadelphous in Pueraria) and monomorphic anthers 42a. Leaflets toothed, at least minutely in the apical portion [Fig. 653] 43a. Stipules entire, 1-nerved, the distinct portion 8 or more times as long as wide; inflorescence (1–) 4–20 cm tall during anthesis, usually 4–15 times as tall as wide [Fig. 647] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melilotus 43b. Stipules entire or toothed, 2- or 3-nerved, the distinct portion commonly less than 8 times as long as wide; inflorescence 0.3–10 cm tall, rarely exceeding 3 times as tall as wide 44a. Leaflets all with petiolules of ± similar length . . . . . . . (in part) Trifolium 44b. Terminal leaflet on a petiolule distinctly longer than the petiolules of the lateral leaflets [Fig. 653] 45a. Calyx actinomorphic; fruit curved to coiled as many as 7 times (rarely straight in M. falcata) [Fig. 645] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Medicago 45b. Calyx zygomorphic; fruit straight or slightly curved 46a. Corolla marcescent; 5 or all of the filaments dilated near the apex; legume shorter than 10 mm, included or barely exserted from the persistent calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Trifolium 46b. Corolla deciduous; all the filaments slender at the apex; legume 10–16 mm long, long-exserted from the persistent calyx . . . . . Trigonella 42b. Leaflets entire or sometimes with 1–3 lobes in Pueraria and Strophostyles 47a. Leaves paripinnate, the terminal leaflet modified into a tendril . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Lathyrus 47b. Leaves imparipinnate, the terminal leaflet present 48a. Terminal leaflet much larger than the orbicular to reniform lateral leaflets (i.e., usually more than 2.5 times as long); fruit a schizocarp, prominently curved, without uncinate hairs . . . . . . . . . . . . . (in part) Coronilla 48b. Terminal leaflet not or only somewhat larger than the lateral leaflets (i.e., usually less than 1.5 times as long, sometimes much larger in a few species of Desmodium and Hylodesmum, but those species with narrower lateral leaflets); fruit a schizocarp with uncinate hairs or a legume, ± straight

574  tricolpate s

49a. Stamens distinct; corolla yellow (white to yellow-white or lightpurple to blue-purple in some Baptisia) 50a. Petioles 10–25 mm long; legume strongly compressed, 40–80 mm long; ovary sessile or borne on a stipe up to 3 mm long in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thermopsis 50b. Petioles 2–10 mm long, becoming nearly obsolete on upper leaves; legume turgid or inflated, 7–50 (–60) mm long; ovary stipitate, the stipe 5–15 mm long in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . Baptisia 49b. Stamens diadelphous, 9 stamens connate into a group with 1 separate stamen (monadelphous in Hylodesmum and submonadelphous in Pueraria); corolla pink to purple, white, or yellow-white 51a. Leaflets without stipels [Fig. 639]; ovary with 1 ovule, maturing as a 1-seeded, indehiscent fruit 52a. Petiolule of terminal leaflet clearly longer than the petiolules of the lateral leaflets (i.e., the leaves pinnately compound); stipules subulate or setaceous, not striate; calyx lobes acuminate; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lespedeza 52b. Petiolule of terminal leaflet ± the same length as the petiolules of the lateral leaflets (i.e., the leaves palmately compound); stipules narrow-ovate, striate [Fig. 639]; calyx lobes blunt; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kummerowia 51b. Leaflets usually with stipels [Fig. 638]; ovary with 2 or more ovules, maturing as a fruit with 2 or more seeds (Amphicarpaea also producing more or less apetalous flowers from near the base of the plant with 1 ovule) 53a. Fruit a schizocarp; plants erect or ascending (prostrate in Desmodium rotundifolium) 54a. Stamens monadelphous; distinct portion of the calyx (i.e., the lobes) less than 50% as long as the connate portion (i.e., the tube); lower margin of schizocarp deeply insiced such that the sinuses extend completely to the upper suture; schizocarp borne on a stipe exceeding the length of the persistent stamens, glabrous on the upper suture . . . . . . . . . . . . . . . . . . . . . Hylodesmum 54b. Stamens diadelphous; distinct portion of the calyx more than 50% as long as the connate portion; lower margin of schizocarp moderately incised such that the sinuses do not extend the entire distance to the upper suture; schizocarp borne on a stipe that is shorter than the persistent stamens, uncinatepubescent on the upper suture . . . . . . . . . . . . . . . . . . . Desmodium 53b. Fruit a legume; plants trailing, twining, or climbing (i.e., vines; with upright stems in Glycine and sometimes in Phaseolus vulgaris) 55a. Bracts of pedicels obtuse to rounded at the apex; flowers without subtending bracteoles; plants with dimorphic flowers— petaliferous flowers with 10 diadelphous stamens and maturing as a 3- or 4-seeded aerial fruit, and more or less apetalous flowers with fewer than 10 distinct stamens and maturing as a 1-seeded, often subterranean, fruit . . . . . . . . . . . . . Amphicarpaea 55b. Bracts of pedicels acute at the apex; flowers with closely subtending bracteoles that overlap the calyx (caution: the bracteoles caducous in Pueraria); plants with monomorphic, petaliferous flowers

Fa bac e a e   575

56a. Corollas 5–7 mm long; plants upright; inflorescence a raceme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glycine 56b. Corollas 8–25 mm long (as short as 3.6 mm in the very rare introduction Strophostyles leiosperma); plants vining (sometimes upright in Phaseolus vulgaris); inflorescence a pseudoraceme (i.e., with 2 or more flowers produced from some nodes) 57a. Keel petal relatively straight; style glabrous; leaflets ovate to suborbicular, 0.8–1.2 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pueraria 57b. Keel petal conspicuously arched or coiled inward; style pubescent; leaflets lanceolate or narrow-elliptic to ovate or rhombic, 1–8 (–10) times as long as wide 58a. Plants pubescent with hairs that are not hooked at the tip and are visible at low magnification; some leaflets often with a lateral lobe on one or both sides; calyx with 4 lobes due to fusion of 2 lobes; keel petals arched inward [Fig. 651]; inflorescence capitate, spike-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strophostyles 58b. Plants with minute, uncinate hairs (note: view at 20× or higher magnification); leaflets unlobed; calyx with 5 lobes; keel petals coiled inward [Fig. 648]; inflorescence elongate, raceme-like . . . . . . . . Phaseolus

Albizia 1. Albizia julibrissin Durazz. E Fig. 630 silk-tree. CT. Waste areas. Sorrie and Somers (1999) noted that planted trees of this species have persisted, but reproduction was not observed. Therefore, this species is not attributed to MA.

Amorpha 1. Amorpha fruticosa L. E false indigo-bush. Amorpha fruticosa L. var. angustifolia Pursh; A. fruticosa L. var. oblongifolia Palmer; A. fruticosa L. var. tennesseensis (Shuttlw. ex Kunze) Palmer • CT, MA, ME, NH, RI, VT. Roadsides, forest fragments, waste areas, areas of habitation.

Amphicarpaea 1. Amphicarpaea bracteata (L.) Fern. N American hog-peanut. Amphicarpaea bracteata (L.) Fern. var. comosa (L.) Fern.; A. comosa (L.) G. Don; A. monoica (L.) Ell.; A. pitcheri Torr. & Gray; Falcata comosa (L.) Kuntze; Glycine bracteata L.; G. comosa L. • CT, MA, ME, NH, RI, VT. Mesic forests, clearings, and shaded roadsides. Amphicarpaea bracteata is sometimes divided into two species—A. bracteata, the slender form with smaller leaflets and white to light purple corollas, and A. comosa, the robust form with larger leaflets and purple corollas. Turner and Fearing (1964) showed that too many intermediate forms exist to regard these as separate taxa.

Anthyllis 1. Anthyllis vulneraria L. E common kidney-vetch. CT, NH, VT; also reported from MA by Kartesz (1999), but specimens are unknown. Fields, roadsides, waste areas.

Fig. 630  Leaf of Albizia julibrissin with detail of asymmetrical leaflets.

576  tricolpates

Apios 1. Apios americana Medik. N common ground-nut. Apios americana Medik. var. turrigera Fern.; A. tuberosa Moench; Glycine apios L. • CT, MA, ME, NH, RI, VT. River banks and terraces, mesic soil along streams and field edges.

Arachis 1. Arachis hypogaea L. E peanut. MA; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Waste areas.

Astragalus 1a. Plants annual; legume usually curved to twisted, often forming a complete circle of less than 10 mm diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. contortuplicatus 1b. Plants perennial; legume straight to somewhat curved, but not twisted 2a. Corollas ochroleucous in life; legume bilocular, usually glabrous 3a. Pubescence of herbage basifixed; legume 30–40 mm long, trigonous in crosssection; principal leaves with 9–13 leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . A. glycyphyllos 3b. Pubescence of herbage mesifixed; legume 10–20 mm long, terete in cross-section; principal leaves with 13–35 leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. canadensis 2b. Corollas white to light purple, blue-purple, or purple in life; legume unilocular (note: one of the sutures may be intruded but does not connect to form 2 separate chambers in the fruit), usually pubescent 4a. Principal leaves with 15–25 leaflets [Fig. 631]; stems from a subterranean, branched caudex, mat-forming; keel petals not or only slightly shorter than the banner petal; upper suture of fruit prominently sulcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. alpinus 4b. Principal leaves with 7–15 (–17) leaflets; stems from a surficial crown of a taproot, clustered but not forming mats; keel petals markedly shorter than the banner petal; upper suture of fruit scarcely sulcate 5a. Legume (5–) 8–11 (–13) mm long, ± sessile; flowers 6–9 mm long . . . . . A. eucosmus 5b. Legume 13–25 mm long, on a stipe 2–5 mm long; flowers (7–) 10–12 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. robbinsii 1. Astragalus alpinus L. var. brunetianus Fern.

N C Fig. 631

alpine milk-vetch. Astragalus alpinus L. var. labradoricus (DC.) Fern. • ME, NH, VT. Open, gravel, cobble, and ledge river shores in regions of high-pH bedrock and/or till. 2. Astragalus canadensis L. var. canadensis

NC

Canada milk-vetch. Astragalus canadensis L. var. carolinianus (L.) M.E. Jones; A. canadensis L. var. longilobus Fassett; A. carolinianus L. • cT, vt; mainly associated with Lake Champlain. Lake beaches and headlands and mountain cliffs in regions of high-pH bedrock. 3. Astragalus contortuplicatus L. E Fig. 631  Inflorescence and leaf of Astragalus alpinus.

Hungarian milk-vetch. MA. Wool waste, waste areas. 4. Astragalus eucosmus B.L. Robins.

NC

elegant milk-vetch. Astragalus parviflorus (Pursh) MacM.; Atelophragma elegans (Hook.) Rydb. • ME; northern portion of the state. River shore ledges in high-pH bedrock and/or till regions. Known populations of this species formerly occurred on the Aroostook River in ME.

Fa b ac e a e   577

5. Astragalus glycyphyllos L. E licorice milk-vetch. MA. Waste areas. 6. Astragalus robbinsii (Oakes) Gray

NC

Robbin’s milk-vetch. 6a. Phaca robbinsii Oakes; 6b. Astragalus jesupii (Egglest. & Sheldon) Britt.; 6c. Astragalus blakei Egglest.; A. robbinsii (Oakes) Gray var. blakei (Egglest.) Barneby • ME, NH, VT. River shore beaches and ledges, cliffs, and talus in regions of high-pH bedrock and/or till. 1a. Flowers 7–9 (–10) mm long, usually with a white corolla; legume 10–15 mm long; plants historically documented from a single station on the Winooski River in vt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6a. A. robbinsii var. robbinsii 1b. Flowers 10–12 mm long, usually with a light purple corolla; legume 15–25 mm long; plants not known from the Winooski River, vt 2a. Legume tipped by a beak 1.5–3 mm long; leaflets glabrous or nearly so abaxially; plants known from the Connecticut River in NH and VT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6b. A. robbinsii var. jesupii Egglest. & Sheldon 2b. Legume scarcely beaked or with a projection up to 1.5 mm long; leaflets strigose abaxially; plants known from the mountains of northern vt and river shores of northern ME . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6c. A. robbinsii var. minor (Hook.) Barneby Variety robbinsii is known from VT. It is now extinct. Variety jesupii is known from NH, VT. Variety minor is known from ME, VT. All the infraspecific taxa are of regional conservation concern.

Baptisia 1a. Stipules subfoliaceous, those of the principal leaves 20–40 mm long; raceme usually secund, initially ascending-spreading, becoming declined, with persistent foliaceous bracts 10–30 mm long; stem villous (sometimes becoming glabrate in later season) . . . . B. bracteata 1b. Stipules small and caducous or ± persistent and 3–15 mm long; racemes not secund, ascending to erect, with caducous bracts less than 10 mm long; stem glabrous 2a. Corollas light purple to blue-purple, 25–30 mm long; pedicels 4–12 mm long; legume 30–50 (–60) mm long; leaflets 15–80 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. australis 2b. Corollas yellow, 12–16 mm long; pedicels 3–5 mm long; legume 7–15 mm long; leaflets 10–25 (–40) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. tinctoria 1. Baptisia australis (L.) R. Br. ex Ait. f. var. australis E blue wild indigo. Sophora australis L. • CT, MA, NH, VT. Riparian terraces, river banks, about dwellings. 2. Baptisia bracteata Muhl. ex Ell. var. leucophaea (Nutt.) Kartesz & Gandhi E long-bracted wild indigo. Baptisia bracteata Muhl. ex Ell. var. glabrescens (Larisey) Isely; B. leucophaea Nutt.; B. leucophaea Nutt. var. glabrescens Larisey • MA. Fields, roadsides, waste areas. 3. Baptisia tinctoria (L.) R. Br. ex Ait. f. N yellow wild indigo. Baptisia tinctoria (L.) R. Br. ex Ait. f. var. crebra Fern.; B. tinctoria (L.) R. Br. ex Ait. f. var. projecta Fern. • CT, MA, ME, NH, RI, VT; confined to southern portion of the northern states. Woodlands, dry fields and sand plains, forest openings.

Caragana 1. Caragana arborescens Lam. E Fig. 632 Siberian pea shrub. MA, ME, VT. Roadsides, forest fragments.

Fig. 632  Leaves and dehisced legumes of Caragana arborescens.

578  tricolpates

Cercis 1. Cercis canadensis L. var. canadensis

nC

redbud. CT, MA. Deciduous woodlands, rocky ridges, roadsides, forest fragments. Known historically (as a native) from a single colony on trap rock in New Haven County, CT (Nichols 1923). Massachusetts populations are non-native.

Chamaecrista 1a. Corolla 25–40 mm wide in life, with subequal petals 10–20 mm long; androecium with 10 stamens; anthers longer than 4 mm, commonly ca. 8 mm long; pedicels 8–15 mm long; petiole gland depressed to stalked, scutelliform, 0.5–1.5 mm in diameter . . . . . . . . . . . . . C. fasciculata 1b. Corolla 8–10 mm wide in life, with very unequal petals, the longer one 6–8 mm long [Fig. 633]; androecium with 5 stamens [Fig. 633]; anthers shorter than 4 mm, commonly ca. 3 mm long; pedicels 1.5–4 mm long; petiole gland stalked, umbraculiform, 0.4–0.8 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nictitans 1. Chamaecrista fasciculata (Michx.) Greene n partridge sensitive-pea. Cassia chamaecrista L.; C. fasciculata Michx. • CT, MA, NH, RI. Dry fields and clearings, sand plains, roadsides. This species is non-native in NH and native elsewhere. 2. Chamaecrista nictitans (L.) Moench ssp. nictitans var. nictitans N Fig. 633 wild sensitive-pea. Cassia nictitans L.; Chamaecrista procumbens (L.) Greene • CT, MA, NH, RI, VT. Dry fields and clearings, sand plains, roadsides. The report of this species in ME by Kartesz (1999) was in error.

Cicer Fig. 633  Flower and leaf of Chamaecrista nictitans.

1. Cicer arietinum L. E chick-pea. CT, MA, NH, VT. Fields, dumps, waste areas.

Cladrastris 1. Cladrastis kentukea (Dum.-Cours.) Rudd E Kentucky yellow-wood. Cladrastis lutea (Michx. f.) K. Koch; Sophora kentukea Dum.-Cours. • CT, MA; also reported from RI by Kartesz (1999), but specimens are unknown. Roadsides,

forest borders and fragments, areas of habitation.

Colutea 1. Colutea arborescens L. E bladder-senna. CT, MA, RI. Roadsides, railroads, forest fragments.

Coronilla 1. Coronilla scorpioides (L.) W.D.J. Koch E yellow false crown-vetch. Ornithopus scorpioides L. • CT, MA. Roadsides.

Crotalaria 1. Crotalaria sagittalis L. N Fig. 634 Fig. 634  Leaves and fruits of Crotalaria sagittalis.

arrow-head rattlebox. Crotalaria sagittalis L. var. oblonga Michx. • CT, MA, RI, VT. Sandy soil of fields, roadsides, borrow pits, and pond shores.

Fa b ac e a e   579

Cytisus 1. Cytisus scoparius (L.) Link var. scoparius E Scotch broom. Spartium scoparium L. • CT, MA, ME, NH, RI; most prevalent along the coastal plain. Roadsides, beaches, dunes, sandy wastes. C. ‌ multiflorus (ĽHér.) Sweet × C. purgans Spach. Cytisus ×praecox W.J. Bean is a very rare broom hybrid in New England known from MA. It can be separated from C. scoparius by its grooved branchlets and yellow-white to pale yellow flowers 9–11 mm long (vs. angled, but not grooved, branchlets and bright yellow flowers 20–25 mm long in C. scoparius).

Dalea 1a. Androecium with 5 stamens; primary leaves with 5–9 leaflets 8–25 mm long; corolla white, with the epistemonous petals (i.e., the wing and keel petals) arising from the apex of the connate stamen column . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. candida 1b. Androecium with 9 or 10 stamens; primary leaves with 15–35 leaflets (3–) 5–12 mm long; corolla white to blue or blue-purple, with the epistemonous petals arising laterally from the conate stamen column . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. leporina 1. Dalea candida Willd. E white prairie-clover. Petalostemum candidum Michx. • NH. Railroads, roadsides. 2. Dalea leporina (Ait.) Bullock E fox-tail prairie-clover. Dalea alopecuroides Willd.; Parosela alopecuroides (Willd.) Rydb.; P. leporina Ait. • MA. Waste areas.

Desmodium Reference: Raveill (2006). 1a. Stems prostrate; inflorescences primarily axillary; terminal leaflet mostly 0.8–1.2 times as long as wide [Fig. 638] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. rotundifolium 1b. Stems erect or ascending; inflorescences terminal or both terminal and axillary; terminal leaflet 1–10 times as long as wide 2a. Terminal leaflets (3–) 4–10 times as long as wide, mostly 5–15 mm wide, narrowlanceolate to oblong or narrow-elliptic [Fig. 637] 3a. Schizocarp with 3–6 segments that are ± triangular and abruptly curved on the lower margin [Fig. 637]; petioles mostly 15–50 mm long; leaflets herbaceous, inconspicuously veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. paniculatum 3b. Schizocarp with 1–3 segments that are ± semicircular and gradually rounded on the lower margin; petioles mostly 1–3 (–4) mm long; leaflets subcoriaceous, conspicuously reticulate-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sessilifolium 2b. Terminal leaflets 1.5–4 (–5) times as long as wide, mostly 15–30 mm wide, oblong to broad-ovate 4a. Leaf blades pubescent abaxially with uncinate hairs . . . . . . . . . . . . . . . . D. canescens 4b. Leaf blades abaxially glabrous or pubescent with hairs that are not hooked near the tip 5a. Stipules 8–20 mm long, lanceolate to ovate, persistent; terminal leaflet acuminate at the apex; schizocarp segments (8–) 9–11 mm long . . . . . . . . . . . . . . . . D. cuspidatum 5b. Stipules 2–6 mm long, linear to broad-lanceolate, generally caducous; terminal leaflet obtuse to acute at the apex; schizocarp segments 3.5–8.5 mm long

5 80   tricolpate s

6a. Schizocarp segments ± triangular, abruptly curved on the lower margin [Fig. 637] 7a. Stem pubescent with pilose hairs (a few uncinate hairs may also be present); petiole pubescent with pilose hairs . . . . . . . . . . . . . . . . . . D. perplexum 7b. Stem pubescent with uncinate hairs (pilose hairs sometimes present at the nodes) [Fig. 636]; petiole glabrous or pubescent with uncinate hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. glabellum 6b. Schizocarp segments ± semicircular, gradually rounded on the lower margin [Fig. 635] 8a. Corolla 8–11 mm long; calyx 3.5–5 mm long; stipe of fruit 2–4 mm long; schizocarp with 3–5 segments, each segment (4–) 5–7 mm long . . . . D. canadense 8b. Corolla 4–5 mm long; calyx 1.5–3 mm long; stipe of fruit 1–3 mm long; schizocarp with (1–) 2 or 3 segments, each segment 3.5–5 mm long 9a. Petioles 12–25 (–27) mm long, those of the medial leaves approximately as long as the associated lateral leaflets; stems usually glabrous; leaf blades glabrous or sparsely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . D. marilandicum 9b. Petioles 2–18 (–20) mm long, those of the medial leaves much shorter than the associated lateral leaflets; stems pubescent with pilose or uncinate hairs or both; leaf blades usually pubescent on one or both surfaces 10a. Terminal leaflet (2.5–) 4–6 (–7.5) cm long, 2.5–4 times as long as wide, distinctly longer than the lateral leaflets; stem uncinate-pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. obtusum 10b. Terminal leaflet 1.5–3 cm long, 1.5–2.3 (–5) times as long as wide, scarcely, if at all, longer than the lateral leaflets; stem pilose-pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. ciliare 1. Desmodium canadense (L.) DC. N Fig. 635 showy tick-trefoil. Hedysarum canadense L.; Meibomia canadensis (L.) Kuntze • CT, MA, ME, NH, RI, VT. Forests, river terraces and banks, rivershore meadows, fields, forest borders, roadsides. 2. Desmodium canescens (L.) DC. N Fig. 635  Schizocarp and leaf of Desmodium canadense.

hoary tick-trefoil. Hedysarum canescens L.; Meibomia canescens (L.) Kuntze • CT, MA, RI; also reported from ME by Isely (1998), but specimens are unknown. Dry-mesic forests, woodlands, roadsides, and fields. 3. Desmodium ciliare (Muhl. ex Willd.) DC. ex Loud. N hairy small-leaved tick-trefoil. Hedysarum ciliare Muhl. ex Willd.; Meibomia ciliaris (Muhl. ex Willd.) Blake • CT, MA, RI. Woodlands, forest borders, dry openings. 4. Desmodium cuspidatum (Muhl. ex Willd.) DC. ex Loud.

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large-bracted tick-trefoil. Desmodium grandiflorum DC.; D. longifolium (Torr. & Gray) Smyth; Hedysarum cuspidatum Muhl. ex Willd.; H. grandiflorum Walt.; Meibomia grandiflora (DC.) Kuntze; M. longifolia (Torr. & Gray) Vail • CT, MA, NH, RI, VT; mainly restricted to the southern portion of the northern New England states in which it occurs. Dry-mesic to mesic forests and woodlands, these usually on rocky slopes, but infrequently on river terraces, also sometimes in dry fields and open rights-of-way. 5. Desmodium glabellum (Michx.) DC.

Fig. 636  Upper stem of Desmodium glabellum with detail of uncinate hairs.

N C Fig. 636

smooth tick-trefoil. Hedysarum glabellum Michx.; Meibomia glabella (Michx.) Kuntze • CT. Woodlands, roadsides, open powerline rights-of-way. The report of this species in MA by Isely (1998) was almost certainly an error considering the manuscript is internally inconsistent (the map and text are not in agreement). The report of this species in RI by Gould et al. (1998) was based on a taxonomy that included Desmodium perplexum as a race of D. glabellum (i.e., D. perplexum does occur in RI, but D. glabellum s.s. does not).

Fa bac e a e   5 8 1

6. Desmodium marilandicum (L.) DC. N smooth small-leaved tick-trefoil. Hedysarum marilandicum L.; Meibomia marilandica (L.) Kuntze • CT, MA, NH, RI. Dry-mesic forests and woodlands, forest borders. 7. Desmodium obtusum (Muhl. ex Willd.) DC. N stiff tick-trefoil. Desmodium rigidum (Ell.) DC.; Hedysarum obtusum Muhl. ex Willd.; Meibomia obtusa (Muhl. ex Willd.) Vail; M. rigida (Ell.) Kuntze • CT, MA, NH, RI. Dry-mesic forests and woodlands, forest borders. 8. Desmodium paniculatum (L.) DC. N Fig. 637 panicled tick-trefoil. Hedysarum paniculatum L.; Meibomia paniculata (L.) Kuntze • CT, MA, ME, NH, RI, VT. Dry-mesic forests and woodlands, forest borders. Infrequent plants show a broader terminal leaflet (up to 25 mm wide) that can cause confusion with Desmodium glabellum and D. perplexum. Desmodium paniculatum usually shows glabrous to glabrate midstems, usually lacks minute uncinate hairs on the adaxial leaflet surfaces, and usually has sparse, very short (shorter than 0.5 mm), appressed hairs on the abaxial leaflet surfaces. Desmodium glabellum and D. perplexum, on the other hand have. pilose and/or uncinate-pubescent midstems, usually have minutely uncinate adaxial leaflet surfaces, and usually have more evident, slightly longer (0.5 mm long or longer), spreading to ascending hairs on the abaxial leaflet surfaces.

Fig. 637  Schizocarp and leaf of Desmodium paniculatum.

‌8 × 10. Desmodium ×humifusum (Muhl. ex Bigelow) Beck is a very rare hybrid ticktrefoil in New England known from CT, MA. This plant is similar to D. rotundifolium in ‌humifusum has its prostrate habit and relatively broad terminal leaflets. Desmodium × stipules 4–8 mm long that are deltate-ovate to subulate, not broadly clasping the stem, and usually caducous, the terminal leaflet is ovate to rhombic and mostly 1.6–1.9 times as long as wide, and the fruiting pedicels are 7–9 mm long. Desmodium rotundifolium, on the other hand, has stipules 5–12 mm long that are ovate, broadly and inequilaterally clasping the stem, and persistent, the terminal leaflet is broad-ovate to suborbicular and mostly 0.8–1.2 times as long as wide [Fig. 638], and the pedicels are (5–) 10–15 mm long. See Raveill (2002) for evidence of a hybrid origin for this nothospecies. 9. Desmodium perplexum Schub. N perplexed tick-trefoil. Desmodium dillenii Darl.; Meibomia dillenii (Darl.) Kuntze • CT, MA, ME, NH, RI, VT. Dry-mesic forests and woodlands, forest borders. 10. Desmodium rotundifolium DC. N Fig. 638 round-leaved trailing tick-trefoil. Desmodium michauxii (Vail) Daniels; Meibomia michauxii Vail; M. rotundifolia (DC.) Kuntze • CT, MA, NH, RI, VT. Dry-mesic forests and woodlands, commonly associated with rocky slopes. 11. Desmodium sessilifolium (Torr. ex M.A. Curtis) Torr. & Gray

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sessile-leaved tick-trefoil. Hedysarum sessilifolium Torr. ex M.A. Curtis; Meibomia sessilifolia (Torr. ex M.A. Curtis) Kuntze • CT, MA, RI. Dry, open, sandy areas, including roadsides and banks.

Galega 1. Galega officinalis L. E professor-weed. CT, MA, ME. Fields, roadsides, waste areas.

Genista 1. Genista tinctoria L. E dyer’s greenweed. CT, MA, ME, RI, VT. Fields, roadsides.

Fig. 638  Inflorescence and leaf of Desmodium rotundifolium borne along a prostrate stem.

5 82   tricolpate s

Gleditsia 1. Gleditsia triacanthos L. E honey-locust. Gleditsia triacanthos L. var. inermis (L.) Schneid. • CT, MA, ME, NH, RI, VT. Roadsides, forest fragments and borders, river banks. Unarmed forms are sometimes planted along town and city streets.

Glycine 1. Glycine max (L.) Merr. E soybean. Phaseolus max L. • MA, VT. Fields, roadsides, areas of cultivation.

Glycyrrhiza 1. Glycyrrhiza lepidota Pursh E American licorice. Glycyrrhiza glutinosa Nutt.; G. lepidota Pursh var. glutinosa (Nutt.) S. Wats. • CT, MA, ME, RI. Fields, dumps, waste areas, wool-waste.

Gymnocladus 1. Gymnocladus dioicus (L.) K. Koch E Kentucky coffee-tree. Guilandina dioica L. • CT, MA, ME. Roadsides, forest borders, areas of habitation.

Hedysarum 1. Hedysarum alpinum L. ssp. americanum (Michx.) Fedtsch. N alpine sweet-vetch. Hedysarum alpinum L. var. americanum Michx.; H. americanum (Michx.) Britt. • ME, VT; also reported from MA by Seymour (1982) and from RI by Isely (1998), but specimens are unknown. Ice-scoured river shores, cliffs, talus, and subalpine ravines in regions of high-pH bedrock and/or till. The reports of this species for MA and RI were almost certainly in error given that habitat is lacking in those states for this calciphile. The report from NH by Magee and Ahles (1999) is based on the following collection: White Mts., Aug, unknown s.n. (MASS!). This specimen, formerly of the Bumstead Herbarium, almost certainly represents a label mix-up.

Hylodesmum Hylodesmum is a recent segregate genus of Desmodium (Ohashi and Mill 2000). In addition to the differences in morphology (see key to the genera), this phylogenetically distinct group also differs ecologically from many Desmodium. Species of Hylodesmum are typical of shaded forests, while most species of Desmodium are usually found in open or partially open sites. 1a. Flowering stems leafless, the inflorescence arising from the ground; stipules 2–4 mm long, caducous; pedicels 10–20 mm long; stipe 10–18 mm long; terminal leaflet broad-elliptic to ovate or broad-ovate, mostly 2–6 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. nudiflorum 1b. Flowering stems with leaves, the inflorescence arising from leaf axils or the apex of the stem; stipules 9–12 mm long, semipersistent; pedicels 3–8 mm long; stipe 4–12 mm long; terminal leaflet broad-ovate to suborbicular, mostly 6–10 cm wide . . . . . . . . . . . H. glutinosum 1. Hylodesmum glutinosum (Muhl. ex Willd.) H. Ohashi & R.R. Mill N pointed-leaved tick-trefoil. Desmodium acuminatum (Michx.) DC.; D. glutinosum (Muhl. ex Willd.) Wood; Hedysarum glutinosum Muhl. ex Willd.; Meibomia acuminata (Michx.) Blake • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic forests and borders, often associated with rocky slopes, ledges, and talus.

Fa bac e a e   5 83

2. Hylodesmum nudiflorum (L.) H. Ohashi & R.R. Mill N naked tick-trefoil. Desmodium nudiflorum (L.) DC.; Hedysarum nudiflorum L.; Meibomia nudiflora (L.) Kuntze • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic forests, woodlands, and borders.

Kummerowia The report of Kummerowia stipulacea (Maxim.) Makino by Tucker (1995) for CT was in error because the specimens are K. striata. 1. Kummerowia striata (Thunb.) Schindl. E Fig. 639 Japanese-clover. Lespedeza striata Thunb. • CT, RI. Roadsides, disturbed soil, edges of small pools, sandy pond shores.

Laburnum 1. Laburnum anagyroides Medik. E golden chain-tree. MA. Fields, roadsides, areas of habitation. ‌1 × Laburnum alpinum Griseb. Laburnum ×watereri Dipp. is a rare chain-tree hybrid known from ME. It can be recognized by its sparsely sericeous leaflets (on the abaxial surface) and racemes and sparsely pubescent, few-seeded legumes (which are rarely developed) with a narrow, wing-like keel (moderately to densely sericeous leaflets and racemes and well-developed, appressed-pubescent fruits with a thick keel in L. anagyroides).

Lathyrus 1a. Leaves lacking leaflets, reduced to a tendril; corollas yellow . . . . . . . . . . . . . . . . . . . L. aphaca 1b. Leaves with 2 or more well-developed leaflets; corollas white to ochroleucous or pinkpurple to purple (yellow only in L. pratensis, a species with racemes of 4–10 flowers vs. solitary or rarely paired in L. aphaca) 2a. Leaves with 4–16 leaflets [Fig. 641]; native plants 3a. Stipules ± symmetrical, with 2 basal lobes, approximately equal in size to the leaflets; leaflets fleshy; legumes mostly 8–11 mm wide . . . . . . . . . . . . . . . . . . . L. japonicus 3b. Stipules asymmetrical, with 1 basal lobe [Fig. 641], definitely smaller than the leaflets; leaflets scarcely fleshy; legumes 4–7 mm wide 4a. Corollas pink-purple to purple; raceme with 3–6 flowers; stipules 1–10 mm long, semisagittate (i.e., with a sharply pointed basal lobe); leaflets elliptic to ellipticoblong, 2.5–9 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. palustris 4b. Corollas white to ochroleucous; raceme with 5–15 flowers; stipules 15–30 mm long, some semicordate (i.e., with a rounded or bluntly pointed basal lobe); leaflets ovate to elliptic, 2–2.4 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . L. ochroleucus 2b. Leaves with 2 leaflets [Fig. 640]; non-native plants originating from cultivation 5a. Plants annual, without rhizomes; inflorescence with 1–3 (–4) flowers 6a. Leaflets linear to broad-linear; legume glabrous; flowers usually solitary, not especially fragrant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sativus 6b. Leaflets ovate or elliptic; legume pubescent; flowers 1–3 (–4) in a raceme, very fragrant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. odoratus 5b. Plants perennial, with rhizomes; inflorescence with (3–) 4–10 flowers

Fig. 639  Leaf of Kummerowia striata showing striate stipule.

5 84   tricolpate s

7a. Corollas 14–25 mm long; legumes 40–100 × 6–10 mm; stems conspicuously winged 8a. Stipules 3–10 mm wide, lanceolate to ovate [Fig. 640]; leaflets ovate to lanceolate; corollas 18–25 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. latifolius 8b. Stipules 1–2 (–4) mm wide, narrow-lanceolate; leaflets oblong-lanceolate to narrow-lanceolate; corollas 13–20 mm long . . . . . . . . . . . . . . . . . . . . . . . L. sylvestris 7b. Corollas 10–15 mm long; legumes 20–50 × 4–7 mm; stems without prominent wing-angles 9a. Corolla yellow; stipules 10–30 mm long; calyx lobes subequal in length, slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. pratensis 9b. Corolla red-purple; stipules 5–12 mm long; calyx lobes unequal, broad and short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. tuberosus 1. Lathyrus aphaca L. E yellow vetchling. MA. Fields, roadsides, waste areas. 2. Lathyrus japonicus Willd. var. maritimus (L.) Kartesz & Gandhi N beach vetchling. Lathyrus japonicus Willd. ssp. maritimus (L.) P.W. Ball; L. japonicus Willd. var. glaber (Ser.) Fern.; L. japonicus Willd. var. pellitus Fern.; L. maritimus (L.) Fries; L. maritimus (L.) Fries var. glaber (Ser.) Eames; L. maritimus (L.) Fries var. pellitus (Fern.) Gleason; Pisum maritimum L. • CT, MA, ME, NH, RI, VT; coastal sites with disjunct populations on Lake Champlain. Atlantic coast beaches and dunes, saline marshes, rarely of sand and cobble lake shores. Recent studies do not support segregation into two taxa based on indument. This trait appears to influenced by degree of exposure—plants from protected microsites are generally glabrous (David Barrington, personal communication). 3. Lathyrus latifolius L. E Fig. 640 Fig. 640  Leaf and winged stem of Lathyrus latifolius.

everlasting vetchling. CT, MA, ME, NH, RI, VT. Roadsides, field and lawn edges, railroads, waste areas. 4. Lathyrus ochroleucus Hook.

NC

cream vetchling. VT; lower (i.e., northern) Lake Champlain. Lake shore bluffs and headlands in regions of high-pH bedrock. 5. Lathyrus odoratus L. E sweet vetchling. CT, VT; also reported from ME by Campbell et al. (1995), but specimens are unknown. Fields, waste areas, roadsides, areas of cultivation. 6. Lathyrus palustris L. N Fig. 641 marsh vetchling. Lathyrus myrtifolius Muhl. ex Willd.; L. palustris L. var. macranthus (White) Fern.; L. palustris L. var. myrtifolius (Muhl. ex Willd.) Gray; L. palustris L. ssp. pilosus (Cham.) Hultén; L. palustris L. var. pilosus (Cham.) Ledeb.; Orobus myrtifolius (Muhl. ex Willd.) A. Hall • CT, MA, ME, NH, RI, VT. Borders of saline and brackish marshes, upper edge of coastal beaches, river shores, low meadows. 7. Lathyrus pratensis L. E Fig. 641  Leaf and slender stem of Lathyrus palustris.

meadow vetchling. CT, MA, ME, NH, VT. Fields, roadsides, open banks, clearings. 8. Lathyrus sativus L. E white vetchling. MA. Dumps, fields. 9. Lathyrus sylvestris L. E narrow-leaved vetchling. CT, MA, NH, RI, VT. Fields, roadsides, waste areas, forest borders. 10. Lathyrus tuberosus L. E tuber vetchling. CT, MA, ME, VT. Fields, grasslands.

Fa b ac e a e   5 8 5

Lens 1. Lens culinaris Medik. E lentil. Ervum lens L.; Lens esculenta Moench • CT, MA, ME. Ballast, fields.

Lespedeza Hybrids are occasionally collected in Lespedeza and are most frequent in larger populations that contain two or more species. Though not all hybrids present an easy manner for detecting the type of plant (i.e., if the plant is an orthospecies or a nothospecies), some plants do provide clues as to their origin. Plants with ascending stems in mixed populations of species with procumbent or erect stems are likely hybrids. Plants with pink flowers in mixed populations of species with white or purple flowers are likely hybrids. Native plants with fruits that are shortly exserted from the calyx (i.e., the calyx is ca. ⅔ as long as the fruit) are likely hybrids (given that native species of Lespedeza have fruits that are up to about as long as the calyx or 2 or more times as long as the calyx). References: Clewell (1966), Raveill (2006). 1a. Shrubs or suffutescent herbs 10–30 dm tall; corollas 8–15 mm long; stipules persistent 2a. Calyx lobes aristate-tipped; leaflets broad-oval, 1–1.3 times as long as wide; inflorescence not or scarcely exceeding the subtending leaves . . . . . . . . . . . L. cyrtobotrya 2b. Calyx lobes acute or even mucronate, but not aristate-tipped; leaflets narrow-elliptic to oval, 1.5–3 times as long as wide; inflorescence much exceeding the subtending leaves 3a. Corolla 8–11 mm long; calyx lobes acute, shorter than or equal to the length of the basal, connate portion of the calyx; leaflets oval, 1.5–2 times as long as wide; stems solitary or few together from the summit of the root crown, the young stems brown; seeds mottled purple or green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. bicolor 3b. Corolla (10–) 12–15 mm long; calyx lobes often mucronate, longer than the basal connate portion of the calyx; leaflets narrow-elliptic to oval, 2–3 times as long as wide; stems many and clustered from the summit of the root crown, the young stems purple; seeds purple-black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. thunbergii 1b. Herbs 2–15 (–20) dm tall or long; corollas 5–9 mm long; stipules usually caducous 4a. Corollas pink-purple to purple; legume (2–) 2.5–4 times as long as the persistent calyx [Fig. 644] 5a. Plants procumbent or trailing 6a. Stem and petiole pilose (i.e., pubescent with spreading hairs); primary leaves with broad-ovate leaflets 1.2–2 times as long as wide; racemes with 8–12 (–14) flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. procumbens 6b. Stem and petiole strigose (i.e., pubescent with ± appressed hairs); primary leaves with elliptic to ovate or obovate leaflets 1.5–2.5 (–3) times as long as wide; racemes with 4–8 (–12) flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. repens 5b. Plants erect to ascending 7a. Chasmogamous inflorescence loose, with 4–6 (–8) flowers, relatively elongate, the peduncle and rachis combined 2–5 times as long as the subtending leaf [Fig. 642]; keel petal 1–2 mm longer than the wing petals; calyx ca. 20% (rarely as much as 25%) as long as the legume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. frutescens 7b. Chasmogamous inflorescence dense, with 4–14 flowers, borne on relatively short peduncles, the peduncle and rachis combined up to 1.5 times as long as the subtending leaf; keel petal up to as long as the wing petals; calyx 20–40% as long as the legume 8a. Leaflets oblong to linear, (4–) 5–7 times as long as wide . . . . . . . . . L. virginica

5 86   tricolpate s

8b. Leaflets elliptic-oblong to broad-elliptic or obovate, 1–3 (–3.5) times as long as wide 9a. Leaflets glabrous on the adaxial surface or sometimes strigose along the midrib; upper 2 calyx lobes connate for ca. 50% or more of their length above the basal, tubular portion of the calyx; stems usually strigose . . . . . . L. violacea 9b. Leaflets strigose to pilose on the adaxial surface; upper 2 calyx lobes distinct or shortly connate above the basal, tubular portion of the calyx; stems usually pilose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. stuevei 4b. Corollas white or ochroleucous, sometimes with a purple spot on the banner petal; legume not or only slightly exceeding the persistent calyx [Fig. 644] 10a. Flowers solitary or in clusters of 2 or 3, borne on very short peduncles, the peduncle and rachis combined shorter than the subtending leaf; upper 2 calyx lobes connate ca. 50% of their length; leaflets truncate or retuse at the apex . . . . . L. cuneata 10b. Raceme with 10–44 flowers, relatively elongate, the peduncle and rachis combined 1–3.5 times as long as the subtending leaf; calyx lobes all distinct; leaflets rounded to obtuse at the apex 11a. Leaflets ovate-elliptic to obovate, 1.2–1.8 times as long as wide; middle stem leaves borne on petioles 10–15 (–20) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. hirta 11b. Leaflets narrow-oblong to oblong or elliptic, 2.5–8 times as long as wide; middle stem leaves borne on petioles 1–3 (–6) mm long 12a. Racemes included in or scarcely exceeding the subtending leaves, with subglobose to short-ovoid clusters of flowers; calyx 8–12 mm long; corolla 8–12 mm long; leaflets elliptic to oblong (rarely narrow-oblong), (2–) 2.5–5 (–8) times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. capitata 12b. Racemes exserted beyong the subtending leaves, with ovoid to cylindric clusters of flowers; calyx 5–8 mm long; corolla 5–7 mm long; leaflets ellipticoblong to narrow-oblong, 4–8 times as long as wide . . . . . . . . . . . . L. angustifolia 1. Lespedeza angustifolia (Pursh) Ell. N narrow-leaved bush-clover. Lespedeza angustifolia (Pursh) Ell. var. brevifolia Britt.; L. capitata Michx. var. angustifolia Pursh • MA; also reported from RI by Magee and Ahles (1999), but specimens are unknown. Sandy areas and pond shores, mostly along the coastal plain. 2. Lespedeza bicolor Turcz. E two-colored bush-clover. CT, MA, VT. Fields, roadsides, waste areas. 3. Lespedeza capitata Michx. N round-headed bush-clover. Lespedeza bicknellii House; L. capitata Michx. var. stenophylla Bissell & Fern.; L. capitata Michx. var. velutina (Bickn.) Fern. • CT, MA, ME, NH, RI, VT. Woodlands, dry fields and clearings, roadsides, sandplains. ‌3 × 7. Lespedeza ×longifolia DC. is a rare hybrid bush-clover that known is from MA, ME, NH. It usually displays one of three morphologies—(1) plants with leaves of L. hirta but the congested inflorescence of L. capitata; (2) plants with leaves intermediate between the parents but with well-exserted inflorescences; and (3) plants generally ‌longifolia may not resembling L. capitata but with elongate petioles. The name L. × apply to this nothospecies, but rather to L. angustifolia × L. hirta or unusual plants of L. capitata (Clewell 1966). ‌3 × 13. Lespedeza ×simulata Mack. & Bush is very rare hybrid bush-clover known from CT. It is characterized by foliage that generally resembles L. virginica but with relatively long calyx lobes. The report for VT by Seymour (1969) is unlikely given the distribution of L. virginica (it is absent from VT). 4. Lespedeza cuneata (Dum.-Cours.) G. Don E Chinese bush-clover. CT, MA. Fields, roadsides.

Fa bac e a e   5 87

5. Lespedeza cyrtobotrya Miq. E leafy bush-clover. CT, MA. Fields, roadsides. 6. Lespedeza frutescens (L.) Hornem. N Fig. 642 violet bush-clover. Hedysarum frutescens L.; Lespedeza violacea, auct. non (L.) Pers. • CT, MA, NH, RI, VT. Dry-mesic woodlands and forests, often on rocky slopes and ridges, openings on balds and ledges. This species has long been called Lespedeza violacea; however, that name properly belongs to the species that has been called L. intermedia (see discussion under L. violacea). 6 × 7. This very rare bush-clover hybrid is known from RI. It has fruits mostly 1.2–1.5 times as long as the sepals and has fewer-flowered and looser inflorescences than Lespedeza hirta.

Fig. 642  Inflorescence and leaves of Lespedeza frutescens.

6 × 10. This very rare bush-clover hybrid is known from RI. It has fruits mostly 1.2–1.5 times as long as the sepals and has fewer-flowered and looser inflorescences than Lespedeza stuevei. The leaf blades have, on average, more pubescence than L. frutescens. ‌6 × 13. Lespedeza ×acuticarpa Mackenzie & Bush is a very rare bush-clover hybrid known from RI in New England. This taxon is morphologically diverse, ranging from plants that resemble L. frutescens with unusually narrow leaflets to plants like L. virginica with a few, well-exserted racemes. 7. Lespedeza hirta (L.) Hornem. ssp. hirta N Fig. 643 hairy bush-clover. Lespedeza capitata (L.) Hornem. var. calycina (Schindl.) Fern.; L. polystachya Michx. • CT, MA, ME, NH, RI, VT; limited in ME to the extreme southern portion of the state. Woodlands, forest clearings, dry openings. Clewell (1966) notes that the plants of Marthas Vineyard and Nantucket Island, MA, are morphologically distinct from other populations of this species in their denser and longer indument. 7 × 8. This very rare bush-clover hybrid is known from MA. It shows many stems ascending (rather than procumbent or erect, as in the parental taxa) and calyx lobes often ca. ⅔ as long as the mature legume. 7 × 10. This very rare bush-clover hybrid is known from MA. It is similar to Lespedeza hirta × L. violacea (7 × 12), but the adaxial leaf surface is moderately pubescent (rather than sparsely), and the abaxial leaf surface is pubescent with dense and/or ascending hairs (rather than moderate and appressed). ‌7 × 12. Lespedeza ×nuttallii Darl. is a rare bush-clover hybrid known from CT, MA, ME, NH, RI, VT. It usually demonstrates the foliage characteristics of L. violacea with the inflorescence characteristics of L. hirta (e.g., well exserted inflorescences). Further, many collections also show longer calyx lobes relative to the length of the fruit than L. violacea (i.e., the fruits are mostly 1.2–1.5 times as long as the calyx). Isely (1998) ‌nuttallii may actually have L. stuevei or stated that some material identified as L. × L. virginica as the purple-flowered parent instead of L. violacea. 7 × 13. This rare bush-clover hybrid is known from CT. It has fruits that shortly exceed the sepals (mostly 1.2–1.5 times as long) and leaf blades that are glabrous adaxially, moderately appressed-pubescent abaxially, and 3.2–3.8 times as long as wide. 8. Lespedeza procumbens Michx. N trailing bush-clover. CT, MA, NH, RI. Dry-mesic woodlands, rocky forests, roadsides, banks, open balds and ledges. 8 × 10. This very rare bush-clover hybrid is known from MA. It shows many stems ascending (rather than procumbent or erect, as in the parental taxa). ‌ × 13. Lespedeza ×brittonii Bickn. is the name for a group of rare bush-clover hybrids 8 known from MA. They are characterized by erect to procumbent stems that are sparsely to evidently pilose, some racemes few-flowered and evidently exserted, and leaflets of intermediate shape or varying toward L. virginica (which is oblong to linear).

Fig. 643  Legume of Lespedeza hirta.

5 88   tricolpate s

9. Lespedeza repens (L.) W. Bart. N creeping bush-clover. Hedysarum repens L. • CT. Woodlands and forests, often on rocky slopes and in dry openings. 10. Lespedeza stuevei Nutt.

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tall bush-clover. CT, MA, RI, VT. Sandy soil of woodlands, roadsides, clearings, and barrens. 11. Lespedeza thunbergii (DC.) Nakai E Thunberg’s bush-clover. MA; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, waste areas. 12. Lespedeza violacea (L.) Pers. N Fig. 644

Fig. 644  Legume of Lespedeza violacea.

wand bush-clover. Hedysarum violaceum L.; Lespedeza intermedia (S. Wats.) Britt.; L. stuevei Nutt. var. intermedia S. Wats. • CT, MA, ME, NH, RI, VT. Woodlands, forest openings, rocky and ledgy ridges, sandy fields and plains, roadsides. This species has long been called both Lespedeza frutescens and L. intermedia (commonly the latter in New England literature). However, the type specimen of Hedysarum violaceam, on which the name L. violacea is based, is actually the plant traditionally called L. intermedia (Reveal and Barrie 1991). Therefore, the name L. intermedia is a superfluous synonym, and L. violacea is the appropriate name for this plant. 13. Lespedeza virginica (L.) Britt. N slender bush-clover. Medicago virginica L. • CT, MA, NH, RI. Dry-mesic woodlands, forests, and clearings.

Lotus 1a. Corolla ochroleucous to pink or red-striate, (4–) 5–9 mm long; basal connate portion of calyx 1–2 mm long; stipules inconspicuous glands; flowers solitary, borne on axillary peduncles; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. unifoliatus 1b. Corolla yellow, 10–14 mm long; basal connate portion of calyx 2.5–3.5 mm long; stipules foliaceous, resembling the leaflets; flowers borne in peduncled umbels; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. corniculatus 1. Lotus corniculatus L. E garden bird’s-foot-trefoil. Lotus corniculatus L. var. arvensis (Schkuhr) Ser. ex DC. • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, waste areas. 2. Lotus unifoliatus (Hook.) Benth. var. unifoliatus E American bird’s-foot-treefoil. Acmispon americanum (Nutt.) Rydb.; Hosackia americana (Nutt.) Piper; Lotus americanus (Nutt.) Bisch.; L. purshianus F.E. & E.G. Clem. ex Ottley; L. purshianus F.E. & E.G. Clem. ex Ottley var. glaber (Nutt.) Munz • CT, MA. Waste areas, yards.

Lupinus Reports of Lupinus albus L. in MA (e.g., Sorrie and Somers 1999) were based on a specimen taken from cultivated material—11 Sep 1917, Kidder s.n. (NEBC!). 1a. Leaves with 5–7 (–11) leaflets 2–5 (–6) cm long; racemes usually 10–20 cm tall; corollas 12–16 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. perennis 1b. Leaves with (8–) 11–17 leaflets 7.5–13 cm long; racemes usually 20–40 cm tall; corollas 15–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. polyphyllus 1. Lupinus perennis L. ssp. perennis

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sundial lupine. Lupinus perennis L. var. occidentalis S. Wats. • CT, MA, ME, NH, RI, VT. Sandy or gravelly soil of fields, roadsides, floodplains, railroads, woodlands, cleared rights-of-ways, and waste areas.

Fa bac e a e   5 8 9

2. Lupinus polyphyllus Lindl. var. polyphyllus E blue lupine. Lupinus pallidipes Heller; L. polyphyllus Lindl. var. albiflorus L.H. Bailey; L. polyphyllus Lindl. var. pallidipes (Heller) C.P. Sm. • CT, MA, ME, NH, VT. Fields, roadsides, gardens.

Medicago Though flowering specimens possess many good characters on which to base identifications, fruits are generally necessary for identification of some of the rarer species of Medicago that have been introduced to New England. Reference: Small and Jomphe (1989). 1a. Inflorescence with 1 or 2 (–3) flowers; legume (30–) 40–100 mm long, unarmed, merely curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. monantha 1b. Inflorescence with (1–) 2–50 flowers; legume 1–10 mm long, armed with thin prickles or unarmed, curved to coiled up to 7 times (rarely straight in M. falcata and M. lupulina) 2a. Corollas 6–12 mm long, blue-purple to purple or rarely yellow; legumes unarmed; plants perennial 3a. Corollas blue-purple to purple, usually 8–11 mm long; legumes coiled (1–) 2–3 times; stems erect to ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. sativa 3b. Corollas yellow, usually 5–8 mm long; legumes falcate-curved (rarely straight or partly coiled); stems prostrate to erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. falcata 2b. Corollas 2–5 (–7) mm long, yellow; legumes armed with thin prickles (always unarmed in M. lupulina, rarely unarmed in M. rigidula) [Fig. 645]; plants annual or biennial 4a. Legume reniform-curved or slightly spiraled near the apex, without prickles [Fig. 646], with a single seed; inflorescence with (10–) 14–24 (–50) flowers . . . . . . . . . . . . . M. lupulina 4b. Legume 2- to 7-times spiral-coiled, with thin prickles (prickles rarely lacking in M. rigidula), with several seeds; inflorescence with (1–) 2–5 (–8) flowers 5a. Legume in a very tight spiral, extremely hard-walled, usually glandular-puberulent, the young legume concealed within the calyx when the petals drop; prickles of legume (when present) stocky, firm, ± round at the base . . . . . . . . . . . . . . . . . M. rigidula 5b. Legume in a somewhat tight spiral, soft-walled, glabrous or pubescent with eglandular hairs (sometimes glandular in M. minima, but that species with small fruits 3–5 mm in diameter), the young legume projecting sideways from the calyx when the petals drop; prickles of legume relatively thin, flexible, with 2 roots, 1 root arising in the dorsal surture, the other root arising in the submarginal vein 6a. Faces of the fruit coils lacking a reticulum of veins over much of the surface; prickles of legume tending to slant toward the base of the fruit . . . . M. disciformis 6b. Faces of the fruit coils with a reticulum of bold veins over much of the surface (the veins sparingly branched in M. laciniata, but still evident); prickles of legume tending to project outward from fruit (i.e., parallel with fruit coil) 7a. Leaflets broad-obovate to broad-obcordate, 0.7–1.1 times as long as wide, usually with a dark blotch or mottles on the adaxial surface; prickles on legume usually curved and somewhat conforming to the outline of the fruit [Fig. 645] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. arabica 7b. Leaflets elliptic to obovate or obcordate, 1–2 (–2.5) times as long as wide, without dark mottles; prickles on legume relatively straight (except near apex of fruit), diverging from outline of fruit 8a. Stipules entire or slightly dentate; legumes sparsely pubescent; stems moderately to densely villous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. minima

5 90   tricolpate s

8b. Stipules lacerate; legumes usually glabrous; stems glabrous or sparsely pubescent 9a. Corolla 2–3 (–4) mm long; coils of fruit loosely appressed to each other (i.e., spaces evident between the coils when the fruit is viewed on edge); margin of fruit coils very broad . . . . . . . . . . . . . . . . . . . . . . M. praecox 9b. Corolla (3–) 4–8 mm long; coils of fruit moderately to tightly appressed to each other; margin of fruit coils narrow 10a. Leaflets 8–20 mm long, apically denticulate; legumes 4–8 (–10) mm in diameter, with transverse veins that anastomose toward the margins, with the submarginal veins separated from the marginal vein by a deep groove that is visible when the legume is viewed on edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. polymorpha 10b. Leaflets 5–10 (–12) mm long, commonly laciniately toothed; legumes 2.5–5 mm in diameter, with tranverse veins that are merely sparingly branched toward the margins, with the submarginal veins separated from the marginal vein by a narrow groove that is ± not visible when the legume is viewed on edge . . . . . . . . . . . . . M. laciniata 1. Medicago arabica (L.) Huds. E Fig. 645 spotted medick. Medicago polymorpha L. var. arabica L. • CT, MA, ME, RI, VT. Fields, waste areas, wool waste, areas of cultivation. 2. Medicago disciformis DC. E Fig. 645  Armed legume of Medicago arabica.

veinless medick. MA. Wool waste. 3. Medicago falcata L. E yellow medick. Medicago sativa L. ssp. falcata (L.) Arcang. • MA. Fields, waste areas, wool waste. This species and Medicago sativa form a difficult complex of intergrading forms, the intergradations likely the result of cultivation and hybridization. Isely (1998) warned that most large, yellow-flowered material identified as Medicago falcata in herbaria are likely various hybrids within the M. sativa complex. He justified this stance by noting that the fruits of these plants are coiled to some degree (those of M. falcata are usually merely falcate-curved). 3 × 11. This variable medick hybrid has been reported from VT but is likely also found in ‌varia Martyn other states where Medicago falcata has been reported. The name M. × is thought to apply to hybrids between pure M. falcata and pure M. sativa, these recognized by the unusual green-blue or purple-black corollas (a novel character state compared with the putative parental taxa). Certain combinations of morphological character states have been given specific names (e.g., M. glomerata Balbis applies to plants with yellow corollas 6–10 mm long and coiled legumes). See Tutin (1968) for additional discussion. 4. Medicago laciniata (L.) P. Mill. E cut-leaved medick. Trigonella laciniata L. • MA, ME. Waste areas, wool waste. 5. Medicago lupulina L. E Fig. 646 black medick. Medicago lupulina L. var. cupaniana (Guss.) Boiss.; M. lupulina L. var. glandulosa Neilr. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas.

Fig. 646  Unarmed legume of Medicago lupulina.

6. Medicago minima (L.) L. E bur medick. Medicago minima (L.) L. var. compacta Neyraut; M. minima (L.) L. var. longiseta DC.; M. polymorpha L. var. minima L. • CT, MA. Wool waste, dumps. 7. Medicago monantha (C.A. Mey.) Trautv. E single-flowered medick. Trigonella monantha C.A. Mey.; T. monantha C.A. Mey. ssp. noeana (Boiss.) Hub.-Mor.; T. noeana Boiss. • MA. Waste areas, dumps.

Fa b ac e a e   5 9 1

8. Medicago polymorpha L. E toothed medick. Medicago hispida Gaertn.; M. hispida Gaertn. var. apiculata (Willd.) Urban; M. hispida Gaertn. var. confinis (W.D.J. Koch) Burnat • CT, MA, ME, VT. Waste areas, wool waste, areas of cultivation. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI and was unaware of any collections. 9. Medicago praecox DC. E Mediterranean medick. MA. Wool waste. 10. Medicago rigidula (L.) All. E Tifton medick. Medicago agrestis Ten. ex DC. • MA. Wool waste. 11. Medicago sativa L. E purple medick. CT, MA, ME, NH, RI, VT. Fields, roadsides, areas of cultivation.

Melilotus Melilotus altissimus Thuill. was reported from ME by Hultén and Fries (1986), but specimens are unknown. Steele and Wojciechowski (2003) found Melilotus to be monophyletic, but its recognition creates a paraphyletic Trigonella. Further work may show these two genera need to be combined under the name Trigonella (it has priority over Meliolotus). 1a. Corollas yellow, 2–3 mm long; inflorescence compact at anthesis, only 1–2 (–3) cm long, elongating in fruit; pedicels ascending, 0.5–1 mm long; legumes 1.5–3 mm long; plants 0.2–0.6 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. indicus 1b. Corollas yellow or white, (3.5–) 4–7 mm long; inflorescence 3–12 cm long at anthesis, elongating in fruit; pedicels decurved, 1–2 mm long; legumes 2.5–5 mm long; plants 0.5–3 m tall 2a. Corollas yellow, 5–7 mm long; wing petals ± as long as the banner petal and usually longer than the keel petals; legumes cross-striate on the outer surface . . . . . . M. officinalis 2b. Corollas white, (3.5–) 4–5 mm long; wing petals shorter than the banner petal and ± as long as the keel petals [Fig. 647]; legumes reticulate-patterened on the outer surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. albus 1. Melilotus albus Medik. E Fig. 647 white sweet-clover. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 2. Melilotus indicus (L.) All. E Indian sweet-clover. Trifolium indicum L. • MA, ME, NH, VT. Fields, waste areas, gardens. 3. Melilotus officinalis (L.) Lam. E yellow sweet-clover. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads.

Onobrychis 1. Onobrychis viciifolia Scop. E Sanfoin. MA, VT. Fields, abandoned homesteads.

Ornithopus 1. Ornithopus sativus Brot. ssp. sativus E common bird’s-foot. MA; eastern portion of state. Waste areas.

Fig. 647  Inflorescence of Melilotus albus with detail of a flower.

5 92   tricolpate s

Oxytropis 1. Oxytropis campestris (L.) DC. var. johannensis Fern.

NC

St. John River oxytrope. Oxytropis johannensis (Fern.) Heller • ME; northern portion of state. Ice-scoured river shores in regions of high-pH bedrock and/or till, on ledge, cobble, and gravel substrate.

Phaseolus 1a. Inflorescence with a slender, flexuous axis; legumes 3–6 cm long; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. polystachios 1b. Inflorescence with a stout, relatively straight axis; legumes 8–30 cm long; plants annual 2a. Corolla 15–20 mm long, red or bright purple (rarely light purple); inflorescence usually exserted beyond the subtending leaves; plants twining . . . . . . . . . . . . . . . . . . . . . P. coccineus 2b. Corolla 10–15 mm long, pink-purple to nearly white; inflorescence often not exceeding the subtending leaves; plants upright or twining . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. vulgaris 1. Phaseolus coccineus L. E scarlet bean. MA. Fields, gardens, dumps. 2. Phaseolus polystachios (L.) B.S.P.

N C Fig. 648

wild bean. Dolichos polystachyus L.; Phaseolus polystachios (L.) B.S.P. var. aquilonius Fern.; P. smilacifolius Pollard • CT. Also reported from ME by Gleason (1963) and RI by George (1992), but specimens are unknown. Rocky forests and woodlands, cliff bases. The report for ME by Gleason (1963) is almost certainly in error given that this species is not known north of CT based on specimens that have been examined. Fig. 648  Flower of Phaseolus polystachios showing inwardly coiled keel petals.

3. Phaseolus vulgaris L. E kidney bean. CT, MA, ME, NH, RI. Gardens, pastures, areas of cultivation. Included in this species are several other cultivated beans including garden bean, pole bean, and string bean.

Pisum 1. Pisum sativum L. E garden pea. Pisum arvense L.; P. sativum L. var. arvense (L.) Poir. • CT, MA, ME, NH. Fields, waste places, areas of cultivation.

Pueraria Pueraria montana was reported from ME by Kartesz (1999) based on comments in Frankel (1989). Frankel based his report on Snyder (1987). However, Snyder never mentioned ME in his distribution reporting (though he does mention CT and MA), and Frankel went on to discredit the ME report in the next paragraph. 1. Pueraria montana (Lour.) Merr. var. lobata (Willd.) Maesen & S. Almeida E kudzu. Dolichos lobatus Willd.; Pueraria lobata (Willd.) Ohwi; P. thunbergiana (Sieb. & Zucc.) Benth. • CT, MA. Yards, roadsides, thickets.

Robinia 1a. Corolla white (rarely tinged with pink), 15–20 (–25) mm long; raceme pendulous; ovary and legume glabrous; upper margin of legume with a narrow wing; branchlets glabrous or sparsely pubescent; trees to 25 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. pseudoacacia

Fa bac e a e   5 9 3

1b. Corolla pink to red-purple, 20–25 mm long; raceme suberect to spreading, sometimes pendulous; ovary and legume hispid; upper margin of legume unwinged; branchlets glandular, the glands sessile or stipitate; shrubs to 5 m tall 2a. Leaves with 7–13 leaflets that become glabrate abaxially at maturity; branchlets bristly glandular [Fig. 649]; bracts not aristate-tipped (note: the bracts are caducous and evident only prior to anthesis); racemes lax and relatively open, with 4–11 flowers . . . . . . R. hispida 2b. Leaves with 13–25 leaflets that remain permanently appressed-pubescent abaxially (sometimes inconspicuously so); branchlets glandular with sessile or subsessile glands; bracts aristate-tipped; racemes suberect, crowded, with 8–20 flowers . . . . . . . . . R. viscosa 1. Robinia hispida L. E Fig. 649 bristly locust.  1a. Robinia fertilis Ashe; R. grandiflora Ashe; 1b. Robinia pallida Ashe; R. speciosa Ashe • CT, MA, ME, NH, RI, VT. Roadsides, field edges, abandoned homesteads. 1a. Plants producing abundant fruit; leaflets elliptic to elliptic-oblong or elliptic-obovate, acute to obtuse at the apex, at least sparsely pubescent abaxially, 7–25 mm wide; flowers 20–25 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. Robinia hispida var. fertilis (Ashe) Clausen 1b. Plants rarely fruiting, and when fruits are produced these are often without seeds; leaflets oval to suborbicular, rounded to obtuse at apex, ± glabrous abaxially, 15–45 mm wide; flowers 25–30 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. Robinia hispida var. hispida Variety fertilis is known from CT. This variety was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI. Variety hispida is known from CT, MA, ME, NH, RI, VT.

Fig. 649  Leaf and branchlet of Robinia hispida.

2. Robinia pseudoacacia L. E black locust. CT, MA, ME, NH, RI, VT. Roadsides, forest fragments, field edges, riparian forests. 3. Robinia viscosa Vent. var. viscosa E clammy locust. CT, MA, ME, NH, RI, VT. Roadsides, abandoned homesteads.

Scorpiurus 1. Scorpiurus muricatus L. E prickly scorpion’s-tail. Scorpiurus subvillosus L.; S. sulcatus L. • MA. Wool waste.

Securigera 1. Securigera varia (L.) Lassen E Fig. 650 purple crown-vetch. Coronilla varia L. • CT, MA, ME, NH, RI, VT. Roadsides, fields, banks.

Senna Senna marilandica (L.) Link was reported from MA by Isely (1998). However, the manuscript is internally inconsistent—the map noted a collection for MA but the text stated this species only reaches southeastern NY. Senna marilandica was also reported from CT by Pollard (1894), a report that was more recently taken up by Kartesz (1999). However, Pollard’s report of this species in New England was prior to the publication of Senna hebecarpa, a species named in 1937 by Fernald (i.e., Pollard’s report of S. marilandica in New England was based on plants of S. hebecarpa before they were recognized as distinct from S. marilandica). Additionally, S. corymbosa (Lam.) Irwin & Barneby was reported from MA or RI (scale of map does not allow precise reporting) and S. occidentalis (L.) Link was reported from RI by Isely (1998), but specimens are unknown. Given that all the above species and S. obtusifolia were mapped in approximately the same location (southeastern MA/southeastern RI), it appears that some or all of these reports were likely errors in mapping (note that a valid report of S. obtusifolia does exist, see below).

Fig. 650  Inflorescence of Securigera varia.

5 94   tricolpate s

1a. Gland located near base of petiole; leaves with 6–8 (–10) pairs of elliptic to oblanceolate leaflets 2.3–3.5 times as long as wide; inflorescence of 9- to 20-flowered axillary and terminal racemes; fruits 7–10 (–12) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hebecarpa 1b. Gland located between or immediately below the lowest pair of leaflets; leaves with (2–) 3 pairs of obovate (rarely to elliptic) leaflets 1.3–1.8 times as long as wide; inflorescence of solitary or paired axillary flowers, these sometimes clustered near stem apex; fruits 10–20 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. obtusifolia 1. Senna hebecarpa (Fern.) Irwin & Barneby N northern wild senna. Cassia hebecarpa Fern. • CT, MA, NH, RI, VT; also reported from ME by Kartesz (1999), but specimens are unknown. Fields, roadsides, forest borders, riparian corridors. 2. Senna obtusifolia (L.) Irwin & Barneby E sickle-pod wild senna. Cassia obtusifolia L. • MA. Railroads, yards.

Sesbania 1. Sesbania herbacea (P. Mill.) McVaugh E tall river-hemp. Darwinia exaltata Raf.; Sesban exaltatus (Raf.) Rydb; Sesbania exaltata (Raf.) Corey; S. macrocarpa Muhl. • MA. Fields, roadsides, ditches.

Strophostyles Reference: Riley-Hulting et al. (2004). 1a. Legumes permanently pubescent, (12.2–) 20–40.7 mm long; peduncles slender, herbaceous, 0.2–0.7 mm thick; corollas 3.6–7 (–8.3) mm long; keel petals mostly concealed by wing petals, only a short segment of the slightly curved beak exposed; basal, connate portion of calyx pubescent; seeds glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. leiosperma 1b. Legumes glabrate at maturity, (26–) 30–96 mm long; peduncles stout, ligneous, 0.8–1.3 mm thick; corollas 8–15 mm long; keel petals with a long, curved beak that protrudes well beyond the wing petals; basal, connate portion of calyx glabrous or essentially so; seeds pubescent 2a. Bracteoles 1.5–4.2 mm long, lanceolate, acute at the apex, usually as long as or longer than the basal connate portion of the calyx; leaflets frequently with a lobe on one or both margins, the terminal one 25–36 mm wide and 1.5–3 times as long as wide; beak of keel petals slender, ca. 1 mm in diameter at the base, projecting away from the banner petal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. helvola 2b. Bracteoles 0.8–2.4 mm long, ovate to oblong, blunt, usually shorter than the basal, connate portion of the calyx; leaflets usually lacking lobes, but the margins sometimes sinuate, the terminal one 3–20 mm wide and 2.2–8 (–10) times as long as wide; beak of keel petals stouter, 1.5–2 mm in diameter at the base, oriented to be closely associated with the banner petal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. umbellata 1. Strophostyles helvola (L.) Ell. N Fig. 651 annual woolly bean. Phaseolus helvolus L.; Strophostyles helvola (L.) Ell. var. missouriensis (S. Wats.) Britt. • CT, MA, ME, RI; mainly on the coastal plain. Sandy soil of beaches, roadsides, woodlands, and clearings, most frequent in near coastal areas. Fig. 651  Flower of Strophostyles helvola showing inwardly arched keel petals.

2. Strophostyles leiosperma (Torr. & Gray) Piper E small-flowered woolly bean. Phaseolus leiospermus Torr. & Gray • CT. Dumps, borrow pits. 3. Strophostyles umbellata (Muhl. ex Willd.) Britt.

NC

perennial woolly bean. Glycine umbellata Muhl. ex Willd.; Strophostyles umbellata (Muhl. ex Willd.) Britt. var. paludigena Fern. • RI. Sandy soil of fields, woodlands, and clearings, mainly near the coast.

Fa bac e a e   5 9 5

Tephrosia 1. Tephrosia virginiana (L.) Pers. N wild goat’s-rue. Cracca latidens Small; C. virginiana L.; Tephrosia latidens (Small) Standl.; T. virginiana (L.) Pers. var. glabra Nutt.; T. virginiana (L.) Pers. var. holosericea (Nutt.) Torr. & Gray • CT, MA, NH, RI. Fields, roadsides, trail edges, clearings, woodlands, mainly on sandy soils.

Thermopsis Thermopsis montana Nutt. was reported from ME by Seymour (1982). The specimen supporting this report—Parlin 16543 (GH!)—is T. villosa. 1. Thermopsis villosa (Walt.) Fern. & Schub. E Blue Ridge false lupine. Thermopsis caroliniana M.A. Curtis • CT, MA, ME, NH, VT. Fields, roadsides, railroads, waste areas. This species has been erroneously called Thermopsis mollis (Michx.) M.A. Curtis ex Gray in various references covering this area (e.g., Seymour 1982, Magee and Ahles 1999).

Trifolium Trifolium macraei Hook. & Arn. was first reported from New England by Knowlton and Deane (1918), and has been reported in other floras since (e.g., Kartesz 1999). The specimen on which this record was based was determined to be T. incarnatum by Sorrie and Somers (1999). Trifolium carolinianum Michx. was reported from VT by Atwood et al. (1973), but specimens are unknown. Reference: Coombe (1968). 1a. Flowers yellow, turning brown after anthesis 2a. Central leaflet sessile or on a petiolule of ± similar length as the lateral leaflets; corollas 5–7 mm long; inflorescence 10–17 (–20) mm tall; stipules ± as long as the petioles; style 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. aureum 2b. Central leaflet borne on a petiolule conspicuously longer than those of the lateral leaflets [Fig. 653]; corollas 3.5–5 (–6) mm long; inflorescence 5–15 mm tall; stipules shorter than the petioles; style less than 1 mm long 3a. Petiolule of the central leaflet up to 1 mm long; inflorescence with usually 5–15 flowers; corollas 3.5–4 mm long; banner petal inconspicuously veined . . . . . . . T. dubium 3b. Petiolule of the central leaflet 1–3 mm long; inflorescence with usually 20–30 flowers; corollas (3.5–) 4–5 (–6) mm long; banner petal with 10 conspicuous veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. campestre 1b. Flowers white to pink to purple 4a. Flowers pedicellate, the pedicels usually longer than 2 mm; petals white or white and pink 5a. Stems creeping along the ground, rooting at the nodes; petals usually concolored; stipules connate to the petiole in the basal portion, then with fused margins for a distance, forming a tube that surrounds the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . T. repens 5b. Stems ascending, not rooting at the nodes; petals bicolored; stipules connate to the petiole in the basal portion, then distinct, the tips completely separate and not forming a tube . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. hybridum 4b. Flowers sessile or subsessile, the pedicels (when present) up to 0.5 mm long; petals white to pink to purple 6a. Basal connate portion of calyx more densely pubescent on the upper side, becoming obliquely inflated in fruit, the upper side gibbous; flowers each subtended by a bracteole, resupinate (i.e., twisted within the calyx such that the banner petal is in a lateral or lower position; not resupinate in T. fragiferum) [Fig. 655]

5 96   tricolpate s

7a. Flowers not resupinate; peduncles (20–) 40–80 (–200) mm long; plants perennial, usually stoloniferous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. fragiferum 7b. Flowers resupinate; peduncles up to 50 mm long; plants annual, not stoloniferous 8a. Inflorescence mostly 10–20 mm in diameter, lobulate in fruit, borne on elongate peduncles 20–50 mm long; stems 10–40 (–60) cm tall . . . T. resupinatum 8b. Inflorescence mostly 5–10 mm in diameter, hemispheroid to spherical in fruit, subsessile or borne on a peduncle shorter than 20 mm; stems 5–15 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. tomentosum 6b. Basal connate portion of calyx uniformly glabrate or pubescent, not becoming obliquely inflated in fruit; flowers lacking bracteoles (with bracteoles in T. glomeratum), not resupinate (i.e., the banner petal uppermost) 9a. Outer 2–4 (–6) flowers of each inflorescence bearing petals and fertile, the remaining flowers on the interior apetalous and sterile; inner flowers eventually elongating and producing a mass of woody, plumose stalks that surround the fruiting calyces and form a compact bur that is pushed against the soil surface or below it by downward growth of the peduncle; stems prostrate, stolon-like . . . . T. subterraneum 9b. All the flowers of each inflorescence bearing petals and fertile; fruiting inflorescence not resembling a compact bur and not pushed against soil surface or below it; stems ascending to erect (sometimes procumbent in T. echinatum and T. glomeratum) 10a. Corollas 3–4 mm long, definitely shorter than the calyx; leaflets narrowoblanceolate to oblanceolate or narrow-oblong, 3–5.5 times as long as wide [Fig. 652] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. arvense 10b. Corollas 4–16 mm long, ± as long as or longer than the calyx; leaflets of lower leaves obovate or obcordate to elliptic, 0.8–3 times as long as wide (sometimes oblanceolate and up to 4 times as long as wide in T. dichotomum) 11a. Corollas 4–6 mm long, ± as long as or shortly exceeding the calyx; inflorescences sessile or subsessile 12a. Flowers subtended by bracteoles; basal connate portion of the calyx glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. glomeratum 12b. Flowers without bracteoles; basal, connate portion of the calx pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. striatum 11b. Corollas (6–) 8–25 mm long, definitely longer than the calyx; inflorescences sessile or pedunculate 13a. Inflorescence sessile or on peduncles to 20 (–30) mm long, subtended either by a pair of opposite leaves or their expanded stipules or by an involucre (i.e., a ring of fused bracts); plants perennial (annual in T. dalmaticum) 14a. Inflorescence subtended by an involucre; corolla 8–10 mm long, pink; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. dalmaticum 14b. Inflorescence subtended by a pair of opposite leaves or their expanded stipules [Fig. 654]; corolla 12–20 mm long, red-purple to purplered (very rarely cream-white in T. pratense); plants perennial 15a. Distinct portion of the stipule broad-triangular, shorter than the connate portion; leaflets ovate or obovate to elliptic, 1.2–2.5 times as long as wide, frequently with light mottles on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. pratense

Fa b ac e a e   5 97

15b. Distinct portion of the stipule lanceolate, longer than the connate portion; leaflets elliptic to oblong, 2–3 times as long as wide, usually without light mottles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. medium 13b. Inflorescence borne on peduncles 10–70 mm long, not subtended by a pair of opposite leaves or an involucre (i.e., all the leaves alternate; except T. echinatum); plants annual 16a. Corollas 16–25 mm long; leaflets linear to elliptic-oblong, 20–50 × 2.5–5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. purpureum 16b. Corollas 5–12 mm long; leaflets oblanceolate or elliptic to obovate or obcordate, 8–30 × 3–12 mm 17a. Peduncles subtended by a pair of opposite leaves; calyx lobes stiff and spinose, especially in fruit . . . . . . . . . . . . . . . . . . . . . . . . . T. echinatum 17b. Peduncles subtended by a single leaf; calyx lobes pliable, not spinose 18a. Corollas red (rarely white); inflorescence conic to cylindric, (20–) 30–60 mm tall, at least the taller ones greater than 3 times as tall as wide; leaflets obovate to obcordate, 0.8–1.2 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. incarnatum 18b. Corollas purple with white near the apex (or with the reverse coloration); inflorescence ovoid to short-cylindric, up to 30 mm tall, less than 3 times as tall as wide; leaflets oblanceolate or obovate to elliptic, 1.5–4 times as long as wide . . . . . . . . . . . . . T. dichotomum 1. Trifolium arvense L. E Fig. 652 rabbit-foot clover. CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, streets, waste areas. 2. Trifolium aureum Pollich E

Fig. 652  Inflorescence and upper leaves of Trifolium arvense.

palmate hop clover. Trifolium agrarium L. • CT, MA, ME, NH, RI, VT; nearly throughout. Roadsides, fields, waste areas. 3. Trifolium campestre Schreb. E Fig. 653 pinnate hop clover. Trifolium procumbens, sensu L. 1755, not L. 1753 • CT, MA, ME, NH, RI, VT. Roadsides, fields, waste areas. 4. Trifolium dalmaticum Vis. E Balkan clover. MA. Dumps, waste areas. 5. Trifolium dichotomum Hook. & Arn. E branched Indian clover. Trifolium albopurpureum Torr. & Gray var. dichotomum (Hook. & Arn.) Isely; T. macraei Hook. & Arn. var. dichotomum (Hook. & Arn.) Brewer ex S. Wats. • MA. Wool waste. 6. Trifolium dubium Sibthorp E lesser hop clover. CT, MA, ME, RI, VT. Fields, roadsides, lawns, waste areas. 7. Trifolium echinatum Bieb. E prickly clover. Trifolium supinum Savi • MA. Wool waste. 8. Trifolium fragiferum L. E strawberry clover. VT. Low fields and other damp, disturbed areas. 9. Trifolium glomeratum L. E clustered clover. MA. Wool waste. 10. Trifolium hybridum L. E alsike clover. Trifolium elegans Savi; T. hybridum L. ssp. elegans (Savi) Aschers. & Graebn.; T. hybridum L. var. elegans (Savi) Boiss.; T. hybridum L. var. pratense Rabenh. • CT, MA, ME, NH, RI, VT; nearly throughout. Fields, lawns, roadsides.

Fig. 653  Leaf of Trifolium campestre.

5 98   tricolpate s

11. Trifolium incarnatum L. E crimson clover. Trifolium incarnatum L. var. elatius Gibelli & Belli • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 12. Trifolium medium L. E zigzag clover. MA, ME, VT. Fields, dumps. 13. Trifolium pratense L. E Fig. 654 red clover. Trifolium pratense L. var. sativum (P. Mill.) Schreb. • CT, MA, ME, NH, RI, VT; throughout. Fields, roadsides, lawns, shorelines, waste areas. 14. Trifolium purpureum Lois. E Fig. 654  Inflorescence and subtending leaves of Trifolium pratense.

purple clover. MA. Wool waste. 15. Trifolium repens L. E white clover. CT, MA, ME, NH, RI, VT; throughout. Fields, roadsides, lawns, waste areas. 16. Trifolium resupinatum L. E Fig. 655 Persian clover. MA, VT. Fields, roadsides, waste areas. 17. Trifolium striatum L. E knotted clover. MA. Fields, roadsides. 18. Trifolium subterraneum L. E burrowing clover. MA. Wool waste. 19. Trifolium tomentosum L. E woolly clover. MA. Yards, waste areas.

Fig. 655  Inflorescence of Trifolium resupinatum with resupinate corolla, so that the banner petal is lowermost.

Trigonella Trigonella ramosa L. was reported from MA by Knowlton and Deane (1918), but specimens are unknown. 1. Trigonella corniculata (L.) L. E cultivated fenugreek. MA. Ballast.

Ulex 1. Ulex europaeus L. E common gorse. MA; coastal plain. Open, sandy areas and disturbed places.

Vicia Steele and Wojciechowski (2003) found Vicia to be paraphyletic in regard to closely related genera (e.g., Lathyrus, Lens, Pisum). Further work may show that Vicia needs to be combined or dismantled in order that monophyletic taxa be recognized. 1a. Flowers solitary or in inflorescences of 2–50, in either case borne on a long peduncle 1–10.5 cm long [Fig. 656]; style pubescent all around the tip or glabrous in V. hirsuta 2a. Corollas 2.5–7 (–8) mm long, white to light purple; flowers solitary or in racemes of 2–5 (–7); legume 6–13 mm long 3a. Legumes hirsute, with (1–) 2 (–3) seeds, obliquely tapering from the sutures and pointed at the tip; lobes of the calyx ± equal length; leaves with (8–) 10–16 leaflets . . . . . V. hirsuta 3b. Legumes glabrous, with usually 4 seeds, equally rounded from the sutures and blunt at the tip; lobes of the calyx distinctly unequal in length; leaves with 4–10 (–12) leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. tetrasperma

Fa b ac e a e   5 9 9

2b. Corollas (9–) 10–13 (–18) mm long, blue, white and blue, or rarely entirely white; flowers in racemes of 10–50 [Fig. 656]; legume 15–40 mm long 4a. Plants perennial; calyx not gibbous or saccate on the upper side, the pedicel appearing to attach to the basal part of the flower; upper calyx lobes broad-triangular; limb of the banner petal ± as long as the claw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. cracca 4b. Plants annual or biennial; calyx gibbous or saccate on the upper side, the pedicel appearing to attach to the underside of the flower; upper calyx lobes lanceolate to linear-triangular; limb of the banner petal ca. half as long as the claw . . . . . . . . V. villosa 1b. Flowers solitary or paired, sessile or borne on a peduncle up to 1 cm long, borne in the axils of leaves [Fig. 657]; style pubescent on the abaxial (i.e., outer or lower) surface near the apex 5a. Terminal leaflet absent, not modified into a tendril; legumes 5–20 (–30) cm long; leaflets 5–10 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. faba 5b. Terminal leaflet of at least the upper leaves modified into a tendril; legume 1.5–6 cm long; leaflets 0.8–3 (–3.5) cm long 6a. Calyx 3.5–4.5 mm long; corollas 5–6 mm long; tendrils unbranched (sometimes absent on some leaves); legumes 15–25 (–30) × 3–4 mm . . . . . . . . . . . . . . . . V. lathyroides 6b. Calyx 4–13 mm long; corollas 10–30 mm long; tendrils generally branched [Fig. 657]; legumes 20–45 × 3.5–11 mm 7a. Banner petal pubescent on abaxial (i.e., outer) surface; stipules entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. pannonica 7b. Banner petal glabrous on abaxial surface; stipules entire, lobed, or toothed 8a. Corollas yellow, commonly with purple streaks, drying drab light purple, 25–30 (–35) mm long; legumes pubescent . . . . . . . . . . . . . . . . . . . . . . . . V. grandiflora 8b. Corollas light purple to white, 10–25 (–30) mm long; legumes glabrous or pubescent 9a. Calyx actinomorphic or nearly so, the lower lobes scarcely longer than the upper lobes; peduncle and inflorescence axis undeveloped, the inflorescence with 1 or 2 (–3) flowers [Fig. 657]; legumes sessile; plants annual . . . . . . V. sativa 9b. Calyx zygomorphic, the lower lobes distinctly longer than the upper lobes; inflorescence subsessile or shortly peduncled, the axis somewhat developed and up to 10 mm long, with 2–7 contiguous flowers; legumes on a stipe ca. 1.5 mm long; plants perennial from rhizomes . . . . . . . . . . . . . . . . . . . . . . . V. sepium 1. Vicia cracca L. ssp. cracca E Fig. 656 bird vetch. CT, MA, ME, NH, RI, VT; nearly throughout. Fields, roadsides, lawn edges. 2. Vicia faba L. E broad vetch. CT, MA, ME, VT. Fields, gardens, areas of cultivation. 3. Vicia grandiflora Scop. E large yellow vetch. Vicia grandiflora Scop. var. kitaibeliana W.D.J. Koch • MA. Fields, roadsides, waste areas. 4. Vicia hirsuta (L.) S.F. Gray E tiny vetch. Ervum hirsutum L. • CT, MA, ME, NH, RI. Fields, roadsides, waste areas, railroads. 5. Vicia lathyroides L. E spring vetch. MA. Fields, roadsides, waste areas. 6. Vicia pannonica Crantz E Hungarian vetch. CT. Fields, roadsides, waste areas.

Fig. 656  Inflorescence and leaf of Vicia cracca.

60 0   tricolpate s

7. Vicia sativa L. E Fig. 657 common vetch.  7a. Vicia angustifolia L.; V. angustifolia L. var. segetalis (Thuill.) W.D.J. Koch; V. sativa L. var. angustifolia (L.) Ser.; V. sativa L. var. nigra L.; V. sativa L. var. segetalis (Thuill.) Ser.; 7b. Vicia sativa L. var. linearis Lange • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, yards, gardens, shorelines, streets. 1a. Leaflets oblong-lanceolate to linear, 1.5–6 (–7) mm wide, 4–10 times as long as wide [Fig. 657]; calyx 7–11 (–12) mm long; corolla pink-purple to white, 10–18 mm long; legume black at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7a. Vicia sativa ssp. nigra (L.) Ehrh. 1b. Leaflets narrow-obovate to oblong, 4–10 mm wide, 2–5 (–7) times as long as wide; calyx 10–15 mm long; corolla pink-purple, 18–25 (–30) mm long; legume light brown to brown (less frequently black) at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7b. Vicia sativa ssp. sativa Variety nigra is known from CT, MA, ME, NH, RI, VT. Variety sativa is known from CT, MA, ME, NH, VT. Fig. 657  Inflorescence and leaf of Vicia sativa ssp. nigra.

8. Vicia sepium L. E hedge vetch. Vicia sepium L. var. montana W.D.J. Koch • ME, NH, VT. Fields, roadsides, waste areas. 9. Vicia tetrasperma (L.) Schreb. E four-seeded vetch. Ervum tetraspermum L.; Vicia tetrasperma (L.) Schreb. var. tenuissima Druce • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, railroads, waste areas. 10. Vicia villosa Roth E hairy vetch.  10a. Vicia dasycarpa Ten.; Vicia varia Host • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 1a. Plants glabrate to pubescent with subappressed to spreading hairs shorter than 1 mm; upper lobe of calyx lanceolate to narrow-lanceolate, 1–2 (–2.4) mm long; racemes common with 10–20 loosely arranged flowers . . . . . . . . . . . . . 10a. Vicia villosa ssp. varia (Host) Corb. 1b. Plants conspicuously villous with hairs 1–2 mm long; upper lobe of calyx acicular, 2–4 mm long; racemes commonly with more than 20 crowded flowers . . . 10b. Vicia villosa ssp. villosa Subspecies varia is known from MA, ME. Subspecies villosa is known from CT, MA, ME, NH, RI, VT.

Wisteria 1a. Ovary and legume glabrous; pedicels 5–10 (­–15) mm long; corollas 15–20 mm long; legumes 10–12 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. frutescens 1b. Ovary minutely villous and legume initially villous, retaining some pubescence at least along one or both sutures; pedicels mostly 15–20 mm long; corollas 15–25 (–27) mm long; legumes 20–25 mm wide 2a. Leaves with 13–19 leaflets; corollas 15–20 mm long; racemes 20–50 (–70) × 6–7 cm, with flowers developing from base to apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. floribunda 2b. Leaves with 7–13 leaflets; corollas 20–25 (–27) mm long; racemes 10–20 (–30) × 7–9 cm, with flowers opening nearly simultaneously . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . W. sinensis 1. Wisteria floribunda (Willd.) DC. E Japanese wisteria. Glycine floribunda Willd.; Kraunhia floribunda (Willd.) Taubert; Rehsonia floribunda (Willd.) Stritch • MA, ME, NH. Forest edges and fragments, areas of habitation. 2. Wisteria frutescens (L.) Poir. E American wisteria. Glycine frutescens L.; Kraunhia frutescens (L.) Greene; K. macrostachya (Torr. & Gray) Small; Wisteria frutescens (L.) Poir. var. macrostachya Torr. & Gray; W. macrostachya (Torr. & Gray) Nutt. ex B.L. Robins. & Fern. • CT, MA. Forest edges and fragments, areas of habitation. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI. 3. Wisteria sinensis (Sims) DC. E Chinese wisteria. Glycine sinensis Sims; Rehsonia sinensis (Sims) Stritch • CT, MA, RI, VT. Forest edges and fragments, areas of habitation.

Fagac e a e   6 0 1

Fagaceae 1a. Nuts triangular in cross-section; pith terete or obscurely angled in cross-section; winter buds slender-fusiform, 8–19 mm long; bark smooth and light gray, even in age (unless roughened by cankers from disease); staminate flowers in dense, spherical clusters . . . . . . . . . . . . . . . . . . Fagus 1b. Nuts terete or compressed; pith with 5 prominent points; winter buds ovoid to globose, up to 10 mm long; bark gray to brown, furrowed in age; staminate flowers in slender aments 2a. Carpellate flowers numbering 2–4 per involucre; involucre covered with spine-like bracts [Fig. 658], enclosing 1–3 nuts at maturity; winter buds not clustered toward distal end of twig . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Castanea 2b. Carpellate flowers 1 per involucre; involucre not appearing spiny, enclosing 1 nut at maturity; winter buds clustered toward distal end of twig [Fig. 663] . . . . . . . . . . . . . . Quercus

Castanea 1a. Leaf blades abaxially with multicellular, scale-like glands that may be embedded on blade surface 2a. Leaf blades abaxially ± glabrous soon after expansion (sometimes with some simple hairs along the veins); winter buds glabrous; petioles mostly 10–30 mm long . . . . C. dentata 2b. Leaf blades abaxially sparsely to densely stellate pubescent (sometimes becoming glabrate later in season); winter buds pubescent; petioles mostly longer than 30 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sativa 1b. Leaf blades abaxially lacking multicellular glands 3a. Involcure subtending 2 or 3 nuts, with pubescent spine-like bracts; nuts mostly 20–30 mm wide; petioles 10–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. mollissima 3b. Involucre subtending a solitary nut (rarely 2), with usually glabrous spine-like bracts; nuts 7–19 mm wide; petioles 3–7 (–10) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pumila 1. Castanea dentata (Marsh.) Borkh. N Fig. 658 American chestnut. Fagus-castanea dentata Marsh. • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic, deciduous to mixed evergreen-deciduous forests. 2. Castanea mollissima Blume E Chinese chestnut. CT, MA, RI. Forest fragments and borders, roadsides, areas of habitation. 3. Castanea pumila P. Mill. E dwarf chestnut. Castanea alnifolia Nutt.; C. alnifolia Nutt. var. floridana Sarg.; C. ashei (Sudworth) Sudworth; C. floridana (Sarg.) Ashe; C. pumila P. Mill. var. ashei Sudworth • MA. Forest fragments and borders, roadsides, areas of habitation. 4. Castanea sativa P. Mill. E Spanish chestnut. MA. Forest fragments and borders, roadsides, areas of habitation.

Fagus 1a. Leaf blades crenate-dentate, with 4–9 pairs of lateral veins; prickles of carpellate involucre ± straight to upcurved, relatively thinner and softer . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. sylvatica 1b. Leaf blades serrate, with 9–14 pairs of lateral veins; prickles of carpellate involucre ± straight to recurved, relatively stouter and firmer . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. grandifolia

Fig. 658  Leaf and carpellate involucre of Castanea dentata.

602  tricolpates

1. Fagus grandifolia Ehrh. N American beech. Fagus grandifolia Ehrh. var. caroliniana (Loud.) Fern. & Rehd. • CT, MA, ME, NH, RI, VT; throughout. Dry-mesic to mesic, deciduous or mixed evergreen-deciduous forests. 2. Fagus sylvatica L. E European beech. CT, MA, ME, RI. Forest fragments and borders, roadsides, abandoned homesteads, areas of habitation.

Quercus The following identification key has been written for mature sun leaves. Due to the differences in shape and pubescence between sun and shade leaves, adjustments may be needed if shade leaves are used. References: Palmer (1948), Miller and Lamb (1985), Jensen (1997), Nixon and Muller (1997). 1a. Leaf blades with bristle-tipped teeth or lobes (entire in Q. imbricaria and Q. phellos, which have a bristle-tipped blade apex) [Fig. 662]; bracts of carpellate involucre flattened, lacking darkened or pubescent calluses at the base; fruits maturing in second year, therefore, 2 sizes of fruit may be present on a given plant; inner surface of endocarp densely tomentose 2a. Leaf blades entire 3a. Leaf blade ovate to elliptic or obovate, 15–75 mm wide; petioles 10–20 mm long; carpellate involucres 5–9 mm tall and 10–18 mm in diameter; nuts 10–18 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. imbricaria 3b. Leaf blade linear to narrow-elliptic, 10–25 mm wide; petioles 2–4 (–6) mm long; carpellate involucres 3–6.5 mm tall and 7.5–11 mm in diameter; nuts 6.5–10 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. phellos 2b. Leaf blades lobed 4a. Petioles (8–) 10–25 mm long; nut 8–11 mm wide, the outer surface pubescent; leaf blades abaxially tomentose, with triangular to ovate lobes; short to tall shrubs up to 5 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. ilicifolia 4b. Petioles 20–70 mm long; nut 9–21 mm wide, the outer surface glabrous or nearly so; leaf blades abaxially glabrous or with hairs along the veins and/or in the axils of veins (infrequently with patches of tomentum on the surface in Q. velutina), with oblong or rectangular lobes; trees to 30 m tall 5a. Carpellate involucre pubescent on the inner surface, the marginal bracts loose and projecting, forming a fringe around the nut; terminal winter buds 7–10 mm long, the scales pubescent on the abaxial surface [Fig. 663] . . . . . . . . . . . . . . . . . Q. velutina 5b. Carpellate involucre glabrous on the inner surface or with a ring of hairs around the nut scar, the marginal bracts covered and concealed, not forming a fringe; terminal winter buds 3–7 mm long, the scales glabrous on the abaxial surface or pubescent only near the apex 6a. Terminal winter bud pubescent in the apical half, noticeably 5-angled in cross-section; carpellate involucre concealing ⅓ to ½ of the nut; nut with 1 or more concentric rings of fine pits near the apex [Fig. 659] . . . . . . . . . . . . . . . . Q. coccinea 6b. Terminal winter bud glabrous or with a few hairs near the apex, terete to obscurely angled in cross-section; carpellate involucre concealing ⅕ to ⅓ of the nut; nut lacking rings of fine pits near apex 7a. Sinuses of leaf blade generally extending more than ½ the distance from the tips of the lobes to the midrib; nut 10–16 mm long; carpellate involucre 9.5–16 mm in diameter; terminal winter buds bluntly pointed at apex, 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. palustris

Fagac e a e   6 03

7b. Sinuses of leaf blade generally extending less than ½ the distance from the tips of the lobes to the midrib; nut 15–30 mm long; carpellate involucre 18–30 mm in diameter; terminal winter buds sharply pointed at apex, 4–7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. rubra 1b. Leaf blades with rounded to acutely pointed teeth or lobes (the teeth with an apical papilla in Q. muehlenbergii and Q. prinoides) [Fig. 660]; bracts of carpellate involucre thickened and/or with darkened or pubescent calluses at the base; fruits maturing in first year, therefore, only 1 size of fruit present on a given plant; inner surface of endocarp glabrate or minutely pubescent near base and apex 8a. Terminal and axillary winter buds with persistent, filiform stipules; carpellate involucre with irregularly spreading scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. cerris 8b. Winter buds without persistent stipules; carpellate involucre with closely appressed scales 9a. Leaf blades with prominent lobes, at least some of the sinuses extending more than ⅓ the distance from the tip of the lobes to the midrib; mature leaf blades abaxially glabrous or pubescent 10a. Mature leaf blades abaxially glabrous or essentially so (at least appearing so without high magnification); branchlets glabrous 11a. Leaf blades cuneate at the base, on petioles 8–25 mm long; nut subsessile or on a peduncle to 25 (–40) mm long; carpellate involucre concealing ¼ (rarely to ⅓) of the nut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. alba 11b. Leaf blades cordate at the base, on petioles 4–8 mm long; nut on a peduncle (25–) 35–65 (–100) mm long; carpellate involucre concealing ⅓ to ½ (or more) of the nut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. robur 10b. Mature leaf blades abaxially with simple and/or stellate hairs; branchlets often pubescent 12a. Leaf blades typically with 1 or more deep sinuses near or below the middle of the blade that extend more than ½ distance from the tips of the lobes to the midrib, the distal portion of the blade with shallower sinuses or merely with tooth-like lobes [Fig. 660]; carpellate involucre (8–) 15–50 mm tall, the scales terminated by soft awns, these forming a loose, marginal fringe; branches often with corky wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. macrocarpa 12b. Leaf blades usually either ± regularly lobed or with deeper sinuses toward the apex, in either case lacking a pandurate shape; carpellate involucre 7–15 (–18) mm tall, the scales not projecting to form a fringe; branches lacking corky wings 13a. Leaf blades often cruciform (i.e., with 2 larger lobes that project at ± right angles from the midrib), abaxially with minute, stellate hairs; peduncle up to 6 (–40) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. stellata 13b. Leaf blades variably lobed, but never cruciform, abaxially with both minute, stellate hairs and taller, erect, 1- to 4-rayed hairs; peduncles (20–) 40–70 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Q. bicolor 9b. Leaf blades with short, tooth-like lobes, the sinuses extending less than ⅓ the distance from the tip of the lobes to the midrib; mature leaf blades abaxially with simple and/or stellate-branched hairs 14a. Peduncle (20–) 40–70 mm long; leaf blades abaxially with both minute, stellate hairs and taller, erect, 1- to 4-rayed hairs across the surface; plants of swamps, riparian forests, and other poorly drained habitats . . . . . . . . . . . . . . . (in part) Q. bicolor 14b. Peduncle essentially absent or up to 20 (–25) mm long; leaf blades abaxially with only minute, stellate hairs across the surface, though taller, erect hairs are found along the veins in Q. montana (taller, erect, red hairs often found across the abaxial

604 tricolpate s

surface in coastal MA populations of Q. prinoides); plants of dry, well-drained ridges, slopes, and plains 15a. Leaf blade abaxially with both minute, stellate hairs and taller, erect simple or fascicled hairs along the veins, each tooth-like lobe lacking a papilla-like point; bark dark gray to dark brown with deep furrows, neither flaky nor scaly; carpellate involucre 18–25 mm in diameter; peduncle 8–20 (–25) mm long . . . . . . Q. montana 15b. Leaf blades abaxially with sparse to dense, minute stellate hairs only (coastal MA populations of Q. prinoides often have erect, reddish hairs in addition the stellate ones), each tooth-like lobe terminating with a papilla-like point [Fig. 661]; bark gray, without deep furrows, flaky or scaly on mature stems; carpellate involucre 8–20 (–22) mm in diameter; peduncle up to 8 mm long 16a. Leaf blades with (9–) 10–14 (–16) pairs of secondary veins; small trees to 18 m tall, usually found on limestone and marble hills and outcrops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. muehlenbergii 16b. Leaf blades with 5–8 (–9) pairs of secondary veins [Fig. 661]; colonial shrubs rarely exceeding 3 m in height, found usually on dry soils of ridges and sandy plains . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q. prinoides 1. Quercus alba L. N eastern white oak. CT, MA, ME, NH, RI, VT; generally absent in extreme northern New England. Dry-mesic to mesic forests and woodlands, infrequently found in wet-mesic soils of seepage swamps and riparian forests. ‌1 × 2. Quercus ×jackiana Schneid. is a rare oak hybrid known from CT, MA, RI. The leaf blades are intermediate in nature, generally with an outline more similar to Q. alba but with some pubescence abaxially (as in Q. bicolor). The nut is sometimes nearly sessile or other times on long peduncles. ‌1 × 7. Quercus ×bebbiana Schneid. is a very rare oak hybrid known from CT, VT. This nothospecies shows varying degrees of intermediacy between the parental taxa, especially in leaf blade outline. ‌ 8. Quercus ×saulii Schneid. is a rare oak hybrid known from CT, MA, RI, VT. The leaf 1 × blades most resemble Q. montana; however, the lobes are more pronounced and the sinuses between them are deeper. ‌ 12. Quercus ×faxonii Trel. is a very rare oak hybrid known from MA. It displays 1 × intermediacy in leaf blade outline and pubescence and peduncle length (see identification key for details). In the absence of fruits, specimens of this hybrid can be ‌saulii (1 × 8). confused with Quercus × 1‌ × 13. Quercus ×bimundorum Palmer is a very rare oak hybrid known from MA. It is intermediate between the parental taxa. The leaf blades are subcordate at the base, and the petioles are 5–10 mm long. ‌1 × 15. Quercus ×fernowii Trel. is a very rare oak hybrid known from MA. It displays intermediacy in leaf blade outline and pubescence and peduncle length (see identification key for details). 2. Quercus bicolor Willd. N swamp white oak. CT, MA, ME, NH, RI, VT; largely absent from extreme northern New England, though extending north in the Lake Champlain Valley of VT. Swamps, riparian forests, lacustrine forests, forest borders. ‌ × 7. Quercus ×schuettei Trel. is a very rare oak hybrid known from VT. It is primarily 2 known from the lacustrine floodplain of Lake Champlain where both parents grow together. The hybrid’s leaves are generally a little smaller and show less prominent lobes than those of Q. macrocarpa. Further, the pubescence of the leaf blades is less dense and not as persistent as that of Q. bicolor.

Fagac e a e   6 0 5

3. Quercus cerris L. E European turkey oak. MA. Forest borders and fragments, roadsides, areas of habitation. 4. Quercus coccinea Muenchh. N Fig. 659 scarlet oak. Quercus coccinea Muenchh. var. tuberculata Sarg. • CT, MA, ME, NH, RI, VT; mainly absent from extreme northern New England, though disjunct in Chittenden County, VT. Drymesic forests and woodlands. ‌4 × 5. Quercus ×robbinsii Trel. is a rare oak hybrid known from MA, RI. It has leaf blades that are sparsely stellate-pubescent abaxially and some blades with somewhat triangular lobes, petioles (8–) 17–31 mm long, nuts mostly 13–14 × 9–13 mm that are sparsely pubescent on the exterior, and pubescent terminal winter buds 3–4.4 mm long. ‌ × 14. Quercus ×benderi Baenitz is a rare oak hybrid known from MA. It is difficult 4 to distinguish morphologically given the close similarity of the parental taxa. Unlike Q. rubra, this plant shows yellow fall foliage. Further, the nuts are frequently sterile.

Fig. 659  Nut of Quercus coccinea showing concentric rings of minute pits around the style remnant.

‌ × 16. Quercus ×fontana Laughlin is a rare oak hybrid known from MA. It has 4 pubescent terminal winter buds and a pubescent inner surface of the carpellate involucre, similar to Q. velutina, but the terminal winter buds are too short (4–6 mm long) and the leaf blades have lobes that are too long (sinuses extending more than half way from the tips of the lobes to the midrib of the leaf). 5. Quercus ilicifolia Wangenh. N scrub oak. CT, MA, ME, NH, RI, VT; generally absent from the northern portion of northern New England states. Xeric to dry-mesic woodlands and barrens. 5 ‌ × 14. Quercus ×fernaldii Trel. is a rare oak hybrid known from MA, ME. It shows intermediacy in leaf blade outline and pubescence, terminal winter bud pubescence (they are sparsely pubescent), and fruit size (see identification key for details). 5 ‌ × 16. Quercus ×rehderii Trel. is a rare oak hybrid known from MA, ME, NH, RI. It is best diagnosed using the leaf blades, which are very different between the parental taxa. The plants are frequently shrubby (as in Q. ilicifolia), but sometimes are more arboreal than is typical for Q. ilicifolia. 6. Quercus imbricaria Michx. I shingle oak. MA. Forest fragments and borders, areas of habitation. ‌6 × 16. Quercus ×leana Nutt. is a very rare oak hybrid known from MA. The leaf blades look much like Q. imbricaria but have broad, undulate lobes that are infrequently tipped by a small bristle (but most leaf blades lack any marginal lobes with a bristle-tip). The report of this nothospecies in VT by Kartesz (1999) was erroneous. 7. Quercus macrocarpa Michx. N Fig. 660 burr oak. Quercus macrocarpa Michx. var. depressa (Nutt.) Engelm.; Q. mandanensis Rydb. • CT, MA, ME, NH, VT; also reported from RI by Seymour (1982), but specimens are unknown. Swamps, riparian and lacustrine forests, also found on dry-mesic to mesic soil of forests in regions of high-pH bedrock (e.g., VT). This species is sometimes confused with Quercus bicolor given they often grown in similar locations. In addition to characters used in the identification key, peduncle length will assist with determinations: (0–) 6–20 (–25) mm long in Q. macrocarpa and (20–) 40–70 mm long in Q. bicolor. 8. Quercus montana Willd. N mountain chestnut oak. Quercus prinus L. • CT, MA, ME, NH, RI, VT; mainly limited to the southern portion of the northern New England states, but extending north in VT in the Lake Champlain Valley. Dry-mesic to mesic forests and woodlands, often associated with ridges and rocky slopes. 9. Quercus muehlenbergii Engelm. N yellow chestnut oak. Quercus acuminata (Michx.) Sargent; Q. prinoides Willd. var. acuminata (Michx.) Gleason; Q. prinus L. var. acuminata Michx. • CT, MA, VT; western New England. Ridges,

Fig. 660  Leaf blade of Quercus macrocarpa showing prominent sinuses near center.

60 6   tricolpate s

hills, and outcrops on high-pH bedrock. Quercus muehlenbergii and Q. prinoides are frequently confused. The former is a small tree and does not begin producing fruits until it reaches 3 m or more of height. Quercus prinoides is a colonial shrub that does produce fruits often when the plants are shorter than 0.5 m. 10. Quercus palustris Muenchh. n pin oak. CT, MA, ME, RI, VT. Swamps, riparian forests, pond shores. This species is native to southern New England but introduced (as a planting) in ME and VT. 10 ‌ × 16. Quercus ×vaga Palmer & Steyermark is a very rare oak hybrid known from MA. It tends to show the bark and growth habit of Q. palustris along with the involucral bracts of Q. velutina. The leaf blades also show tufts of pubescence in the axils of the primary lateral veins. 11. Quercus phellos L. Fig. 661  Leaf blade of Quercus prinoides showing lobes terminated by a papilla.

willow oak. CT. Forest fragments, roadsides. Dowhan (1979) reported a single mature tree was found in New London County, CT. The history of the tree is unknown (i.e., whether or not it was planted). Given the proximity of native occurrences (Long Island, NY), it is possible the tree is native to New England. 12. Quercus prinoides Willd. N Fig. 661 dwarf chestnut oak. Quercus prinoides Willd. var. rufescens Rehd. • CT, MA, NH, RI, VT; also reported from ME by Fernald (1950b), but specimens are unknown. Dry, sandy fields and roadsides, woodlands, rocky hillsides and ridges. An uncommon form of this species (formerly treated as var. rufescens) is known from New England that has red hairs on the leaf blades and branchlets. ‌12 × 15. Quercus ×stelloides Palmer is a very rare oak hybrid known from MA, RI. It is best diagnosed by using the leaf blades, which resemble most Q. stellata with 3 or 4 pairs of short, rounded or pointed lobes that are sometimes expanded and retuse at the apex and stellate-pubescent abaxially. It is normally a shrubby plant 1–2 m tall but sometimes grows to 4 m. 13. Quercus robur L. E English oak. MA, ME, NH, RI. Roadsides, forest fragments, abandoned homesteads, areas of habitation. 14. Quercus rubra L. N Fig. 662

Fig. 662  Leaf blade of Quercus rubra showing bristle-tipped lobes.

northern red oak. Quercus borealis Michx. f.; Q. borealis Michx. f. var. maxima (Marsh.) Ashe; Q. rubra L. var. ambigua (Gray) Fern.; Q. rubra L. var. borealis (Michx. f.) Farw. • CT, MA, ME, NH, RI, VT; nearly throughout. Dry-mesic to mesic forests and woodlands. ‌ × 16. Quercus ×hawkinsii Sudworth is rare oak hybrid known from CT, MA, ME. 14 However, given the frequency of sympatry of the parental taxa, this nothospecies may be more common than has been reported. The plant generally resembles Q. rubra; however, the winter bud scales are pubescent and the leaf blades often with some persistest pubescence along the veins or in the axils of the veins. The carpellate involucre generally is closer to Q. velutina in its shape and pubescence on the inner surface. 15. Quercus stellata Wangenh. N post oak. CT, MA, RI; coastal plain. Dry-mesic woodlands, fields, and barrens, often associated with rocky ridges and slopes. 16. Quercus velutina Lam. N Fig. 663

Fig. 663  Pubescent winter buds of Quercus velutina.

black oak. CT, MA, ME, NH, RI, VT; absent from much of northern and eastern ME. Dry-mesic forests and woodlands. This species is very similar to Quercus rubra. In addition to the characters mentioned in the key, Q. velutina has slightly darker, more furrowed bark and often has shallower leaf blade sinues (relative to the blade width) than Q. rubra.

F r a n ke ni ac e a e   6 07

Frankeniaceae Frankenia 1. Frankenia pulverulenta L. E European sea-heath. MA. Wool waste.

Gentianaceae Reference: Gillett (1963). 1a. Leaves of the stem reduced to scales 0.5–4.5 mm long; flowers 2.5–4 (–7) mm long; lobes of the corolla imbricate in bud . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bartonia 1b. Leaves of the stem foliaceous [Fig. 669], (5–) 10–150 mm long; flowers 5–55 mm long; lobes of the corolla convolute in bud 2a. Corolla rotate, with lobes much longer than the basal, connate portion [Figs. 668, 669] 3a. Style absent, the stigmas decurrent on the sides of the ovary along the carpel sutures; stamens epipetalous; anthers basifixed . . . . . . . . . . . . . . . . . . . . . . Lomatogonium 3b. Style present, the stigmas elevated above the ovary; stamens free from the petals; anthers versatile 4a. Corolla with (4–) 5–12 lobes [Figs. 668, 669], the lobes lacking a prominent glandular spot; anthers recurving or coiling after anthesis . . . . . . . . . . . . . . . . . Sabatia 4b. Corolla with 4 (rarely 5) lobes, each lobe with a prominent, fringed, glandular spot; anthers remaining straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Frasera 2b. Corolla tubular, funnelform, campanulate, or salverform, with lobes shorter than the basal, connate portion 5a. Style evident, slender; corolla salverform, red-purple to pink to nearly white; anthers becoming spirally coiled 6a. Inflorescence a panicle-like or corymb-like cyme; style slightly bifid, divided for a short distance below the stigma lobes; stigma lobes semicircular to bluntly triangular from side view, fleshy; capsules narrow-cylindrical . . . . . . . . . . . . . . . . . . . . . Centaurium 6b. Inflorescence a spike-like cyme; style not divided beneath the stigma lobes; stigma lobes flabellate from side view, only slightly fleshy; capsules ellipsoid . . . . . . . Schenkia 5b. Style absent, inconspicuous, or short and stout; corolla cylindric, funnelform, or campanulate, blue to white or green to purple-green; anthers not spirally coiled 7a. Lobes of the corolla with conspicuous, basal spurs in the larger flowers; corolla green to purple-green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halenia 7b. Lobes of the corolla lacking spurs; corolla blue to white 8a. Sinuses of the corolla folded or plaited, often of a color or texture different from those of the corolla lobes; plants perennial; calyx with a continuous membrane or rim on the inside at the summit; nectary glands located at base of ovary . . . . . . Gentiana

60 8   tricolpate s

8b. Sinuses of the corolla neither folded nor plaited, of a color and texture similar to those of the lobes; plants annual or biennial; calyx without a membrane or with a membrane confined to the base of each sinus; nectary glands located at base of corolla 9a. Margins of the corolla lobes conspicuously fringed in the apical portion [Fig. 667]; corolla 35–55 mm long, usually 4-merous; ovules covering much of the inner surface of the ovary; calyx with a membrane on the inside at the base of each sinus; seeds papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gentianopsis 9b. Margins of the corolla lobes not fringed; corolla 10–23 mm long, commonly 5-merous; ovules confined to bands along each of the 2 sutures of the ovary; calyx without an internal membrane; seeds smooth . . . . . . . . . . . . . . Gentianella

Bartonia Leaf arrangement is an important character in Bartonia, but it has been given too much weight. It is important to realize that it does display variation, and rare individuals within a population will show different character states (e.g., B. virginica with alternate leaves). Other characters, such as corolla and anther morphology, need to be considered as well. When measuring the length of the stigmas, be sure to measure the basal portion that is decurrent onto the ovary. Timing is also important as the stigma does not reach its full length until midto late August. At this time, the ovary is rather plump and most of the anthers will have fallen from the stamens. Reference: Gillet (1959). 1a. Stigmas, including the basal lobes, (1.3–) 1.5–2.3 mm long at maturity; petals oblong, rounded to obtuse at the apex and with a short, but prominent, apiculus; anthers apiculate at the apex; capsules dehiscing from the apex to a point proximal to the stigmas; stems with opposite leaves [Fig. 664], the nodes becoming progressively more crowded near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. virginica 1b. Stigmas 0.8–1.5 mm long at maturity; petals lanceolate to oblong-lanceolate, acute at the apex and usually without a clear apiculus; anthers rounded at the apex or rarely with an apiculus in B. iodandra; capsules dehiscing from the apex to the base of the stigmas; stems with alternate leaves, the nodes not or scarcely becoming more crowded near the base 2a. Petals lanceolate and acute at the apex; filaments green to yellow-brown, except sometimes near attachement of anther where purple; anthers usually yellow, 0.3–0.5 (–0.6) mm long; branches of inflorescence ascending to spreading; stems often slender and lax, usually green; calyx essentially distinct  . . . . . . . . . . . . . B. paniculata 2b. Petals oblong-lanceolate, acute at the apex or infrequently with an obscure apiculus; filaments purple (rarely green to yellow-brown); anthers usually purple, (0.4–) 0.5–0.6 (–1) mm long; branches of inflorescence erect to strictly ascending [Fig. 664]; stems stouter, strict, usually purple or tinged with purple; calyx often shortly connate near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. iodandra 1. Bartonia iodandra B.L. Robins.

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purple screwstem. Bartonia paniculata (Michx.) Muhl. var. intermedia Fern.; B. paniculata (Michx.) Muhl. ssp. iodandra (B.L. Robins.) J. Gillett; B. paniculata (Michx.) Muhl. var. iodandra (B.L. Robins.) Fern.; B. paniculata (Michx.) Muhl. var. sabulonensis (Fern.) Fern.; B. virginica (L.) B.S.P. var. sabulonensis (Fern.) Boivin • MA, ME, NH, RI. Swamps, peatlands, abandoned borrow pits, meadows. Bartonia iodandra has been treated as a subspecies or variety of B. paniculata by most authors, likely due to similarities in leaf arrangement and stigma length. However, it possesses many character states that are transitional to B. virginica, including stem stature and coloration, petal shape and apex, anther apex, stamen color, and seed size. Gillett (1959) noted its probable hybrid origin. As such, it is inappropriate to ally it with one of its putative parents. Its occurrence in Newfoundland (beyond the range of its parents) and production of wellformed seeds support its recognition as a hybrid-derived species (rather than as an F₁ hybrid).

G e nt i a n ac e a e   6 0 9

2. Bartonia paniculata (Michx.) Muhl. N twining screwstem. Bartonia lanceolata Small; B. virginica (L.) B.S.P. var. paniculata (Michx.) Boivin; Centaurella paniculata Michx. • CT, MA, RI. Swamps, peatlands, abandoned borrow pits, meadows. A specimen determined as this species is problematic—Weatherby 6879 (NEBC!). It shows the alternate leaves, short stigmas, and short yellow anthers of B. paniculata but has oblong to oblong-lanceolate petals, purple filaments, and relatively stout stems. Further, the apex of the petals shows a clear apiculus. The specimen requires further study and may be of recent hybrid origin. 3. Bartonia virginica (L.) B.S.P. N Fig. 664 Virginia screwstem. Sagina virginica L. • CT, Ma, Me, NH, RI, VT. Fields, meadows, swamps, wet sand and peat about temporary pools.

Centaurium See Mansion (2004) for the dismantling of this group (the segregate genus Schenkia also known from New England). 1a. Flowers on pedicels mostly (1.5–) 2–5 mm long; calyx nearly equaling the length of the basal, connate portion of the corolla; corolla lobes 2–4 mm long; plants lacking a basal rosette of leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pulchellum 1b. Flowers sessile or subsessile; calyx usually ca. ½ as long as the basal, connate portion of the corolla; corolla lobes usually 5–6 mm long; plants with a basal rosette of leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. erythraea 1. Centaurium erythraea Raf. E European centaury. Centaurium umbellatum, auct. non Gilib. • MA, VT; also reported from RI by George (1992), but specimens are unknown. Fields, roadsides, waste areas. 2. Centaurium pulchellum (Sw.) E.H.L. Krause E branched centaury. Gentiana pulchella Sw. • CT, MA, ME, VT. Fields, roadsides, waste areas.

Frasera 1. Frasera albicaulis Dougl. ex Griseb. ssp. nitida (Benth.) C.L. Hitchc. E white-stemmed elkweed. Frasera albicaulis Dougl. ex Griseb. var. nitida (Benth.) C.L. Hitchc.; F. nitida Benth.; Swertia albicaulis (Dougl. ex Griseb.) Kuntze var. nitida (Benth.) Jepson; S. nitida (Benth.) Jepson • MA. Dry soil of hillsides, forest fragments, and clearings.

Gentiana 1a. Leaf blades and lobes of the calyx prominently ciliolate (note: the cilia visible at low magnification); lobes of the corolla about as long as or shorter than the appendages in the sinuses of the corolla 2a. Appendages in the sinuses of the corolla 2- or 3-cleft, about as long as the corolla lobes [Fig. 666]; lobes of the corolla broad-rounded at the apex and about as wide as the plaits [Fig. 666]; calyx lobes usually broad-lanceolate to obovate . . . . . . . . . . . . . . . G. clausa 2b. Appendages in the sinuses of the corolla fimbriate, longer than the corolla lobes [Fig. 665]; lobes of the corolla truncate and often apiculate, narrower than the plaits [Fig. 665]; calyx lobes usually lanceolate to elliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. andrewsii 1b. Leaf blades and lobes of the calyx lacking cilia or with scattered, minute cilia; lobes of the corolla 1.5–5 times as long as the appendages in the sinuses of the corolla 3a. Involucral leaf blades ascending, linear to linear-lanceolate, 3–15 mm wide; lobes of the corolla 1.5–3 times as long as the appendages in the sinuses of the corolla; leaf blades narrow-lanceolate, dark green, not clasping at the base . . . . . . . . . . . . . . . . . . . . . . G. linearis

Fig. 664  Inflorescence and upper leaves of Bartonia virginica.

610  tricolpates

3b. Involucral leaf blades spreading, narrow-ovate to ovate, 15–30 mm wide; lobes of the corolla 3–5 times as long as the appendages in the sinuses of the corolla; leaf blades broad-lanceolate, light green, somewhat clasping at the base . . . . . . . . . . . . . G. rubricaulis 1. Gentiana andrewsii Griseb. var. andrewsii

Fig. 665  Unrolled corolla of Gentiana andrewsii showing truncate lobes and appendages with a truncate and fimbriate apex.

N C Fig. 665

Andrew’s bottle gentian. Dasystephana andrewsii (Griseb.) Small; Pneumonanthe andrewsii (Griseb.) W.A. Weber • CT, MA, VT; also reported from NH by Magee and Ahles (1999), but specimens are unknown. Meadows, fields, ditches. Many records of this plant (including at least some of those from NH) are referable to Gentiana clausa (personal observation and James Pringle, personal communication). 2. Gentiana clausa Raf. N Fig. 666 meadow bottle gentian. CT, MA, ME, NH, RI, VT. Meadows, mesic forests, clearings, riparian forests, roadside banks. 3. Gentiana linearis Froel. N narrow-leaved gentian. Dasystephana linearis (Froel.) Britt.; Gentiana saponaria L. var. linearis (Froel.) Griseb. • MA, ME, NH, VT. River banks and riverside meadows, fields, open, often peaty, wetlands, meadows. 4. Gentiana rubricaulis Schwein.

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red-stemmed gentian. Dasystephana grayi (Kusnez.) Britt.; Gentiana linearis Froel. ssp. rubricaulis (Schwein.) J. Gillett; G. linearis Froel. var. lanceolata Gray; G. linearis Froel. var. latifolia Gray • ME. Meadows, clearings, graminoid marshes, open rights-of-way. The report of Gentiana rubricaulis in VT by Seymour (1982) was erroneous. It was based on a specimen of G. clausa (Jenkins and Zika 1995). Fig. 666  Unrolled corolla of Gentiana clausa showing rounded lobes and appendages with a cleft apex.

Gentianella Reference: Gillett (1957). 1a. Corolla 10–15 mm long, with a fringe of hairs on the inside around the base of the lobes; lobes of the corolla 3–5 mm long, obtuse to acute at the apex . . . . . . . . . . . . . . . . . . G. amarella 1b. Corolla 15–23 mm long, without a fringe at the base of the lobes; lobes of the corolla 4–7 mm long, acute to acuminate at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. quinquefolia 1. Gentianella amarella (L.) Boerner ssp. acuta (Michx.) J. Gilbert

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northern dwarf-gentian. Amarella acuta (Michx.) Raf.; Gentiana acuta Michx.; G. amarella L. ssp. acuta (Michx.) Hultén; G. amarella L. var. acuta (Michx.) Herder; G. amarella L. var. stricta (Griseb.) S. Wats.; Gentianella acuta (Michx.) Hiitonen; G. amarella (L.) Boerner var. acuta (Michx.) Herder • ME, VT; northern portion of states. Open and forested river banks, subalpine gullies and brook sides, occurring in regions of high-pH bedrock and/or till. 2. Gentianella quinquefolia (L.) Small ssp. quinquefolia

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stiff dwarf-gentian. Gentiana amarelloides Michx.; G. quinquefolia L. • CT, MA, ME, NH, VT. Fields, pastures, roadsides, banks, pond shores, commonly in regions of high-pH bedrock.

Gentianopsis 1. Gentianopsis crinita (Froel.) Ma N Fig. 667 greater fringed-gentian. Anthopogon crinitum (Froel.) Raf.; Gentiana crinita Froel.; Gentianella crinita (Froel.) G. Don • CT, MA, ME, NH, RI, VT. Fields, meadows, roadsides, clearings.

Fig. 667  Corolla of Gentianopsis crinita.

G e nt i a nac e a e   6 1 1

Halenia 1. Halenia deflexa (Sm.) Griseb. ssp. deflexa N American spurred-gentian. Halenia heterantha Griseb.; H. deflexa (Sm.) Griseb. var. heterantha (Griseb.) Fern.; Swertia deflexa Sm.; Tetragonanthus deflexus (Sm.) Kuntze • MA, ME, NH, VT. Mesic forests and forest edges, fields, river banks and shores, clearings, trail edges. Rare forms of this species in New England lack spurs.

Lomatogonium 1. Lomatogonium rotatum (L.) Fries ex Fern.

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marsh-felwort. Gentiana rotata (L.) Froel.; Pleurogyne rotata (L.) Griseb.; Swertia rotata L. • ME; eastern coastal portion of the state. Coastal islands, pools, turfs, and open, rocky areas.

Sabatia References: Fernald (1916), Wilbur (1955). 1a. Perianth (8–) 9- to 12-merous [Fig. 668]; plants perennial, with elongate rhizomes terminating in basal rosettes of leaves 2a. Calyx lobes oblanceolate to spatulate (rarely linear), without a hyaline margin, 1–3 mm wide, with 3–5 distinct nerves; basal, connate portion of the calyx distinctly nerved or corrugated; plants primarily of saline and brackish marshes . . . . . . . . . . . . . . S. dodecandra 2b. Calyx lobes linear, with a thin, hyaline margin, 0.5–1.5 mm wide, with 1–3 obscure nerves; basal, connate portion of the calyx smooth or inconspicuously nerved; plants primarily of sandy and peaty pond shores of the coastal plain . . . . . . . . . . . . S. kennedyana 1b. Perianth (4–) 5 (–7)-merous [Fig. 669]; plants annual, biennial, or perennial from a short, branched caudex 3a. Basal, connate portion of the calyx prominently 5-nerved, each nerve rib-like and thinly membranous-winged; lateral veins of the calyx lobes developed more prominently than the midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. campestris 3b. Basal, connate portion of the calyx inconspicuously and/or finely nerved, the nerves neither rib-like nor thinly winged; lateral veins of calyx lobes developed equally or less prominently than the midvein 4a. Cyme dichasial (i.e., the axis appearing trichotomously branched because each node has two lateral branches in addition to the central axis); leaf blades of the middle and upper stem ovate to narrow-ovate, broad-rounded to cordate and often clasping at the base; stem strongly tetragonal, with conspicuous wing-angles 0.1–0.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. angularis 4b. Cyme monochasial (i.e., the axis appearing dichotomously branched because each node has a single, lateral branch in addition to the central axis) [Fig. 669]; leaf blades of the middle and upper stem oblong or linear to oblanceolate, narrowed but not clasping at the base; stem terete or irregularly ridge-angled 5a. Principal leaf blades linear to oblong or narrow-oval, widest at or below the middle, broadly sessile at the base, not darkening in drying; corolla usually not or only slightly exceeding the calyx; plants perennial from a short, branched caudex, primarily of sandy and peaty pond shores of the coastal plain . . . . . . S. campanulata 5b. Principal leaf blades elliptic to oblanceolate, widest at or above the middle, narrowly tapering to the base [Fig. 669], often darkening in drying; corolla usually distinctly surpassing the calyx; plants annual, usually from fibrous roots, primarily of saline and brackish marshes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. stellaris

61 2   tricolpate s

1. Sabatia angularis (L.) Pursh E common rose-gentian. Chironia angularis L. • CT, MA. Fields, clearings. 2. Sabatia campanulata (L.) Torr.

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slender rose-gentian. Chironia campanulata L.; Sabatia campanulata (L.) Torr. var. gracilis (Michx.) Fern.; S. gracilis (Michx.) Salisb. • MA. Coastal plain pond shores. 3. Sabatia campestris Nutt. E western rose-gentian. Sabatia nervosa Raf. • CT; also reported from ME by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, clearings. 4. Sabatia dodecandra (L.) B.S.P. var. dodecandra

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perennial rose-gentian. Chironia chloroides Michx.; C. dodecandra L.; Sabatia chloroides (Michx.) Pursh • CT; southern portion of state. Saline to brackish marshes. 5. Sabatia kennedyana Fern. Fig. 668  Flower of Sabatia kennedyana.

N C Fig. 668

Plymouth rose-gentian. Sabatia dodecandra (L.) B.S.P. var. kennedyana (Fern.) Ahles • MA, RI. Coastal plain pond shores. 6. Sabatia stellaris Pursh

N C Fig. 669

annual rose-gentian. Chironia amoena Raf.; C. stellata Muhl.; Sabatia amoena (Raf.) G. Don; S. maculata (Benth.) Benth. ex Gray; S. maritima Raf.; S. simulata Britt. • CT, MA, RI; coastal region of states. Saline and brackish marshes, brackish ponds, meadows adjacent to coastal marshes.

Schenkia 1. Schenkia spicata (L.) Mansion E spiked centaury. Centaurium spicatum (L.) Fritsch; Gentiana spicata L. • MA. Saline marshes.

Geraniaceae Fig. 669  Flowers and upper leaves of Sabatia stellaris.

1a. Leaf blades pinnately lobed or pinnately divided [Fig. 670]; stamens 10 per flower but the outer series of filaments sterile and lacking anthers; seeds smooth (though usually pubescent); carpel beaks separating from the style column, dehiscing from apex to base, the free portions of the carpel beaks becoming spirally coiled; carpel beaks pubescent on the adaxial (i.e., inner) surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erodium 1b. Leaf blades palmately lobed or palmately divided; stamens 10 per flower, all fertile (except G. pusillum with 3–5 sterile filaments); seeds reticulate or striate (smooth in 2 species); carpel beaks remaining partially adnate to the style column, dehiscing from base to apex [Fig. 672], the free portions of the carpel beaks merely outwardly curving; carpel beaks glabrous or nearly so on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geranium

Erodium A genus very similar to Geranium in floral and fruiting characteristics, differing primarily by features discussed in the key to the genera. Our species of Erodium are much easier to identify when collected in fruit. The sepals are generally accrescent, and measurements provided in the key are for fruiting specimens. The apical pits of the mericarp bodies are crucial for identification. The presence/absence of glands can often be assessed on immature fruiting material. However, the prescence/absence of furrows below the apical pits requires mature fruits. The furrows (when present) in most species are paired (but see E. moschatum), forming a transverse groove below the apical pit, and are separated from each other by a narrow septum (i.e., the paired furrows are on the same latitudinal line). Reference: Taylor (1993).

Ge r a ni ac e a e   6 13

1a. Leaf blades pinnately lobed, the blade segments not completely divided to the midrib or the midrib with a prominent wing of tissue (rarely divided near base of leaf, but then with only 1 or, very rarely, 2 pairs of distinct leaflets) 2a. Sepals 5–7 mm long; mericarps 3–5 mm long; carpel beaks 18–35 mm long; bracts of umbel ovate-orbicular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. malacoides 2b. Sepals 8–15 mm long; mericarps 7.5–11 mm long; carpel beaks (25–) 50–113 mm long; bracts of umbel lanceolate to ovate or triangular 3a. Apical pits of mericarp bodies stipitate-glandular within, without furrows below them; umbel with 3–10 flowers 4a. Carpel beaks 60–100 mm long; sepals 12–15 mm long; mericarp bodies 9–11 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) E. ciconium 4b. Carpel beaks 25–42 mm long; sepals 8–12 mm long; mericarp bodies 8–9 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) E. stephanianum 3b. Apical pits of mericarp bodies eglandular within, with 2 furrows below them (lacking furrows in E. cygnorum); umbel with 1–6 flowers 5a. Base of stem, as well as upper portion of stem, pubescent with glandular hairs; leaf blades oblong to ovate in outline, with usually 5 or more principal lobes; mericarp bodies with 2 furrows below the apical pits . . . . . . . . . . . . . . . . . . . . . E. botrys 5b. Base of stem pubescent with eglandular hairs or sometimes glabrate (upper portion of stem glandular-pubescent); leaf blades ovate in outline, with usually 3 principal lobes; mericarp bodies without furrows below the apical pits . . . . . . . . . . . . . . . . . . E. cygnorum 1b. Leaf blades pinnately divided into distinct leaflets [Fig. 670] 6a. Carpel beaks (56–) 60–100 mm long; mericarp bodies 9–11 mm long; sepals 12–15 mm long; leaf blades often with small, intercalary lobes/leaflets between the major lobes or leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) E. ciconium 6b. Carpel beaks 10–40 mm long; mericarp bodies 4–9 mm long; sepals 5–12 mm long; leaf blades without intercalary leaflets between the major leaflets 7a. Mericarp bodies 8–9 mm long; apical pits of mericarp bodies stipitate-glandular within, without furrows below them; sepals 8–12 mm long . . . . (in part) E. stephanianum 7b. Mericarp bodies 4–7 mm long; apical pits of mericarp bodies eglandular or sessileglandular within, with 2 furrows or a pit-like furrow below them; sepals 5–9 mm long 8a. Leaflets prominently lobed, with sinuses extending ½ to nearly the entire distance from the tips of the lobes to the leaflet midrib [Fig. 670]; anther-bearing filaments without basal teeth; apical pits of mericarp bodies eglandular, with 2 furrows below them . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. cicutarium 8b. Leaflets toothed to lobed, the sinuses rarely extending more than ½ the distance from the tips of the teeth or lobes to the leaflet midrib; anther-bearing filaments with 2 teeth at the base; apical pits of mericarp bodies sessile-glandular, with a pit-like furrow below them (the additional furrow also sessile-glandular) . . . . E. moschatum 1. Erodium botrys (Cav.) Bertol. E long-beaked stork’s-bill. Geranium botrys Cav. • MA, ME, VT. Wool waste, disturbed soil, lawns. 2. Erodium ciconium (L.) ĽHér. ex Ait. E common stork’s-bill. Geranium ciconium L. • MA. Wool waste, dumps. 3. Erodium cicutarium (L.) ĽHér. ex Ait. ssp. cicutarium E Fig. 670 red-stemmed stork’s-bill. Geranium cicutarium L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, sandy lawns, waste areas, wool waste.

Fig. 670  Flowers and leaves of Erodium cicutarium.

61 4   tricolpate s

4. Erodium cygnorum Nees E Australian stork’s-bill. MA. Wool waste. Reports of Erodium laciniatum (Cav.) Willd. var. bovei (Delile) Murbeck from MA are based on collections of E. cygnorum (i.e., the original determination was incorrect)—5 Sep 1917, Fletcher s.n. (GH!, NEBC!). 5. Erodium malacoides (L.) ĽHér. ex Ait. E Mediterranean stork’s-bill. Geranium malacoides L. • MA. Waste areas, yards. The apical pits of the mericarp bodies are similar to Erodium botrys (i.e., eglandular within and with 2 furrows below them). 6. Erodium moschatum (L.) ĽHér. ex Ait. var. praecox Lange E musky stork’s-bill. Geranium moschatum L. • CT, MA, ME. Fields, roadsides, waste areas, wool waste. 7. Erodium stephanianum Willd. E Stephen’s stork’s-bill. MA. Wool waste.

Geranium The gynoecium of Geranium is composed of 5 connate carpels that form a compound ovary. Each carpel is made up of a basal, thickened portion called the mericarp body (which contains the ovules) and an elongate, apical portion called the carpel beak (which is adnate to and conceals the style column prior to fruit maturation). The carpel beaks separate from the style column and outwardly curve at maturity, remaining attached to the style column near or at its apex. In most species of Geranium, the carpel beaks do not split all the way to the style remains. The region between the apex of the dehisced carpel beaks and the style remains is called the style beak and can be seen as a narrowed portion near the apex of the intact carpel beaks prior to separation. Above the carpel beaks project the styles. References: Aedo (2000, 2001). 1a. Leaves palmately compound, the terminal segment petiolulate; mericarp bodies separating from the carpel beaks but remaining attached to the style column for a period of time by 2 subapical filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. robertianum 1b. Leaves palmately lobed, the leaf segments confluent; mericarp bodies remaining attached to the carpel beaks, without subapical filaments 2a. Petals 12–20 mm long [Fig. 673]; anthers 2–2.6 mm long; style beaks 4–8 mm long; plants perennial from a stout rhizome 3a. Each cymule of the inflorescence usually 1-flowered; stems with several leaves; petals retuse at the apex, deep red (rarely white) . . . . . . . . . . . . . . . . . . . . . G. sanguineum 3b. Each cymule of the inflorescence usually 2-flowered; stems with a single pair of leaves, the other leaves basal; petals entire or very slightly retuse at the apex, bluepurple or pink (rarely white) 4a. Sepals, pedicels, and style beaks with stipitate glands [Fig. 673]; pedicels deflexed after anthesis; petals blue to blue-purple, 12–13 mm wide, 1.5–2.1 times as long as wide [Fig. 673] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. pratense 4b. Sepals, pedicels, and style beaks pubescent but not glandular; pedicels suberect after anthesis; petals pink to pink-purple (rarely white), 8.5–11.6 mm wide, 1.3–1.4 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. maculatum 2b. Petals 2.5–10 mm long; anthers 0.4–1.5 mm long; style beaks 0–6 mm long; plants annual or biennial, commonly from a taproot, or perennial with rhizomes in G. pyrenaicum, G. sibiricum, and G. thunbergii 5a. From 3–5 of the 5 outer stamens sterile and lacking anthers; seeds smooth; petals 2.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. pusillum

Ge r a ni ac e a e   6 15

5b. All of the stamens fertile and bearing anthers; seeds reticulate or striate, sometimes obscurely so (+/- smooth in G. aequale and G. pyrenaicum); petals (3–) 3.2–10 mm long 6a. Sepals obtuse to acute at the apex, sometimes with a short mucro 0.1–0.3 mm long; mericarp bodies glabrous on most of the surfaces (pubescent in G. pyrenaicum, ciliate near the base in G. aequale), cross-wrinkled or smooth; seeds ± smooth or striate 7a. Mericarp bodies pubescent; plant perennial from a short, vertical rhizome; seeds 2.2–2.7 mm long; petals 7–11 mm long . . . . . . . . . . . . . . . . . . . . . G. pyrenaicum 7b. Mericarp bodies glabrous (though ciliate near the base in G. aequale); plants annual; seeds 1.4–1.8 mm long; petals (3–) 3.5–8.5 (–10) mm long 8a. Mericarp bodies cross-wrinkled, eciliate; seeds striate; anthers 0.7–1.5 mm long; petals (3–) 4.5–8.5 (–10.5) mm long; pedicels 2.3–4.2 times as long as the sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. molle 8b. Mericarp bodies smooth, densely ciliate near the base; seeds ± smooth; anthers 0.4–0.6 mm long; petals 3.5–4.5 mm long; pedicels 1.5–2.5 times as long as the sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. aequale 6b. Sepals acuminate at the apex, tipped with an awn 0.5–2 mm long [Figs. 671, 672]; mericarp bodies at least sparsely hirsute (sometimes nearly glabrous in G. columbinum), smooth; seeds reticulate 9a. Pedicels 3–13 mm long, 0.6–2.2 times as long as the sepals; style beak 1–2 mm long 10a. Mericarp bodies pubescent with antrorsely oriented, eglandular hairs ca. 1 mm long; seeds inconspicuously reticulate-patterned with irregular, thin-walled aereolae; ultimate segments of middle and upper stem leaves usually narrow-oblong and obtuse at the apex . . . . . . . . . . . G. carolinianum 10b. Mericarp bodies pubescent with spreading, often glandular, hairs 0.2–0.6 mm long; seeds conspicuously reticulate-patterened with ± uniform, thick-walled aereolae; ultimate segments of middle and upper stem leaves usually linear and acute at the apex . . . . . . . . . . . . . . . . . . . . . . . . G. dissectum 9b. Pedicels 10–60 mm long, 1.8–8.2 times as long as the sepals [Fig. 671]; style beak 0–0.5 (–1) mm long or 3.5–4.5 mm long 11a. Plants perennial, from rhizomes; style beak 0–0.5 (–1) mm long 12a. Each cymule of the inflorescence usually 2-flowered; petals 7–10 × 5–5.5 mm, 1.4–1.8 times as long as wide, pubescent on the adaxial surface near the base; eglandular hairs of the pedicels spreading; seeds 1.9–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. thunbergii 12b. Each cymule of the inflorescence usually 1-flowered; petals 5–7 × 1.5–2 mm, ca. 2.5 times as long as wide, glabrous on the adaxial surface (though ciliate near the base); eglandular hairs of the pedicels retrorsely appressed; seeds 2.6–2.7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . G. sibiricum 11b. Plants annual or biennial, without rhizomes; style beak 3.5–4.5 mm long 13a. Pedicels spreading-pubescent with both longer eglandular hairs and shorter glandular hairs [Fig. 671]; petals 4–6 × 2–3 mm; seeds 1.6–1.7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. bicknellii 13b. Pedicels retrorsely appressed-pubescent with eglandular hairs; petals 8–10 × 4–5 mm; seeds 2.2–2.4 mm long . . . . . . . . . . G. columbinum 1. Geranium aequale (Bab.) Aedo E smooth Dove’s-foot crane’s-bill. Geranium molle L. var. aequale Bab. • MA, VT. Fields, lawns, areas of habitation.

616 tricolpate s

2. Geranium bicknellii Britt. N Fig. 671 northern crane’s-bill. Geranium bicknellii Britt. var. longipes (S. Wats.) Fern.; Geranium carolinianum L. var. longipes S. Wats. • CT, MA, ME, NH, RI, VT. Roadsides, clearings, forests, rocky slopes, ledges, waste areas, dry banks. 3. Geranium carolinianum L. N Fig. 672 Carolina crane’s-bill. Geranium carolinianum L. var. confertiflorum Fern.; G. carolinianum L. var. sphaerospermum (Fern.) Breitung; G. dissectum L. var. carolinianum (L.) Hook. f.; G. sphaerospermum Fern. • CT, MA, ME, NH, RI, VT. Dry-mesic forests, openings, roadsides, waste areas. Fig. 671  Flowers of Geranium bicknellii borne on relatively elongate pedicels.

4. Geranium columbinum L. E long-stalked crane’s-bill. MA. Fields. 5. Geranium dissectum L. E cut-leaved crane’s-bill. Geranium laxum Hanks ex Hanks & Small • CT, MA. Dumps, roadsides, waste areas. 6. Geranium maculatum L. N spotted crane’s-bill. CT, MA, ME, NH, RI, VT. Mesic to wet-mesic forests, meadows, roadsides, and clearings. 7. Geranium molle L. E dove’s-foot crane’s-bill. CT, MA, ME, NH, RI, VT. Fields, lawns, roadsides, waste areas. 8. Geranium pratense L. E Fig. 673 meadow crane’s-bill. CT, MA, ME, VT. Fields, roadsides. 9. Geranium pusillum L. E small-flowered crane’s-bill. CT, MA, ME, NH, VT. Fields, lawns, areas of cultivation.

Fig. 672  Calyx and fruit of Geranium carolinianum.

10. Geranium pyrenaicum Burm. f. E hedgerow crane’s-bill. VT. Fields, roadsides, waste areas. 11. Geranium robertianum L. N mountain crane’s-bill. Robertiella robertiana (L.) Hanks • CT, MA, ME, NH, RI, VT. Rocky forests, talus slopes, trail edges, ledges, coastal beaches and headlands. 12. Geranium sanguineum L. E bloody crane’s-bill. MA, ME, VT; also reported from NH and RI by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, railroads, waste areas. 13. Geranium sibiricum L. E

Fig. 673  Flowers of Geranium pratense showing stipitateglandular pedicels.

Siberian crane’s-bill. CT, MA. Fields, roadsides, waste areas, dumps. 14. Geranium thunbergii Sieb. & Zucc. ex Lindl. & Paxton E dew-drop crane’s-bill. Geranium nepalense Sweet var. thunbergii (Sieb. & Zucc. ex Lindl. & Paxton) Kudo • CT, MA. Fields, roadsides, waste areas, gardens.

Grossulariaceae Ribes Ribes oxyacanthoides L. was reported from ME by Campbell et al. (1995); however, the supporting specimens (at MAINE!) were all R. hirtellum, as evidenced by stamens exceeding the petals, cuneate to truncate leaf blade bases, and absence of internodal prickles. References: Sinnott (1985), Morin (2009).

G rossul a r i ac e a e   6 17

1a. Inflorescence a raceme, with (3–) 4–20 or more flowers [Figs. 674, 676]; pedicels articulated at the summit, the fruits falling from them; branchlets unarmed (armed in R. lacustre) 2a. Ovary and fruit pubescent with bristly, gland-tipped hairs [Fig. 676] 3a. Stems with thin prickles; racemes spreading to drooping; petals broad-flabellate to orbicular; berries purple or black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. lacustre 3b. Stems unarmed; racemes ascending to erect [Fig. 676]; petals cuneate-obovate; berries dark red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. glandulosum 2b. Ovary and fruit glabrous or with sessile, resinous glands [Fig. 674] 4a. Hypanthium above the ovary tubular, 9–15 mm long; sepals gold-yellow; mature leaf blades 20–35 (–50) mm long, each lobe with 0–3 (–7) apical teeth . . . . . . . . . . R. aureum 4b. Hypanthium above the ovary campanulate or scutelliform, up to 4.5 mm long; sepals green-purple or green-white to yellow; mature leaf blades (25–) 30–90 mm long, abundantly toothed, the teeth not restricted to the apical portion of the lobe 5a. Leaf blades, winter buds, and branchlet apices resin-dotted [Fig. 674]; berries black; hypanthium above the ovary cupuliform 6a. Pedicels 0.1–2 mm long, much exceeded by the conspicuous, lanceolate bracts [Fig. 674]; racemes with 6–15 flowers; calyx lobes glabrous or very sparsely pubescent on the abaxial surface; ovary without resin glands; branchles with prominent decurrent ridges extending from the nodes that are noticeably raised above the branch surface, without a fetid odor when bruised . . . . R. americanum 6b. Pedicels 2–8 mm long, much longer than the minute, ovate bracts; racemes with 4–10 flowers; calyx lobes pubescent on the abaxial surface; ovary with at least a few sessile resin glands; branchles with obscure decurrent ridges that are barely raised above the branch surface, with a fetid odor when bruised . R. nigrum 5b. Leaf blades, winter buds, and branchlet apices without resin glands; berries red (rarely translucent white in R. rubrum); hypanthium above the ovary scutelliform 7a. Axis of raceme and pedicels stipitate-glandular; anther sacs adjacent to one another; lateral leaf lobes commonly directed forward; terminal leaf lobe deltate; calyx lobes green-purple; stems straggling to ascending . . . . . . . . . . . . . . . . R. triste 7b. Axis of raceme and pedicels without glands; anther sacs widely separated by a broad connective; lateral leaf lobes spreading; terminal leaf lobe ovate; calyx lobes yellow to green; stems ascending to erect . . . . . . . . . . . . . . . . . . . . . . . . . . . R. rubrum 1b. Inflorescence a solitary flower or a corymb-like cluster of 2–4 (–5) flowers [Fig. 675]; pedicels not articulated at the summit, the fruits remaining on them; branchlets often armed with slender spines, prickles, and/or bristles, especially at the nodes 8a. Berry covered with slender prickles at maturity, these appearing as stalked glands on the ovary during flowering [Fig. 675]; tubular portion of the hypanthium above the ovary longer than the calyx lobes in most flowers [Fig. 675] . . . . . . . . . . . . . . . . . . . . . . R. cynosbati 8b. Berry glabrous, glandular, or bristly, but without prickles, the ovary not stipitateglandular during flowering (except usually in R. uva-crispa); tubular portion of the hypanthium above the ovary equaling or shorter than the calyx lobes 9a. Flowers in anthesis with stamens evidently longer than the sepals; peduncles 5–20 mm long; styles 7–14 mm long 10a. Stamens 6–8 mm long; hypanthium purple to green-purple; calyx lobes 4–5 mm long; nodal prickles 2–5 (–11) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . R. rotundifolium 10b. Stamens 9–12 mm long; hypanthium white to cream-white; calyx lobes 5–7.5 mm long; nodal prickles 7–17 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. missouriense

618  tricolpates

9b. Flowers in anthesis with stamens shorter than to barely longer than the sepals; peduncles 2–6 mm long; styles 2.5–8 mm long 11a. Hypanthium, sepals, and berry pubescent (often also with some gland-tipped bristles); sepals 5–7 mm long, round-obovate; nodal spines stout, 10–15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. uva‑crispa 11b. Hypanthium, sepals, and fruit glabrous (rarely the hypanthium and fruit sparsely pubescent); sepals 3–4 mm long, oblong to broad-oblong; nodal spines slender, 3–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. hirtellum Fig. 674  Inflorescence and resindotted leaf of Ribes americanum.

1. Ribes americanum P. Mill. N Fig. 674 eastern black currant. Ribes floridum L’Hér. • CT, MA, ME, NH, RI, VT. Forests and forest fragments, swamps, rocky slopes, riparian forests, stream banks. 2. Ribes aureum Pursh var. villosum DC. E golden currant. Ribes odoratum H. Wendl. • CT, MA, VT. Abandoned homesteads, roadsides, forest fragments. The reports of this species from ME by Seymour (1982) and others were based on specimens collected from cultivated plants (i.e., this species is not naturalized in ME). 3. Ribes cynosbati L. N Fig. 675 eastern prickly gooseberry. Grossularia cynosbati (L.) P. Mill.; Ribes cynosbati L. var. atrox Fern.; R. cynosbati L. var. glabratum Fern.; R. cynosbati L. var. inerme (Rehd.) Bailey • CT, MA, ME, NH, VT. Deciduous forests, rocky slopes, forest borders, partially forested talus. Ribes cynosbati has a distinctive inflorescence with elongate peduncles (7–25 mm) and pedicels (5–16 mm). 4. Ribes glandulosum Grauer N Fig. 676

Fig. 675  Inflorescence of Ribes cynosbati.

skunk currant. Ribes prostratum ĽHér.; R. resinosum Pursh • CT, MA, ME, NH, VT; becoming rare in southern New England. Swamps, wetland borders, forests, rocky slopes, ledges, ascending to high elevation in the mountains and even found in sheltered locales near treeline. 5. Ribes hirtellum Michx. N hairy-stemmed gooseberry. Grossularia hirtella (Michx.) Spach; Ribes hirtellum Michx. var. calcicola (Fern.) Fern.; R. hirtellum Michx. var. saxosum (Hook.) Fern.; R. oxyacanthoides L. var. calcicola Fern.; R. oxyacanthoides L. var. hirtellum (Michx.) Scoggan; R. oxyacanthoides L. var. saxosum (Hook.) Coville • CT, MA, ME, NH, RI, VT. Swamps, forests, roadsides, fields, stream banks, pond shores, wetland borders, peatlands. 6. Ribes lacustre (Pers.) Poir. N

Fig. 676  Inflorescence of Ribes glandulosum.

bristly swamp currant. Limnobotrya lacustris (Pers.) Rydb.; Ribes oxyacanthoides L. var. lacustre Pers. • CT, MA, ME, NH, VT; also reported from RI by Gould et al. (1998), but specimens are unknown; confined to the western portion of southern New England states. Swamps, wetland borders, forests, rocky slopes, shorelines, ascending to high elevation in the mountains. This shrub is very distinctive in its branches being densely beset with slender prickles. Only Ribes cynosbati rarely shows this feature, but that species has gray to graybrown branchlets (vs. yellow-brown or red-brown branchlets). 7. Ribes missouriense Nutt. E Missouri gooseberry. Grossularia missouriensis (Nutt.) Coville & Britt. • CT. Fields, roadsides, areas of habitation. 8. Ribes nigrum L. E garden black currant. CT, MA, ME, NH, VT. Fields, roadsides, abandoned homesteads, gardens. Jenkins and Zika (1995) erroneously excluded this species from VT. It does not require flowering material to separate this species from Ribes americanum (see identification key). 9. Ribes rotundifolium Michx.

NC

Appalachian gooseberry. Grossularia rotundifolia (Michx.) Coville & Britt.; Ribes stamineum Hornem.; R. triflorum Willd. • CT, MA. Deciduous forests, rocky slopes, talus.

Grossul a r i ac e a e   6 19

10. Ribes rubrum L. E garden red currant. Ribes sativum Syme; R. sylvestre (Lam.) Mert. & Koch; R. vulgare Lam. • CT, MA, ME, NH, RI, VT. Fields, roadsides, abandoned homesteads, gardens, hedge rows,

river banks, forest fragments. Some collection of this plant have translucent white berries (as opposed to the usual red fruit). 11. Ribes triste Pallas N swamp red currant. Ribes rubrum L. var. alaskanum (Berger) Boivin; R. triste Pallas var. albinervium (Michx.) Fern. • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Forests, forested talus, swamps, wetland margins, stream banks. 12. Ribes uva-crispa L. var. sativum DC. E European gooseberry. Grossularia reclinata (L.) P. Mill.; Ribes grossularia L.; R. uva-crispa L. ssp. reclinatum (L.) Reichenb. • CT, MA, NH, VT; also reported from ME by Campbell et al. (1995), but specimens are unknown. Fields, roadsides, forest fragments, abandoned homesteads.

Haloragaceae Reference: Crow and Hellquist (1983). 1a. Perianth 3-merous; flowers with 3 stamens and 3 carpels; leaves alternate, the blades of the emersed ones foliaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Proserpinaca 1b. Perianth 4-merous; flowers with 4 or 8 stamens and 4 carpels; leaves opposite or alternate or both on the same plant, the blades of the emersed ones commonly reduced and bract-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myriophyllum

Myriophyllum The ability of certain species to form turions (i.e., winter buds) is a useful identification character. The turions appear as a mass of crowded, highly reduced leaves. Of the species with dissected leaves, only M. alternifolium, M. humile, and M. spicatum do not form turions. The other species do, these generally present on the plant from September or October through April. Reference: Aiken (1981). 1a. Leaf blades (when present) entire, represented by small scales or scale-like bumps on the erect stems [Fig. 679]; bracts entire [Fig. 679] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. tenellum 1b. Leaf blades pinnately divided [Figs. 677, 678, 680]; bracts entire or, more commonly, toothed or divided [Figs. 677, 678, 680] 2a. Foliage leaves alternate, subopposite, or irregularly subverticillate; staminate flowers commonly with 4 stamens 0.4–1.1 mm long; flowers in the axils of submersed leaves or in emersed spikes 3a. Leaves alternate (sometimes some subopposite, especially in submersed plants) [Fig. 677]; fruit 0.7–1.2 mm long; mericarps rounded on the outer surface, without ridges or projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. humile 3b. Leaves typically both alternate and whorled on the same stem; fruit 1.3–2.5 mm long; mericarps with 2 tuberculate, longitudinal ridges on the outer surface 4a. Flowers in emersed spikes, subtended by bract-like leaves; anthers 0.8–1.1 mm long; fruits 1.3–1.8 mm long; turions (i.e., winter buds) not formed . . . . . . M. pinnatum 4b. Flowers in the axils of submersed leaves; anthers ca. 0.4 mm long; fruits 2–2.5 mm long; turions present, formed in autumn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. farwellii

620  tricolpates

2b. Foliage leaves regularly whorled [Figs. 678, 680]; staminate flowers commonly with 8 stamens 1.2–2 mm long (only 4 stamens in M. heterophyllum or the staminate flowers missing); flowers in emersed spikes 5a. Flowers subtended by bracts mostly 25–35 mm long with uniform linear divisions 4–8 mm long, the bracts ± similar in size and division to the submersed leaves; submersed leaves with petioles 5–7 mm long; plants dioecious, ours always carpellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. aquaticum 5b. Flowers subtended by bracts less than 20 mm long, with marginal teeth or divisions (when present) shorter than 4 mm, the bracts conspicuously reduced in size and less divided than the submersed leaves; submersed leaves with petioles 0–2 mm long; plants monoecious 6a. Flowers and their subtending bracts alternate (sometimes the lowest opposite); leaf blades with usually 3–7 pairs of narrow segments; spikes 2–5 cm tall; turions not formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. alterniflorum 6b. Flowers and their subtending bracts whorled [Fig. 680]; leaf blades with usually 6–20 pairs of narrow segments; spikes 4–15 (–37) cm tall; turions formed in late season (not formed in M. spicatum) 7a. Carpellate bracts less than 2 times as long as the flowers or fruits [Fig. 678]; staminate bracts (i.e., the upper ones) with entire to denticulate margins 8a. Middle leaves with mostly 6–12 segments on each side of the rachis; upper leaves rounded at the apex; stems of ± similar diameter below the inflorescence and near the stem base, green near the apex of vegetative shoots (except sometimes in late season when turions are being formed), ± erect; turions (i.e., winter buds) formed in the fall . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. sibiricum 8b. Middle leaves with mostly 12–20 segments on each side of the rachis; uppermost leaves truncate at the apex; stems thicker below the inflorescence than at the stem base, up to twice as thick, usually red near the apex of vegetative shoots, curved to lie parallel with the water [Fig. 678]; turions not formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. spicatum 7b. Carpellate bracts usually 2 or more times as long as the flowers or fruits [Fig. 680]; staminate bracts with serrulate to pinnatifid margins 9a. Carpellate bracts prominently serrate to pinnately lobed [Fig. 680]; fruits 2–2.5 mm long; turions formed in the late summer and fall on the stems, clavate in outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. verticillatum 9b. Carpellate bracts merely serrulate (sometimes pectinately lobed near the water line); fruits 1–1.5 mm long; turions formed in the fall at the base of the stems or on the rhizomes, not clavate in outline . . . . . . . . . . . M. heterophyllum 1. Myriophyllum alterniflorum DC. N alternate-flowered water-milfoil. Myriophyllum alterniflorum DC. var. americanum Pugsley • CT, MA, ME, NH, RI, VT; generally rare outside of ME. Still or slow-moving, circumneutral to basic water of lakes and rivers. 2. Myriophyllum aquaticum (Vell.) Verdc. E Parrot’s-feather water-milfoil. Enydria aquatica Vell.; Myriophyllum brasiliense Camb.; M. proserpinacoides Gillies ex Hook. & Arn. • CT. Ponds, often those in areas of habitation. 3. Myriophyllum farwellii Morong N Farwell’s water-milfoil. CT, MA, ME, NH, RI, VT. Still or slow-moving, circumneutral to basic water of lakes and rivers. 4. Myriophyllum heterophyllum Michx. E Fig. 677  Leaves and fruits of Myriophyllum humile.

variable-leaved water-milfoil. CT, MA, ME, NH, RI, VT. Still or slow-moving, circumneutral to basic water of lakes and rivers.

H a lo r agac e a e   62 1

4 × Myriophyllum laxum Shuttlw. ex Chapman is a rare hybrid water-milfoil known from CT, ME. It tends to show some scattered (i.e., alternate) leaves at the base of the inflorescence (the others whorled), and the blades are often ± lobed (those of M. heterophyllum are whorled and serrulate). This hybrid could be confused with Myriophyllum pinnatum; however, the latter is smaller and has bracts that vary from pinnately lobed to prominently serrate. 5. Myriophyllum humile (Raf.) Morong N Fig. 677 low water-milfoil. Burshia humilis Raf.; Myriophyllum ambiguum Nutt. var. natans DC.; M. capillaceum Torr.; M. humile (Raf.) Morong forma capillaceum (Torr.) Fern.; M. humile (Raf.) Morong forma natans (DC.) Fern. • CT, MA, ME, NH, RI, VT. Still or slow-moving, acidic to circumneutral water of lakes, rivers, and pools, pond shores, often found on shorelines of receding waters. Myriophyllum humile is extremely variable and shows different morphologies depending on water level. Stems generally become longer and branches bear more and finer segments as water depth increases, with terrestrial forms appearing strikingly different from aquatic forms. 6. Myriophyllum pinnatum (Walt.) B.S.P.

NC

cut-leaved water-milfoil. Myriophyllum scabratum Michx.; Potamogeton pinnatum Walt. • CT, MA, RI; coastal plain. Fresh or brackish ponds and pond shores, muddy or peaty pools and depressions.

Fig. 678  Habit of Myriophyllum spicatum showing short carpellate bracts.

7. Myriophyllum sibiricum Komarov N northern water-milfoil. Myriophyllum exalbescens Fern.; M. spicatum L. ssp. exalbescens (Fern.) Hultén; M. spicatum L. var. exalbescens (Fern.) Jepson • CT, MA, ME, NH, VT; reported from RI by George (1992), but specimens are unknown. Slow or still-moving, circumneutral to basic water of lakes and rivers. This native species is sometimes difficult to separate from the introduced Myriophyllum spicatum. In addition to characters used in the key, M. sibiricum has knob-like vegetative shoot apices and stems that usually become distinctly whitened in drying, while M. spicatum has tassel-like vegetative shoot apices and stems that remain green or brown (rarely whitening) in drying. 8. Myriophyllum spicatum L. E Fig. 678 Eurasian water-milfoil. CT, MA, ME, NH, VT. Slow or still-moving, circumneutral to basic water of lakes and rivers. 9. Myriophyllum tenellum Bigelow N Fig. 679 slender water-milfoil. CT, MA, ME, NH, RI, VT. Shallow, acidic to circumneutral water of lakes, pond shores.

Fig. 679  Habit of Myriophyllum tenellum.

10. Myriophyllum verticillatum L. N Fig. 680 whorled water-milfoil. Myriophyllum verticillatum L. var. intermedium W.D.J. Koch; M. verticillatum L. var. pectinatum Wallr. • CT, MA, ME, NH, VT; generally becoming rare in southern and western New England. Slow or still-moving, circumneutral to basic water of lakes and rivers.

Proserpinaca 1a. Flowers subtended by serrate leaf-like bracts (the lower flowers sometimes subtended by nearly pinnatifid bracts); leaf blades (15–) 20–40 (–60) mm long, the submersed ones with 7–14 pairs of segments 5–30 mm long; fruits 2.3–6 mm wide . . . . . . . . . . . . . . . . . . . P. palustris 1b. Flowers subtended by pinnately lobed or divided leaf-like bracts [Fig. 681]; leaf blades 10–30 mm long, the lower and/or submersed ones with (4–) 6–9 (–12) pairs of segments 2–7.5 mm long; fruits 2–2.8 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pectinata 1. Proserpinaca palustris L. N marsh mermaid-weed. Proserpinaca amblyogona (Fern.) Small; P. intermedia Mackenzie; P. palustris L. var. amblyogona Fern.; P. palustris L. var. crebra Fern. & Grisc.; P. palustris L. var. latifolia Schindl. • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving, basic to circumneutral water of lakes and rivers, pond shores, swamps, fens, mucky pools.

Fig. 680  Habit of Myriophyllum verticillatum showing prominent carpellate bracts.

622  tricolpates

1‌ × 2. Proserpinaca ×intermedia Mackenzie is a rare mermaid-weed hybrid known from MA, RI. It can be recognized by the pinnately lobed submsersed leaves with 7–12 ‌intermedia is further characterized pairs of divisions 2–3.5 mm long. Proserpinaca × by leaves with a central rachis 1–4 mm wide (vs. 0.2–1 mm in P. pectinata) and fruits 2.3–3.6 mm wide. 2. Proserpinaca pectinata Lam. N Fig. 681 comb-leaved mermaid-weed. MA, ME, NH, RI; mainly along the coastal plain. Pond shores, pools, ditches, wet, peaty depressions. In southwestern ME and eastern NH this species occurs inland from the coastal plain. However, those areas harbor ponds that are hydrologically and/or floristically similar to coastal plain pond shores.

Hamamelidaceae Fig. 681  Leaves and fruits of Proserpinaca pectinata.

Hamamelis 1. Hamamelis virginiana L. N American witch-hazel. Hamamelis virginiana L. var. parvifolia Nutt. • CT, MA, ME, NH, RI, VT. Deciduous to mixed evergreen-deciduous forests, swamps, riparian forests, rocky slopes, forested talus.

Hydrangeaceae 1a. Filaments toothed near apex, dimorphic, the outer ones longer; pith hollow; leaf blades and young branchlets pubescent with stellate hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Deutzia 1b. Filaments without apical teeth, monomorphic; pith solid; plants pubescent with simple hairs or glabrous 2a. Fertile flowers with 5 petals 1–3 mm long and 10 stamens; marginal flowers of inflorescence often enlarged, sterile, with only 3 or 4 petaloid sepals (enlarged flowers sometimes absent) [Fig. 682]; fruit with 10–15 ribs, dehiscing by pores; winter buds axillary to the petioles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrangea 2b. Flowers with 4 petals 15–30 mm long and 20–40 stamens [Fig. 683]; marginal flowers of inflorescence neither enlarged nor sterile [Fig. 683]; fruit without ribs, dehiscing by longitudinal valves; winter buds embedded in the petiole bases . . . . . . . . . . . Philadelphus

Deutzia 1. Deutzia scabra Thunb. E fuzzy pride-of-Rochester. Deutzia scabra Thunb. var. plena (Maxim.) Rhed. • CT, MA, RI, VT. Forest fragments, roadsides, areas of habitation. Some forms of this shrub that escape in New England have double corollas (the “plena” cultivar).

Hydrangea 1a. Leaf blades with (3–) 5–7 coarsely serrate lobes . . . . . . . . . . . . . . . . . . . . . . . . . . H. quercifolia 1b. Leaf blades unlobed, the margins dentate or serrate to serrulate

H y d r a n g e ac e a e   623

2a. Cyme panicle-like, pyramidal or ovoid; ovary half-inferior; petioles 0.6–2.4 cm long; winter buds appressed and pointing forward . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. paniculata 2b. Cyme corymb-like, flat-topped or slightly convex [Fig. 682]; ovary completely inferior; petioles (0.8–) 2–8 (–11.5) cm long; winter buds projecting outward from twig 3a. Leaf blades nearly glabrous abaxially or sparsely hirsute along the midrib and sometimes the main lateral veins as well . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. arborescens 3b. Leaf blades uniformly pubescent across the abaxial surface . . . . . . . . . . . . H. cinerea 1. Hydrangea arborescens L. E Fig. 682 American hydrangea. Hydrangea arborescens L. var. oblonga Torr. & Gray; H. arborescens L. var. sterilis Torr. & Gray • CT, MA, VT. Roadsides, forest borders and fragments, river banks, areas of cultivation. 2. Hydrangea cinerea Small E

Fig. 682  Inflorescence of Hydrangea arborescens.

ashy hydrangea. Hydrangea arborescens L. ssp. discolor (Ser. ex DC.) McClintock; H. arborescens L. var. discolor Ser. ex DC. • MA. Roadsides, forest borders and fragments, areas of cultivation. 3. Hydrangea paniculata Sieb. E panicled hydrangea. CT, MA, ME, NH; also reported from RI by George (1992), but specimens are unknown. Roadsides, forest borders and fragments, areas of cultivation. 4. Hydrangea quercifolia Bartr. E oak-leaved hydrangea. CT. Roadsides, forest borders and fragments, areas of cultivation.

Philadelphus 1a. Branchlets gray; branches with close bark; leaf blades abaxially softly pubescent on and between the veins; pedicels, hypanthium, and abaxial sepal surfaces softly pubescent (sometimes one or more of these surfaces glabrate) [Fig. 683] . . . . . . . . . . . . . . . . P. pubescens 1b. Branchlets brown; branches with exfoliating bark; leaf blades abaxially glabrous or pubescent only on the veins and in the axils of veins; pedicels, hypanthium, and abaxial sepal surface usually glabrous (rarely one or more surface sparsely pubescent) 2a. Inflorescence a raceme with 5–7 flowers [Fig. 683]; flowers 25–35 mm wide; most leaf blades remotely denticulate or remotely dentate (entire blades may also be present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. coronarius 2b. Inflorescence a solitary flower or cymule with 3 flowers; flowers 35–55 mm wide; most leaf blades entire (remotely denticulate leaf blades may also be present) . . . . . P. inodorus 1. Philadelphus coronarius L. E sweet mock-orange. CT, MA, ME, NH, RI, VT. Roadsides, forest fragments, abandoned homesteads, areas of cultivation. Specimens determined as Philadephus caucasicus Koehne have been collected from MA (Seymour 1982). However, these plants differ from P. coronarius mainly on subtle differences in pubescence that appear to be variable at the population level (i.e., they may not be indicative of species-level distinctions; Alan Weakley, personal communication). Therefore, these plants are tentatively placed here in P. coronarius until further study is performed. 2. Philadelphus inodorus L. E scentless mock-orange. Philadelphus gloriosus Beadle; P. grandiflorus Willd.; P. inodorus L. var. grandiflorus (Willd.) Gray • CT, MA, RI. Roadsides, forest fragments, abandoned homesteads, areas of cultivation. 3. Philadelphus pubescens Loisel. E Fig. 683 hoary mock-orange. Philadelphus intectus Beadle; P. latifolius Schrad. ex DC.; P. pubescens Loisel. var. intectus (Beadle) A.H. Moore • CT, MA. Roadsides, forest fragments, abandoned homesteads, areas of cultivation.

Fig. 683  Inflorescence of Philadelphus pubescens.

624  tricolpates

Hypericaceae 1a. Petals pink to flesh-colored; stamens 9, connate into 3 distinct groups, alternating with conspicuous, orange glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triadenum 1b. Petals yellow to orange-yellow; stamens 5–many, distinct or connate into 3–5 groups, without hypogynous glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypericum

Hypericum Reference: Adams and Robson (1961). 1a. Flowers with 4 sepals in unequal groups of 2 and 4 petals [Fig. 687]; styles 2; low, compact, mat- or mound-forming shrubs up to 3 dm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. stragulum 1b. Flowers with 5 sepals and 5 petals [Figs. 684, 685, 686]; styles 3 or 5 (rarely 4); shrubs 5–30 dm tall or herbs 2a. Plants woody; capsules 3-locular (rarely 4-locular) 3a. Inflorescence of solitary, terminal flowers (rarely as many as 3 flowers together); longer sepals 10–15 mm long; petals 10–20 mm long; placentae of fruit not meeting in the center . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. frondosum 3b. Inflorescence a terminal cyme of (1–) 3 or more flowers (and sometimes also of axillary cymules); longer sepals 2–7 mm long; petals 4.5–10 mm long; placentae of fruit meeting in center 4a. Terminal cyme with (1–) 3–7 flowers; leaf blades (4–) 7–15 mm wide; capsules 7–14 (–15) × 3–5 mm; seeds 1.5–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . H. prolificum 4b. Terminal cyme with 7 or more flowers; leaf blades 3–7 (–11) mm wide; capsules 4.5–6 × 1.5–3 mm; seeds 0.8–1.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . H. densiflorum 2b. Plants herbaceous; capsules 1-, 3-, or 5-locular (rarely 4-locular) 5a. Flowers with 5 styles; capsules 5-locular, 15–30 mm tall; petals 25–30 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. ascyron 5b. Flowers with 3 (–4) styles; capsules 1- or 3-locular (rarely 4-locular), 2–7 mm tall; petals 2–12 mm long 6a. Styles connate or closely connivent, persistent in fruit and appearing as a single, straight beak at the summit of the capsule [Fig. 686]; stigmas minute 7a. Sepals distinctly unequal, 5–13 (–15) × 2–3 (–8) mm; petals 10–13 mm long, strongly asymmetrical, relatively straight on one margin and srongly convex on the other margin; stamens numbering 120–200, deciduous; seeds 1.5–1.8 mm long; stems branched near or base or throughout and usually clustered together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. dolabriforme 7b. Sepals subequal to somewhat unequal, (2–) 4–7 × 1–3 mm; petals 5–8 mm long, ± symmetrical to weakly asymmetrical; stamens numbering 60–95, persistent; seeds 0.7–0.8 mm long; stems commonly unbranched until inflorescence and occurring singly 8a. Leaf blades with flat margins, elliptic to elliptic-oblong, 1–3 (–4) cm long, 2–3 times as long as wide; sepals oblanceolate to narrow-obovate, widest above the middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. ellipticum 8b. Leaf blades with revolute margins, narrow-lanceolate to narrow-oblong or narrow-elliptic, 3–6 cm long, 4–6 times as long as wide; sepals lanceolate to ovate, widest near or below the middle . . . . . . . . . . . . . . . . . . . . . . . H. adpressum

Hy p e r i c ac e a e   625

6b. Styles distinct nearly to the base and often divergent, deciduous in fruit and not appearing as a single beak at the summit of the capsule; stigmas capitate 9a. Ovary and capsule 3-locular (rarely 4-locular); flowers with 20–40 stamens 10a. Petals conspicuously dotted and streaked with black; sepals with few or no black glands, obtuse to broad-acute at the apex; seeds 0.6–1.1 mm long; branches terete, without decurrent ridges . . . . . . . . . . . . . . . . . . . . . H. punctatum 10b. Petals dotted only at margins; sepals copiously dotted and lined with black glands, acuminate at the apex; seeds 1–1.3 mm long; branches with sharp, decurrent ridges below the leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . H. perforatum 9b. Ovary and capsule unilocular; flowers with 5–22 stamens 11a. Leaf blades subulate to linear-subulate, 1-nerved, appressed, 1–4 mm long; inflorescence frequently monochasial (i.e., with alternate branches) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. gentianoides 11b. Leaf blades linear to elliptic, 1- to 7-nerved, spreading to ascending, 3–45 mm long; inflorescence clearly dichasial (i.e., with opposite branches) 12a. Capsules ellipsoid, abruptly tapered and rounded at the summit [Fig. 684]; sepals linear to linear-lanceolate, broadest at or near the middle; stems decumbent and leafy-bracted at the base 13a. Bracts of the cyme subulate, not resembling the leaves; sepals acute at the apex, subequal in length to the capsule . . . . . . . . . . . . . . . H. mutilum 13b. Bracts of the cyme elliptic, leaf-like [Fig. 684]; sepals obtuse at the apex, conspicuously shorter than the capsule [Fig. 684] . . . . . . H. boreale 12b. Capsules ellipsoid-conic to conic, gradually tapered and pointed at the summit [Fig. 685]; sepals lanceolate, broadest below the middle; stems erect, not leafy-bracted at the base (though with leafy stolons) 14a. Leaf blades linear to narrow-oblanceolate, 1–4 (–6) mm wide, 1- or 3-nerved (rarely 5-nerved), narrowly tapering to the base [Fig. 685]; petals 2.5–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. canadense 14b. Leaf blades lanceolate to narrow-elliptic or narrow-oblong, 3–9 (–15) mm wide, 5- or 7-nerved, broadly tapering to rounded at the base; petals 3.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. majus 1. Hypericum adpressum Raf. ex Bart. NC creeping St. John’s-wort. CT, MA, RI; coastal plain. River shores, coastal plain pond shores, sandy and peaty depressions.

Fig. 684  Inflorescence of Hypericum boreale showing abruptly tapering apex of capsules.

2. Hypericum ascyron L. N great St. John’s-wort. Hypericum pyramidatum Ait. • CT, MA, ME, NH, VT; found primarily in western New England. Riparian forests, river banks, low fields. 3. Hypericum boreale (Britt.) Bickn. N Fig. 684 northern St. John’s-wort. Hypericum mutilum L. ssp. boreale (Britt.) J. Gillett • CT, MA, ME, NH, RI, VT. Shorelines, swamps, wetland edges. ‌ × 4. Hypericum ×dissimulatum Bickn. is an uncommon St. John’s-wort hybrid known 3 from CT, MA, ME, RI, VT. It is frequently sterile (at least partially, the capsules not filled with well-formed seeds). It generally resembles H. canadense, but the leaf blades average wider (1.5–10 mm vs. 1–4 (–6) mm) and the capsule is usually shorter (3–4.5 mm vs. 4–6 (–6.5) mm) and relatively blunter (i.e., intermediate in shape between the two parents). 4. Hypericum canadense L. N Fig. 685 lesser Canada St. John’s-wort. Hypericum canadense L. var. galiiforme Fern.; H. canadense L. var. magninsulare Weatherby • CT, MA, ME, NH, RI, VT. Shorelines, swamps, wetland edges, small pools and other ephemeral wet areas.

Fig. 685  Inflorescence of Hypericum canadense showing gradually tapering apex of capsules.

626  tricolpate s

4 × 10. This rare St. John’s-wort hybrid is known from CT, MA, NH. It is intermediate between its parental taxa in leaf blade venation (3- or 5-veined), base shape (broad-cuneate), and width (the principal ones mostly 2–3.5 mm). It is often collected with both parents. ‌dissimulatum 4 × 11. This rare St. John’s-wort hybrid is known from ME. It is similar to H. × (3× 4) except that the sepals tend to be more pointed at apex (sharply acute vs. merely acute; side by side comparison of specimens is useful here). 5. Hypericum densiflorum Pursh E bushy St. John’s-wort. Hypericum glomeratum Small • MA. Fields, waste areas. 6. Hypericum dolabriforme Vent. E straggling St. John’s-wort. Hypericum bissellii B.L. Robins. • CT. Roadsides. 7. Hypericum ellipticum Hook. N Fig. 686 pale St. John’s-wort. CT, MA, ME, NH, RI, VT. Shorelines, marshes, low fields, wetland edges. 8. Hypericum frondosum Michx. E cedar glade St. John’s-wort. Hypericum aureum Bartr. • CT, MA. Roadsides, fields, areas of cultivation. Fig. 686  Flower of Hypericum ellipticum showing connate styles.

9. Hypericum gentianoides (L.) B.S.P. N orange-grass St. John’s-wort. Sarothra gentianoides L. • CT, MA, ME, NH, RI, VT. Sand plains, dry fields, roadsides, railroads, and other open, sandy areas. 10. Hypericum majus (Gray) Britt. N greater Canada St. John’s-wort. Hypericum canadense L. var. majus Gray • CT, MA, ME, NH, RI, VT. Shorelines, low fields, wetland edges, ditches. 10 × 11. This very rare St. John’s-wort hybrid is known from ME. It can be recognized by its intermediate capsule shape, similar to H. × (3× 4). It differs from ‌dissimulatum ‌dissimulatum H. × in that the leaf blades average broader (principal ones 4.8–10.8 mm wide vs. 1.5–10 mm). 11. Hypericum mutilum L. N dwarf St. John’s-wort. Hypericum mutilum L. var. parviflorum (Willd.) Fern. • CT, MA, ME, NH, RI, VT. Shorelines, low fields, wetland edges. 12. Hypericum perforatum L. ssp. perforatum E common St. John’s-wort. CT, MA, ME, NH, RI, VT. Fields, roadsides, open banks, railroads, shorelines, waste areas. 13. Hypericum prolificum L. E shrubby St. John’s-wort. Hypericum spathulatum (Spach) Steud. • CT, MA, ME, NH, RI, VT. Fields, forest edges, roadsides. 14. Hypericum punctatum Lam. N spotted St. John’s-wort. Hypericum subpetiolatum Bickn. ex Small • CT, MA, ME, NH, RI, VT. Fields, meadows, forest openings. 15. Hypericum stragulum P. Adams & Robson

Fig. 687  Flowers of Hypericum stragulum.

N C Fig. 687

multi-stemmed St. Andrew’s-cross. Ascyrum hypericoides L. var. multicaule (Michx. ex Willd.) Fern.; Hypericum hypericoides (L.) Crantz ssp. multicaule (Michx. ex Willd.) Robson; H. hypericoides (L.) Crantz var. multicaule (Michx. ex Willd.) Fosberg • MA; Nantucket Island. Coastal plain pond shores, sand plains.

Triadenum 1a. Sepals 5–8 mm long, acute to acuminate at the apex; petals 8–10 mm long; styles 2–3 mm long; capsules gradually tapering to the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. virginicum

H y p e r i c ac e a e   627

1b. Sepals 3–5 mm long, obtuse to rounded at the apex; petals 5–8 mm long; styles 0.5–1.5 mm long; capsules somewhat abruptly tapering to the apex . . . . . . . . . . . . . . . . T. fraseri 1. Triadenum fraseri (Spach) Gleason N Fraser’s marsh-St. John’s-wort. Elodea fraseri Spach; Hypericum virginicum L. var. fraseri (Spach) Fern.; Triadenum virginicum (L.) Raf. ssp. fraseri (Spach) J. Gillett; T. virginicum (L.) Raf. var. fraseri (Spach) Cooperrider • CT, MA, ME, NH, VT; becoming rare in southern New England. Shorelines, wetland edges, swamps. Triadenum fraseri is a northern sister species of T. virginicum. 2. Triadenum virginicum (L.) Raf. N Virginia marsh-St. John’s-wort. Hypericum virginicum L. • CT, MA, ME, NH, RI, VT. Shorelines, wetland edges, swamps.

Juglandaceae Though the fruits of New England representatives of the family are usually described as a nut, but they are in fact a drupe with a fibrous exocarp that is either dehiscent (Carya) or indehiscent (Juglans). 1a. Leaves with 11–19 leaflets, the middle lateral leaflets the largest; branchlets with chambered pith; drupe with an indehiscent exocarp (i.e., husk); each staminate ament borne separately; staminate flowers with 8–40 stamens and glabrous anthers . . . . . . . . . . . . . Juglans 1b. Leaves with 5–9 (–11) leaflets, the terminal leaflet the largest; branchlets with solid pith; drupe with a 4-valved, dehiscent exocarp; staminate aments borne in clusters of 3; staminate flowers with 3–10 stamens and pilose anthers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carya

Carya Manning (1950) gave one of the most thorough accounts of the morphological details of Carya that are important for identification, including their variation through the growing season. Anyone serious about learning the genus Carya should spend time studying this work. Details of the terminal buds are important for identification. Caution is needed because some branchlets are terminated by pseudoterminal buds, especially those bearing fruit. Pseudoterminal buds, which can identified by the presence of a twig scar proximal to the bud, differ both in size and in the shape of outer scales compared with terminal buds. Similarly, details of leaf pubescence are especially important for identification. Though technically there exists more than one type of hair in Carya that appears to be compound (specifically, it appears to be a stellate hair with multiple rays emerging from the same point), they are not distinguished here due to practical difficulties in separating them. Therefore, hairs of the leaf blades surfaces are described here only as simple or compound. References: Manning (1950), Stone (1997). 1a. Terminal bud yellow or orange-yellow to yellow-brown, with 4–6 valvate scales; leaves with (5–) 7–9 (–13) leaflets; exocarp (i.e., husk) 2–3 mm thick, the sutures narrowly winged; staminate aments borne, in part, on reduced, ± leafless shoots that originate on the branches of the previous season . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cordiformis 1b. Terminal bud light brown to red-brown or dark brown, with 6–15 imbricate scales; leaves with 3–9 (–11) leaflets; exocarp 3–15 mm thick (as thin as 2 mm in C. glabra), the sutures usually unwinged; staminate aments borne only at base of leafy shoots on branchlets of the current season 2a. Leaflets strongly ciliate with tufts of hairs borne below the apex of the marginal teeth [Fig. 688]; leaves with (3–) 5 (–7) leaflets; pedicels and bracts of the staminate flowers glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ovata

628  tricolpate s

2b. Leaflets at maturity becoming eciliate or retaining some cilia, but without subapical tufts of hairs on the marginal teeth; leaves with (3–) 5–9 (–11) leaflets; pedicels and/or bracts of staminate flowers pubescent 3a. Leaflets abaxially glabrous to pubescent with mostly simple hairs, some compound hairs with 2–4 rays also present; terminal bud 5–15 mm long, the bud scales abaxially glabrous to hirsute; branchlets relatively slender, (1.8–) 2–4.1 (–4.4) mm thick, glabrous or nearly so; exocarp 2–5 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. glabra 3b. Leaflets abaxially pubescent with simple hairs and abundant compound hairs with 2–8 (–17) rays; terminal bud 8–20 mm long, the bud scales abaxially tomentose; branchlets stout, (3.5–) 4–9 mm thick, hirsute (sometimes becoming subglabrous in C. laciniosa); exocarp 4–13 mm thick 4a. Petiole and rachis densely hirsute; leaflets usually acute at the apex, pubescent predominately with compound hairs, the simple hairs scarce; branchlets red-brown; bark ridged, not exfoliating; exocarps rough but glabrous . . . . . . . . . . . . C. tomentosa 4b. Petiole and rachis sparely pubescent to subglabrous; leaflets usually acuminate at apex, pubescent with both simple and compound hairs; branchlets light orangebrown to light yellow-brown; bark separating into long strips or broad plates; exocarps minutely hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. laciniosa 1. Carya cordiformis (Wangenh.) K. Koch N bitternut hickory. Carya cordiformis (Wangenh.) K. Koch var. latifolia Sarg.; Hicoria cordiformis (Wangenh.) Britt.; Juglans cordiformis Wangenh. • CT, MA, ME, NH, RI, VT; limited in ME to the southern portion of the state. Forests, rocky hillsides, riparian terraces, especially those influenced by rich colluvium or high-pH bedrock and/or till. ‌1 × 2. Carya ×demareei Palmer is a very rare hickory hybrid known from VT. It is identified by its smooth bark, leaves with mostly 7 leaflets, drupes with thin exocarp (2–3 mm thick) and exocarp sutures with very narrow wings, and terminal winter buds with weakly imbricate, yellow-brown to light red-brown scales. ‌1 × 4. Carya ×laneyi Sarg. is a rare hickory hybrid known from NH, VT. This tree generally resembles C. glabra as to the leaves and fruits, but the endocarp shell is very thin (1–1.5 mm thick). More specifically, it has a pointed terminal bud with valvate bud scales (like C. cordiformis), but the bud scales are brown. The bark is slightly roughened (not separating in plates as in C. ovata). The leaflets usually number 7 per leaf (5 is less common) and lack the subapical tufts of hairs on the marginal teeth. 2. Carya glabra (P. Mill.) Sweet N pignut hickory. Carya glabra (P. Mill.) Sweet var. hirsuta (Ashe) Ashe; C. glabra (P. Mill.) Sweet var. megacarpa (Sarg.) Sarg.; C. ovalis (Wangenh.) Sarg. var. hirsuta (Ashe) Sarg.; Hicoria glabra (P. Mill.) Britt.; H. glabra (P. Mill.) Britt. var. hirsuta Ashe • CT, MA, NH, RI, VT; reaching its northern limit in southern NH, but extending north along western VT into northern counties. Mainly dry-mesic, sometimes thin, soils of forests, woodlands, and ridges. This species is polymorphic in New England and consists of two intergrading morphologies. Plants with bark exfoliating in longitudinal plates, frequently red petioles, and drupes with exocarps promptly dehiscing to the base have been called Carya ovalis. Plants with close bark, frequently green petioles, and drupes with exocarps tardily splitting (or not splitting at all) represent C. glabra s.s. Though Manning (1950) maintained these two as separate species, he stated that trees with fruits intermediate between C. glabra and C. ovalis are almost as frequent as trees with characteristic fruits. Note that he also wrote these two species could only be separated by examination of mature fruits. 3. Carya laciniosa (Michx. f.) Loud. E shellbark hickory. Hicoria laciniosa (Michx. f.) Sarg.; Juglans laciniosa Michx. f. • MA. Roadsides, areas of habitation.

Ju g l a n dac e a e   62 9

4. Carya ovata (P. Mill.) K. Koch N Fig. 688 shagbark hickory. Carya ovata (P. Mill.) K. Koch var. pubescens Sarg. Hicoria ovata (P. Mill.) Britt.; Juglans alba L., pro parte • CT, MA, ME, NH, RI, VT; becoming rare to the north and generally absent from northern counties. Dry-mesic to mesic soils of forests, woodlands, ridges, and outcrops. 5. Carya tomentosa (Poir. in Lam.) Nutt. N mockernut hickory. Carya alba (L.) Nutt. ex Ell., pro parte; C. tomentosa (Poir. in Lam.) Nutt. var. subcoriacea (Sarg.) Palmer & Steyermark; Hicoria tomentosa (Poir. in Lam.) Raf.; Juglans alba L., pro parte; Juglans tomentosa Poir in Lam. • CT, MA, RI; also reported from NH Kartesz (1999), but specimens are unknown. Mesic to dry-mesic soils of forests, woodlands, and rocky slopes. Early reports of this species in VT (e.g., Fernald 1950b) were discredited by Jenkins and Zika (1995), the reports apparently based on specimens of Carya ovata labeled as C. alba (this latter name having been used for both species and favored for rejection by several authors).

Fig. 688  Tufts of hairs along margin of leaf blade of Carya ovata.

Juglans 1a. Carpellate inflorescence with 2–5 flowers; leaf scar lacking a pubescent band along the upper margin; branchlets with light brown pith; drupes solitary or paired, globose to subglobse; leaflets usually somewhat glossy and ± glabrous when mature . . . . . . . . . . J. nigra 1b. Carpellate inflorescence with 5–20 flowers; leaf scar with a transverse, pubescent band along the upper margin; branchlets with light or dark brown pith; drupes in racemes of (2–) 3–7 or more, ovoid to ovoid-obloid (sometimes subglobose in J. ailantifolia) [Fig. 689]; leaflets usually dull and at least sparingly pubescent adaxially 2a. Branchlets with dark brown pith, with relatively thick diaphragms that are nearly as broad as the chambers they alternate with; upper margin of leaf scar straight or scarcely notched; staminate aments 5–8 cm long; exterior of endocarp (i.e., shell) strongly 6- to 8-ridged and with irregular, ± sharp crests and projections . . . . . . . . . . . . . . . . . . . . J. cinerea 2b. Branchlets with light brown pith, with thin diaphragms that are much narrower than the chambers they alternate with; upper margin of leaf scar with a conspicuous notch; staminate aments 15–30 cm long; exterior of endocarp rugose . . . . . . . . . . . . . J. ailantifolia 1. Juglans ailantifolia Carr. E Japanese walnut. Juglans sieboldiana Maxim. • MA, RI. Roadsides, forest fragments, areas of habitation. This species was reported for ME by Magee and Ahles (1999), but the collections were taken from planted trees—7 Jul 1948, Bicknell s.n. (MAINE!). ‌1 × 2. Juglans ×bixbyi Rehd. This infrequent walnut hybrid is known from ct, mA (and likely found in other states). It is most common in the planted setting along streets and in yards, but naturalized populations have been observed. The hybrid is intermediate between its two parental species. This is perhaps best evidenced in the exterior of the endocarp (i.e., the shell), which is ridged, but not as prominently as in J. cinerea. 2. Juglans cinerea L. N Fig. 689 white walnut. Wallia cinerea (L.) Alef. • CT, MA, ME, NH, RI, VT. Rich, mesic forests, riparian forests, roadsides. This tree is sometimes cultivated and found along roadsides and near homes. 3. Juglans nigra L.

nC

black walnut. Wallia nigra (L.) Alef. • CT, MA, ME, NH, RI, VT. Roadsides, yards, field edges, deciduous forests. This species is certainly not native to most of New England; however, it is possibly native to some CT counties (fide Dowhan 1979; including Litchfield County) and should receive conservation focus. Fig. 689  Drupes of Juglans cinerea.

630  tricolpates

Lamiaceae 1a. Calyx with a small, but evident, tranverse ridge on the upper side, the upper lip deciduous in fruit [Fig. 704] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scutellaria 1b. Calyx without a protuberance on the upper side, persistent in fruit 2a. Corolla apparently composed of a lower lip only, the upper lip either very small or displaced and part of the lower lip [Figs. 690, 706] 3a. Lower lip 5-lobed, the 2 small, lateral lobes representing the laterally displaced lobes of the upper lip; inflorescence elongate, raceme-like, 5–20 cm tall, leafy-bracted only at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Teucrium 3b. Lower lip 3- or 4-lobed, the upper lip present, but very small; inflorescence either of verticillasters in the axils of foliage leaves or of flowers in leafy-bracted, spike-like inflorescences 4–8 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ajuga 2b. Corolla either clearly zygomorphic and with an evident upper and lower lip or nearly actinomorphic [Figs. 691, 694, 698] 4a. Flowers with only 2 stamens, the other 2 stamens absent or represented by small staminodes (caution: the pollen sacs are separated by a connective in Salvia, creating the appearance of 4 anthers in some species) [Figs. 692, 698] 5a. Corolla yellow to pale yellow, the distal margin of the lower lip fringed [Fig. 692]; inflorescence a terminal panicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Collinsonia 5b. Corolla white, ochroleucous, pink, red to purple, or blue (yellow and spotted with purple in Mondarda punctata), lacking a prominently fringed lower lip; inflorescence of verticillasters in axils of foliage or bracteal leaves, sometimes the verticillasters congested to form a dense cluster of flowers 6a. Calyx strongly zygomorphic; corolla blue to purple-blue (often pale blue to pale purple in Blephilia, which has red-purple to purple spots) 7a. Verticillasters with pedicellate flowers in the axils of foliage leaves; corolla 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hedeoma 7b. Verticillasters with sessile or short-pedicellate flowers in the axils of reduced, bract-like leaves, sometimes congested and forming terminal clusters of flowers; corolla 8–30 mm long 8a. Inflorescence usually composed of 5–18 or more separate verticillasters of flowers; anthers attached to a narrow connective that is articulated to the filament, the upper end of the connective bearing a single pollen sac, the lower end of the connective either with a deformed pollen sac or without a pollen sac . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Salvia 8b. Inflorescence composed of 1–5 closely crowded verticillasters of flowers, collectively forming dense, terminal clusters; anthers without a narrow connective, with 2 pollen-bearing sacs that are contiguous . . . . . . . Blephilia 6b. Calyx actinomorphic or nearly so; corolla white, ochroleucous, pink, or red to purple (often blue-purple in Monarda fistulosa) 9a. Corolla white, weakly zygomorphic, 1.8–4.4 mm long; flowers in dense but separate verticillasters in the axils of foliage leaves . . . . . . . . . . . . . . . . . Lycopus 9b. Corolla only rarely white, strongly zygomorphic, 15–45 mm long; flowers in dense, crowded verticillasters at the tips of the stems and branches [Fig. 701], sometimes also in dense, axillary clusters . . . . . . . . . . . . . . . . . . . . . . . . . . Monarda

L a m i ac e a e   63 1

4b. Flowers with 4 stamens [Figs. 694, 705] 10a. Basal, connate portion of the corolla (i.e., the tube) longer than and somewhat concealing the stamens 11a. Inflorescence terminal, spike-like, the flowers subtended by reduced, bractlike leaves and separated by conspicuously shortened internodes relative to the vegetative portion of the stem; leaf blades linear to oblong or lanceolate 12a. Corolla blue to purple, exceeding the calyx in lenth; calyx lobes without awns, the upper lobe with a obcordate appendage at the apex; plants woody near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lavandula 12b. Corolla yellow or black to dark red-brown with yellow on the lower lip, shorter than to approximately equaling the calyx; calyx lobes with prominent awns 1–2 mm long, the upper lobe lacking an appendage; plants herbaceous throughout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sideritis 11b. Inflorescence axillary, composed of few to many flowers in the axils of foliage leaves and separated by internodes of normal length (only the uppermost flowers with reduced bracts and/or internodes); leaf blades broad-elliptic to cordate-reniform 13a. Stems conspicuously white-tomentose; corolla white; lobes of the calyx numbering 10, with reflexed and spinulose apices; plants from a taproot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marrubium 13b. Stems retrorse-scabrous, hirsute, or glabrous; corolla blue-purple; lobes of the calyx numbering 5, awned at the apex; plants from rhizomes and stolons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Glechoma 10b. Basal, connate portion of the corolla shorter than the stamens, the stamens readily visible during anthesis [Figs. 693, 705] 14a. Inflorescence axillary, composed of few to many flowers in the axils of foliage leaves and separated by internodes of normal length (only the uppermost flowers with reduced bracts and/or internodes) [Fig. 700] 15a. Calyx actinomorphic or nearly so 16a. Corolla nearly actinomorphic, the 4 or 5 lobes nearly equal in length (i.e., not bilabiate; the upper lobe slightly wider than the others and often emarginate at the apex in Mentha) 17a. Each verticillaster with numerous, crowded flowers; corolla with usually 4 lobes, the upper usually emarginate; ovary lobed to the point of appearing as 4 nearly distinct segments . . . . . . . . . . . . . . . (in part) Mentha 17b. Each verticillaster with 2–6 loosely disposed flowers; corolla with 5 lobes; ovary lobed, but the segments clearly connate ca. ⅓ their length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Trichostema 16b. Corolla zygomorphic, definitely bilabiate 18a. Flowers with short pedicels 19a. Stems prostrate or trailing; each flower bearing axil with (2–) 3 flowers; upper pair of stamens longer than the lower pair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Glechoma 19b. Stems erect; each flower-bearing axil with 4–8 flowers; upper pair of stamens shorter than the lower pair . . . . . . . . . . . . . . . . . . . . . . Ballota 18b. Flowers sessile or subsessile 20a. Principal leaf blades palmately lobed; corolla 8–12 mm long, the upper lip conspicuously villous . . . . . . . . . . . . . . . . . . . . . . . . . . . Leonurus

632  tricolpate s

20b. Leaf blades simple; corolla 10–30 mm long, the upper lip glabrous or sparsely pilose 21a. Apex of sepals terminated by short, stiff spines; anthers dehiscing by latitudinal valves; mericarps broad-obovoid, rounded at the apex; lower lip of the corolla with a pair of projections near the sinuses on the adaxial (i.e., upper) surface [Fig. 694] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Galeopsis 21b. Apex of sepals acuminate, but not spiny; anthers dehiscing by longitudinal valves; mericarps three-angled, truncate or subtruncate at the apex; lower lip of the corolla without a pair of projections near the base of the lobes . . . . . . . . . . . . . . . . . . . . . . (in part) Lamium 15b. Calyx zygomorphic, with an upper and lower lip 22a. Corolla 4-lobed, nearly actinomorphic, the upper lobe slightly wider than the others and often emarginate at the apex (rarely with 5 subequal lobes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Mentha 22b. Corolla zygomorphic, definitely bilabiate 23a. Upper calyx lip with broad-triangular lobes, the lobes wider than long; corolla white or yellow (rarely becoming pink), the connate portion curved upward; leaf blades ovate to deltate-ovate; plants citrus-scented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melissa 23b. Upper calyx lip with narrow-triangular lobes, the lobes longer than wide; corolla pale purple, red-purple, or pink (rarely white), the connate portion straight; leaf blades ovate, elliptic, oblong, or narrower; plants without a citrus scent . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Clinopodium 14b. Inflorescence terminal or axillary, spike-like, raceme-like, or panicle-like, the flowers subtended by reduced, bract-like leaves and/or separated by conspicuously shortened internodes relative to the vegetative portion of the stem 24a. Inflorescence an open, panicle-like cyme 25a. Stamens with arcuate-curved filaments [Fig. 707]; leaf blades linear to rhombic-lanceolate; ovary lobed but the segments clearly connate ca. ⅓ their length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Trichostema 25b. Stamens relatively straight; leaf blades elliptic or ovate to broad-ovate; ovary lobed to the point of appearing as 4 nearly distinct segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Origanum 24b. Inflorescence spike-like (sometimes capitate) or raceme-like 26a. Each node of the inflorescence with 2 flowers, 1 each in the axil of a bract 27a. Leaf blades lanceolate or oblanceolate to elliptic, 1–2.5 cm wide, green; petioles absent on all but the lowest leaves; corolla (14–) 16–35 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physostegia 27b. Leaf blades ovate-oblong to broad-ovate, 2.8–10 cm wide, tinged or suffused with purple; petioles 30–50 mm long; corolla 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Perilla 26b. Each node of the inflorescence with 4 or more flowers 28a. Upper lip of the corolla concavely arched, appearing as a small hood (i.e., the corolla galeate) [Figs. 695, 705] 29a. Apex of sepals terminated by short, stiff spines; anthers dehiscing by latitudinal valves; lower lip of the corolla with a pair of projections near the sinuses on the adaxial (i.e., upper) surface [Fig. 694] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Galeopsis

La m i ac e a e   63 3

29b. Apex of sepals acuminate, but not spiny; anthers dehiscing by longitudinal valves; lower lip of the corolla without a pair of projections near the base of the lobes 30a. Calyx zygomorphic; filaments bifid near the apex, the lower of the tooth-like branches bearing the anther; floral bracts broadly rounded at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prunella 30b. Calyx actinomorphic or nearly so; filaments not bifid at the summit; floral bracts obtuse to acuminate at the apex 31a. Inner (i.e., upper) pair of stamens slightly to noticeably longer than the outer pair; calyx with 15 nerves 32a. Corolla 8–12 mm long, usually spotted on the lower lip; upper and lower pair of stamens ± parallel, both ascending under the upper lip of the corolla; anther sacs divergent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nepeta 32b. Corolla 4–9 mm long, unspotted; upper pair of stamens (i.e., the longer pair) downwardly directed, the lower pair of stamens upwardly oriented; anther sacs ± parallel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Agastache 31b. Inner pair of stamens slightly shorter than the outer pair; calyx with 5 or 10 nerves 33a. Median portion of inflorescence with conspicuously reduced bracteal leaf blades compared with the upper vegetative leaves; mericarps ovoid or obloid, rounded to obtusely pointed at the apex . . . . . . . . . . . . . . . . . . . . . . . Stachys 33b. Median portion of the inflorescence with scarcely reduced bracteal leaf blades compared with the upper vegetative leaves; mericarps three-angled, truncate or subtruncate at the apex . . . . . . . . . . . . . . . . . . . (in part) Lamium 28b. Upper lip of the corolla flat or upwardly curved, not hood-like [Fig. 691] 34a. Stamens, at least the longer pair, upwardly curved, closely ascending under the upper corolla lip and not surpassing it [Fig. 691] 35a. Uppermost lobe of the calyx wider than the other 4; calyx with 15 nerves; floral bracts spinose-serrate at least near the base (entire in D. thymiflorum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dracocephalum 35b. Upper 3 lobes of the calyx differing from the other 2; calyx with 10–13 nerves; floral bracts entire to crenate-serrate 36a. Corolla 5–7 mm long; leaf blades linear to narrowoblanceolate; each flower subtended by minute bracts shorter than the calyx; plants annual, without stolons, 1–3 dm tall . . . . . Satureja 36b. Corolla 12–15 mm long; leaf blades narrow-ovate or ovate to ovate-oblong; each flower subtended by ciliate, linear-subulate bracts about as long as the calyx; plants perennial from short stolons, 2–5 dm tall . . . . . . . . . . . . . . . . . . . . . (in part) Clinopodium 34b. Stamens individually straight or nearly so, at least the longer pair protruding from the corolla (except in carpellate plants of Mentha, which have smaller flowers and included anthers) [Fig. 693] 37a. Middle lobe of upper calyx lip suborbicular, with winged margins that are decurrent on the calyx tube, forming a concave cap over the top of the calyx; corolla with a 4-lobed upper lip and a single-lobed lower lip . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ocimum

634 tricolpate s

37b. Middle, upper lobe of the calyx subulate to triangular, not forming a cap over the entire calyx; corolla nearly actinomorphic or zygomorphic and then with a 3-lobed lower lip 38a. Leaf blades with entire margins 39a. Plants diffusely branched, mat-forming; leaf blades 5–10 mm long; calyx pubescent inside the connate tube . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thymus 39b. Plants upright; leaf blades 10–95 mm long; calyx ± glabrous inside the connate tube 40a. Calyx 10- to 13-nerved; anthers with parallel pollen sacs; verticillasters with densely crowded flowers, forming capitate cymes at the apex of the stems and branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Pycnanthemum 40b. Calyx 15-nerved; anthers with divergent pollen sacs; verticillasters with 3–7 flowers in the axils of bracts, forming a terminal, spike-like inflorescence . . . . . . . . . . . . . Hyssopus 38b. Leaf blades with toothed margins 41a. Inflorescence secund [Fig. 693]; anthers with divergent pollen sacs; plants annual, without rhizomes . . . . . . Elsholtzia 41b. Inflorescence not secund; anthers with ± parallel pollen sacs; plants rhizomatous perennials (sometimes from a woody caudex in Agastache) 42a. Corolla nearly actinomorphic, the lower lip unspotted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Mentha 42b. Corolla zygomorphic, the lower lip spotted with purple in commonly observed species 43a. Calyx 15-nerved; upper lip of corolla somewhat convex and subgaleate; lower lip of corolla unspotted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Agastache 43b. Calyx 10- to 13-nerved; upper lip of corolla relatively flat or upwardly curved, not forming a hood; lower lip of corolla commonly spotted with purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Pycnanthemum

Agastache Reference: Lint and Epling (1945). 1a. Leaf blades abaxially whitened by dense, minute, white hairs; calyx minutely pubescent; petioles of middle leaves 10–15 (–20) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. foeniculum 1b. Leaf blades green abaxially, the pubescence (when present) not altering the background color of the blade; calyx glabrous; petioles of middle leaves (15–) 20–50 mm long 2a. Calyx lobes 1–1.5 mm long, narrow-ovate; corolla green-yellow; inflorescence 10–15 mm thick; floral bracts broad-ovate and tipped by a short, acuminate point; stems glabrous or minutely pubescent on middle internodes; leaf blades pubescent across the surface abaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. nepetoides 2b. Calyx lobes 2–2.5 mm long, narrow-triangular; corolla purple; inflorescence 15–20 mm thick; floral bracts round-ovate and tipped by an elongate, caudate-acuminate point; stems with some longer hairs on the middle internodes; leaf blades pubescent mainly on the major veins abaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. scrophulariifolia

L a m i ac e a e   63 5

1. Agastache foeniculum (Pursh) Kuntze E lavender giant-hyssop. Agastache anethiodora (Nutt.) Britt.; Hyssopus anethiodorus Nutt. • CT, NH, RI. Fields, roadsides, gardens, lawn edges, dumps. This species has a pubescent calyx

with blue lobes. 1 × Agastache rugosa (Fisch. & C.A. Mey.) Kuntze is a rare giant-hyssop hybrid that is known from MA. It distinguished by its pubescent calyx, connate portion of corolla 7–8 (–10) mm long, and leaf blades prominently cordate at base. It lacks the dense, white pubescence on the abaxial leaf surface of Agastache foeniculum and has a longer connate portion of the corolla (this 6.5–7.5 mm long in A. foeniculum). This hybrid is a planted species and sometimes escapes from the garden setting. 2. Agastache nepetoides (L.) Kuntze

NC

catnip giant-hyssop. Hyssopus nepetoides L. • CT, VT; also reported from MA by Lint and Epling (1945), but specimens are unknown. Forest borders and fragments, shaded roadsides, rocky banks. Sorrie and Somers (1999) also have found no evidence that this species occurs in MA. 3. Agastache scrophulariifolia (Willd.) Kuntze

NC

purple giant-hyssop. Agastache scrophulariifolia (Willd.) Kuntze var. mollis (Fern.) Heller; Hyssopus scrophulariifolius Willd. • CT, MA, VT; also reported from NH by Lint and Epling (1945), but specimens are unknown. Forests, frequently dry-mesic, rocky types, forest fragments, roadsides, river banks, riparian forests.

Ajuga 1a. Plants upright, not forming mats; upper lip of the corolla with 2 teeth; stems usually evenly pubescent on each face (varying to subglabrous); flowers in verticillasters of 6 or more at each node . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. genevensis 1b. Plants with stolons, mat-forming, the flowering stems upright; upper lip of the corolla entire; stems usually pubescent on opposite faces in lines of decurrence from the leaves, alternating faces at each node; flowers in verticillasters of usually 6 at each node . . . . . . . . . . . . . A. reptans 1. Ajuga genevensis L. E standing bugle. CT, MA, ME, RI, VT. Lawns, fields, open rights-of-way, outcrops, areas of habitation. 2. Ajuga reptans L. E Fig. 690 carpet bugle. CT, MA, ME, RI, VT. Lawns, fields, gardens, waste areas, areas of habitation.

Ballota See Morton (1966) for nomenclature of our taxa of Ballota. 1. Ballota nigra L. E black-horehound. 1a. Ballota alba L.; B. borealis Schweig.; B. nigra L. ssp. foetida (Koch) Hayek; 1b. Ballota nigra L. var. ruderalis (Sw.) Koch • CT, MA. Dumps, waste areas. 1a. Calyx lobes 1.5–3 mm long excluding the mucros, ovate to triangular-ovate, with a tiny mucro at the apex 0.2–0.5 (–1.7) mm long . . . . . . . . . . . . . . . . . . . . . 1a. B. nigra var. alba (L.) Sm. 1b. Calyx lobes (3–) 4–6.5 mm long excluding the awns, triangular-lanceolate to subulatelanceolate, with a prolonged awn at the apex 1.5–3 mm long . . . . . . . . . 1b. B. nigra var. nigra Variety alba is known from CT. It has often been referred to by the epithet foetida, but this name does not have priority at the varietal rank. Variety nigra is known from MA.

Blephilia 1a. Lower calyx lobes 1–1.7 mm long, extending beyond the sinuses of the upper calyx lobes; stems puberulent with recurved hairs mostly 0.1–0.5 mm long; leaf blades subentire or with low teeth; lowest node of inflorescence subtended by leaves with petioles shorter than 5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. ciliata

Fig. 690  Flowers of Ajuga reptans.

636  tricolpates

1b. Lower calyx lobes up to 1 mm long, not extending beyond the sinuses of the upper calyx lobes; stems hirsute with spreading hairs mostly 1–2 mm long (very rarely glabrous); leaf blades serrate, often saliently so; lowest node of inflorescence subtended by leaves with petioles 10 mm long or longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. hirsuta 1. Blephilia ciliata (L.) Benth.

NC

downy wood-mint. Monarda ciliata L. • CT, MA, VT; also reported from RI by George (1992), but specimens are unknown. Mesic, deciduous forests, roadsides, meadows, banks, dry fields and pastures. 2. Blephilia hirsuta (Pursh) Benth.

NC

hairy wood-mint. 2b. Monarda hirsuta Pursh • CT, MA, VT; western New England. Rich, sometimes rocky, upland and riparian forests, cliff bases, dry fields. 1a. Stems and leaf blades ± glabrous . . . . . . . . . . . . . . . . . . . . 2a. B. hirsuta var. glabrata Fern. 1b. Stems and leaf blades conspicuously pubescent . . . . . . . . . . . . . 2b. B. hirsuta var. hirsuta Variety glabrata is known from VT (and is endemic to that state). It was collected in a dry field (atypical habitat compared with var. hirsuta). Variety hirsuta is known from CT, MA, VT.

Clinopodium See Cantino and Wagstaff (1998) for circumscription of this genus. Clinopodium nepeta (L.) Kuntze was reported from RI by George (1992), but specimens are unknown. 1a. Verticillasters with 2–6 flowers in the axils of many of the foliage leaves, subtended by minute bracts shorter than the calyx; calyx pubescent inside the connate tube; plants annual or short-lived perennial, without stolons, 10–20 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . C. acinos 1b. Verticillasters with numerous flowers in dense clusters at the stem tip and 1–3 of the upper axils, each flower subtended by ciliate, linear-subulate bracts about as long as the calyx [Fig. 691]; calyx glabrous inside the connate tube; plants perennial from short stolons, 20–50 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. vulgare 1. Clinopodium acinos (L.) Kuntze E Fig. 691  Inflorescence of Clinopodium vulgare.

basil-thyme. Acinos arvensis (Lam.) Dandy; Calamintha acinos (L.) Clairville ex Gaud.; Satureja acinos (L.) Scheele; Thymus acinos L. • CT, MA, ME, VT. Fields, roadsides, trail edges, waste areas. 2. Clinopodium vulgare L. N Fig. 691 wild basil. Calamintha vulgaris (L.) Halácsy; Satureja vulgaris (L.) Fritsch; S. vulgaris (L.) Fritsch var. neogaea Fern. • CT, MA, ME, NH, RI, VT. Fields, roadsides, ledges, rocky forests and woodlands.

Collinsonia Collinsonia punctata Ell. was reported from CT by Dowhan (1979), presumably based on the following annotated sheet—31 Jul 1884, Woodward s.n. (NCBS; image seen!). The specimen is C. canadensis, as evidenced by the smaller calyx and corolla (and as considered by the collector). Reference: Peirson et al. (2006). 1. Collinsonia canadensis L. N Fig. 692 northern horsebalm. CT, MA, NH, RI, VT. Mesic, deciduous, often rich, forests and riparian terraces.

Fig. 692  Flowers of Collinsonia canadensis.

Dracocephalum 1a. Corolla (15–) 20–25 (–30) mm long, the basal, connate portion distinctly exserted from the calyx; each node of the inflorescence with 4–6 (–10) flowers . . . . . . . . . . . . . . . . . D. moldavica 1b. Corolla 7–15 mm long, the basal, connate portion not or only slightly exserted from the calyx; each node of the inflorescence with 6–12 flowers

L a m i ac e a e   637

2a. Floral bracts spinose-serrate; lower stem leaves with blades 25–80 mm long; calyx 6–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. parviflorum 2b. Floral bracts entire or ciliate; lower stem leaves with blades 10–35 mm long; calyx 10 mm long or longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. thymiflorum 1. Dracocephalum moldavica L. E Moldavian dragonhead. Moldavica moldavica (L.) Britt. • CT, VT. Fields, roadsides, waste areas. 2. Dracocephalum parviflorum Nutt.

nC

American dragonhead. Moldavica parviflora (Nutt.) Britt. • CT, MA, ME, NH, RI, VT. Fields, railroads, yards, lawns, gardens, waste areas, woodlands. This species is non-native to most of New England, but a recent collection from Rutland County, VT, is likely native (and that population is of regional conservation concern). 3. Dracocephalum thymiflorum L. E thyme-leaved dragonhead. Moldavica thymiflora (L.) Rydb. • MA. Wool waste.

Elsholtzia 1. Elsholtzia ciliata (Thunb.) Hyl. E Fig. 693 crested late-summer-mint. Elsholtzia cristata Willd.; Sideritis ciliata Thunb. • CT, MA, ME, VT. Roadsides, yards, railroads, disturbed sites.

Galeopsis Reference: Townsend (1972). 1a. Stems pubescent with minute, recurved, eglandular hairs (longer glandular hairs may also be present as well), not swollen at the nodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. ladanum

Fig. 693  Secund inflorescence and upper leaves of Elsholtzia ciliata.

1b. Stems pubescent with long, rigid, spreading to slightly recurved, eglandular hairs (shorter glandular hairs may also be present as well), swollen at the nodes 2a. Central lobe of lower corolla lip emarginate, the sides deflexed so that the lobe is somewhat convex [Fig. 694]; corolla 13–15 (–16) mm long, usually largely pink . . . . G. bifida 2b. Central lobe of lower corolla lip entire, flat; corolla 15–23 (–28) mm long, usually largely white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. tetrahit 1. Galeopsis bifida Boenn. E Fig. 694 split-lipped hemp-nettle. Galeopsis tetrahit L. var. bifida (Boenn.) Lej. & Court. • CT, MA, ME, NH, VT. Fields, roadsides, waste areas, gardens, disturbed soil. 2. Galeopsis ladanum L. E red hemp-nettle. Galeopsis ladanum L. var. latifolia (Hoffman) Wallr. • CT, MA, ME, RI, VT. Waste areas, ballast, sandy locations near the Atlantic coast. This species has usually a largely pinkred corolla 15–28 mm long. 3. Galeopsis tetrahit L. E brittle-stemmed hemp-nettle. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, yards.

Glechoma 1. Glechoma hederacea L. E Gill-over-the-ground. Glechoma hederacea L. var. micrantha Moric.; G. hederacea L. var. parviflora (Benth.) House; Nepeta hederacea (L.) Trevisan • CT, MA, ME, NH, RI, VT. Fields, lawns, dumps, about dwellings.

Fig. 694  Flower of Galeopsis bifida.

638  tricolpate s

Hedeoma 1a. Leaf blades lanceolate to ovate or obovate, 4–15 mm wide, the principal ones petiolate; upper calyx lobes triangular and eciliate, the lower ones subulate and ciliate; mericarps subglobose, 0.7–1 mm long, not glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. pulegioides 1b. Leaf blades linear to narrow-oblong, 1–3 mm wide, sessile or subsessile; all the calyx lobes subulate and ciliate; mericarps narrow-ovoid, 1–1.3 mm long, glaucous . . . . . . . . . . . . H. hispida 1. Hedeoma hispida Pursh N rough false pennyroyal. CT, MA, ME, NH, RI, VT; rare in the northeastern states (e.g., ME, NH). Dry fields and banks, high-pH rocks and ledges, railroads, waste areas. 2. Hedeoma pulegioides (L.) Pers. N American false pennyroyal. Cunila pulegioides L.; Melissa pulegioides (L.) L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, ledges, trail edges.

Hyssopus 1. Hyssopus officinalis L. E hyssop. CT, MA, ME, VT. Roadsides, abandoned homesteads.

Lamium Reference: Mennama (1989). 1a. Corolla yellow; lower lip of the corolla with three evident lobes (i.e., the lateral lobes welldeveloped and nearly as long as the middle lobe); anthers glabrous . . . . . . . . L. galeobdolon 1b. Corolla white to pink to purple; lower lip of the corolla with a prominent middle lobe that is much larger than the reduced or wanting lateral lobes; anthers pubescent 2a. Corolla (18–) 20–35 mm long, with an upper lip 7–14 mm long, the basal, connate portion curved at the base [Fig. 695]; plants perennial from rhizomes 3a. Corolla pink to brown-purple (rarely white), with a ring of spreading hairs on the adaxial (i.e., inside) surface 2–3 mm from the base; leaf blades commonly with a white blotch bordering each side of the midrib; terminal tooth of the upper leaf blades usually obtuse at the apex; pollen orange. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. maculatum 3b. Corolla white (rarely pink), with a ring of backward pointing hairs on the adaxial surface 4–5 mm from the base; leaf blades without a white blotch; terminal tooth of the upper leaf blades acute at the apex; pollen light yellow . . . . . . . . . . . . . . . . . . . . . L. album 2b. Corolla 10–20 (–25) mm long, with an upper lip 3–5 mm long, the basal, connate portion straight; plants annual, without rhizomes 4a. Upper leaves and bracts sessile and clasping; calyx lobes erect (i.e., oriented straight forward); flowers without a subtending bractlet; connate portion of the corolla (i.e., the tube) glabrous on the inner surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. amplexicaule 4b. All the leaves petiolate, the upper bracts sessile but not clasping; calyx lobes spreading; flowers commonly with a subtending bractlet; connate portion of the corolla sometimes pubescent on the inner surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. purpureum 1. Lamium album L. ssp. album E white henbit. MA, ME, RI. Roadsides, waste areas, gardens. 2. Lamium amplexicaule L. var. amplexicaule E common henbit. CT, MA, ME, NH, RI, VT. Fields, lawns, gardens, about dwellings.

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3. Lamium galeobdolon (L.) L. ssp. galeobdolon E yellow henbit. Galeobdolon argentatum Smejkal; G. luteum Huds.; Lamiastrum galeobdolon (L.) Ehrend. & Polatschek • MA, ME. Gardens, waste areas, forest borders. 4. Lamium maculatum L. E Fig. 695 spotted henbit. CT, MA, ME, NH, VT. Roadsides, lawns, waste areas, gardens, about dwellings. 5. Lamium purpureum L. E red henbit.  5b. Lamium dissectum With.; L. hybridum, auct. non Vill. • CT, MA, ME, NH, RI, VT. Fields, waste areas. 1a. Leaf blades and bract blades crenate to crenate-serrate, not decurrent along the petioles, those subtending flowers with teeth shorter than 2 mm; mericarps (2–) 2.2–2.5 (–2.8) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5a. L. purpureum var. purpureum 1b. Leaf blades and bract blades prominently incise-crenate and decurrent along the petioles, particularly the upper ones, those subtending flowers with teeth longer than 2 mm; mericarps (2.5–) 2.7–3 (–3.3) mm long . . . . . . . . . . . . . . . . . . . . . 5b. L. purpureum var. incisum (Willd.) Pers. Variety purpureum is known from CT, MA, ME, NH, RI, VT. Variety incisum is known from CT, MA, ME, NH, VT.

Lavandula 1. Lavandula angustifolia P. Mill. ssp. angustifolia E English lavender. Lavandula officinalis Chaix; L. spica L.; L. vera DC. • VT. Dry fields.

Leonurus 1. Leonurus cardiaca L. E motherwort. Leonurus cardiaca L. ssp. villosus (Desf. ex Spreng.) Hyl.; L. cardiaca L. var. villosus (Desf. ex Spreng.) Benth. • CT, MA, ME, NH, RI, VT. Fields, banks, forest borders, waste areas, about dwellings.

Lycopus Lycopus superficially resembles some species of Mentha. The two genera can be separated using a number of characters, including stamen number (2 in Lycopous, 4 in Mentha), mericarp shape (truncate at apex in Lycopus and sometimes with a crest or tubercles, rounded at apex in Mentha), and scent of fresh foliage (weakly or not aromatic in Lycopus, strongly mint- or fruit-scented in Mentha). Reference: Henderson (1962). 1a. Lobes of the calyx broad-triangular, shorter than 1 mm, less than 2 times as long as wide, obtuse to subacute at the apex, equaled or surpassed by the mericarps at maturity; lower leaf blades serrate to coarsely serrate 2a. Corolla 5-lobed, with flaring lobes; calyx with 5 lobes; plants with tubers; interior angles of the mericarps shorter than the 2 lateral angles, therefore, the apex of the combined mericarps concave [Fig. 696]; leaf blades and stem glabrous to sparsely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. uniflorus 2b. Corolla 4-lobed (sometimes the upper lobed notched), with forward-oriented lobes [Fig. 698]; calyx with 4 lobes; plants without tubers; interior angles of the mericarps as long as the 2 lateral angles, therefore, the apex of the combined mericarps flat across the apex [Fig. 697]; leaf blades and stem pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. virginicus 1b. Lobes of the calyx narrow-triangular, 1–2.1 (–2.5) mm long, greater than 2 times as long as wide as long, acuminate at the apex, surpassing the mericarps at maturity [Fig. 696]; lower leaf blades serrate to pinnately lobed

Fig. 695  Flowers of Lamium maculatum showing the upward curved basal portion of the corolla.

640 tricolpate s

3a. Corolla 4-lobed; interior angles of the mericarps shorter than the 2 lateral angles, therefore, the apex of the combined mericarps concave [Fig. 696]; plants without tubers (except in L. asper, a rare introduction) 4a. Leaf blades sessile or subsessile; rhizomes tuberous-thickened at tip; mericarps (1.3–) 1.4–1.8 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. asper 4b. Leaf blades borne on evident petioles; rhizomes without tubers; mericarps 0.7–1.3 mm wide 5a. Leaf blades pubescent abaxially with hairs 0.01–0.5 mm long; fresh anthers 0.26–0.5 mm long; calyx 2–3.3 mm long; mericarps 1–1.37 mm long, the collar-like corky crest separated by a distance of 0.1–0.3 mm at the base on the adaxial (i.e., inner) surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. americanus 5b. Leaf blades pubescent abaxially with hairs 0.5–1.6 mm long; fresh anthers 0.5–0.7 mm long; calyx 3–4.5 mm long; mericarps 1.3–1.73 mm long, the collar-like corky crest completely encircling the mericarp or separated by up to 0.2 mm at the base on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. europaeus 3b. Corolla 5-lobed or 4-lobed with the upper lobe prominently notched; interior angles of the mericarps as long as the 2 lateral angles, therefore, the apex of the combined mericarps flat across the apex [Fig. 697]; plants with tubers 6a. Leaf blades lanceolate to elliptic, either concavely narrowed from the basal-most teeth to a narrow base or with an evident petiole . . . . . . . . . . . . . . . . . . . . . . . . . L. rubellus Fig. 696  Schizocarps of Lycopus americanus showing concave apex of fruit.

6b. Leaf blades oblong to oblong-lanceolate, convexly narrowed from the basal-most teeth to a broad-cuneate to rounded base . . . . . . . . . . . . . . . . . . . . . . . . . . . L. amplectens 1. Lycopus americanus Muhl. ex W. Bart. N Fig. 696 American water-horehound. Lycopus americanus Muhl. ex W. Bart. var. longii Benner; L. americanus Muhl. ex W. Bart. var. scabrifolius Fern.; L. europaeus L. var. sinuatus (Ell.) Gray; L. pennsylvanicus Muhl. ex Raf.; L. pinnatifidus Raf.; L. sinuatus Ell.; Phytosalpinx americanus (Muhl.) Lunell • CT, MA, ME, NH, RI, VT; nearly throughout. Swamps, shorelines, wetland margins. 2. Lycopus amplectens Raf. N Fig. 697 clasping water-horehound. Lycopus amplectens Raf. var. pubens (Britt.) Fern.; L. pubens Britt.; L. sessilifolius Gray • CT, MA, RI; primarily along the coastal plain. Swamps, pond shores, often on sandy and/or peaty soils. 3. Lycopus asper Greene E

Fig. 697  Schizocarps of Lycopus amplectens showing relatively plane apex of fruit.

western water-horehound. Lycopus lucidus Turcz. ex Benth. ssp. americanus (Gray) Hultén; L. lucidus Turcz. ex Benth. var. americanus Gray; L. maritimus Greene; Phytosalpinx aspera (Greene) Lunell • MA. Marshes, shorelines. 4. Lycopus europaeus L. E European water-horehound. Lycopus europaeus L. ssp. mollis (Kern.) Rothm. ex Skalickÿ; L. europaeus L. var. mollis (Kern.) Briq.; L. mollis Kern.; L. vulgaris Pers. • MA. Roadsides, waste areas. 5. Lycopus rubellus Moench

NC

stalked water-horehound. Lycopus rubellus Moench var. arkansanus (Fresen.) Benner; L. rubellus Moench var. lanceolatus Benner; L. velutinus Rydb. • CT, MA, RI; primarily along the coastal plain, though disjunct in Berkshire County, MA. Swamps, pond shores, stream banks. This species was reported by Seymour (1982) to occur in NH and VT. All collections seen by me that were originally determined as Lycopus rubellus from those states were based on specimens of Lycopus americanus. The report of this species from Oxford County, ME, by Henderson (1962) is discredited, as it is far too north for this taxon. 6. Lycopus uniflorus Michx. N northern water-horehound. Lycopus parviflorus Maxim.; L. virginicus L. var. pauciflorus Benth. • CT, MA, ME, NH, RI, VT. Swamps, shorelines, wetland margins.

L a m i ac e a e   6 41

7. Lycopus virginicus L. N Fig. 698 Virginia water-horehound. Lycopus membranaceus Bickn. • CT, MA, ME, NH, RI, VT. Swamps, shorelines, wetland margins.

Marrubium 1. Marrubium vulgare L. E white horehound. CT, MA, ME, RI, VT. Waste areas, dumps, about dwellings.

Melissa 1. Melissa officinalis L. E lemon-balm. CT, MA, RI, VT. Roadsides, field edges, gardens, waste areas. Report of this species by Haines and Vining (1999) was based on an incorrectly identified specimen. The collection is Hedeoma pulegioides— early Aug 1992, unknown s.n. (MAINE!).

Mentha A difficult genus complicated by hybridization and polyploidy. Misapplication of names has caused tremendous confusion, and often the names that authors have used actually apply to different species or hybrids (the correct name can sometimes be determined by examining the characters they used to identify the taxa). Many species of Mentha produce smallflowered, carpellate plants in addition to the normal ones with bisexual flowers. The bisexual flowers have exserted stamens at anthesis, the carpellate ones do not. Mentha suaveolens Ehrh. was reported from ME by Gleason and Cronquist (1991), but specimens are unknown. Further, reports of this species are complicated by the fact that usage of this name and its ‌rotundifolia) hybrid with M. longifolia (M. × are frequently confused. References: Harley (1972), Tucker and Naczi (2007). 1a. Inflorescence consisting largely of verticillasters of flowers in the axils of normal foliage leaves separated by internodes of normal length [Fig. 700] 2a. Leaf blades narrow-oblanceolate to rhombic-elliptic (infrequently to ovate), usually cuneate at the base [Fig. 700], (1.5–) 1.7–3.3 (–4.1) times as long as wide, gradually decreasing in size toward the apex of the plant; calyx 2–3 mm long, the subulate to narrow-triangular (rarely triangular) lobes (0.6–) 0.7–0.9 (–1.1) mm long; fresh plants with pennyroyal or peppermint odor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. canadensis 2b. Leaf blades ovate to suborbicular, rounded at the base, 1.3–2.3 times as long as wide, not gradually descreasing in size toward apex of plant, the uppermost 1 or 2 nodes with conspicuoulsy reduced leaves; calyx 1.5–2.5 mm long, the narrow-triangular to broadtriangular lobes 0.5–0.8 (–0.88) mm long; fresh plants with sweet fruit odor (rarely pennyroyal-scented) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. arvensis 1b. Inflorescence consisting of congested (at least in part) verticillasters of flowers that are subtended by reduced, bracteal leaves and separated by highly shortened internodes [Fig. 699] 3a. Inflorescence composed of 2 or 3 closely aggregated verticillasters of flowers, collectively forming a globose to short-ovoid cluster of flowers 15–25 (–30) × 12–20 mm (sometimes also with 1–3 additional separate verticillasters of flowers from the upper axils of foliage leaves) [Fig. 699] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. aquatica 3b. Inflorescence composed of 4–19 or more verticillasters of flowers, elongate and cylindric, (30–) 40–100 (–150) × 5–10 (–15) mm 4a. Leaf blades ovate to suborbicular, 1–2 times as long as wide, mostly shorter than 45 mm, rugose-veiny abaxially, usually strongly so . . . . . . . . . . . . . . . . (in part) M. spicata 4b. Leaf blades lanceolate or oblong-lanceolate to oblong-ovate, 2–3.5 times as long as wide, mostly longer than 45 mm, not or scarcely rugose-veiny abaxially

Fig. 698  Flower of Lycopus virginicus.

642   tricolpate s

5a. Leaf blades oblong-lanceolate to oblong-elliptic or oblong-ovate, widest near the middle, pubescent entirely with simple hairs; stems pubescent; pollen-bearing anthers 0.28–0.38 mm long; mericarps 0.54–0.79 mm long; fresh plants with musty odor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. longifolia 5b. Leaf blades lanceolate to narrow-ovate, widest near the base, sometimes with a few scattered dendroid hairs on the abaxial surface; stems glabrous (rarely sparsely pubescent); pollen-bearing anthers 0.38–0.58 mm long; mericarps 0.74–0.94 mm long; fresh plants with spearmint odor . . . . . . . . . . . . . . . . . . . . . . . . (in part) M. spicata 1. Mentha aquatica L. E Fig. 699 water mint. 1a. Mentha hirsuta Huds.; M. palustris P. Mill.; 1b. Mentha adspersa Moench; ‌piperita M. ‌piperita citrata Ehrh.; M. × L. ssp. citrata (Ehrh.) Briq.; M. × L. var. citrata (Ehrh.) Rouy • CT, MA, ME, RI. Wet-mesic to hydric soil of roadsides and ditches.

Fig. 699  Inflorescence of Mentha aquatica.

1a. Calyx pubescent; leaf blades usually pubescent, tending to be truncate at the base; fertile plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. M. aquatica var. aquatica 1b. Calyx glabrous; leaf blades glabrous or nearly so, tending to be subcordate at the base; sterile plants (i.e., not producing fruit) . . . . . . . . . . . 1b. M. aquatica var. citrata (Ehrh.) Fresen. Variety aquatica is known from CT, MA, ME. Variety citrata is known from CT, RI. 1‌ × 2. Mentha ×verticllata L. is a rare mint hybrid known from CT, VT. It is variable (like many mint hybrids), and the inflorescence varies from subverticillate to subcapitate. However, the verticillasters are most often subtended by bracts that ± resemble foliage leaves (at least the lower), so the hybrid is most often confused with species like M. arvensis and M. canadensis. It is normally a sterile, pubescent plant with a calyx 2.5–3.5 (–4) mm long. ‌1 × 5. Mentha ×piperita L. is an infrequent mint hybrid known from CT, MA, ME, NH, RI, VT. It most closely resembles M. aquatica. The hybrid has a relatively dense, obloid inflorescence (25–) 45–80 × 12–20 mm, calyx with pubescence only on the lobes (i.e., the basal, connate portion is glabrous), usually glabrous pedicels, and the abaxial leaf blade surface is glabrous or with hairs along only the main veins (vs. basal, connate portion of the calyx pubescent, pedicels pubescent, and abaxial leaf blade surface sparsely to densely pubescent in M. aquatica var. aquatica). Further, M. × ‌piperita is sterile (vs. fertile in M. aquatica var. aquatica). From M. aquatica var. citrata it is separated by its pubescent calyx lobes and lanceolate to narrow-ovate leaf blades that are acute at the apex (vs. glabrous calyx lobes and ovate leaf blades that are obtuse at the apex in M. aquatica var. citrata). 2. Mentha arvensis L. ssp. parietariaefolia (Becker) Briq. E ginger mint. Mentha arvensis L. var. parietariaefolia Becker; M. gentilis L.; M. variegata Sole • CT, MA, ME, RI, VT. Swamps, wetland borders, shorelines, about dwellings. The name Mentha gentilis has long been considered to apply to hybrids between M. arvensis and M. spicata. However, the type of that name has been shown to be a pollen-sterile clone of M. arvensis (treated as ssp. parietariaefolia). This plant sometimes has white-spotted leaves due to a viral infection. 2 ‌ × 5. Mentha ×gracilis Sole is an infrequent mint hybrid known from CT, MA, ME, NH, RI, VT. It has an inflorescence somewhat similar to Mentha canadensis in that the lower verticillasters of flowers are subtended by scarcely reduced leaves; however, the bracts become more highly reduced and the inflorescence often becomes somewhat congested in the upper 2 or 3 verticillasters. It differs in that the calyx is 2–3.5 mm long, with lobes 0.8–1.2 mm long, with a glabrous basal, connate portion, and the plants are sterile (vs. calyx 2–3 mm long, with lobes mostly 0.7–0.9 mm long, with a pubescent basal, connate portion, and plants fertile in M. canadensis). Further, the hybrids are often tinged with red. This plant has long gone by the names M. gentilis and M. cardiaca. The former name applies to a species (see above) and the latter name is a superfluous synonym of this hybrid.

L a m i ac e a e   6 43

3. Mentha canadensis L. N Fig. 700 American wild mint. Mentha arvensis L. ssp. canadensis (L.) H. Hara; M. arvensis L. var. canadensis (L.) Kuntze; M. arvensis L. var. glabrata (Benth.) Fern.; M. arvensis L. ssp. haplocalyx Briq.; M. arvensis L. var. villosa (Benth.) S.R. Stewart • CT, MA, ME, NH, RI, VT. Swamps, wetland borders, shorelines, low fields, disturbed areas. The status of Mentha canadensis has long been debated and what rank, if any, to afford this native North American species has been controversial. Though morphological differences exist that allow separation of most North American and European populations, these differences are not always constant and overlap occurs in some individuals. However, other data (e.g., cytology, geography, phytochemicals, hybridization experiments) show that two taxa should be recognized (Gill et al. 1973, Tucker and Chambers 2002) and that M. candensis is an ancient hybrid between M. arvensis and M. longifolia. Most collections determined as M. arvensis are in fact M. canadensis. 4. Mentha longifolia (L.) L. E horse mint. Mentha mollissima Borkh. ex P. Gaertn. et al.; M. spicata L. var. longifolia L.; M. sylvestris L. • CT, MA. Roadsides, shorelines, about dwellings. ‌4 × M. suaveolens Ehrh. Mentha ×rotundifolia (L.) Huds. is a fertile mint hybrid known from CT, MA, ME. It is capable of forming hybrid swarms with its parental taxa. Morphologically very diverse and the extremes approach each parent. It usually has pubescent stems, tall slender spike-like inflorescences with largely or wholly green bracts, leaf blades 1–2 times as long as wide that are pubescent and rugose-veiny abaxially, and small pollen-bearing anthers (mostly 0.28–0.38 mm long). 5. Mentha spicata L. ssp. spicata E spearmint. Mentha viridis (L.) L. • CT, MA, ME, NH, RI, VT. Roadsides, ditches, wet fields, shorelines. Mentha spicata is thought to be an allopolyploid derived from M. longifolia and M. suaveolens (Harley 1972). It shows variation in leaf blade dimensions, prominence of leaf veins, and pubes­ cence. Rare plants of M. spicata show short, broad, and rugose leaf blades. These plants, which are transitional to M. suaveolens, are best identified by examination of anthers and fruits (M. suaveolens has pollen-bearing anthers 0.28–0.38 mm long and mericarps 0.57–0.75 mm long). ‌ × M. suaveolens Ehrh. Mentha ×villosa Huds. nothovar. alopecuroides (Hull) 5 Briq. is a morphologically variable, sterile mint hybrid known from CT, MA, VT. It is most similar to M. suaveolens, especially in leaf blade shape and indument characters (i.e., leaf blades ovate to suborbicular, 1–2 times as long as wide, pubescent and rugoseveiny abaxially) but tends to be more robust, standing 60–140 cm tall (vs. 40–100 cm), with a thicker inflorescence 12–15 mm wide (vs. 5–10 mm), larger leaf blades 40–80 × 30–60 mm (vs. (15–) 30–45 × (10–) 20–40 mm), a pink corolla (vs. white or pink), and has a spearmint odor as in M. spicata (rather than sweet fruit odor).

Monarda Monarda citriodora Cerv. ex Lag. was reported from MA by Knowlton and Deane (1923a), but specimens are unknown. Reference: Scora (1967). 1a. Inflorescence consisting of 2–5 separate verticillasters of flowers (or a solitary verticillaster in depauperate plants); corolla yellow with red-purple spots, the upper lip strongly arcuate; stamens not exserted beyond the upper corolla lip; leaf blades (5–) 10–15 (–20) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. punctata 1b. Inflorescence consisting of a single terminal verticillaster of flowers (sometimes with a second verticillaster below the terminal one) [Fig. 701]; corolla pink to purple, red, red-purple, or white to ochroleucous, unspotted or spotted, the upper lip straight or slightly curved; stamens exserted beyond the upper lip by (1–) 2–4 mm; leaf blades (8–) 15–80 mm wide 2a. Corolla 30–45 mm long, red (rarely red-purple), not long-villous near the apex of the upper lip (though minute hairs may be present); bract blades usually deeply tinged with red; leaf blades 7–15 cm long, borne on petioles 10–40 mm long . . . . . . . . . . . . . . M. didyma

Fig. 700  Inflorescence of Mentha canadensis.

644   tricolpate s

2b. Corolla 15–35 mm long, pink to purple, red-purple, or white to ochroleucous, longvillous near the apex of the upper lip or not; bract blades green or tinged with white, gray, or purple; leaf blades 6–12 cm long, borne on petioles 10–20 (–30) mm long 3a. Corolla light purple (rarely purple to red-purple or white), softly puberulent on the abaxial (i.e., outer) surface as well as densely villous near the apex of the upper lip with hairs that have pale septa; calyx densely hirsute within with erect hairs ⅓ to ½ as long as the calyx lobes (short-villous within in var. rubra); leaf blades lanceolate or triangularlanceolate to narrow-ovate, (8–) 15–38 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . M. fistulosa 3b. Corolla red-purple or white to ochroleucous (rarely pink), glabrous or softly puberulent on the abaxial surface, not or only sparsely villous near the apex of the upper lip with hairs that have red-purple septa; calyx glabrous within or sparsely villous with hairs shorter than ⅓ the length of the calyx lobes; leaf blades broadlanceolate to ovate or triangular-ovate, 20–60 (–75) mm wide 4a. Corolla red-purple, unspotted, minutely puberulent on the abaxial surface and usually the upper lip sparsely villous with hairs that have red-purple septa; upper lip of corolla 13–16 mm long; bract blades tinged or completely suffused with purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. media 4b. Corolla white to ochroleucous and spotted with purple on the lower lip (rarely the corolla pink), glabrous or minutely puberulent on the abaxial suface, the upper lip without long, villous hairs; upper lip of corolla 5–8 mm long; bract blades often tinged with white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. clinopodia 1. Monarda clinopodia L. E basil bee-balm. CT, MA, RI, VT. Fields, roadsides, waste areas. The flowers of Monarda clinipodia are fragrant as they are adapted for insect pollination. 2. Monarda didyma L. E scarlet bee-balm. Monarda coccinea Michx. • CT, MA, ME, NH, VT; also reported from RI by George (1997), but specimens are unknown. Roadsides, waste ares, gardens, about dwellings. The flowers of Monarda didyma are inodorous as they are primarily pollinated by birds. 3. Monarda fistulosa L. n Fig. 701 wild bee-balm.  3b. Monarda fistulosa L. forma albescens Farw.; 3c. Monarda mollis L.; M. scabra Beck • CT, MA, ME, NH, RI, VT. Fields, roadsides, forests, clearings, forest fragments. 1a. Corolla purple to red-purple; middle lobe of the lower corolla lip 4–6 mm long; bract blades tinged with purple . . . . . . . . . . . . . . . . . . 3a. M. fistulosa ssp. fistulosa var. rubra Gray 1b. Corolla light purple (rarely white); middle lobe of the lower corolla lip 2–4 mm long; bract blades green or tinged with white (rarely with miveins tinged with purple) Fig. 701  Inflorescence of Monarda fistulosa.

2a. Petioles and abaxial surface of leaf blades pubescent, in part, with spreading hairs (0.5–) 1–3 mm long . . . . . . . . . . . . . . . . . . . . . . . 3b. M. fistulosa ssp. fistulosa var. fistulosa 2b. Petioles and abaxial surface of leaf blades pubescent with appressed hairs shorter than 1 mm . . . . . . . . . . . . . . . . . . . . . . . . 3c. M. fistulosa ssp. fistulosa var. mollis (L.) Benth. Variety rubra is non-native and known from CT, MA, ME, NH. Fernald (1901) over-reported this taxon because he considered Monarda media to be synonymous with this name. Variety fistulosa is native and known from CT, MA, ME, NH, VT. Variety mollis is native and known from CT, MA, ME, NH, RI, VT. 4. Monarda media Willd. E purple bee-balm. Mondarda fistulosa L. var. media (Willd.) Gray • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest fragments. 5. Monarda punctata L. ssp. punctata var. villicaulis (Pennell) Palmer & Steyermark N C spotted bee-balm. Monarda punctata L. ssp. villicaulis Pennell • CT, MA, VT. Fields, roadsides, clearings. Reports of Monarda punctata ssp. punctata var. punctata from VT (e.g., Seymour 1982) are referable to this taxon.

La m i ac e a e   6 45

Nepeta 1. Nepeta cataria L. E catnip. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, deciduous forests, clearings.

Ocimum 1. Ocimum basilicum L. E sweet basil. CT, MA, RI. Fields, roadsides, gardens.

Origanum 1a. Calyx ± actinomorphic, with 5 subequal teeth; smaller bracts of inflorescence usually tinged with red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. vulgare 1b. Calyx zygomorphic, with 1 prominent lip, the lower lip poorly developed, and with a slit on one side; smaller bracts of inflorescence usually green . . . . . . . . . . . . . . . . . . . . . . . O. majorana 1. Origanum majorana L. E sweet marjoram. Majorana hortensis Moench; M. majorana (L.) Karst. • MA. Fields, roadsides, gardens. 2. Origanum vulgare L. E wild marjoram. CT, MA, VT. Fields, roadsides, lawns, gardens.

Perilla 1. Perilla frutescens (L.) Britt. E perilla-mint.  1a. Ocimum frutescens L.; Perilla ocymoides L.; 1b. Ocimum crispum Thunb.; Perilla ocymoides L. var. crispa Benth. • CT, MA. Roadsides, waste areas, gardens. 1a. Leaf blades crenate-serrate to low-serrate, sometimes green on the adaxial surface, otherwise purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. P. frutescens var. frutescens 1b. Leaf blades prominently serrate, with long, sharp teeth, purple on both surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. Perilla frutescens var. crispa (Benth.) Deane Variety frutescens is known from CT. Variety crispa is known from CT, MA, RI.

Physostegia 1. Physostegia virginiana (L.) Benth. ssp. virginiana n obedient false dragonhead. Dracocephalum virginianum L.; Physostegia virginiana (L.) Benth. var. elongata Boivin; P. virginiana (L.) Benth. var. granulosa (Fassett) Fern.; P. virginiana (L.) Benth. var. speciosa (Sweet) Gray • CT, MA, ME, NH, RI, VT. Fields, roadsides, gardens, river shores, lake shores. Most populations of this species are introduced to New England, especially those found in fields and along roadsides. However, a form with abundant glandular hairs on the calyx and smaller leaves with fewer, shorter teeth is thought to be native. This relatively rare form is known from the Kennebec River in ME and the Connecticut River and Lake Champlain in VT. It may be of conservation concern.

Prunella 1a. At least the upper leaf blades pinnately lobed; corolla yellow-white (rarely pink or purple); lower calyx lobes narrow-lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. laciniata 1b. Leaf blades entire to crenulate; corolla light purple (rarely white); lower calyx lobes lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. vulgaris

646  tricolpate s

1. Prunella laciniata (L.) L. E cut-leaved selfheal. Prunella alba Pallas ex Bieb. • MA. Fields, roadsides, waste areas. 2. Prunella vulgaris L. n common selfheal.  2a. Prunella vulgaris L. var. elongata Benth.; P. vulgaris L. var. lanceolata (W. Bart.) Fern. • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, shorelines. 1a. Middle stem leaf blades lanceolate to narrow-oblong, usually cuneate at the base, 2–5 times as long as wide . . . . . . . . . . . . . . . . . 2a. P. vulgaris ssp. lanceolata (W. Bart.) Hultén 1b. Middle stem leaf blades ovate to ovate-oblong, usually rounded at the base, 1.5–2.5 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. P. vulgaris ssp. vulgaris Subspecies lanceolata is native and known from CT, MA, ME, NH, RI, VT. Subspecies vulgaris is non-native and known from CT, MA, ME, NH, RI, VT.

Pycnanthemum Reference: Grant and Epling (1943). 1a. Inflorescence congested but many of the branches within the clusters visible, especially in fruit [Fig. 702], the individual clusters 15–35 (–45) mm wide; calyx zygomorphic, the upper lip definitely shorter than the lower lip, the lobes tipped with a tuft of long, septate hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. incanum 1b. Inflorescence very dense, only the lowest branches, if any, visible [Fig. 703], the individual clusters 3–15 (–20) mm wide; calyx ± actinomorphic, the lobes lacking a tuft of long, septate hairs (though other hairs may be present) 2a. Principal leaf blades lanceolate to ovate, 12–30 (–40) mm wide, mostly less than 2.5 times as long as wide, rounded at the base, with a few low teeth on each margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. muticum 2b. Principal leaf blades linear to lanceolate, 2–12 (–15) mm wide, mostly more than 2.5 times as long as wide, cuneate at the base, with or without teeth 3a. Leaf blades 2–4 (–5.5) mm wide, sessile, glabrous [Fig. 703]; stem glabrous or rarely with a few upcurved hairs on the angles; calyx lobes 1–1.5 mm long . . . . . P. tenuifolium 3b. Leaf blades 3–12 (–15) mm wide, short-petioled to subsessile, pubescent on at least the abaxial surface; stem pubescent on the angles and sometimes the faces as well; calyx lobes 0.5–1 mm long (up to 1.5 mm long in P. torrei) 4a. Bracts, and often also the leaf blades, pubescent on the adaxial surface with minute, soft hairs (view at 20×); stems pubescent on the faces as well as the angles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. verticillatum 4b. Bracts and leaf blades glabrous on the adaxial surface (sometimes with minute, stiff scabrules on the adaxial bract surface); stems pubescent primarily on the angles or also pubescent on the faces in P. torrei 5a. Calyx lobes triangular, 0.5–1 mm long; pubescence of stem confined chiefly to the angles; leaf blades entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. virginianum 5b. Calyx lobes narrow-lanceolate, 1–1.5 mm long; pubescence of stem uniformly distributed; leaf blades usually with a few low teeth . . . . . . . . . . . . . . . . . . . . P. torrei 1. Pycnanthemum incanum (L.) Michx. var. incanum N Fig. 702

Fig. 702  Inflorescence of Pycnanthemum incanum showing branches visible in lower portion.

hoary mountain-mint. Koellia dubia (Gray) Small; K. incana (L.) Kuntze • CT, MA, NH, RI, VT; confined to southern portion of northern New England states. Woodlands, forest openings, ridges, balds. ‌1 × 3. Pycnanthemum ×clinopodioides Torr. & Gray is a very rare mountain-mint hybrid in New England, known from CT, MA. Though usually treated as a species, research

L a m i ac e a e   6 47

by Chambers and Chambers (1971) and Kesseli and Dole (ca. 1998) showed this taxon to be of recent hybrid origin. It is identified by its congested but visible inflorescence branches and slightly zygomorphic calyx, similar to P. incanum. However, it differs from that taxon in its principal leaf blades that are narrow-lanceolate, 10–25 mm wide, usually more than 3 times as long as wide, and borne on petioles 3–6 mm long, and its bracts that are green or at most slightly whitened by pubescence (the principal leaf blades of P. incanum are oblong to ovate, 15–35 mm wide, usually less than 3 times as long as wide, and borne on petioles 5–15 mm long, and its bracts are conspicuously whitened, especially adaxially, by dense, fine pubescence) 2. Pycnanthemum muticum (Michx.) Pers. N clustered mountain-mint. Koellia mutica (Michx.) Kuntze • CT, MA, ME, NH, RI, VT; primarily confined to southern half of northern New England states. Woodlands, forest openings, fields, open rights-of-way, ridges, balds. Some collections of this species are transitional to Pycnanthemum verticillatum and can be difficult to separate. The species are close morphologically and share the character of minute pubescence on the adaxial surface of bracts and, sometimes, upper leaf blades. Collecting multiple individuals will better capture leaf blade variability (the critical character) and make identifications less difficult. 3. Pycnanthemum tenuifolium Schrad. N Fig. 703 narrow-leaved mountain-mint. Koellia flexuosa, auct. non (Walt.) MacM.; Pycnanthemum flexuosum, auct. non (Walt) B.S.P. • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic fields, roadsides, and clearings. 4. Pycnanthemum torrei Benth.

NC

Torrey’s mountain-mint. CT, NH. Woodlands, forest fragments. 5. Pycnanthemum verticillatum (Michx.) Pers. N whorled mountain-mint.  5a. Koellia pilosa (Nutt.) Britt.; Pycnanthemum pilosum Nutt.;  5b. Koellia verticillata (Michx.) Kuntze; Pycnanthemum virginianum (L.) T. Dur. & B.D. Jackson ex B.L. Robins. & Fern. var. verticillatum (Michx.) Boivin • CT, MA, RI, VT; also reported from ME by Magee and Ahles (1999), but specimens are unknown. Mesic to wet-mesic forests, fields, meadows, and roadsides. 1a. Faces of upper stem internodes conspicuously pilose with spreading hairs; pubescence of abaxial leaf blade conspicuous and uniform across the surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5a. P. verticillatum var. pilosum (Nutt.) Cooperrider 1b. Faces of upper stem internodes moderately to sparsely pubescent with short, pilose hairs; pubescence of abaxial leaf blade chiefly or entirely confined to the veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5b. P. verticillatum var. verticillatum Variety pilosum is known from CT, MA. Variety verticillatum is known from CT, MA, RI, VT; also reported from ME by Magee and Ahles (1999), but specimens are unknown. 6. Pycnanthemum virginianum (L.) T. Dur. & B.D. Jackson ex B.L. Robins. & Fern. N Virginia mountain-mint. Koellia virginiana (L.) MacM. • CT, MA, ME, NH, RI, VT. Fields, banks, roadsides, clearings.

Salvia Salvia farinacea Benth. was reported from CT by Magee and Ahles (1999), but specimens are unknown. 1a. Upper lip of calyx ± entire 2a. Corolla 5–7 mm long, only shortly exceeding the calyx; leaf blades broad-ovate to triangular-ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tiliifolia 2b. Corolla 12–20 mm long, greatly exceeding the calyx; leaf blades linear to oblong or lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. azurea

Fig. 703  Inflorescences and leaves of Pycnanthemum tenuifolium.

648   tricolpate s

1b. Upper lip of calyx with 3 lobes or teeth 3a. Upper lip of calyx provided with 3 minute and closely approximate teeth, the distance between the lateral teeth shorter than 1 mm 4a. Leaves basally disposed; stem with 1–3 leaf-bearing nodes, pubescent with glandular hairs in the apical portion; corolla (15–) 20–30 mm long . . . . . . . . . S. pratensis 4b. Leaves chiefly cauline; stem with 3 or more leaf-bearing nodes, pubescent with eglandular hairs; corolla 8–12 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nemorosa 3b. Upper lip of calyx provided with 3 conspicuous lobes that are separated by rectangular or U-shaped sinuses, the distance between the lateral lobes longer than 1 mm 5a. Corolla red (rarely purple or white), 40–42 mm long; calyx red; verticillasters with 2–6 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. splendens 5b. Corolla white or blue to purple, sometimes marked with yellow, 8–30 (–35) mm long; calyx green and tinged to varying degrees with white, red, or purple, sometimes the green background color difficult to detect; verticillasters with 4–30 flowers 6a. Verticillasters with 12–30 flowers; corolla 8–15 mm long; upper calyx lip up to ⅓ as long as the basal, connate portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. verticillata 6b. Veticillasters with 4–12 flowers; corolla 20–30 mm long; upper calyx lip more than ½ as long as the basal, connate portion 7a. Stems and leaf blades densely and minutely white-pubescent; leaf blades 1–8 cm long, borne on petioles 0–3 cm long . . . . . . . . . . . . . . . . . . . . . . S. officinalis 7b. Stems and leaf blades with sparser and coarser pubescence; lower leaf blades 7–20 cm long, borne on petioles (2–) 3–10 cm long 8a. Leaves chiefly cauline, the lower toothed; upper lip of corolla longer than the the basal, connate portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sclarea 8b. Leaves basally disposed, the lower pinnately lobed (sometimes merely toothed); upper lip of corolla shorter than the basal, connate portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lyrata 1. Salvia azurea Michx. ex Lam. var. grandiflora Benth. E azure-blue sage. Salvia azurea Michx. ex Lam. ssp. pitcheri (Torr. ex Benth.) Epling; S. pitcheri Torr. ex Benth. • CT. Fields, roadsides, waste areas. 2. Salvia lyrata L. lyre-leaved sage. CT; southern portion of state. Meadows.

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3. Salvia nemorosa L. E woodland sage. MA. Fields, waste areas. 4. Salvia officinalis L. E West Indian sage. CT, ME, RI, VT. Fields, roadsides, waste areas, gardens. 5. Salvia pratensis L. E meadow sage. Salvia tenorii Spreng. • CT, MA. Fields, roadsides, waste areas, gardens. 6. Salvia sclarea L. E cleareye sage. CT, MA. Fields, roadsides, waste areas, dumps. 7. Salvia splendens Sellow ex Roemer & J.A. Schultes E scarlet sage. CT, MA. Fields, roadsides, waste areas. 8. Salvia tiliifolia Vahl. E linden-leaved sage. MA. Dumps, waste areas. 9. Salvia verticillata L. E whorled sage. ct, mA, VT. Fields, roadsides, waste areas, yards.

L a m i ac e a e   6 49

Satureja 1. Satureja hortensis L. E summer savory. CT, MA, ME, VT. Waste areas, gardens.

Scutellaria Reference: Epling (1942). 1a. Bracts near the middle of the inflorescence scarcely differentiated from the upper foliage leaves (i.e., the flowers occurring in the axils of leaves) [Fig. 704]; principal leaf blades sessile or borne on short petioles up to 4 mm long 2a. Corolla 12–20 mm long; leaf blades 20–60 mm long, lanceolate to ovate-oblong, commonly 2–4 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. galericulata 2b. Corolla 7–9 mm long; leaf blades 10–16 mm long, narrow-ovate or triangular to suborbicular, usually less than 2 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . S. parvula 1b. Bracts near the middle of the inflorescence conspicuously reduced in size compared with the upper foliage leaves (i.e., the flowers in leafy-bracteate racemes); principal leaf blades on petioles 5–25 (–33) mm long 3a. Racemes not secund; median leaf blades lanceolate or oblanceolate, 4–15 mm wide, entire or low-crenate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. integrifolia 3b. Racemes secund; median leaf blades narrow-ovate to ovate, 15–50 (–80) mm wide, dentate or serrate 4a. Corolla 5–8 mm long, ± straight and not curved upward out of the calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lateriflora 4b. Corolla 12–16 (–18) mm long, evidently curved upward out of the calyx . . . . S. altissima 1. Scutellaria altissima L. E tall skullcap. MA. Gardens, areas of cultivation. 2. Scutellaria galericulata L. N Fig. 704 hooded skullcap. Scutellaria epilobiifolia A. Hamilton; S. galericulata L. var. epilobiifolia (A. Hamilton) Jordal; S. galericulata L. ssp. pubescens (Benth.) A. & D. Löve; S. galericulata L. var. pubescens Benth. • CT, MA, ME, NH, RI, VT; nearly throughout. Swamps, shorelines, hydric ditches, marshes. American plants have been distinguished as separate from Eurasian ones using the specific epithet epilobiifolia or infraspecific epithet pubescens (e.g., Fernald 1950b). Though mean differences do occur in corolla length and mericarp surface pattern, there is a significant morphological overlap and no clear distinctions can be drawn (Epling 1942). ‌2 × 4. Scutellaria ×churchilliana Fern. is an infrequent skullcap hybrid (in northern New England) that is known from ME, VT. It is intermediate between its two parent species. From S. lateriflora it can be separated by the corolla 8–12 mm long that curves upward out of the calyx (vs. corolla 5–8 mm long and nearly straight). From S. galericulata it can be separated by its petioles 3–10 mm long and smaller corollas (vs. petioles up to 4 mm long and corolla 12–20 mm long). The inflorescence of S. × ‌churchilliana is variable, sometimes appearing axillary (as in S. galericulata) and other times appearing racemose (as in S. lateriflora). 3. Scutellaria integrifolia L.

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entire-leaved skullcap. Scutellaria incana Biehler ssp. hispida (Benth.) Epling; S. integrifolia L. var. hispida Benth. • CT, MA; mainly along the coastal plain, but also extending up the Connecticut River Valley in CT. Mesic to hydric fields and pastures. 4. Scutellaria lateriflora L. E mad dog skullcap. CT, MA, ME, NH, RI, VT. Swamps, shorelines, marshes, low fields.

Fig. 704  Inflorescence of Scutellaria galericulata.

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5. Scutellaria parvula Michx.

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little skullcap.  5a. Scutellaria ambigua Nutt.; S. leonardii Epling; S. nervosa Pursh var. ambigua (Nutt.) Fern.; S. parvula Michx. var. leonardii (Epling) Fern.; 5b. Scutellaria australis (Fassett) Epling • CT, Ma, Me, VT. Woodlands, ledges, balds, river banks, meadows, fields, disturbed soil, often associated with thin soils over bedrock. 1a. Stem pubescent with minute, curved-ascending, eglandular hairs; calyx pubescent with eglandular hairs; principal leaf blades with (1–) 2 (–3) pairs of lateral veins . . . . . . . . . . . . . . . . . . . . . . . . . 5a. S. parvula var. missouriensis (Torr.) Goodman & C.A. Lawson 1b. Stem pubescent with minute, retrorse (or curved-ascending in var. australis) eglandular hairs and longer, spreading, glandular hairs; calyx pubescent, in part, with glandular hairs; principal leaf blades with (2–) 3–5 pairs of lateral veins 2a. Lateral veins of leaf blades arched and anastomosing to form a continuous, submarginal vein; eglandular hairs of the stem curved-ascending; leaf blades abaxially with glandular hairs only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5b. S. parvula var. australis Fassett 2b. Lateral veins of the leaf blades not anastomosing or inconspicuously so; eglandular hairs of stem retrorsely oriented; leaf blades abaxially with both glandular and eglandular hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5c. S. parvula var. parvula Variety missouriensis is known from CT, Ma, Me, VT. Variety australis is known from CT. Variety parvula is known from VT. Previous reports of var. parvula from York County, ME, were based on specimens of S. parvula var. missouriensis that were only partially identified (i.e., the specimens were identified to the species level only and they were taken to be the typical variety without review of the sheets). Epling (1942) also cited a specimen of var. parvula from ME with simply the word “Fulton”, which was meant to be a place name. However, I am unaware of any town or village in the state by that name. Further, Epling did not map this species as being from ME. All varieties are of conservation concern.

Sideritis 1. Sideritis montana L. E mountain ironwort. CT. Fields, roadsides, waste areas.

Stachys Stachys aspera Michx. was reported from ME (e.g., Fernald 1950b) based on transitional specimens of S. hyssopifolia (annotated by Mulligan and Munro; see reference). Stachys officinalis (L.) Trev. was reported from MA (as S. betonica Benth.) based on a single plant (Knowlton and Deane 1923a). This is not taken here as evidence of naturalization. A similar stance was taken by Mulligan and Munro (1989). References: Epling (1934), Mulligan and Munro (1989). 1a. Stems, leaf blades (at least abaxially), and calyces densely and softly pilose-tomentose to lanate-tomentose with white to white-gray hairs 2a. Leaf blades cuneate at the base, the marginal teeth obscured by dense pubescence, the upper surface white to gray-white with dense tomentum; calyx lobes largely concealed by pubescence of inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. byzantina 2b. Leaf blades truncate to cordate at the base, the marginal teeth evident, the upper surface gray-green with merely moderately dense tomentum; calyx lobes projecting beyond pubescence of inflorescence and more evident . . . . . . . . . . . . . . . . . . . S. germanica 1b. Stems, leaf blades, and calyces glabrous to pubescent, the pubescence not both white and woolly, the hairs generally not obscuring the surfaces 3a. Plants annual, from taproots 4a. Leaf blades ovate to broad-ovate, truncate to cordate at the base; corolla 6–9 mm long, pink to purple or white; calyx lobes approximately equaling the length of the basal, connate portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. arvensis

L a m i ac e a e   6 5 1

4b. Leaf blades lanceolate or ovate to oblong or elliptic, cuneate at the bae; corolla 10–16 mm long, pale yellow to white; calyx lobes approximately ½ as long as the basal, connate portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. annua 3b. Plants perennial, from rhizomes 5a. Leaf blades linear to narrow-oblong, 3–7 (–12) mm wide, entire or partially serrate with low teeth; calyx glabrous (rarely with a few hairs on the upper side), with triangular lobes that are not prolonged into a spine-like point . . . . . . . . . . . . . . . . . . . S. hyssopifolia 5b. Leaf blades narrow-oblong or lanceolate to triangular or narrow-ovate, 10–40 mm wide, crenate to sharply serrate; calyx pubescent, with lanceolate lobes that are prolonged into a spine-like point 6a. Petioles (5–) 10–30 (–45) mm long, some at least 20% as long as the associated leaf blade; calyx lobes ca. ½ as long as the basal, connate portion . . . . S. clingmanii 6b. Petioles 1–10 mm long, all shorter than 20% as long as the associated leaf blade; calyx lobes ⅔ to fully as long as the basal, connate portion 7a. Faces of stem glabrous; calyx pubescent with eglandular hairs . . . . . S. hispida 7b. Faces of stem pubescent; calyx pubescent, in part, with glandular hairs 8a. Calyx pubescent with eglandular hairs rarely longer than 1 mm that are ± of similar length as the glandular hairs; calyx lobes abruptly tapering to a slender tip; corolla usually purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. palustris 8b. Calyx pubescent with eglandular hairs mostly 1.5–3 mm long that are conspicuously longer than the glandular hairs; calyx lobes ± evenly tapering to a slender tip [Fig. 705]; corolla usually pink to light purple . . . . . . . . . . . S. pilosa 1. Stachys annua L. E annual hedge-nettle. MA, ME, RI. Fields, roadsides, waste areas. 2. Stachys arvensis L. E field hedge-nettle. MA, ME. Fields, waste areas. 3. Stachys byzantina K. Koch E woolly hedge-nettle. Stachys lanata Jacq.; S. olympica Poir. • CT, RI, VT. Fields, roadsides, waste areas, gardens. 4. Stachys clingmanii Small E Clingman’s hedge-nettle. VT. Fields, roadsides, clearings. 5. Stachys germanica L. E German hedge-nettle. RI. Fields, roadsides, waste areas, gardens. 6. Stachys hispida Pursh N hispid hedge-nettle. Stachys palustris L. var. hispida (Pursh) Boivin; S. tenuifolia Willd. var. hispida (Pursh) Fern.; S. tenuifolia Willd. var. platyphylla Fern. • CT, MA, ME, NH, RI, VT. Mesic to wet-mesic soil of swamps, shorelines, riparian forests, and roadsides. 7. Stachys hyssopifolia Michx. n hyssop-leaved hedge-nettle. Stachys atlantica Britt.; S. hyssopifolia Michx. var. lythroides (Small) J. Nelson; S. lythroides Small • CT, MA, ME, NH, RI. Coastal plain pond shores, roadsides. This species is introduced in ME. 8. Stachys palustris L. E marsh hedge-nettle. Stachys palustris L. var. elliptica Clos; S. palustris L. var. petiolata Clos; S. palustris L. var. segetum (Mutel) Grogn. • CT, MA, ME, NH, RI, VT. Shorelines, fields, roadsides, waste areas. 9. Stachys pilosa L. N C Fig. 705 hairy hedge-nettle.  9a. Stachys borealis Rydb.; S. homotricha (Fern.) Rydb.; S. palustris L. var. homotricha Fern.; S. palustris L. var. nipigonensis Jennings; S. palustris L. var. phaneropoda

Fig. 705  Flowers of Stachys pilosa var. pilosa showing long eglandular hairs on calyx.

65 2   tricolpate s

Weatherby; S. palustris L. ssp. pilosa (Nutt.) Epling; S. palustris L. var. pilosa (Nutt.) Fern.; S. palustris L. var. puberula Jennings; 9b. Stachys arenicola Britt.; S. brevidens Rydb.; S. palustris L. ssp. arenicola (Britt.) Gill • CT, MA, ME, NH, RI, VT. Fields, meadows, roadsides, shorelines, wetland borders, forest edges. 1a. Hairs on the stem angles soft, not pustulose-based, predominantly spreading, not much thicker than the hairs of the stem faces (though often longer) . . . . . . 9a. S. pilosa var. pilosa 1b. Hairs of the stem angles stiff, often pustulose-based, reflexed, noticeably thicker than the hairs of the stem faces . . . . . . . . . . . 9b. S. pilosa var. arenicola (Britt.) G. Mulligan & D. Munro Variety pilosa is known from CT, MA, ME, NH, RI, VT. Variety arenicola is known from RI; also reported from CT, MA, ME NH, and VT by Kartesz (1999), but specimens are unknown. It is of conservation concern.

Teucrium 1a. Leaf blades once or twice pinnately lobed into ± oblong segments; pedicels 4–8 mm long; calyx conspicuously saccate at the base; plants annual, 10–30 cm tall . . . . . . . . . . . . . . T. botrys 1b. Leaf blades merely toothed; pedicels 1–3 mm long; calyx scarcely saccate at the base; plants rhizomatous perennials, (15–) 30–100 cm tall 2a. Corolla green-yellow, 9–10 mm long; verticillasters with 1 or 2 flowers, collectively forming what appears to be a lax, secund raceme; leaf blades truncate to cordate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. scorodonia 2b. Corolla pink-purple, 11–18 mm long; verticillasters with 3 or more flowers, collectively forming a more crowded, terete raceme; leaf blades cuneate to rounded at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. canadense 1. Teucrium botrys L. E cut-leaved germander. CT, MA. Fields, waste areas. 2. Teucrium canadense L. N Fig. 706 American germander.  2a. Teucrium canadense L. var. angustatum Gray; T. canadense L. var. littorale (Bickn.) Fern.; T. canadense L. var. virginicum (L.) Eat.; T. littorale Bickn.; 2b. Teucrium boreale Bickn.; T. canadense L. var. boreale (Bickn.) Shinners; T. canadense L. ssp. occidentale (Gray) W.A. Weber; T. occidentale Gray; T. occidentale Gray var. boreale (Bickn.) Fern. • CT, MA, ME, NH, RI, VT. Coastal beaches and tidal marshes, river shores, lake shores, wetland margins. Fig. 706  Flowers of Teucrium canadense.

1a. Pubescence of the plant entirely eglandular; leaf blades abaxially pubescent with curled or bent hairs; upper 3 calyx lobes acute at the apex (the middle lobe sometimes acuminate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a. T. canadense var. canadense 1b. Much of the pubescence of the plant glandular, especially in the inflorescence and on surface of the calyx; leaf blades abaxially pubescent with ± straight hairs; upper 3 calyx lobes obtuse at the apex (the middle lobe sometimes acute) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. T. canadense var. occidentale (Gray) McClintock & Epling Variety canadense is known from CT, MA, ME, NH, RI, VT. Variety occidentale is known from CT, MA, ME, NH, VT. 3. Teucrium scorodonia L. E woodland germander. MA. Fields, roadsides, waste areas.

Thymus Thymus praecox Opiz ssp. arcticus (Dur.) Jalas was reported from MA, NH, and RI by Kartesz (1999), but specimens are unknown. Thymus serpyllum L. has long been reported from various states in New England; however, these reports were based on collections of T. pulegioides (Seymour 1982). Reference: Jalas (1972).

La m i ac e a e   6 5 3

1a. Leaf blades revolute, not ciliate; upper calyx lobes usually eciliate; corolla usually white to pale purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. vulgaris 1b. Leaf blades plane or barely revolute, usually ciliate at least at the base; upper calyx lobes usually ciliate; corolla usually pink-purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. pulegioides 1. Thymus pulegioides L. E lemon thyme. Thymus ovatus P. Mill.; T. serpyllum L. ssp. chamaedrys (Fries) Voll. • CT, MA, ME, NH, RI, VT. Dry fields, roadsides, waste areas, lawns, railroads. 2. Thymus vulgaris L. E garden thyme. MA. Fields, banks, waste areas.

Trichostema Trichostema is often defined as excluding Isanthus based on floral differences. However, phylogentic research showed that Isanthus is properly part of the genus Trichostema (Huang 2002). 1a. Calyx ± actinomorphic; stamens nearly straight or weakly arched, shorter than 10 mm; 2 principal lateral veins of leaf blade reconnecting to midvein . . . . . . . . . . . . . . . . T. brachiatum 1b. Calyx zygomorphic; stamens conspicuously arched [Fig. 707], 12–20 mm long; 2 principal veins of leaf blade not reconnecting to midvein or the lateral veins absent altogether 2a. Leaf blades oblong or elliptic to ovate, 5–25 mm wide, 2.5–4 times as long as wide; longer hairs of upper stem (0.3–) 0.5–2 mm long; mericarps 1.9–2.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. dichotomum 2b. Leaf blades linear, 1–5 mm wide, 5–15 times as long as wide; longer hairs of upper stem 0.1–0.3 (–0.4) mm long; mericarps 1.6–1.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . T. setaceum 1. Trichostema brachiatum L.

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pennyroyal bluecurls. Isanthus brachiatus (L.) B.S.P.; I. brachiatus (L.) B.S.P. var. linearis Fassett • CT, MA, VT. Ledges, fields, river banks, woodlands, forested hillsides. 2. Trichostema dichotomum L. N Fig. 707 forked bluecurls. Trichostema dichotomum L. var. puberulum Fern. & Grisc. • CT, MA, ME, NH, RI, VT. Sandy open areas such as fields, roadsides, banks, sand plains, and waste areas. 3. Trichostema setaceum Houtt.

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narrow-leaved bluecurls. Trichostema dichotomum L. var. lineare (Walt.) Pursh; T. lineare Walt. • CT; southwestern portion of state. Dry, sandy soil. Fig. 707  Flower of Trichostema dichotomum.

Lardizabalaceae Akebia 1. Akebia quinata (Houtt.) Dcne. E chocolate-vine. CT, MA; also reported from RI by George (1992), but specimens are unknown. Roadsides, forest fragments, open ledges, waste areas, fields. 1‌ × Akebia trifoliata Koidz. Akebia ×pentaphylla (Makino) Makino is a rare chocolatevine hybrid known from MA. It is similar to A. quinata but shows leaves with 3–5 leaflets that have prominent crenations along the margins. Akebia quinata, on the other hand, has leaves with (3–) 5 (–7) leaflets that have entire to undulate margins (rarely with 1 or 2 obscure teeth).

65 4   tricolpate s

Lentibulariaceae 1a. Leaves simple, with a viscid adaxial surface, without traps [Fig. 708]; flowers solitary [Fig. 708], without bracts; calyx with 5 lobes; plants terrestrial, of wet cliffs and gullies . . . . . . . . . Pinguicula 1b. Leaf-like branches simple to highly dissected, not viscid, bearing bladder-like traps on the divisions or on separate branches [Figs. 709, 711]; flowers solitary or borne in racemes, each subtended by a bract; calyx with 2 lobes; plants aquatic or of wetlands . . . . . . . . . . Utricularia

Pinguicula 1. Pinguicula vulgaris L. ssp. vulgaris

N C Fig. 708

violet butterwort. ME, NH, VT. Boreal and subalpine cliffs, gullies, and rock slabs, often associated with high-pH bedrock.

Utricularia Trap measurements were performed on dried specimens. References: Crow and Hellquist (1985), Taylor (1989). Fig. 708  Habit of Pinguicula vulgaris.

1a. Peduncle subtended by a whorl of conspicuously inflated branches that serve as floats to hold the peduncle above the surface of the water [Fig. 710]; leaf-like branches repeatedly dichotomous, forking 6 or more times into two equal divisions and lacking a main axis 2a. Raceme with 1–5 (–7) flowers [Fig. 710]; peduncle above the inflated branches 3–10 cm long; inflated branches 10–40 mm long, with nearly parallel margins for much of the proximal half, abruptly tapering to the peducle, bearing dissected branches only near the tip [Fig. 710]; bracts 2–3 mm long, sometimes ± 3-lobed at the apex; sepals 2.5–3.5 mm long; spur entire at tip (very rarely bifid); leaf-like branches divided at the base into 2 ± equal branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. radiata 2b. Raceme with (4–) 9–14 (–17) flowers; peduncle above the inflated branches 10–30 cm long; inflated branches 30–80 mm long, gradually tapering to the peduncle from a point distal to the middle, bearing dissected branches in the distal half; bracts 3–5 (–7) mm long, entire at the apex; sepals 3–5 mm long; spur bifid at apex; leaf-like branches divided at the base into 2 unequal branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. inflata 1b. Peduncle not subtended by a whorl of inflated branches; leaf-like branches simple, branched fewer than 6 times, or repeatedly branched but then with a flexuous main axis with branches emerging from the sides 3a. Leaf-like branches whorled; traps located at the tips of branches; corolla red-purple with a yellow spot on the lower lip (rarely white) . . . . . . . . . . . . . . . . . . . . . . . . . . . U. purpurea 3b. Leaf-like branches alternate (note: the branches are divided at the very base in some species and appear opposite) [Fig. 711]; traps scattered along the branches [Figs. 709, 711]; chasmagamous (i.e., petaliferous) corolla yellow or purple with cream to yellow spot on the lower lip in U. resupinata 4a. Leaf-like branches mostly simple; traps 0.2–1.1 mm long; plants with stems anchored in the substrate 5a. Corolla purple with a cream to yellow spot on the lower lip; peduncle with a single flower subtended by a tubular bract; leaf-like branches with 1 or more transverse cross-septa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. resupinata 5b. Corolla yellow (white to red-purple in cleistogamous flowers of U. subulata); peduncle with 1–10 flowers, each flower subtended by non-tubular bracts; leaf-like branches without cross-septa

L e nt i bu l a r i ac e a e   6 5 5

6a. Inflorescence with yellow, chasmogamous flowers, these with a downward directed spur 7–14 mm long and a lower corolla lip 9–16 mm long; pedicels 1–2 mm long; each flower subtended by a basifixed bract and a pair of bractlets; capsules ovoid, 3.8–5 mm long; intact leaf-like branches with a mass of brown cells at the apex, often forming a callus tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. cornuta 6b. Inflorescence either with white to red-purple cleistogamous flowers or with yellow chasmogamous flowers, the latter with a spur 2.4–5 mm long that is appressed to the lower corolla lip 4–7 mm long; pedicels 4–15 mm long; each flower subtended by a peltate bract; capsules globose, 1–1.5 mm long; intact leaflike branches without an apical mass of brown cells . . . . . . . . . . . . . . . . U. subulata 4b. Leaf-like branches divided 1–8 or more times [Figs. 709, 711]; traps (0.5–) 1–5 mm long (U. gibba frequently with many traps less than 1 mm long); plants free-floating or partially anchored in substrate (sometimes stranded during low water periods) 7a. Ultimate segments of leaf-like branches flat, scarcely narrower than the primary branches; branches dimorphic—chlorophyllous ones above the substrate with few or no traps and achlorophyllous ones buried in the substrate with abundant traps [Fig. 709] 8a. Corolla 10–16 mm long, the lower lip wider than long and with a welldeveloped palate; ultimate segments of chlorophyllous leaf-like branches with an apical seta and numerous lateral setae; chlorophyllous branches without traps [Fig. 709]; traps 1.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. intermedia 8b. Corolla 6–8 mm long, the lower lip longer than wide and with a poorly developed palate; ultimate segments of chlorophyllous leaf-like branches with only an apical seta, if any at all; chlorophyllous branches with traps; traps (0.6–) 0.9–1.8 (–2) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. minor 7b. Ultimate segments of leaf-like branches capillary, much finer than the primary branches; branches monomorphic (i.e., all chlorophyllous and trap-bearing) or dimorphic in U. striata 9a. Flowers dimorphic—petaliferous, chasmogamous flowers in an erect, emersed, 2- to 8-flowered raceme and nearly apetalous, cleistogamous flowers that are borne singly at the tips of solitary or paired, submersed peduncles 5–20 mm long; lower lip of chasmogamous corolla with 3 short, but evident, rounded lobes; peduncle without scales below the lowest flowers; seeds circular in outline, unwinged; turions 2–5 (–6) mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . U. geminiscapa 9b. Flowers monomorphic, all petaliferous and borne in erect, emersed racemes with (1–) 2–14 flowers; lower lip of corolla ± entire or retuse, but without 3 lobes; peduncles with 1–3 (–4) scales below the lowest flowers (usually lacking scales in U. gibba); turions either 5.5–13.5 mm thick or wanting and then less than 1 mm thick; seeds either with a pronounced, encircling wing or prismatically angled in outline 10a. Leaf-like branches 15–60 mm long, divided 6 or more times [Fig. 711]; traps 1.5–5 mm long; corolla without stipitate glands; raceme with (3–) 6–14 flowers; fruits infrequently produced, a pyxis; seeds prismatically 4- to 6-angled, surrounded by a very narrow wing; turions coarse, 5.5–13.5 mm thick [Fig. 711], the component branches ciliate with stout, gray hairs . . . . . . . . . . . . . U. vulgaris 10a. Leaf-like branches 5–15 mm long, few-branched, divided 1–4 times; traps 0.5–2.5 mm long; corolla with scattered, minute stipitate-glands (at least on the spur); raceme with (1–) 2–6 flowers; fruit usually produced, a capsule dehiscing by 2 lateral valves; seeds irregularly circular or elliptic in outline, surrounded by a conspicuous wing; turions wanting, less than 1 mm thick 11a. Leaf-like branches monomorphic, all chlorophyllous and trap-bearing; corolla 6–12 mm long (up to 25 mm long in some coastal plain populations from MA south); upper surface of seeds ± smooth or with some bluntly conical processes that are confined to the central area of the seed . . . . . . . . . . U. gibba

65 6   tricolpate s

11b. Leaf-like branches dimorphic—chlorophyllous ones above the substrate divided 3 or 4 times with few traps and achlorophyllous buried in the substrate divided 1–4 times with abundant traps; corolla 15–25 mm long; upper surface of seeds covered with numerous, bluntly conical processes that extend even onto the wings of the seed . . . . . . . . . . . . . . . . . . . . U. striata 1. Utricularia cornuta Michx. N horned bladderwort. Stomoisia cornuta (Michx.) Raf. • CT, MA, ME, NH, RI, VT; nearly throughout. Peatlands, peaty and mucky shores, boggy pools and seeps. 2. Utricularia geminiscapa Benj. N mixed bladderwort. Utricularia clandestina Nutt. ex Gray • CT, MA, ME, NH, RI, VT. Shallow water of lakes, ponds, and peatland pools. This species can be confused with smaller plants of Utricularia macrorhiza. It has ultimate branch segments with 0–2 (–4) latera setae and its smaller turions lack hairs (vs. (0–) 2–4 (–8) lateral setae on ultimate segments and larger turions with abundant hairs in U. macrorhiza). 3. Utricularia gibba L. N creeping bladderwort. Trilobulina fibrosa (Walt.) Raf.; Utricularia biflora Lam.; U. fibrosa Walt.; U. obtusa Sw.; U. pumila Walt. • CT, MA, ME, NH, RI, VT. Shallow water of lakes and pools, often becoming stranded as water level recedes. Forms of this species with larger corollas and longer spurs have been called Utricularia biflora. Though this form appears to be a distinctive, coastal plain element here in the northeast, there are intermediate forms known from northern and western states (i.e., there is no morphological gap between small-flowered and large-flowered forms; Taylor 1989). 4. Utricularia inflata Walt. E swollen bladderwort. Plectoma inflata (Walt.) Raf.; Utricularia ceratophylla Michx. • MA. Shallow water of ponds. 5. Utricularia intermedia Hayne N Fig. 709 Fig. 709  Dimorphic branches of Utricularia intermedia.

flat-leaved bladderwort. Lentibularia intermedia (Hayne) Nieuwl. & Lunell • CT, MA, ME, NH, RI, VT. Shallow water of lakes, pools in swamps and peatlands, often becoming stranded when water level recedes. 6. Utricularia minor L. N lesser bladderwort. Lentibularia minor (L.) Raf.; Xananthes minor (L.) Raf. • CT, MA, ME, NH, RI, VT. Still or slow-moving, shallow water of lakes, ponds, streams, and sloughs. 7. Utricularia purpurea Walt. N eastern purple bladderwort. Vesiculina purpurea (Walt.) Raf. • CT, MA, ME, NH, RI, VT; limited in VT to the northeastern portion of the state. Shallow water of lakes and ponds. 8. Utricularia radiata Small N Fig. 710 floating bladderwort. Utricularia inflata Walt. var. minor Chapman; U. inflata Walt. var. radiata (Small) Stone • CT, MA, ME, NH, RI, VT; limited in VT to the southeastern portion of the state. Shallow water of lakes and ponds, infrequently associated with still sections of streams and inlets. 9. Utricularia resupinata B.D. Greene ex Bigelow N

Fig. 710  Inflated branches and inflorescence of Utricularia radiata.

resupinate bladderwort. Lecticula resupinata (B.D. Greene ex Bigelow) Barnh. • CT, MA, ME, NH, RI, VT; absent or rare in far northern New England. Shallow water of lakes and ponds, often associated with sandy substrate, though also occurring on mud and muck. 10. Utricularia striata Le Conte ex Torr.

NC

striped bladderwort. Trilobulina striata (Le Conte ex Torr.) Raf.; Utricularia fibrosa Britt. • CT, MA; reported from RI by Kartesz (1999), but specimens are unknown. Acid ponds, boggy pools. This species, though always considered rare, has been over-reported. Many specimens believed to be this species were large-flowered forms of Utricularia gibba (i.e., the species is even more rare than previously believed).

Le n t i bu l a r i ac e a e   6 57

11. Utricularia subulata L.

NC

slender bladderwort. Setiscapella cleistogama (Gray) Barnh.; S. subulata (L.) Barnh.; Utricularia cleistogama (Gray) Britt. • MA, RI; restricted to the coastal plain. Sandy and peaty pond shores, bogs, swamps, boggy pools. 12. Utricularia vulgaris L. ssp. macrorhiza (Le Conte) Clausen N Fig. 711 greater bladderwort. Utricularia macrorhiza Le Conte; U. vulgaris L. var. americana Gray • CT, MA, ME, NH, RI, VT. Still or slow-moving water of lakes, rivers, and streams.

Limnanthaceae

Fig. 711  Branches and turion of Utricularia vulgaris.

Floerkea 1. Floerkea proserpinacoides Willd.

NC

false mermaidweed. CT, MA, VT; western portion of states. Riparian forests, swamps, forested seeps.

Linaceae Linum See McDill et al. (2009) for rationale of including Radiola in Linum. Reference: Rogers (1963). 1a. Flowers with 4 sepals, petals, stamens, and styles; sepals with 3 apical lobes; leaf blades 2–3 mm long; plants diminutive annuals, 3–10 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. radiola 1b. Flowers with 5 sepals, petals, stamens, and styles; sepals without apical lobes, though sometimes ciliate or toothed along the margin; leaf blades 3–35 mm long; plants annual or perennial, 10–100 cm tall 2a. Petals blue, 10–23 mm long; capsules 5–10 mm wide; sepals without glands 3a. Leaf blades with 3 nerves; sepals 7–9 mm long at maturity, the inner ciliolate; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. usitatissimum 3b. Leaf blades with 1 nerve, sometimes with 2 additional, obscure nerves; sepals 5–7 mm long at maturity, eciliolate; plants perennial . . . . . . . . . . . . . . . . . . . . . . L. perenne 2b. Petals white or yellow, 3–10 mm long; capsules 2–2.5 mm wide; at least the inner sepals glandular-ciliate, glandular-toothed, or with sessile marginal glands (sometimes entire in L. striatum) 4a. Petals white; lower pedicels 10–25 mm long; staminodes present; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. catharticum 4b. Petals yellow; pedicels up to 8 (–10) mm long; staminodes absent; plants perennial or annual in L. sulcatum 5a. Styles connate at the base for a distance of 0.3–1.5 mm; leaf blades with a pair of dark glands at the base (i.e., stipular glands); plants annual . . . . . . . . . . . . L. sulcatum

65 8   tricolpate s

5b. Styles distinct; leaf blades lacking a pair of basal glands; plants perennial 6a. Fruit obturbinoid, as tall as or taller than wide, pointed at the apex [Fig. 712], 2–3 mm tall; false septa of fruit incomplete (though the fruit still separating into 10 mericarps at maturity) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. intercursum 6b. Fruit depressed globose, shorter than wide, rounded to bluntly pointed at the apex [Fig. 713], 1.3–2 mm tall; false septa of fruit nearly complete 7a. Inner sepals evidently glandular-toothed [Fig. 713]; leaf blades linear to narrow-lanceolate or lanceolate-elliptic, the larger ones (1–) 1.5–3.5 (–4) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. medium 7b. Inner sepals entire or with sessile, marginal glands; leaf blades elliptic to oblanceolate or obovate, the larger ones 3–10 mm wide 8a. Branches striate-angled; inflorescence elongate, panicle-like; true septa of fruit, as well as the false septa, eciliate . . . . . . . . . . . . . . . . . . . . . . L. striatum 8b. Branches ± terete; inflorescence broad, corymb-like; true septa of fruit usually sparsely ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. virginianum 1. Linum catharticum L. E white flax. Cathartolinum catharticum (L.) Small • MA, ME, NH, VT. Fields, roadsides, ditches, seepy banks. Fig. 712  Fruit of Linum intercursum showing pointed apex.

2. Linum intercursum Bickn. N Fig. 712 sandplain yellow flax. Cathartolinum intercursum (Bickn.) Small • CT, MA, RI. Open, sandy areas such as beaches, sand plains, fields, and barrens. 3. Linum medium (Planch.) Britt. ssp. texanum (Planch.) A. Haines

N C Fig. 713

common yellow flax. Cathartolinum curtissii (Small) Small; C. medium (Planch.) Small var. texense (Planch.) Moldenke; Linum curtissii Small; L. medium (Planch.) Britt. var. texanum (Planch.) Fern.; L. striatum Walt. var. texanum (Planch.) Boivin; L. virginianum L. var. texanum Planch. • CT, MA, ME, RI, VT. Dry-mesic to, less commonly, wet-mesic soils of fields, roadsides, clearings, woodlands, railroads, and disturbed ground. 4. Linum perenne L. E blue flax. ME. Sandy soil. 5. Linum radiola L. E all-seed flax. Millegrana radiola (L.) Druce; Radiola linoides Roth • ME. Fields, pastures. Fig. 713  Fruit of Linum medium showing blunt apex.

6. Linum striatum Walt. N ridged yellow flax. Cathartolinum striatum (Walt.) Small; Linum striatum Walt. var. multijugum Fern.; Nezera striata (Walt.) Nieuwl. • CT, MA, RI. Low meadows, bogs, pond shores, wet-mesic clearings. Coastal plain and island populations of this species in New England tend to show a higher proportion of leaves and inflorescence branches with opposite arrangement. 7. Linum sulcatum Riddell var. sulcatum

NC

grooved yellow flax. Cathartolinum sulcatum (Riddell) Small; Mesynium sulcatum (Riddell) A. & D. Löve • CT, MA, NH, RI, VT. Dry fields, roadsides, and clearings. 8. Linum usitatissimum L. E cultivated flax. Linum humile P. Mill.; L. usitatissimum L. var. humile (P. Mill.) Pers. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads. Collected more frequently in the late 1800s and first half of the 1900s. 9. Linum virginianum L. N woodland yellow flax. Cathartolinum virginianum (L.) Reichenb.; Nezera virginiana (L.) Nieuwl. • CT, MA, RI; also reported from ME by Kartesz (1999), but specimens are unknown. Dry fields,

roadsides, woodlands, and clearings.

Li nac e a e   6 5 9

Linderniaceae Lindernia 1. Lindernia dubia (L.) Pennell N yellow-seeded false pimpernel.  1a. Ilysanthes inequalis (Walt.) Pennell; Lindernia anagallidea (Michx.) Pennell; 1b. Gratiola dubia L.; Ilysanthes attenuata (Spreng.) Small; I. dubia (L.) Barnh.; Lindernia dubia (L.) Pennell var. inundata (Pennell) Pennell; L. dubia (L.) Pennell var. riparia (Raf.) Fern. • CT, MA, ME, NH, RI, VT. Sandy or muddy shorelines, including fresh-tidal river shores, wet ditches, accretion bars. 1a. Principal foliage leaf blades mostly 6–15 (–20) mm long, much larger than the bracts, all broadly rounded at the base; pedicels 10–25 mm long, conspicuously surpassing the bracts (except sometimes the lowest); seeds 1.5–2 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. L. dubia var. anagallidea (Michx.) Cooperrider 1b. Principal foliage leaf blades 10–30 mm long, often not much larger than the bracts, at least the lower ones narrowed to the base; pedicels 1–15 mm long, not conspicuously surpassing the subtending bracts; seeds 2–3 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. L. dubia var. dubia Variety anagallidea is known from CT, MA, ME, NH, RI, VT. Variety dubia is known from CT, MA, ME, NH, RI, VT.

Loasaceae Mentzelia 1. Mentzelia oligosperma Nutt. ex Sims. E stickleaf mentzelia. MA. Wool waste.

Lythraceae 1a. Leaves dimorphic—submersed ones that are opposite and finely dissected and floating ones that are alternate, borne in a crowded rosette, and have rhombic blades borne on inflated petioles [Fig. 717]; fruit indehiscent, caltrop-shaped, with 4 stout horns . . . . . . . . Trapa 1b. Leaves monomorphic, neither dissected nor with inflated petioles [Figs. 714, 715, 716]; fruit a capsule 2a. Corolla zygomorphic; hypanthium swollen or spurred near the base on one side; flowers with (6–) 11 or 12 (–14) stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cuphea 2b. Corolla actinomorphic; hypanthium neither asymmetrically swollen nor spurred; flowers with 4–10 stamens or commonly 12 in Lythrum salicaria 3a. Hypanthium cylindric to narrow-cupuliform, 2 or more times taller than wide, with 6 or 12 conspicuous veins; perianth (5–) 6 (–7)-merous; leaf blades sessile or subsessile (narrowed to the base in L. virgatum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lythrum

66 0   tricolpate s

3b. Hypanthium hemispheric to broad-cupuliform, ± as tall as wide, without evident veins or with 4–8 evident veins; perianth (3–) 4 or 5 (–7)-merous; leaf blades borne on short petioles or narrowly tapering to an ill-defined petiole 4a. Petals evident, 10–15 mm long, deciduous [Fig. 714]; flowers with 8 or 10 stamens; intersepalar appendages linear, much narrower and longer than the sepals; leaf blades 50–150 mm long; plants perennial, with ± woody bases surrounded by loose, spongy tissue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Decodon 4b. Petals minute, 0.5–1 mm long, caducous [Fig. 716]; flowers with (3–) 4 (–6) stamens; intersepalar appendages triangular, ± similar in size and shape to the sepals; leaf blades 10–30 (–50) mm long; plants annual . . . . . . . . . . . . . . . . . . . . . Rotala

Cuphea 1a. Corolla narrower than 25 mm; calyx 7–12 mm long in flower . . . . . . . . . . . . . C. viscosissima 1b. Corolla wider than 25 mm; calyx (10–) 14–18 mm long in flower . . . . . . . . . . C. procumbens 1. Cuphea procumbens Ortega E creeping waxweed. Parsonsia procumbens (Ortega) Heller • MA. Wet-mesic to hydric fields and pastures. 2. Cuphea viscosissima Jacq.

NC

clammy waxweed. Cuphea petiolata (L.) Koehne; Parsonsia petiolata (L.) Rusby • CT, MA, RI; also reported from NH by Gleason and Cronquist (1991). Hydric to dry-mesic open areas such as fields, pastures, roadsides, and waste areas. Reports of this species in VT (e.g., Magee and Ahles 1999) are based on a cultivated specimen—Norwich, wild garden, Loveland s.n. (VT).

Decodon 1. Decodon verticillatus (L.) Ell. N Fig. 714 swamp-loosestrife. Decodon verticillatus (L.) Ell. var. laevigatus Torr. & Gray; Lythrum verticillatum L.; Nesaea verticillata (L.) Kunth. • CT, MA, ME, NH, RI, VT. Lake and river borders, swamps, frequently growing in shallow water.

Lythrum Reference: Graham (1975). Fig. 714  Flowers and leaves of Decodon verticillatus.

1a. Inflorescence an elongate thyrse borne at the summit of the stem and branches, the primary one mostly 10–40 cm tall; flowers with 12 stamens, trimorphic; petals 7–12 mm long; leaf blades 3–10 cm long, 2 or 3 at each node 2a. Intersepalar appendages 2.5–3 mm long, usually longer than the sepals; plants generally pubescent on the stem and/or leaf blades (very rarely glabrous); leaf blades truncate to subclasping at the base (very rarely tapering) . . . . . . . . . . . . . . . . . . . L. salicaria 2b. Intersepalar appendages ca. 1 mm long, about as long as or shorter than the sepals; plants glabrous throughout; leaf blades tapering to the base . . . . . . . . . . . . . . . . L. virgatum 1b. Inflorescence composed of solitary or paired flowers in the axils of leaves; flowers with 4–6 stamens, mono- or dimorphic (with 12 stamens and trimorphic in L. junceum); petals 2–6 mm long; leaf blades 0.8–4 cm long, usually 1 at each of the middle and upper nodes (regularly opposite in L. lineare) 3a. Flowers with 12 stamens; hypanthium spotted with red near the base; petals 5–6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. junceum

Ly t h r ac e a e   6 6 1

3b. Flowers with 4–6 stamens; hypanthium without red spots; petals 2–5 mm long (up to 6 mm long in L. alatum) 4a. Leaf blades narrow-oblong to lanceolate or narrow-ovate, 4–10 mm wide, rounded to truncate or subcordate at the base; hypogynous disk present, appearing as a thickened ring around the base of the ovary; plants perennial . . . . . . . . . . . . . . . L. alatum 4b. Leaf blades linear to narrow-oblong, 1–6 mm wide, cuneate at the base [Fig. 715]; hypogynous disk absent; plants annual or perennial 5a. Vegetative leaves chiefly alternate (the lower nodes sometimes with opposite leaves), mostly longer than the associated internodes; plants annual, usually bearing flowers from near the stem base and upward; intersepalar appendages definitely longer than the calyx lobes; flowers monomorphic as to style length . . . . . . . . . . L. hyssopifolia 5b. Vegetative leaves chiefly opposite, mostly shorter than the associated internodes; plants perennial, usually without flowers at the lower nodes; intersepalar appendages ± as long as the calyx lobes; flowers dimorphic—pin flowers with long styles and thrum flowers with short styles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. lineare 1. Lythrum alatum Pursh ssp. alatum winged loosestrife. Lythrum dacotanum Nieuwl. • CT, MA, ME, NH, RI, VT. Low fields and meadows, river banks, accretion bars, marshes, open swamps.

NC Fig. 715  Flowers and leaves of Lythrum hyssopifolia.

2. Lythrum hyssopifolia L. N Fig. 715 hyssop-leaved loosestrife. Lythrum adsurgens Greene • MA, ME, NH, RI; mainly along the coastal plain. Borders of saline marshes, back dune hollows and other wet depressions near the Atlantic coast. 3. Lythrum junceum Banks & Sol. E rose loosestrife. Lythrum flexuosum, auct. non Lag.; L. graefferi Ten. • MA. River banks. 4. Lythrum lineare L. NC saltmarsh loosestrife. CT; southwestern portion of state. Borders of saline and brackish marshes. 5. Lythrum salicaria L. E purple loosestrife. Lythrum salicaria L. var. gracilior Turcz.; L. salicaria L. var. tomentosum (P. Mill.) DC. • CT, MA, ME, NH, RI, VT. Swamps, marshes, shorelines, wet ditches, low fields. 6. Lythrum virgatum L. E European wand loosestrife. MA. Marshes, waste areas. The report of this species from NH by Fernald (1950b) is based on a specimen of Lythrum salicaria with nearly glabrous herbage and tapering leaf blade bases (the “gracilior” form)—2 Sep 1926, Batchelder s.n. (NEBC!).

Fig. 716  Fruits and leaves of Rotala ramosior.

Rotala 1. Rotala ramosior (L.) Koehne N C Fig. 716 toothcup. Ammannia ramosior L.; Rotala ramosior (L.) Koehne var. interior Fern. & Grisc.; Rotala ramosior (L.) Koehne var. typica Fern. & Grisc. • CT, MA, NH, RI; mainly along the coastal plain. Pond shores, on mud, sand, and, rarely, ledge substrate, frequently in areas with drawdown in late season.

Trapa 1. Trapa natans L. E Fig. 717 water-chestnut. CT, MA, NH, VT. Lakes, rivers.

Fig. 717  Floating leaves of Trapa natans.

662  tricolpate s

Malvaceae 1a. Plants woody, trees up to 35 m tall; peduncle of the inflorescence adnate to a conspicuous, elongate bract [Figs. 722, 723]; stamens pentadelphous; fruit a nut-like drupe . . . . . . . . . . . . . Tilia 1b. Plants herbaceous or shrubs to 6 m tall in Hibiscus syriacus; inflorescence without an adnate bract; stamens monadelphous; fruit a capsule or schizocarp 2a. Gynoecium with 5 wholly connate carpels [Fig. 719]; fruit a capsule; filaments connate, forming a tube, bearing anthers along the sides of the tube [Fig. 719], crowned with 5 teeth at the summit of the filament tube 3a. Epicalyx with 3, ovate, laciniately toothed bractlets; styles short, the stigmas almost sessile on the ovary; petals white; seeds covered with long, white hairs . . . . Gossypium 3b. Epicalyx with (6–) 8–15 linear, entire bractlets; style branches elongate, clearly elevating the stigmas above the ovary; petals yellow, pink, red-pink, blue, light purple, or white (when white usually with a red or purple base); seeds without long hairs 4a. Calyx connate for most of its length, splitting on one side and early deciduous; mature fruit 6–25 cm long, more than 2 times as long as wide . . . . . . . . Abelmoschus 4b. Calyx basally connate for ca. ½ its length, not splitting, persistent in fruit; mature fruit 1–3.5 cm long, less than 2 times as long as wide . . . . . . . . . . . . . . . . . . . . . Hibiscus 2b. Gynoecium with 5–40 basally connate carpels [Figs. 718, 721]; fruit a schizocarp, the carpels separating at maturity into indehiscent or dehiscent mericarps; filaments connate, forming a tube, bearing anthers primarily near the apex (along the sides in Lavatera) [Fig. 720], without teeth at summit of filament tube 5a. Epicalyx absent 6a. Gynoecium with (5–) 10–15 (–30) carpels, mature carpels each with 3–9 seeds and terminated by a slender, solitary beak 3–6 mm long; petals 7–13 mm long, yellow; leaf blades entire to crenate, with or without low, obscure lobes 7a. Gynoecium with (5–) 10–15 (–30) carpels, the mature ones terminated by a slender beak 3–6 mm long and with 3–9 seeds; leaf blades (5–) 10–15 (–20) cm long, broad-ovate to orbicular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Abutilon 7b. Gynoecium with 6–8 carpels, the mature ones terminated by a slender apiculus mostly 0.5–1 mm long and with 3 seeds; leaf blades 3–7 cm long, ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudabutilon 6b. Gynoecium with 5–12 (–15) carpels, mature carpels each with 1 seed and either lacking a beak or terminated by 1 or 2 beaks up to 3 mm long; petals 3–10 mm long, white, yellow, or pink to light purple or light blue-purple; leaf blades conspicuously lobed (merely toothed in Sida spinosa) 8a. Plants dioecious (i.e., with unisexual flowers on separate plants); styles with the stigmatic surface along the inner surface; mature carpels apically beakless or minutely apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Napaea 8b. Plants synoecious (i.e., with bisexual flowers); styles with the stigmatic surface at the capitate apex; mature carpels apically terminated by 1 or 2 erect beaks 9a. Petals white or yellow; lateral walls of separating mericarps persistent . . . Sida 9b. Petals pink to light purple or light blue-purple; lateral walls of separating mericarps eventually disintegrating . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anoda

Ma lvac e a e   6 63

5b. Epicalyx of 3–9 bractlets present 10a. Styles with the stigmatic surface at the capitate apex; mature carpels 2-beaked at the apex, each with 2 seeds (1 seed in Malvastrum) 11a. Petals yellow; leaf blades coarsely toothed or with inconspicuous lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malvastrum 11b. Petals red to red-purple; leaf blades evidently lobed 12a. Petals 3–8 mm long; flowers mostly solitary or paired in the axils of leaves; terminal lobe of leaf blade only slightly exceeding the adjacent lateral lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Modiola 12b. Petals 8–13 mm long; flowers (1–) 2–12 from the axils of leaves {confirm}; terminal lobe of leaf blade elongate, more than 2 times as long as the adjacent lateral lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sphaeralcea 10b. Styles with the stigmatic surface along the inner surface; mature carpels beakless, each with a single seed 13a. Epicalyx with 3 bractlets that are wholly distinct (connate in Lavatera); gynoecium with 8–15 (–20) carpels 14a. Bractlets of epicalyx distinct; mericarps not covered by an expansion of the central axis of the fruit at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malva 14b. Bractlets of epicalyx connate at base; mericarps covered by a disk-like expansion of the central axis of the fruit at maturity (i.e., in fruit) . . . . . Lavatera 13b. Epicalyx with 6–9 bractlets that are connate at the base; gynoecium with 15–40 carpels 15a. Petals 30–50 mm long; staminal tube pentagonal in cross-section, glabrous; mericarps divided by an internal septum into an upper and lowr locule, the upper locule empty . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alcea 15b. Petals 15–20 mm long; staminal tube terete in cross-section, pubescent; mericarps unilocular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Althaea

Abelmoschus 1. Abelmoschus esculentus (L.) Moench E okra. Hibiscus esculentus L. • CT; southwestern portion of state. Fields, waste areas.

Abutilon 1a. Plants woody; leaf blades palmately 3- to 5-lobed; petals orange, with purple lines, 30–50 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pictum 1b. Plants herbaceous; leaf blades unlobed (except for the basal lobes); petals yellow to orange-yellow, 7–13 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. theophrasti 1. Abutilon pictum (Gill. ex Hook. & Arn.) Walp. E painted Indian-mallow. Abutilon striatum Dicks. ex Lindl.; Sida picta Gill. ex Hook. & Arn. • CT. Waste areas, railroads. This species, doubtfully naturalized in New England, was collected in northwestern CT. 2. Abutilon theophrasti Medik. E Fig. 718 velvetleaf Indian-mallow. CT, MA, ME, NH, RI, VT. Fields, waste areas, compost heaps, gardens, about buildings.

Fig. 718  Fruits of Abutilon theophrasti.

66 4   tricolpate s

Alcea 1. Alcea rosea L. E hollyhock. Althaea rosea (L.) Cav. • CT, MA, NH, RI, VT. Fields, waste areas, yards, dumps.

Althaea 1. Althaea officinalis L. E common marsh-mallow. CT, MA. Borders of saline and brackish marshes, dumps.

Anoda 1. Anoda cristata (L.) Schlecht. E crested anoda. Anoda cristata (L.) Schlecht. var. brachyantha (Reichenb.) Hochr.; A. cristata (L.) Schlecht. var. digitata (Gray) Hochr.; A. triangularis (Willd.) DC.; Sida cristata L. • CT, MA. Yards, waste areas, wool waste.

Gossypium 1. Gossypium hirsutum L. var. hirsutum E upland cotton. Gossypium herbaceum, auct. non L.; G. hirsutum L. var. punctatum (K. Schum. & Thonn.) Roberty; G. purpurascens Poir. • MA. Waste areas.

Hibiscus The genus Hibiscus has been determined to be paraphyletic, due, in part, to reliance on shared ancestral characters (i.e., symplesiomorphies) for defining the genus (Pfeil et al. 2002). It may be necessary to utilize many splinter genera to create a monophyletic Hibiscus. If this option is followed, it means that both H. moscheutos and H. trionum would be transferred to other genera. 1a. Plants woody; leaf blades glabrous on the abaxial surface; seeds ciliate . . . . . . H. syriacus 1b. Plants herbaceous; leaf blades stellate-pubescent on the abaxial surface; seeds eciliate 2a. Leaf blades unlobed or with sinuses extending shortly beyond the midpoint of the blade, moderately to densely stellate-pubescent abaxially; petals 40–100 mm long, white to pink or red-pink, sometimes with a red to purple base . . . . . . . . . . . . . . . . . H. moscheutos 2b. Leaf blades conspicuously 3-lobed with deep sinuses extending to near the leaf base, sparsely stellate-pubescent to ± glabrous abaxially; petals 15–40 mm long, yellow with a purple base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. trionum 1. Hibiscus moscheutos L. ssp. moscheutos N Fig. 719

Fig. 719  Flower of Hibiscus moscheutos with connate filaments bearing anthers along the side of the tube.

swamp rose-mallow. Hibiscus incanus Wendl. f.; H. moscheutos L. ssp. incanus (Wendl. f.) Ahles; H. moscheutos L. ssp. palustris (L.) Clausen; H. oculiroseus Britt.; H. palustris L. • CT, MA, NH, RI. Borders of saline and brackish marshes. Hibiscus moscheutos ssp. palustris, a taxon said to be characterized by pink corollas and three-lobed leaf blades, has been traditionally recognized apart from H. moscheutos ssp. moscheutos, said to be characterized by white corollas with a red central region and unlobed leaf blades. Flower color and leaf blade morphology has been shown to intergrade and not reliably distinguish taxa (Blanchard 1976). Therefore, H. moscheutos ssp. palustris is placed in synonymy. 2. Hibiscus syriacus L. E shrubby rose-mallow. CT; also reported from RI by George (1997), but specimens are unknown. Waste areas, disturbed places.

Ma lvac e a e   6 6 5

3. Hibiscus trionum L. E Venice rose-mallow. Trionum trionum (L.) Woot. & Standl. • CT, MA, ME, NH, RI, VT. Fields, roadsides, gardens, railroads, waste areas, wool waste.

Lavatera 1. Lavatera trimestris L. E rose-mallow. Althaea trimestris Kuntze; Stegia trimestris (L.) T. Luque & Devesa • VT; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Gardens, fields, roadsides.

Malva The genus Malva, as currently defined, is not monophyletic (Ray 1995, 1998). Specifically, some species may need to be transferred to other genera to create non-arbitrary genera. For example, the musk mallows (e.g., M. alcea and M. moschata) need to be removed from the core of Malva. This could be accomplished by transferring these species to Lavatera or by segregating them into the narrowly defined Bismalva. 1a. Lower flowers borne singly on long peduncles from the axils of leaves, the peduncles exceeding the subtending leaves; petals 20–35 mm long; plants perennial; leaf blades with deep sinuses to below the middle 2a. Stem pubescent with stellate hairs; bractlets of epicalyx ovate to obovate, densely stellate-pubescent on the abaxial surface; mature carpels keeled on the abaxial (i.e., outer) surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. alcea 2b. Stem pubescent with simple hairs; bractlets of epicalyx linear to narrow-oblanceolate, ± glabrous on the abaxial surface; mature carpels rounded on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. moschata 1b. Lower flowers borne in clusters of 2 or more on short pedicels, fascicled in the axils of leaves, the peduncle (when present) shorter than the subtending leaves; petals 6–25 (–30) mm long; plants annual or biennial; leaf blades with shallow sinuses, rarely as deep as the middle 3a. Petals red-purple, (12–) 20–25 (–30) mm long; bractlets of epicalyx oblong-ovate, usually less than 3 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. sylvestris 3b. Petals white or tinged with pink or purple, 6–13 (–14) mm long; bractlets of epicalyx linear to narrow-lanceolate, 3 or more times as long as wide 4a. Petals 6–13 (–14) mm long, ca. twice as long as the sepals; mericarps smooth or with a few slightly raised veins on the lateral surfaces, smooth or with a few raised veins on the abaxial (i.e., outer) surface; staminal tube pubescent; leaf blades with usually obscure lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. neglecta 4b. Petals 4–7 (–8) mm long, ca. as long as or slightly longer than the sepals; mericarps with conspicuous veins on the lateral surfaces that radiate outward from the attachment point, faintly to conspicuously rugose-reticulate on the abaxial surface; staminal tube glabrous or subglabrous; leaf blades with short, but evident, lobes 5a. Flowers sessile or subsessile in axillary fascicles (sometimes 1 of the flowers of each fascicle on an evident, though still short, pedicel); mericarps faintly rugosereticulate on the abaxial surface, without a wing on each margin (though each vein on the lateral suface prolonged as a small, tooth-like ridge on each margin); stems erect, 50–100 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. verticillata 5b. Flowers conspicuously pedicellate, the pedicels mostly 5–30 mm long; mericarps conspicuously rugose-reticulate on the abaxial surface [Fig. 721], with a thin, entire to toothed wing on each margin 0.1–0.7 (–1.1) mm wide; stems prostrate to erect, 20–50 cm tall

66 6   tricolpate s

6a. Sepals eciliate or short-ciliate with hairs mostly 0.1–0.5 mm long (scattered longer hairs sometimes also present), the entire calyx accrescent and becoming strongly reticulate-veiny in fruit, usually widely spreading under the fruit; fruiting pedicels mostly shorter than 10 mm; mature carpels with a denticulate wing margin 0.3–0.7 (–1.1) mm long at its widest point (note: widest point is typically ± opposite the attachment point) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. parviflora 6b. Sepals long-ciliate with hairs mostly 0.5–1.3 mm long, the entire calyx only slightly accrescent, not becoming reticulate-veiny, usually closed over the fruit; fruiting pedicels mostly longer than 10 mm; mature carpels with a very narrow, entire to undulate wing margin 0.1–0.2 (–0.3) mm wide at its widest point . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. rotundifolia 1. Malva alcea L. E vervain mallow. Bismalva alcea (L.) Medik. • CT, MA, ME, NH, RI, VT. Fields, roadsides, gardens, waste areas. 2. Malva moschata L. E Fig. 720 Fig. 720  Flower of Malva moschata with connate filaments bearing anthers mainly near the apex of the tube.

musk mallow. Bismalva moschata (L.) Medik. • CT, MA, ME, NH, RI, VT. Fields, roadsides, gardens, waste areas. 3. Malva neglecta Wallr. E common mallow. Malva rotundifolia, auct. non L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, gardens, lawns, dumps. 4. Malva parviflora L. E Fig. 721 little mallow. MA, ME. Waste areas, dumps. 5. Malva rotundifolia L. E low mallow. Malva pusilla Sm. • MA. Waste areas, gardens. 6. Malva sylvestris L. E high mallow. Malva erecta C. Presl.; M. mauritiana L.; M. sylvestris L. ssp. mauritiana (L.) Thellung; M. sylvestris L. var. mauritiana (L.) Boiss. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 7. Malva verticillata L. E

Fig. 721  Schizocarp of Malva parviflora.

whorled mallow. Malva crispa (L.) L.; M. verticillata L. var. crispa L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. The mature mericarps of Malva verticillata have lateral veins that are prolonged into small teeth along the margin of the mericarp, but there is no continuous marginal wing. The calyx is accrescent in this species (similar to M. parviflora). More work is needed to confirm the distribution of this species. Some records may actually refer to M. parviflora or M. rotundifolia.

Malvastrum 1. Malvastrum coromandelianum (L.) Garcke E three-lobed false mallow. Malva coromandeliana L. • MA. Wool waste.

Modiola 1. Modiola caroliniana (L.) G. Don E Carolina bristle-mallow. Modiola multifida Moench • MA. Waste areas, about buildings.

Napaea 1. Napaea dioica L. E glade-mallow. ME, VT. Fields, roadsides, railroads.

Ma lvac e a e   6 67

Pseudabutilon 1. Pseudabutilon stuckertii R.E. Fries E South American false velvetleaf. Pseudabutilon callimorphum (Hochr.) R.E. Fries var. friesii (Hassl.) R.E. Fries; P. pedunculatum (R.E. Fries) Krapov.; Sida callimorpha Hochr.; Wissadula callimorpha (Hochr.) Hassl. var. friesii Hassl.; W. pedunculata R.E. Fries • MA. Wool waste.

Sida The genus Sida has been shown to be polyphyletic (Aguilar et al. 2003). Specifically, S. hermaphrodita does not belong to the core of Sida and needs to be transferred to a new genus. 1a. Leaf blades ovate to orbicular, 3- to 7-lobed with deep sinuses; petals white; calyx terete at the base; gynoecium with usually 10 carpels; plants 10–20 (–40) dm tall . . . . . . S. hermaphrodita 1b. Leaf blades narrow-ovate to oblong or elliptic, unlobed; petals yellow; calyx angled at the base; gynoecium with 5 carpels; plants 2–6 (–10) dm tall . . . . . . . . . . . . . . . . . . . . . . . . S. spinosa 1. Sida hermaphrodita (L.) Rusby E Virginia fanpetals. Napaea hermaphrodita L. • MA. Fields, gardens, waste areas. 2. Sida spinosa L. E prickly fanpetals. CT, MA, ME. Roadsides, dumps, waste areas.

Sphaeralcea Sphaeralcea munroana (Dougl. ex Lindl.) Spach ex Gray was reported from MA by Fletcher (1916), but specimens are unknown. 1. Sphaeralcea mendocina (Phil.) K. Schum. E Mendoza globe-mallow. Malva mendocina Phil. • MA. Wool waste. Reports of Sphaeralcea fendleri Gray from MA are based on collections of this species (specimens at GH).

Tilia Reference: Hardin (1990). 1a. Leaf blades 7–20 cm long, sometimes pubescent abaxially, especially when young, with stellate hairs; flowers with staminodes 2a. Petioles less than 50% as long as the blade, glabrous; mature branchlets glabrous; fruit pericarp ± smooth; branches ascending to spreading . . . . . . . . . . . . . . . . . . . . . . T. americana 2b. Petioles more than 50% as long as the blade, pubescent; mature branchlets usually at least sparsely pubescent; fruit pericarp verrucose and 5-grooved; branches pendulous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. petiolaris 1b. Leaf blades 3–9 (–12) cm long, pubescent with simple hairs only; flowers without staminodes 3a. Fruit 8–10 mm long, with a woody pericarp that is conspicuously 5-ribbed; cymes pendent, with 2–5 flowers [Fig. 723]; leaf blades frequently pubescent on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. platyphyllos 3b. Fruit 4.5–6.6 mm long, with a membranaceous pericarp that is weakly, if at all, ribbed; cymes obliquely erect, with 4–15 flowers [Fig. 722]; leaf blades glabrous on the abaxial surface except for some tufts of red-brown hairs in the axils of the primary veins . . . . . . . . . T. cordata 1. Tilia americana L. n American linden.  1a. Tilia americana L. var. neglecta (Spach) Fosberg; T. glabra Vent.; T. neglecta Spach; 1b. Tilia heterophylla Vent.; T. heterophylla Vent. var. michauxii (Nutt.) Sarg.;

66 8   tricolpate s

T. michauxii Nutt.; T. monticola Sarg. • CT, MA, ME, NH, RI, VT. Mesic, deciduous forests, including rich and riparian types. Tilia neglecta is a phase of T. americana var. americana in which the abaxial surface of the leaf blades has a sparse covering of stellate hairs; however, other features (e.g., peduncle and pedicel indumentum) are as in var. americana. 1a. Leaf blades pubescent with bulbous glands, slender hairs, and sometimes scattered stellate hairs on the abaxial surface; peduncles and pedicels glabrous or sparsely puberulent in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. T. americana var. americana 1b. Leaf blades densely pubescent with stellate hairs on the abaxial surface (rarely becoming puberulent in age, but stellate hairs persisting along the major veins); peduncles and pedicels pubescent with stellate hairs in fruit . . . . . . 1b. T. americana var. heterophylla (Vent.) Loud. Variety americana is native and known from CT, MA, ME, NH, RI, VT. Variety heterophylla is nonnative and known from MA, ME. 2. Tilia cordata P. Mill. E Fig. 722 small-leaved linden. CT, MA, ME, RI. Roadsides, forest fragments and edges, areas of habitation. Fig. 722  Inflorescence of Tilia cordata.

2 ‌ × 4. Tilia ×vulgaris Hayne is a planted linden hybrid that sometimes escapes cultivation. It is known from CT, MA, ME, RI. It is recognized by its leaf blades 6–10 cm long with prominent tertiary veins (the tertiary veins of T. cordata are faint), the blades abaxially glabrous except for tufts of white hairs in the axils of veins. The pendent cymes have 5–10 flowers and the fruits are 6.7–7.9 mm long with smooth or only slightly ribbed pericarps. 3. Tilia petiolaris DC. E pendent silver linden. MA. Roadsides, forest fragments, yards. 4. Tilia platyphyllos Scop. E Fig. 723 large-leaved linden. ME. Roadsides, forest fragments, yards. Most reports of this species in New England are based on collections taken from planted trees, including those from CT, MA, and, in part, ME.

Melastomataceae Fig. 723  Inflorescence of Tilia platyphylla.

Rhexia 1a. Stem with 1 opposing pair of angles larger, darker, and convex, the other pair of angles smaller, paler, and flat to concave; petals dull light purple to white, usually glabrous; apical, tubular portion of the hypanthium as tall as or taller than the basal, globose portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. mariana 1b. Stem with all 4 angles subequal; petals red-purple, usually bristly on the abaxial surface; apical, tubular portion of the hypanthium shorter than the basal, globose portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. virginica 1. Rhexia mariana L. var. mariana

NC

Maryland meadow-beauty. Rhexia lanceolata Walt.; R. mariana L. var. exalbida Michx.; R. mariana L. var. leiosperma Fern. & Grisc. • MA. Coastal plain pond shores, often on dry, upper beach. 2. Rhexia virginica L. N Virginia meadow-beauty. Rhexia stricta Pursh; R. virginica L. var. purshii (Spreng.) C.W. James • CT, MA, ME, NH, RI, VT. Pond shores, including coastal plain types, peaty meadows.

M e n i s p e r m ac e a e   6 6 9

Menispermaceae Menispermum 1. Menispermum canadense L. N Canada moonseed. CT, MA, NH, VT; mainly western New England, but disjunct on Cape Cod. Rich forests and woodlands, riparian forests, rocky ridges.

Menyanthaceae 1a. Leaf blades compound, with 3 leaflets, emersed [Fig. 724]; petals pubescent on the adaxial surface [Fig. 724], lacking glands; inflorescence a raceme or panicle . . . . . . . . . . . Menyanthes 1b. Leaf blades simple, floating [Fig. 725]; petals glabrous on the adaxial surface [Fig. 725], bearing a yellow gland near the base of each lobe; inflorescence an umbel . . . . . . Nymphoides

Menyanthes 1. Menyanthes trifoliata L. N Fig. 724 buck-bean. Menyanthes trifoliata L. var. minor Raf. • CT, MA, ME, NH, RI, VT. Fens, peaty meadows, lakeside fens.

Nymphoides

Fig. 724  Habit of Menyanthes trifoliata.

1a. Corolla yellow, 20–25 (–30) mm wide in life, the lobes short-fringed along part of the margin; flowers rarely replaced by spur-like roots; reproductive stem with a pair of opposite leaves subtending the umbel, sometimes extended beyond the umbel and producing additional inflorescences; fruit 12–25 mm long; seeds ciliate on the margin . . . . . . . . . . . . . . . . . . . N. peltata 1b. Corolla white, 5–8 (–10) mm wide in life, the lobes not fringed on the margin; some flowers usually replaced by a cluster of spur-like roots [Fig. 725]; reproductive stem with a single, alternate leaf subtending the umbel; fruit 4–5 mm long; seeds not ciliate . . . . . . . . . . N. cordata 1. Nymphoides cordata (Ell.) Fern. N Fig. 725 little floating-heart. Limnanthemum lacunosum (Vent.) Griseb.; Nymphoides lacunosa (Vent.) Kuntze; Villarsia cordata Ell.; V. lacunosa Vent. • CT, MA, ME, NH, RI, VT. Still or slow-moving water of lakes and rivers. 2. Nymphoides peltata (Gmel.) Kuntze E yellow floating-heart. Limnanthemum peltatum Gmel.; Nymphoides nymphaeoides (L.) Britt. • CT, MA, VT. Still or slow-moving water of lakes and rivers.

Molluginaceae Mollugo 1. Mollugo verticillata L. E green carpetweed. CT, MA, ME, NH, RI, VT. Roadsides, yards and lots, gardens, waste areas, cracks in pavement and concrete.

Fig. 725  Habit of Nymphoides cordata showing a cluster of spur-like roots.

670  tricolpate s

Moraceae References: Mitchell (1988), Wunderlin (1997). 1a. Plants herbaceous, annual; proximal portion of first pair of primary lateral veins at the very margin of the leaf blade (i.e., the basiscopic side of the vein exposed for a short distance) [Fig. 726] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fatoua 1b. Plants woody, perennial; first pair of primary lateral veins not at the margin of the leaf blade (i.e., both the basiscopic and acroscopic sides bordered by green leaf tissue) 2a. Leaf blades entire and unlobed, pinnately veined; branches often with axillary spines; fruit yellow-green, spherical, 6–12 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Maclura 2b. Leaf blades toothed, usually at least some on the plant lobed, either with obvious palmate venation or with 3 main veins from the base of the blade; branches without axillary spines; fruit short-cylindric, spherical, or pyriform in outline, 1–8 cm wide 3a. Flowers enclosed in a fleshy receptacle and accessible only by a small apical opening; terminal winter bud surrounded by a pair of stipules . . . . . . . . . . . . . . . . . . . . Ficus 3b. Flowers not enclosed in the receptacle, visible without dissection [Fig. 727]; terminal winter bud not concealed by stipules 4a. Style bifid [Fig. 727]; carpellate aments and fruits short-cylindric; petioles 2–5 cm long; branchlets glabrous or pubescent, but the hairs not long and spreading . . . Morus 4b. Style undivided; carpellate aments and fruits spherical; petioles 5–10 cm long; branchlets densely hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Broussonetia

Broussonetia 1. Broussonetia papyrifera (L.) ĽHér. ex Vent. E paper-mulberry. Morus papyrifera L.; Papyrius papyriferus (L.) Kuntze • CT, MA, RI. Fields, roadsides.

Fatoua 1. Fatoua villosa (Thunb.) Nakai E Fig. 726 hairy crabweed. Urtica villosa Thunb. • CT. Compost heaps, waste areas. Fatoua villosa was reported from MA by Miller and Wood (2003). However, the individuals are confined to plant pots and the cracks of a brick patio. This is not considered evidence of naturalization. Fatoua villosa bears superficial similarity to Urtica; however, its leaves are alternate, its hairs are not stinging, and it is very distinctive in that the proximal portion of the lowest pair of lateral veins is the margin of the blade. Fig. 726  Inflorescence and basal portion of leaf of Fatoua villosa.

Ficus 1. Ficus carica L. E common fig. MA. Roadsides, clearings, disburbed places.

Maclura 1. Maclura pomifera (Raf.) Schneid. E Osage-orange. Ioxylon pomiferum Raf.; Toxylon pomiferum Raf. ex Sarg. • CT, MA, RI. Roadsides, fields.

M o r ac e a e   671

Morus 1a. Leaf blades adaxially with short, stiff, appressed hairs providing a scabrous texture, abaxially sparsely to densely pubescent across the surface, (7–) 10–18 (–36) × (3–) 8–12 (–15.5) cm; multiples usually pink-red to red-purple at maturity; style branches white, mostly 1.5–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. rubra 1b. Leaf blades adaxially glabrous or sparsely and softly pubescent, abaxially glabrous or with hairs along the main veins and/or tufted in the axils of veins, (4–) 6–12 (–18) × 3–6 (–12) cm; multiples usually green-white (sometimes tinged with pink) or purple to black; style branches red-brown, 0.5–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. alba 1. Morus alba L. E Fig. 727 white mulberry. Morus alba L. var. tatarica (L.) Ser.; M. tatarica L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, areas of habitation. 2. Morus rubra L.

nC

red mulberry. Morus rubra L. var. tomentosa (Raf.) Bureau • CT, MA, RI, VT; mainly western New England. Rocky forests, ridges, edge of talus slopes, base of ledges. This species is occasionally planted, and the RI occurrences are likely non-native.

Fig. 727  Inflorescence of Morus alba.

Myricaceae References: Baird (1968), Bornstein (1997). 1a. Leaves stipulate, with pinnately lobed blades; bracts exceeding the fruit, persistent, forming an enclosing bur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Comptonia 1b. Leaves exstipulate, the blades entire or toothed but without lobes [Fig. 728]; bracts shorter than to equaling the fruit, not forming a bur 2a. Carpellate bracts 2, accrescent, clasping the achene in fruit; achene flattened, not concealed with a thick layer of wax; staminate aments borne at the apex of the previous year’s branchlet; leaf blades serrate in the distal ⅓; winter buds dark brown, ovoid, acute at the apex; preformed staminate aments present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myrica 2b. Carpellate bracts 4–6, small and inconspicuous, even in fruit; achene subglobose, covered with a thick layer of blue-white wax; staminate aments borne below the apex of the previous year’s branchlet; leaf blades variably serrate in the distal ½; winter buds red, subglobose, obtuse at the apex; preformed staminate aments absent . . . . . . . . . . . . Morella

Comptonia 1. Comptonia peregrina (L.) Coult. N sweet-fern. Comptonia peregrina (L.) Coult. var. aspleniifolia (L.) Fern.; Myrica aspleniifolia L.; M. peregrina (L.) Kuntze; Liquidambar peregrina L. • CT, MA, ME, NH, RI, VT. Dry fields, roadsides, woodlands, sand plains, clearings, abandoned borrow pits.

Morella Floral and fruiting morphology, as well as stem and leaf anatomy, support recognition of Morella as distinct from Myrica. 1. Morella caroliniensis (P. Mill.) Small n Fig. 728 small bayberry. Cerothamnus caroliniensis (P. Mill.) Tidestrom; Morella pensylvanica (Mirbel in Duhamel) Kartesz; Myrica caroliniensis P. Mill.; M. heterophylla Raf.; Myrica pensylvanica Mirbel

Fig. 728  Leaves and waxencrusted fruits of Morella caroliniensis.

67 2 tricolpates

in Duhamel • CT, MA, ME, NH, RI, VT. Fields, forest borders, roadsides, ledges, coastal headlands and beach borders, dunes. This species is commonly planted and some populations represent introductions, including all those in VT. It has long been known by the name Myrica pensylvanica (Wilbur 2002).

Myrica 1. Myrica gale L. N sweetgale. Gale palustris Chev.; Myrica gale L. var. subglabra (Chev.) Fern.; M. gale L. var. tomentosa C. DC. • CT, MA, ME, NH, RI, VT. Peaty and/or gravelly lake and river shores, peatlands, open swamps.

Myrsinaceae See references provided under the Primulaceae for rationale of recognizing of this family.

Lysimachia Recent phylogenetic research shows that Anagallis, Centunculus, Glaux, and Trientalis are nested within Lysimachia and, therefore, must be included in it (other options exist, but in this case they would lead to the formation of numerous, poorly defined genera; Hao et al. 2004, Anderberg et al. 2007, Manns and Anderberg 2009). References: Ray (1956), Cholewa (2009). 1a. Corolla pink, red to orange-red, or blue (rarely white); plants annual; fruit a pyxis 2a. Leaves alternate; pedicels up to 1 mm long; calyx 1.5–3 mm long; corollas 1.4–1.7 (–2) mm long; capsules 1.5–2 mm long, with 5–13 seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. minima 2b. Leaves opposite; pedicels 3–35 mm long; calyx 3.5–5 mm long; corollas (2–) 3–7 (–10) mm long; capsules 4–6 mm long, with 12–45 seeds . . . . . . . . . . . . . L. arvensis 1b. Corolla white or yellow; plants perennial; fruit a longitudinally dehiscent or disintegrating capsule 3a. Stems with 1 or more small, scale-like leaves below and a single whorl of leaves at the apex; flowers usually 7-merous, with white corollas . . . . . . . . . . . . . . . . . . . . . . . . . . L. borealis 3b. Stems with many nodes of opposite or whorled leaves; flowers 5- or 6-merous, with yellow corollas or with a white, petaloid calyx in the halophytic L. maritima 4a. Calyx petaloid; corolla absent; flowers axillary and subsessile; plants coastal halophytes with fleshy leaf blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. maritima 4b. Calyx sepaloid; corolla present; flowers axillary or in terminal inflorescences, borne on short to elongate pedicels; plants usually not of saline habitats, with herbaceous leaf blades 5a. Leaf blades not punctate; androecium with staminodes alternating with stamens, the filaments distinct; lobes of the corolla erose on the apical portion of the margin, cuspidate at the apex [Fig. 729] 6a. Leaf blades linear to narrow-lanceolate, 2–6 (–7) mm wide, with obscure lateral veins, the margins eciliolate and revolute . . . . . . . . . . . . . . . . . . . . . . . . L. quadriflora 6b. Leaf blades broad-linear to ovate, 2–65 mm wide, the lower blades usually wider than 6 mm, with evident lateral veins, the margins ciliolate and flat

My rs i n ac e a e   673

7a. Leaf blades broad-lanceolate to ovate, rounded (infrequently to subcordate) at the base, the principal ones 15–60 mm wide; petioles usually ciliate their entire length; plants with long, slender rhizomes . . . . . . . . . . . . . . . . . . . . L. ciliata 7b. Leaf blades broad-linear to narrow-ovate, cuneate (infrequently to rounded) at the base, the principal ones 10–20 mm wide; petioles ciliate only in the basal half, sometimes lacking cilia altogether; plants with short, stout rhizomes (very rarely with slender, elongate rhizomes) . . . . . . . . . . . . . . . . . . . . . . . . . . . L. hybrida 5b. Leaf blades minutely punctate; androecium without staminodes, the filaments connate near the base (filaments distinct in L. thyrsiflora); lobes of the corolla entire (glandular-ciliate in L. punctata), without a cuspidate apex 8a. Flowers borne in short, dense racemes originating from the middle leaf axils [Fig. 730]; filaments distinct, usually of equal length; corolla 6- or 7-merous, the lobes 4–5 mm long, linear, and much shorter than the stamens . . . . L. thyrsiflora 8b. Flowers solitary in the axils of leaves or in racemes from the apex of the stem or branches; filaments connate at the base, often of unequal length; corolla 5-merous, the lobes 5–16 mm long, lanceolate to ovate, and as long as or longer than the stamens 9a. Leaf blades quadrate to suborbicular, 1–2.5 cm long; stems extensively creeping over the ground . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. nummularia 9b. Leaf blades narrow-lanceolate to narrow-ovate, 3–12 cm long; stems erect or ascending 10a. Corolla white; leaves alternate . . . . . . . . . . . . . . . . . . . . . . . . L. clethroides 10b. Corolla yellow; leaves opposite or whorled 11a. Petals not dotted or streaked with black or red; stems pubescent 12a. Calyx lobes 3–5 mm long, dark-margined with red; lobes of the corolla 8–12 mm long, eciliate; flowers borne in terminal or axillary racemes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. vulgaris 12b. Calyx lobes 5–8 mm long, entirely green; lobes of the corolla 12–16 mm long, ciliate with stipitate glands; flowers borne in whorls from the axils of the leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. punctata 11b. Petals dotted or streaked with black or red; stems glabrous or sparsely pubescent 13a. Flowers borne in a terminal raceme; leaves normally opposite, commonly developing bulbils in the axils; pedicels 8–15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. terrestris 13b. Flowers borne in whorls from the axils of the leaves; leaves normally whorled, without bulbils; pedicels 20–50 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. quadrifolia 1. Lysimachia arvensis (L.) U. Manns & A. Anderb. E scarlet pimpernel. Anagallis arvensis L.; A. arvensis L. var. caerulea (Schreb.) Gren. & Godr.; A. caerulea Schreb. • CT, MA, ME, NH, RI, VT. Fields, gardens, coastal turfs, beach and tidal marsh borders, and headlands. Lysimachia foemina (P. Mill.) U. Manns & Anderb. (synonym: Anagallis foemina) has been attributed to New England on the basis of rare plants of L. arvensis with blue corollas. These two species differ in petal margin morphology. Lysimachia arvensis has petals ciliate with (usually) abundant marginal hairs comprised of 3 cells, the terminal cell globose, whereas L. foemina has petals with few or no marginal hairs, the hairs (when present) with (3–) 4 cells and the terminal cell elongate. Examination of New England collections of Lysimachia with blue-flowered corollas show these plants to be L. arvensis.

674 tricolpate s

2. Lysimachia borealis (Raf.) U Manns & A. Anderb. N starflower. Trientalis americana Pursh; Trientalis borealis Raf. ssp. borealis • CT, MA, ME, NH, RI, VT. Deciduous to evergreen forests. 3. Lysimachia ciliata L. N fringed yellow-loosestrife. Steironema ciliatum (L.) Baudo • CT, MA, ME, NH, RI, VT. River banks, riparian forests, lake shores, wet ditches, wetland borders, low fields. 4. Lysimachia clethroides Duby in DC. E goose-neck yellow-loosestrife. CT, ME. Fields, gardens, around dwellings. 5. Lysimachia hybrida Michx. N Fig. 729 lowland yellow-loosestrife. Lysimachia lanceolata Walt. ssp. hybrida (Michx.) J.D. Ray; L. lanceolata Walt. var. hybrida (Michx.) Gray; Steironema hybridum (Michx.) Raf. ex B.D. Jackson; S. lanceolatum (Walt.) Gray var. hybridum (Michx.) Gray • CT, MA, ME, NH, RI, VT. River banks, riparian forests, lake shores, wetland borders, wet ditches, low fields. Reports of Lysimachia lanceolata from New England are based on this species. Fig. 729  Flower with antepetalous stamens of Lysimachia hybrida.

6. Lysimachia maritima (L.) Galasso, Banfi, & Soldano N sea-milkwort. Glaux maritima L.; G. maritima L. var. angustifolia Boivin; G. maritima L. ssp. obtusifolia (Fern.) Boivin; G. maritima L. var. obtusifolia Fern. • MA, ME, NH, RI. Saline and brackish marshes and shorelines. 7. Lysimachia minima (L.) U. Manns & A. Anderb. E chaffweed pimpernel. Anagallis minima (L.) Krause; Centunculus minimus L. • MA. Roadsides, waste areas. 8. Lysimachia nummularia L. E creeping yellow-loosestrife. CT, MA, ME, NH, RI, VT. Riparian forests, abandoned homesteads, lawns, fields, roadsides, waste areas, most often on mesic soils in the shade. 9. Lysimachia punctata L. E large yellow-loosestrife. Lysimachia punctata L. var. verticillata (Bieb.) Klatt • CT, MA, ME, NH, VT. Fields, roadsides, gardens, areas of habitation. 10. Lysimachia quadriflora Sims

NC

four-flowered yellow-loosestrife. Lysimachia longifolia Pursh; L. revoluta Nutt.; Nummularia longifolium (Pursh) Kuntze; Steironema longifolium Gray; S. quadriflorum (Sims) A.S. Hitchc.; S. revolutum (Nutt.) Raf. ex Steud. • Riverbanks, riparian forests, wetland borders, wet ditches. In New England, this species was reported from “dry woods” on one collection (a very anomalous habitat for this species). It is plausible that the Middlesex County, MA, plant was introduced. 11. Lysimachia quadrifolia L. N whorled yellow-loosestrife. CT, MA, ME, NH, RI, VT. Woodlands, sandplains, dry fields and roadsides, clearings. ‌ × 12. Lysimachia ×producta (Gray) Fern. is an uncommon yellow-loosestrife hybrid 11 that is known from CT, MA, ME, NH, RI, VT. It most closely resembles L. terrestris; however, the lower bracts are larger, even foliaceous, and the hybrid shows longer pedicels (up to 30 mm long). 12. Lysimachia terrestris (L.) B.S.P. N swamp yellow-loosestrife. Lysimachia terrestris (L.) B.S.P. var. ovata (Rand & Redf.) Fern. • CT, MA, ME, NH, RI, VT; throughout. Marshes, shorelines, low fields and meadows, swamps. ‌12 × 13. Lysimachia ×commixta Fern. is a rare yellow-loosestrife hybrid known from MA, ME, VT. It has distinct filaments (similar to L. thyrsiflora); however, the racemes are both terminal and axillary (instead of just axillary) and the filaments are equal to slightly longer than the corolla (instead of much longer than the corolla)

My r s in ac e a e   675

13. Lysimachia thyrsiflora L. N Fig. 730 tufted yellow-loosestrife. Naumburgia thyrsiflora (L.) Duby • CT, MA, ME, NH, RI, VT. Swamps, peatlands, lake shores. 14. Lysimachia vulgaris L. E garden yellow-loosestrife. CT, MA, ME, NH, RI, VT. Fields, roadsides, gardens, clearings, areas of habitation.

Nelumbonaceae Fig. 730  Inflorescences of Lysimachia thyrsiflora.

Nelumbo 1. Nelumbo lutea Willd. n Fig. 731 American lotus. Nelumbium luteum Willd.; Nymphaea pentaphylla Walt. • CT, MA, ME. Shallow, still or slow-moving water of lakes and rivers, including fresh-tidal waters. Some populations of this plant in New England are certainly introduced. The ME population and many of the MA populations are believed to be non-native.

Nyctaginaceae Mirabilis 1a. Involucres 1-flowered, cup-shaped, not accrescent in fruit; calyx salverform, with a prolonged, tubular, connate portion 20–30 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. jalapa 1b. Involucres 3- to 5-flowered, broad and open, accrescent in fruit [Fig. 732]; calyx campanulate, with a short, tubular, connate portion 1–2 mm long 2a. Leaf blades ovate to broad-ovate, the larger ones rounded to cordate at the base and borne on petioles 10–30 mm long; inflorescence eglandular . . . . . . . . . . . . . . M. nyctaginea 2b. Leaf blades linear to lanceolate or oblong, tapering to the base, sessile or on short petioles up to 5 mm long; inflorescence usually glandular-pubescent 3a. Leaf blades linear, usually 5 mm wide or narrower; stem glabrous . . . . . . . M. linearis 3b. Leaf blades narrow-lanceolate to lanceolate or oblong, 5–20 mm wide; stem pubescent 4a. Stem pubescent in 2 strips on each internode with short, curved hairs up to 0.5 mm long; anthocarp coarsely tuberculate on the ridges . . . . . . . . . . . . . . M. albida 4b. Stem pubescent, at least at the nodes, with spreading hairs 1–2 mm long; anthocarp relatively smooth on the ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. hirsuta 1. Mirabilis albida (Walt.) Heimerl E pale umbrella-wort. Allionia albida Walt.; Mirabilis albida (Walt.) Heimerl var. lata Shinners; Oxybaphus albidus (Walt.) Sweet • MA. Fields, roadsides, gardens. 2. Mirabilis hirsuta (Pursh) MacM. E hairy umbrella-wort. Allionia hirsuta Pursh; Oxybaphus hirsutus (Pursh) Sweet • CT, MA, ME, VT. Fields, roadsides, waste areas, railroads.

Fig. 731  Flower, immature fruit, and leaf of Nelumbo lutea.

67 6  tricolpates

3. Mirabilis jalapa L. E four o’clock umbrella-wort. Mirabilis jalapa L. ssp. lindheimeri Standl.; M. lindheimeri (Standl.) Shinners • CT, VT. Fields, roadsides, gardens. 4. Mirabilis linearis (Pursh) Heimerl var. linearis E narrow-leaved umbrella-wort. Allionia linearis Pursh; Mirabilis lanceolata (Rydb.) Standl.; M. linearis (Pursh) Heimerl var. subhispida Heimerl; Oxybaphus angustifolius Sweet; O. lanceolatus (Rydb.) Standl.; O. linearis (Pursh) B.L. Robins.; O. linearis (Pursh) B.L. Robins. var. subhispidus (Heimerl) Dayton • CT. Waste areas. 5. Mirabilis nyctaginea (Michx.) MacM. E Fig. 732 heart-leaved umbrella-wort. Allionia nyctaginea Michx.; Oxybaphus nyctagineus (Michx.) Sweet • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads.

Fig. 732  Flowers of Mirabilis nyctaginea.

Oleaceae 1a. Leaf blades pinnately compound with 5–13 (–15) leaflets; corolla absent; fruit a samara . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fraxinus 1b. Leaf blades simple or sometimes pinnately compound with 3 leaflets in Forsythia; corolla present and conspicuous; fruit a drupe or capsule 2a. Flowers precocious, with yellow corollas; leaf blades toothed; branchlets with hollow or chambered pith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Forsythia 2b. Flowers not appearing well in advance of leaf emergence, with white or light bluepurple corollas (infrequently red-blue or red-purple); leaf blades entire; branchlets with solid pith 3a. Corolla not salverform, with elongate lobes that are much longer than the basal, connate portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chionanthus 3b. Corolla salverform, the lobes not conspicuously longer than the basal, connate, tubular portion 4a. Fruit a capsule; anthers included within the connate, tubular portion of the corolla (exserted in S. reticulata); branchlets relatively stout, usually thicker than 2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Syringa 4b. Fruit a drupe; anthers shortly exserted from the tubular, connate portion of the corolla [Figs. 735, 736]; branchlets slender, usually thinner than 2 mm . . . . Ligustrum

Chionanthus 1. Chionanthus virginicus L. E white fringe-tree. Chionanthus virginicus L. var. maritimus Pursh • MA; also reported from RI by George (1997), but specimens are unknown. Fields, forests borders and fragments, roadsides.

Forsythia Reports of Forsythia ovata Nakai from New England are based on collections of F. suspensa; e.g., 29 Aug 1947, Bean s.n. (NEBC!). Cultivated members of this genus were reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated they were not naturalized in RI. 1a. Calyx lobes (5–) 6–7 mm long; branchlets spreading to drooping; pith hollow through internodes and solid at the nodes; leaf blades broad-cunate to rounded at the base, sometimes 3-lobed or 3-foliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. suspensa

O l e ac e a e   67 7

1b. Calyx lobes 2–4.5 mm long; branchlets mostly ascending to arching; pith chambered though the internodes and chambered or solid at the nodes; leaf blades mostly cuneate to broad-cunate at the base, simple (sometimes 3-lobed in F. intermedia) 2a. Pith chambered through the nodes; branchlets green to yellow-green; petals yellow with a green tinge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. viridissima 2b. Pith solid at the nodes; branchlets yellow-brown to brown; petals pale to deep yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. intermedia 1. Forsythia intermedia Zabel E border forsythia. MA, RI. Forest fragments and edges, roadsides, areas of habitation. Kim (1999) analyzed the cpDNA of several species and cultivars of Forsythia. The results revealed that Forsythia intermedia is not the hybrid of F. suspensa and F. viridissima as suggested by many authors. 2. Forsythia suspensa (Thunb.) Vahl E weeping forsythia. CT, MA, VT. Roadsides, forest borders, fields, areas of habitation. 3. Forsythia viridissima Lindl. E green-stemmed forsythia. CT, MA, NH. Roadsides, forest borders, fields, areas of habitation.

Fraxinus 1a. Leaflets borne on short petiolules 3–15 mm long, 5–9 per leaf; flowers with a calyx, which is persistent on the fruit and recognizable as a lobed cupule at the base of the samara; wing of samara decurrent on ca. ½ of the body or less; terminal winter bud adjacent to the distalmost pair of lateral buds [Figs. 733, 734] 2a. Branchlets brown to blue-brown, glabrous; leaflets papillose on the abaxial surface, glabrous; terminal winter bud rounded [Fig. 733]; leaf scar prominently concave on distal margin [Fig. 733]; wing of samara extending ca. ⅓ of the distance to the base of the body; leaf scars with a concave upper margin; petiolules not winged . . . . . . . . . . . . . . F. americana 2b. Branchlets gray-brown, pubescent; leaflets not papillose on the abaxial surface, pubescent (sometimes only sparsely so); terminal winter bud acute [Fig. 734]; leaf scar straight or slightly concave on distal margin [Fig. 734]; wing of samara extending ca. ½ of the distance to the base of the body; leaf scars with a straight or slightly concave upper margin; petiolules commonly winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. pennsylvanica 1b. Leaflets sessile, 7–13 (–15) per leaf; flowers without a calyx; wing of samara decurrent nearly to the base of the body; terminal winter bud separated from the distal-most pair of lateral buds 3a. Leaf rachis with thick patches of tomentum at the nodes where the leaflets attach; leaflets 7–14 (–16) × (2.5–) 3–6.5 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. nigra 3b. Leaf rachis glabrous to puberulent, but without patches of tomentum; leaflets (3–) 5–11 × 1–3 (–4) cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. excelsior 1. Fraxinus americana L. N Fig. 733 white ash. Fraxinus americana L. var. biltmoreana (Beadle) J. Wright ex Fern.; F. americana L. var. microcarpa Gray; F. biltmoreana Beadle • CT, MA, ME, NH, RI, VT; nearly throughout. Upland, deciduous and mixed evergreen-deciduous forests, riparian forests. 2. Fraxinus excelsior L. E European ash. CT, MA. Roadsides, areas of habitation. 3. Fraxinus nigra Marsh. N black ash. CT, MA, ME, NH, RI, VT. Forested swamps and low, riparian forests with a canopy of deciduous or mixed evergreen-deciduous trees.

Fig. 733  Winter branchlet of Fraxinus americana.

67 8 tricolpate s

4. Fraxinus pennsylvanica Marsh. N Fig. 734 green ash. Fraxinus pennsylvanica Marsh. var. austinii Fern.; F. pennsylvanica Marsh. var. integerrima (Vahl) Fern.; F. pennsylvanica Marsh. var. lanceolata (Borkh.) Sarg.; F. pennsylvanica Marsh. var. subintegerrima (Vahl) Fern. • CT, MA, ME, NH, RI, VT. Swamps, shorelines, riparian forests, less frequently in upland forests.

Ligustrum Fig. 734  Winter branchlet of Fraxinus pennsylvanica.

Species of Ligustrum are commonly planted for hedges in New England. Some of the species found in the area are native to Asia and, in those regions, are subdivided into varieties. However, no detailed systematic work has been done to determine which infraspecific taxa are planted here. Therefore, the following keys separate privets only to the rank of species. Reference: Nesom (2009a). 1a. Branchlets either glabrous or minutely and uniformly puberulent; leaf blades glabrous 2a. Basal, connate portion of corolla 5–8 mm long, 1.5–3 times longer than the lobes [Fig. 735]; anthers surpassing the tips of the corolla lobes; panicle and branchlets glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. ovalifolium 2b. Basal, connate portion of corolla 2.5–3 mm long, approximately as long as the lobes or a little shorter [Fig. 736]; anthers not surpassing the tips of the corolla lobes; panicle and usually the branchlets minutely and uniformly puberulent . . . . . . . . . . . . . . . . . . . . . L. vulgare 1b. Branchlets pilose and/or pubescent with hairs of distinctly different lengths; leaf blades pubescent at least on the abaxial midrib or sometimes glabrous in L. obtusifolium 3a. Basal, connate portion of corolla 2.5–3 mm long, slightly shorter to approximately as long as the lobes [Fig. 736], usually barely exserted from calyx; anthers 1–2 mm long, typically exserted beyond the tips of the corolla lobes; panicles 6–10 cm tall, relatively open, often interspersed with non-flowering branchlets; pedicels 1–5 mm long . . . . . . . . . . . . . . L. sinense 3b. Basal, connate portion of corolla 5–8 mm long, 1.5–3 times as long as the lobes (rarely only as long as lobes) [Fig. 735], evidently exserted from calyx; anthers 2–3 mm long, not surpassing the tips of the corolla lobes; panicles 1.5–4 cm tall, relatively compact, not usually interspersed with non-flowering branchlets; pedicels (0–) 0.5–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. obtusifolium 1. Ligustrum obtusifolium Sieb. & Zucc. E Fig. 735 border privet.  1a. Ligustrum obtusifolium Sieb. & Zucc. var. leiocalyx (Nakai) H. Hara;  1b. Ligustrum amurense Carr.; L. ibota Sieb. & Zucc. var. suave Kitagawa; L. obtusifolium Sieb. & Zucc. ssp. suave (Kitagawa) Kitagawa • CT, MA, ME, NH, RI, VT. Roadsides, forest edges and fragments, areas of habitation.

Fig. 735  Inflorescence of Ligustrum obtusifolium var. obtusifolium.

1a. Leaf blades usually sparsely pubescent abaxially; calyx and pedicels densely to moderately pubescent (varying rarely to sparsely pubescent or nearly glabrous) [Fig. 735]; basal, connate portion of corolla 2–3 times as long as the lobes; lateral winter buds with acute to obtuse scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. L. obtusifolium var. obtusifolium 1b. Leaf blades usually glabrous abaxially; calyx and pedicels glabrous to sparsely pubescent; basal, connate portion of corolla 1.5–2 times as long as the lobes; lateral winter buds with acuminate scales . . . . . . . . . . . . . . . . . . . . . . . 1b. L. obtusifolium var. suave (Kitagawa) H. Hara Variety obtusifolium is known from CT, MA, NH, RI, VT. Variety suave is known from MA, ME. The collection from ME—Wise & Hodgdon 517 (NHA!)—is atypical in its pubescent leaf blades (but otherwise appears to be a good fit for the taxon). Variety suave is usually treated at the species level as Ligustrum amurense; however, Nesom (2009a) outlines rationale for recognizing this taxon as a variety. 2. Ligustrum ovalifolium Hassk. E California privet. CT, MA. Roadsides, forest edges and fragments, areas of habitation.

Ol e ac e a e   67 9

3. Ligustrum sinense Lour. E Chinese privet. Ligustrum villosum May • CT, MA, RI. Roadsides, forest edges and fragments, areas of habitation. 4. Ligustrum vulgare L. E Fig. 736 European privet. CT, MA, ME, NH, RI, VT. Roadsides, forest edges and fragments, areas of habitation.

Syringa Reference: Li et al. (2001). 1a. Basal, connate portion of the corolla about as long as or slightly longer than the calyx lobes [Fig. 737]; filaments relatively long, exserted beyond the connate, tubular portion of the corolla [Fig. 737]; corolla white to cream . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. reticulata 1b. Basal, connate portion of the corolla 2.5–8 times as long as the calyx lobes; filaments very short, not exserted beyond the connate, tubular portion of the corolla, the anthers subsessile; corolla usually light purple, though sometimes white, blue, purple, or red-purple

Fig. 736  Inflorescence of Ligustrum vulgare.

2a. Terminal winter bud absent, the main axis of the branchlet not continuing growth in subsequent seasons; panicles produced only from lateral buds . . . . . . . . . . . . . . . S. vulgaris 2b. Terminal winter bud present, the main axis of the branchlet continuing growth the following season; panicles produced, in part, from terminal buds 3a. Corolla with a ± cylindrical connate portion and spreading lobes; panicle tending to arch or even droop; anthers inserted near opening of corolla . . . . . . . . . . . . . . . S. villosa 3b. Corolla with a slender-funnelform connate portion and ascending lobes; panicle upright; anthers inserted well below opening of corolla . . . . . . . . . . . . . . . . . . . S. josikaea 1. Syringa josikaea Jacq. f. ex Reichenb. E Hungarian lilac. MA. Fields, roadsides. 2. Syringa reticulata (Blume) Hara ssp. reticulata E Fig. 737 Japanese tree lilac. Syringa amurensis Rupr. var. japonica (Maxim. ex Dcne.) Franch. & Sav.; S. japonica Maxim. ex Dcne. • MA, NH, VT. Fields, roadsides, forest borders, abandoned homesteads. Reports of Syringa reticulata ssp. amurensis (Rupr.) P.S. Greene from New England are based on specimens of Syringa reticulata ssp. reticulata. 3. Syringa villosa Vahl E late lilac. MA. Fields, roadsides. 4. Syringa vulgaris L. E common lilac. CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, abandoned homesteads. ‌ × Syringa protolaciniata P.S. Greene & M.C. Chang. Syringa ×persica L. is a 4 distinctive lilac hybrid that is known from ME. It rarely escapes cultivation and is frequently referred to as S. chinensis Willd. It is similar to S. vulgaris in many features, including lack of terminal buds and relatively long basal, connate portion of the corollas, but the leaf blades are lanceolate to broad-lanceolate and cuneate at the base (rather than ovate to broad-ovate and truncate to cordate at the base). Further, the leaf blades and flowers average smaller in S. × ‌persica (blades 3–8 cm long and connate portion of corolla mostly 7–10 mm long, rather than 5–12 cm long and 8–12 mm long).

Fig. 737  Flowers of Syringa reticulata with exserted stamens.

68 0   tricolpate s

Onagraceae 1a. Perianth 2-merous [Fig. 738]; fruit indehiscent, covered with uncinate bristles; leaves opposite; petals white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Circaea 1b. Perianth 4- to 6-merous; fruit dehiscent (indehiscent in some Oenothera), without uncinate bristles; leaves opposite or alternate; petals absent, or white to pink to purple, or yellow 2a. Sepals persistent in fruit; androecium with 4 stamens; hypanthium not prolonged beyond the summit of the ovary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ludwigia 2b. Sepals deciduous in fruit; androecium with 8 stamens; hypanthium prolonged beyond the summit of the ovary (except in Chamerion) 3a. Flowers zygomorphic, the petals all oriented toward one side of the flower [Fig. 744]; fruit indehiscent, with 2–6 seeds 1.5–3.5 mm long . . . . . . . . . . . . . . . . . (in part) Oenothera 3b. Flowers actinomorphic [Fig. 745]; fruit dehiscent, with many seeds 0.3–2 mm long 4a. Seeds comose; leaves opposite (especially the lower ones) or alternate throughout in Chamerion; hypanthium not, or only shortly, extending beyond the ovary; petals white to pink to pink-purple 5a. Hypanthium shortly extending beyond the summit of the ovary; petals 2–16 mm long, notched at the apex; stamens in 2 series, erect to ascending; stigma unlobed or with 4 lobes; new shoots produced from turions, rosettes, or sobols . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epilobium 5b. Hypanthium not extending beyond the summit of the ovary; petals 10–20 mm long, ± entire at the apex; stamens in 1 series, declined; stigma with 4 conspicuous lobes; new shoots produced from lateral roots (or sometimes by sobols) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chamerion 4b. Seeds not comose; leaves alternate; hypanthium extending beyond the summit of the ovary; petals yellow (sometimes fading to red) or pink to light purple or redpurple (rarely white) in Clarkia 6a. Petals pink to light purple or red-purple (rarely white), with a pair of small lobes or recurved teeth on the claw; anthers basifixed . . . . . . . . . . . . . . . . . . Clarkia 6b. Petals yellow (sometimes fading to red), lacking lobes or teeth on the claw; anthers mesifixed (i.e., versatile) 7a. Petals with 1 or 2 red dots near the base; stigma capitate or hemispherical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camissonia 7b. Petals without basal red dots; stigma with 4 lobes (caution: the lobes short in O. serrulata) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Oenothera

Camissonia 1a. Fruit 0.7–2 mm wide, terete in cross-section; plants 5–25 (–50) cm tall, generally lacking a basal rosette of leaves; seeds shiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. campestris 1b. Fruit 1.5–2.5 mm wide, quadrangular in cross-section; plants 50–80 cm tall, with a basal rosette of leaves; seeds dull . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bistorta 1. Camissonia bistorta (Nutt. ex Torr. & Gray) Raven E California suncup. Oenothera bistorta Nutt. ex Torr. & Gray • MA. Wool waste. 2. Camissonia campestris (Greene) Raven ssp. campestris E Mojave suncup. Oenothera campestris Greene; O. dentata Cav. var. campestris (Greene) Jepson • MA. Wool waste.

On ag r ac e a e   6 8 1

Chamerion 1. Chamerion angustifolium (L.) Holub ssp. circumvagum (Mosq.) Kartesz N narrow-leaved fireweed. Chamaenerion angustifolium (L.) Scop. ssp. circumvagum (Mosq.) Moldenke; Chamerion platyphyllum (Daniels) A. & D. Löve; Epilobium angustifolium L. var. abbreviatum (Lunell) Munz; E. angustifolium L. ssp. circumvagum Mosq.; E. angustifolium L. var. macrophyllum (Hausskn.) Fern.; E. angustifolium L. var. platyphyllum (Daniels) Fern. • CT, MA, ME, NH, RI, VT; more frequent in northern New England. Fields, roadsides, clearings.

Circaea Reference: Boufford (1982). 1a. Ovary and fruit bilocular, the fruit 2.8–4.5 mm long; rhizomes slender; flowers borne on spreading pedicels [Fig. 738], opening after elongation of the raceme axis, therefore the inflorescence relatively open; petals (1.3–) 1.6–3.9 mm long; anthers (0.5–) 0.7–1 mm long; plants 15–70 (–100) cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. canadensis 1b. Ovary and fruit unilocular, the fruit 1.6–2.6 mm long; rhizomes tuberous-thickened at the end; flowers borne on ascending to erect pedicels, beginning to open prior to elongation of raceme axis, therefore the inflorescence congested near the apex; petals 0.6–2 mm long; anthers 0.2–0.3 mm long; plants 5–25 (–30) cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. alpina 1. Circaea alpina L. ssp. alpina N small enchanter’s-nightshade. CT, MA, ME, NH, RI, VT. Mesic to wet-mesic, deciduous to mixed evergreen-deciduous forests, less frequently in evergreen forests, stream banks, forested seeps. 1‌ × 2. Circaea ×sterilis Boufford is an infrequent, sterile enchanter’s-nightshade hybrid known from CT, MA, ME, NH, VT. Though it has rhizomes that sometimes terminate in enlargements or small tubers, the hybrids most generally resemble C. canadensis given the open inflorescence. However, the fruits abort prior to maturity and usually fall from the inflorescence or can be easily detached post anthesis. Those fruits that do persist on the plant are bilocular, with one locule larger than the other, and are smooth or have low ribs and shallow grooves (vs. with prominent ribs and deep grooves in C. canadensis). One of the best morphological features to identify this hybrid is anther size (0.4–0.7 mm long), which is intermediate between the parental taxa. 2. Circaea canadensis (L.) Hill ssp. canadensis N Fig. 738 broad-leaved enchanter’s-nightshade. Circaea canadensis (L.) Hill var. virginiana Fern.; C. latifolia Hill; C. lutetiana var. canadensis L.; C. quadrisulcata (Maxim.) Franch. & Savigny ssp. canadensis (L.) A. & D. Löve; C. quadrisulcata (Maxim.) Franch. & Savigny var. canadensis (L.) Hara • CT, MA, ME, NH, RI, VT. Mesic to hydric forests, including riparian types, when in uplands frequently found on or at the base of forested, rocky hillsides.

Clarkia 1a. Petals 15–30 mm long, the blade prominently 3-lobed; leaf blades linear to narrowlanceolate, 2–10 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pulchella 1b. Petals 7–14 mm long, the blade simple; leaf blades lanceolate to ovate, 5–20 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rhomboidea 1. Clarkia pulchella Pursh E elk horn clarkia. CT, MA, VT. Fields, wool waste. 2. Clarkia rhomboidea Dougl. ex Hook. E common clarkia. MA. Wool waste.

Fig. 738  Inflorescence of Circaea canadensis.

68 2   tricolpate s

Epilobium Vegetative perennating structures are very important for species determination in Epilobium. A thorough account of these structures is provided by Keating et al. (1982). 1a. Stigma 4-lobed; plants evidently long-villous; petals 10–16 mm long (only 4–9 mm in the very rare E. parviflorum) 2a. Petals 10–16 mm long, with a shallow notch; leaves sessile and somewhat clasping the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. hirsutum 2b. Petals 4–9 mm long, with a deep notch; leaves sessile or subsessile, but not clasping the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. parviflorum 1b. Stigma entire or nearly so; plants ± glabrous to short-pubescent; petals 2–10 mm long 3a. Leaf blades with entire or undulate, usually revolute, margins [Fig. 741]; pubescence of the stem not in decurrent lines from the leaf bases; plants perennating from turions at the tips of stolons 4a. Plants pubescent with divergent hairs (note: this best viewed along the main axis of the stem) [Fig. 741] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. strictum 4b. Plants pubescent with appressed hairs 5a. Leaf blades essentially glabrous on the adaxial surface, 2–15 mm wide; inflorescence nodding in bud . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. palustre 5b. Leaf blades strigose-puberulent on the adaxial surface, 1–7 mm wide; inflorescence erect to arching in bud . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. leptophyllum 3b. Leaf blades with usually toothed, flat margins (entire in E. anagallidifolium); pubescence of the stem in decurrent lines from the leaf bases; plants perennating from sobols, rosettes, or sessile turions 6a. Stems matted, decumbent to erect, soboliferous, usually unbranched above the base of the plant, 0.5–4.5 dm tall; plants boreal and alpine 7a. Leaf blades 10–25 × 3–7 mm, ± entire (rarely very weakly toothed); outer surface of seed smooth; capsules mostly 23–40 mm long; stems mostly 2–10 cm tall; petals pale purple in life, 3–4.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. anagallidifolium 7b. Leaf blades 19–40 × (6–) 8–15 mm, weakly serrate; outer surface of seed minutely papillose or minutely pitted; capsules mostly 35–75 mm long; stems 6–40 cm tall; petals white to pale purple, 2.5–8 mm long 8a. Petals white, 2.5–4 mm long; outer surface of seed minutely pitted; inflorescence erect prior to anthesis; sobols epigeal (though often short and inconspicuous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. lactiflorum 8b. Petals pale purple in life (rarely white; often fading to white with pink, red, or blue tinging in drying), 4–8 mm long; outer surface of seed minutely papillose; inflorescence nodding prior to anthesis; sobols hypogeal . . . . . . . . E. hornemannii 6b. Stems usually solitary, erect, without sobols, usually branched above the base of the plant, (0.5–) 3–10 dm tall; plants of various habitats, commonly found at lower elevations 9a. Coma brown; flower buds pointed at the apex due to the projecting sepal tips; seeds minutely papillose, without a beak; leaf blades gray-green, rugose-veiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. coloratum 9b. Coma white or nearly so; flower buds obtuse to rounded at the apex; seeds longitudinally striate, with a broad, short beak; leaf blades without gray color, not or only slightly rugose-veiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. ciliatum

On ag r ac e a e   6 83

1. Epilobium anagallidifolium Lam.

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pimpernel willow-herb. Epilobium alpinum L., pro parte; E. anagallidifolium Lam. var. pseudoscaposum (Hausskn.) Hultén • ME, NH; northern portion of states. Alpine gullies, slides, and ledges, borders of subalpine brooks. 2. Epilobium ciliatum Raf. N Fig. 739, 740 fringed willow-herb. 2a. Epilobium adenocaulon Hausskn.; E. adenocaulon Hausskn. var. ecomosum (Fassett) Munz; E. americanum Hausskn.; E. ciliatum Raf. var. ecomosum (Fassett) Boivin; E. glandulosum Lehm. var. adenocaulon (Hausskn.) Fern.; E. glandulosum Lehm. var. ecomosum Fassett; 2b. Epilobium adenocaulon Hausskn. var. cinerascens (Piper) M.E. Peck; E. adenocaulon Hausskn. var. occidentale Trel.; E. boreale Hausskn.; E. ciliatum Raf. ssp. glandulosum (Lehm.) Hoch & Raven; E. glandulosum Lehm.; E. glandulosum Lehm. var. cardiophyllum Fern.; E. glandulosum Lehm. var. occidentale (Trel.) Fern.; E. tetragonum L. var. glandulosum (Lehm.) Torr. & Gray • CT, MA, ME, NH, RI, VT. Swamps, wet fields, forested seeps, wetland margins, shorelines, stream banks, ledges.

Fig. 739  Basal rosettes of Epilobium ciliatum var. ciliatum.

1a. Plants usually with leafy basal rosettes (turions sometimes formed in high evelvation habitats) [Fig. 739]; stems well branched apically or throughout (rarely simple); inflorescence relatively open, with much reduced bract blades compared with the leaf blades; petals white or infrequently pink, 2–6 (–9) mm long; seeds (0.6–) 0.8–1.2 (–1.5) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a. E. ciliatum var. ciliatum 1b. Plants usually with fleshy underground turions [Fig. 740], these often near the surface and resembling compact rosettes of short, fleshy leaves; stems simple throughout or sparsely branched apically; inflorescence crowded, with scarcely reduced bract blades compared with the leaf blades; petals rose-purple to pink or rarely white, 4.5–12 (–14) mm long; seeds 1.1–1.6 (–1.9) mm long . . . . . . . . . . . . . . . . . . . . . . . 2b. E. ciliatum var. glandulosum (Lehm.) Dorn Variety ciliatum is known from CT, MA, ME, NH, RI, VT. Variety glandulosum is known from ME, NH, VT. 3. Epilobium coloratum Biehler N eastern willow-herb. CT, MA, ME, NH, RI, VT. Swamps, marshes, seepy areas, river shores, wet ditches. 4. Epilobium hirsutum L. E hairy willow-herb. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 5. Epilobium hornemannii Reichenb. ssp. hornemannii

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Hornemann’s willow-herb. Epilobium alpinum L. var. nutans Hornem. • ME, NH; northern portions of states. Boreal to alpine gullies, cliffs, stream banks, and seeps, frequently growing in wet moss. 6. Epilobium lactiflorum Hausskn.

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white-flowered willow-herb. Epilobium alpinum L. var. lactiflorum (Hausskn.) C.L. Hitchc.; E. hornemannii Reichenb. var. lactiflorum (Hausskn.) D. Löve • ME, NH; northern portions of states. Alpine gullies, cliffs, stream banks, and snowbank communities. 7. Epilobium leptophyllum Raf. N bog willow-herb. Epilobium nesophilum (Fern.) Fern.; E. nesophilum (Fern.) Fern. var. sabulonense Fern.; E. palustre L. var. gracile (Farw.) Dorn; E. palustre L. var. sabulonense (Fern.) Boivin • CT, MA, ME, NH, RI, VT. Bogs, fens, swamps, peaty meadows and marshes, wet ditches. 8. Epilobium palustre L. N marsh willow-herb. Epilobium lineare Muhl.; E. oliganthum Michx.; E. palustre L. var. grammadophyllum Hausskn.; E. palustre L. var. labradoricum Hausskn.; E. palustre L. var. oliganthum (Michx.) Fern. • CT, MA, ME, NH, RI, VT; becoming rare in southern New England. Bogs, fens, swamps, these frequently dominated by evergreen trees, sometimes ascending to wet, mossy, subalpine places. 9. Epilobium parviflorum Schreb. E small-flowered hairy willow-herb. VT. Fields, roadsides, clearings, ditches.

Fig. 740  Underground turions of Epilobium ciliatum var. glandulosum.

68 4   tricolpate s

10. Epilobium strictum Muhl. ex Spreng. N Fig. 741 downy willow-herb. Epilobium densum Raf. • CT, MA, ME, NH, RI, VT. Bogs, fens, swamps, peaty meadows, marshes, shorelines.

Ludwigia References: Munz (1944), Peng et al. (2005). 1a. Leaves opposite, the blades 5–30 mm long, abruptly narrowed to a broad petiole; flowering stems prostrate or creeping, often rooting at the nodes . . . . . . . . . . . . . . L. palustris 1b. Leaves alternate, the principal blades 40–105 mm long, gradually tapering to the stem or a subpetiolar base; stems upright 2a. Flowers borne on pedicels (2–) 3–5 mm long, with conspicuous yellow petals 5–7 mm long; roots usually tuberous-thickened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. alternifolia Fig. 741  Stem and leaves of Epilobium strictum, which are pubescent with spreading hairs.

2b. Flowers sessile or subsessile, lacking petals or sometimes with minute, green petals; roots not thickened 3a. Fruit 4–7 mm long, longer than wide, glabrous; bracteoles 2–5 mm long, attached at or above the base of the fruit [Fig. 742]; branches wing-angled . . . . . . . . . . . L. polycarpa 3b. Fruit 2.5–4 mm long, subglobose, pubescent; bracteoles wanting or shorter than 1 mm, when present attached at the base of or below the fruit; branches scarcely, if at all, wing-angled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sphaerocarpa 1. Ludwigia alternifolia L. N square-pod water-primrose. Ludwigia alternifolia L. var. linearifolia Britt.; L. alternifolia L. var. pubescens Palmer & Steyermark; L. alternifolia L. var. typica Munz • CT, MA, RI. River shores, lake shores, marshes, low fields. 2. Ludwigia palustris (L.) Ell. N common water-primrose. Isnardia palustris L.; Ludwigia palustris (L.) Ell. var. americana (DC.) Fern. & Grisc.; L. palustris (L.) Ell. var. nana Fern. & Grisc. • CT, MA, ME, NH, RI, VT. Shorelines, stream beds, wet ditches, swamps, frequently found on wet silt or mud or in shallow water. ‌ × Ludwigia brevipes (Long) Eames. Ludwigia ×lacustris Eames is a rare water2 primrose hybrid known from CT, RI. It is most likely to be confused with L. palustris due to its overall similarity (including opposite leaves). This hybrid has shortly pedicellate flowers (pedicels 1–2.5 mm long) with sepals 2.5–3 mm long and petals 1.3–3 mm long (rarely the petals absent). Ludwigia palustris is characterized by sessile flowers with sepals 1.1–2 mm long and lacking petals. Also, L. palustris has a longitudinal green band in the center of each of the four faces of the capsule. This feature is lacking in the hybrid. 3. Ludwigia polycarpa Short & Peter

N C Fig. 742

many-fruited water-primrose. CT, MA, ME, VT. Pond shores, rivers shores and accretion bars, pools in riparian forests. 4. Ludwigia sphaerocarpa Ell.

Fig. 742  Capsule of Ludwigia polycarpa with bracteoles positioned at or above the base of the fruit.

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round-pod water-primrose. Ludwigia sphaerocarpa Ell. var. deamii Fern. & Grisc.; L. sphaerocarpa Ell. var. jungens Fern. & Grisc.; L. sphaerocarpa Ell. var. macrocarpa • CT, MA, RI. River shores, pond shores, including coastal plain types, marshes.

Oenothera The circumscription of Oenothera has broadened in recent years by the inclusion of Calylophus and Gaura (in our area) due to phylogenetic evidence that shows these genera to be nested within Oenothera (Levin et al. 2004). References: Raven and Gregory (1972), Straley (1977), Dietrich et al. (1997).

On ag r ac e a e   6 8 5

1a. Flowers zygomorphic, the petals all oriented toward one side of the flower [Fig. 744]; fruit indehiscent, with 2–6 seeds 1.25–3.5 mm long 2a. Sepals 2–3.5 mm long; petals 1.5–3 mm long; anthers 0.5–1 mm long . . . . . O. curtiflora 2b. Sepals 6–18 mm long; petals 6–15 mm long; anthers 1.5–5 mm long 3a. Stems densely villous; inflorescence villous and stipitate-glandular; leaf blades short-villous along veins and margins; bracts 1–2.25 × 0.5–1 mm . . . . . . . . . . . . . . O. gaura 3b. Stems densely strigulose, often with a mixture of glandular or villous pubescence as well; inflorescence densely strigulose, hirtellous, and/or stipitate-glandular; leaf blades, at least in part, strigulose; bracts 1–6 × 0.5–2 mm . . . . . . . . . . . . . . . . . . . . . . . . O. filiformis 1b. Flowers actinomorphic [Fig. 745]; fruit dehiscent, with many seeds 0.3–2 mm long 4a. Ovary quadrangular in cross-section (at least in the apical portion); capsules sharply quadrangular to 4-winged, tapering to the base 5a. Petals 3–10 mm long; style 3–10 mm long; anthers 1.5–2.5 mm long; inflorescence of scattered flowers, the axis and apex commonly drooping in bud . . . . . . . . . . . O. perennis 5b. Petals 10–25 mm long; style 10–20 mm long; anthers 4–8 mm long; inflorescence relatively more compact, the axis and apex commonly erect in bud 6a. Sepal appendages mostly shorter than 1 (–1.4) mm and erect; capsules clavate to obpyramidal in outline, stipitate; plants subglabrous to sparsely pubescent with hairs mostly shorter than 1 (–1.5) mm; hypanthium 5–15 (–17) mm long . . . . . . . . O. fruticosa 6b. Sepal appendages mostly 1–4 mm long and divergent; capsules linear-elliptic, elliptic, to narrow-clavate in outline, sessile or short-stipitate; plants conspicuously pubescent with hairs mostly 1–3 mm long; hypanthium 15–25 mm long . . . . . O. pilosella 4b. Ovary ± terete in cross-section; capsules terete to bluntly quadrangular, abruptly rounded to truncate at the base 7a. Stigmas peltate with 4 short, rounded lobes, stigmatic on the upper and inside surfaces; hypanthium funnel-shaped, quadrangular in cross-section; calyx lobes prominently keeled; stamens alternately of different size . . . . . . . . . . . . . . . . O. serrulata 7b. Stigmas with 4 linear lobes [Fig. 745], stigmatic all around; hypanthium tubular terete in cross-section; calyx lobes scarcely or not at all keeled; stamens all of similar size 8a. Leaf and bract blades prominently dentate to pinnatifid; capsules not tapering to the apex, essentially of uniform width; ovules and seeds vertically oriented in the locules, the seeds without angular faces, conspicuously pitted; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. laciniata 8b. Leaf and bract blades entire to dentate; capsules slightly tapering to the apex, widest at the base; ovules and seeds horizontally oriented in the locules, the seeds with angular faces, not pitted; plants biennial or short-lived perennial 9a. Petals (25–) 30–45 mm long; sepal appendages (2–) 5–9 mm long; anthers 10–15 mm long; stigmas exserted beyond the anthers [Fig. 745] 10a. Apical half of stem, hypanthium, sepals, and ovary conspicuously pubescent, some of the hairs red and pustulose-based; bracts of inflorescence green and persistent into fruit; sepals yellow-green and usually tinged with red or striped with red (sometimes entirely dark red) . . . . . . . . . . . . . . O. glazioviana 10b. Apical half of stem, hypanthium, sepals, and ovary glabrous or appearing so without magnification, pustulose hairs (when rarely present) translucent; bracts of inflorescence sometimes pale green and deciduous; sepals yellowgreen or sometimes tinged with red . . . . . . . . . . . . . . . . . . . . . . . . . O. grandiflora

68 6   tricolpate s

9b. Petals 7–25 mm long; sepal appendages 0.5–6 mm long; anthers 3–10 mm long; stigmas surrounded by or shorter than the anthers 11a. Sepal appendages subterminal, separate in bud, the true apex represented by a small lobe [Fig. 746]; inflorescence sometimes curved near the apex 12a. Plants densely and minutely strigose, sometimes with a few spreading, pustulose hairs on the stem or glandular hairs in the inflorescence; seeds 1.1–1.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. oakesiana 12b. Plants variously pubescent with pustulose-based hairs, gland-tipped hairs, and/or sparse, minute, strigose hairs, or nearly glabrous; seeds 1.1–1.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. parviflora 11b. Sepal appendages terminal, closely connivent in bud [Fig. 743]; inflorescence usually straight and erect 13a. Plants densely and minutely strigose, sometimes with a few spreading, pustulose hairs on the stem or glandular hairs in the inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. villosa 13b. Plants variously pubescent with pustulose-based hairs, gland-tipped hairs, and/or sparse, minute, strigose hairs, or nearly glabrous 14a. Axis of the inflorescence definitely pubescent; bracts of the inflorescence 12–50 × 8–25 mm, green, deciduous . . . . . . . . . . . O. biennis 14b. Axis of the inflorescence glabrous or usually appearing so without magnification; bracts of the inflorescence 10–25 × 2–8 mm, pale green, caducous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. nutans 1. Oenothera biennis L. N Fig. 743 common evening-primrose. Oenothera biennis L. ssp. centralis Munz; O. biennis L. var. pycnocarpa (Atkinson & Bartlett) Wieg.; O. muricata L.; O. pycnocarpa Atkinson & Bartlett • CT, MA, ME, NH, RI, VT; throughout. Fields, roadsides, lawns, river banks, clearings, railroads. 2. Oenothera curtiflora W.L. Wagner & Hoch E

Fig. 743  Flower of Oenothera biennis with terminal sepal appendages.

small-flowered bee-blossom. Gaura mollis James.; G. parviflora Dougl. ex Lehm.; G. parviflora Dougl. ex Lehm. var. lachnocarpa Weatherby; G. parviflora Dougl. ex Lehm. var. typica Munz • MA. Fields, roadsides, waste areas. 3. Oenothera filiformis (Small) W.L. Wagner & Hoch E bee-blossom. Gaura biennis L. var. pitcheri Torr. & Gray; G. filiformis Small; G. longiflora Spach • CT, MA. Fields, roadsides, waste areas, railroads.

4. Oenothera fruticosa L.

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narrow-leaved evening-primrose.  4a. Kneiffia arenicola Small; K. fruticosa (L.) Raimann; K. linearis (Michx.) Spach; Oenothera arenicola (Small) Coker; O. fruticosa L. var. eamesii (B.L. Robins.) Blake; O. fruticosa L. var. linearis (Michx.) S. Wats.; O. fruticosa L. var. microcarpa Fern.; O. tetragona Roth var. brevistipata (Pennell) Munz; O. tetragona Roth var. longistipata (Pennell) Munz; O. tetragona Roth var. riparia (Nutt.) Munz; O. tetragona Roth var. velutina (Pennell) Munz; 4b. Kneiffia glauca (Michx.) Spach; K. latifolia Rydb.; K. tetragona (Roth) Pennell; Oenothera tetragona Roth; O. tetragona Roth ssp. glauca (Michx.) Munz; O. tetragona Roth var. hybrida (Michx.) Fern.; O. tetragona Roth var. latifolia (Rydb.) Fern. • CT, MA, ME, RI. Edges and high areas in saline marshes, shores of brackish ponds, brackish-tidal river shores, fields, roadsides, meadows, eskers, waste areas. 1a. Pubescence of ovary mostly or entirely of eglandular hairs; capsules widest above the middle; leaf blades pubescent and often subentire . . . . . . . . . 4a. O. fruticosa ssp. fruticosa 1b. Pubescence of ovary mostly or entirely of glandular hairs; capsules widest near middle; leaf blades often subglabrous and obscurely dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. O. fruticosa ssp. glauca (Michx.) Straley

On ag r ac e a e   6 87

Subspecies fruticosa is known from CT, MA; also reported from RI by George (1992), but specimens are unknown. Subspecies glauca is known from CT, MA, ME, NH, RI. Subspecies glauca is more often found in dry, open, sandy, inland habitats compared with subspecies fruticosa, which is commonly found in tidal habitats. 5. Oenothera gaura W.L. Wagner & Hoch E Fig. 744 biennial bee-blossom. Gaura biennis L. • CT, MA, VT. Fields, roadsides, waste areas, railroads. Oenothera gaura is a self-compatible species that was probably derived from Oenothera filiformis (a self-incompatible species). This likely explains their close morphological similarity. Oenothera gaura completely lacks tiny strigae on various surfaces (a type of hair possessed by O. filiformis).

Fig. 744  Flower of Oenothera gaura.

6. Oenothera glazioviana Micheli in Mart. E Fig. 745 garden evening-primrose. Oenothera erythrosepala Borbás • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, gardens. Most previous reports of Oenothera grandiflora from New England were referable to this species. 7. Oenothera grandiflora L’Hér. in Ait. E large-flowered evening-primrose. Oenothera biennis L. var. grandiflora (L’Hér. in Ait.) Lindl.; Oenothera grandiflora (L’Hér. in Ait.) var. glabra Seringe in DC. • VT. Gardens, waste areas. This species has long been considered native to Europe by some authors and has been reported from throughout much of New England. However, it is actually native to the southeastern United States and is a very rare introduction. The introduced plant from Europe is Oenothera glazioviana, which is much more commonly planted in the region. 8. Oenothera laciniata Hill E

Fig. 745  Flower of Oenothera glazioviana showing stigmas positioned above the anthers.

cut-leaved evening-primrose. Raimannia laciniata (Hill) Rose • CT, MA, ME, VT. Fields, roadsides, waste areas. 9. Oenothera nutans Atkinson & Bartlett

NC

nodding evening-primrose. Oenothera austromontana (Munz) Raven, W. Dietr., & Stubbe; O. biennis L. ssp. austromontana Munz; O. biennis L. var. austromontana (Munz) Cronq., O. biennis L. var. nutans (Atkinson & Bartlett) Wieg. • CT, MA, ME, NH, VT. Fields, roadsides, river banks, forest borders. 10. Oenothera oakesiana (Gray) J.W. Robbins ex S. Wats. & Coult. N Oakes’ evening-primrose. Oenothera angustissima R.R. Gates; O. biennis L. var. oakesiana Gray; O. cruciata Nutt. ex G. Don var. stenopetala (Bickn.) Fern.; O. parviflora L. ssp. angustissima (R.R. Gates) Munz; O. parviflora L. var. angustissima (R.R. Gates) Wieg.; O. parviflora L. var. oakesiana (Gray) Fern. • CT, MA, ME, NH, RI, VT; most common on the coastal plain. Fields, roadsides, railroads, coastal dunes, upper beaches, and open areas. 11. Oenothera parviflora L. N Fig. 746 small-flowered evening-primrose. Oenothera cruciata Nutt. ex G. Don; O. cruciata Nutt. ex G. Don var. sabulonensis Fern.; O. rubricapitata R.R. Gates • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, river banks, railroads. 12. Oenothera perennis L. N little evening-primrose. Kneiffia perennis (L.) Pennell; K. pumila (L.) Spach; Oenothera pumila L. • CT, MA, ME, NH, RI, VT; throughout. Fields, roadsides, waste areas, meadows, clearings. 13. Oenothera pilosella Raf. ssp. pilosella E meadow evening-primrose. Kneiffia pratensis Small • CT, MA, ME, NH, VT. Fields, roadsides, waste areas, gardens. 14. Oenothera serrulata Nutt. E yellow evening-primrose. Calylophus serrulatus (Nutt.) Raven; Meriolix serrulata (Nutt.) Walp.; Oenothera serrulata Nutt. var. typica Munz • VT. Railroads and other dry, open places.

Fig. 746  Flower of Oenothera parviflora with subterminal sepal appendages.

68 8   tricolpate s

15. Oenothera villosa Thunb. ssp. villosa N hairy evening-primrose. Oenothera biennis L. var. canescens Torr. & Gray; O. canovirens Steele; O. strigosa (Rydb.) Mackenzie & Bush ssp. canovirens (Steele) Munz; O. villosa Thunb. ssp. canovirens (Steele) W. Dietr. & Raven • CT, MA, ME, NH, VT; most common on the coastal plain. Fields, roadsides, coastal dunes, beaches, and open areas.

Orobanchaceae 1a. Plants without chlorophyll, the stems yellow to brown or purple; leaf blades reduced to scales; ovary unilocular, with parietal placentation 2a. Stems branched, usually many times; flowers dimorphic—the upper with a welldeveloped corolla and appearing bisexual, the lower carpellate with a small and unopening corolla that is forced off by the developing ovary; plants parasitic on Fagus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epifagus 2b. Stems simple or rarely few-branched; flowers monomorphic; plants parasitic on various species of plants 3a. Calyx with 5 subequal lobes or with 2 lateral lobes separated by a deep apical and basal sinus and each lobe usually bifurcate (therefore, the calyx usually 4 lobed); stems either subterranean or above-ground and sparsely leafy . . . . . . . . . . . . . . . . . . Orobanche 3b. Calyx tubular and irregularly toothed, split by a deep basal sinus only; stems aboveground and ± concealed by overlapping leaves . . . . . . . . . . . . . . . . . . . . . . . . . . Conopholis 1b. Plants with chlorophyll, with green stems and/or leaves in life; leaf blades foliaceous [Figs. 752, 754]; ovary bilocular, with axile placentation 4a. Foliage leaves of the stem alternate [Fig. 750]; anthers glabrous (sparsely pubescent near apex in Schwalbea) 5a. Flowers subtended by a pair of bractlets situated at the base of the calyx; calyx with 5 lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schwalbea 5b. Flowers without a pair of bractlets; calyx with 2 or 4 lobes (5 lobes in Pedicularis furbishiae) 6a. The two anther sacs on each filament similar; corolla 2 or more times as long as the calyx; bracts herbaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Pedicularis 6b. The two anther sacs on each filament dissimilar—one attached to the filament near its middle, the other suspended from near its apex; corolla less than 2 times as long as the calyx; bracts petaloid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Castilleja 4b. Foliage leaves of the stem opposite [Fig. 754]; anthers pubescent (glabrous in Odontites) 7a. Corolla weakly zygomorphic, not galeate [Figs. 747, 748], 10–50 mm long 8a. Corolla yellow (sometimes marked with brown or red-brown), 25–50 mm long; leaf blades evidently lobed; capsules ovoid to ellipsoid, acute at the apex; anther locules awned at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aureolaria 8b. Corolla pink to purple (rarely white), 10–20 mm long; leaf blades entire; capsules globose to subglobose, rounded at the apex (except sometimes for a mucro); anther locules obtuse to cuspidate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agalinis 7b. Corolla zygomorphic and galeate (i.e., with an upper, hooded lip that invests the stamens) [Figs. 753, 754], 2.5–14 mm long 9a. Leaf and bract blades palmately veined; corolla white, blue, or purple, usually with darker purple lines, the sides of the upper lip reflexed . . . . . . . . . . . . . . Euphrasia

Oro b an c h ac e a e   6 8 9

9b. Leaf and bract blades pinnately veined; corolla white, yellow, or red, the sides not reflexed 10a. Corolla yellow; calyx laterally compressed (i.e., taller than thick), with 2 prominent lateral lobes separated by a deep apical and basal sinus (each lateral lobe bifurcate in Rhinanthus) [Fig. 754] 11a. Each lateral half of the calyx bifurcate (i.e., with 4 total lobes), without foliaceous appendages; calyx accrescent in fruit and becoming very inflated; fruit symmetrical, dehiscing evenly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhinanthus 11b. Lateral halves of calyx unlobed, tipped by a foliaceous appendage; calyx not accrescent in fruit; fruit asymmetrical, dehiscing only or more prominently on the upper side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Pedicularis 10b. Corolla red or both white and yellow; calyx not laterally compressed, subequally 4-lobed 12a. Corolla red, pubescent; anthers glabrous; capsule shorter than the persistent calyx, with more than 4 seeds . . . . . . . . . . . . . . . . . . . . . . . . . Odontites 12b. Corolla white with a yellow palate, glabrous; anthers pubescent; capsule longer than the persistent calyx, with 1–4 seeds . . . . . . . . . . . . . . . Melampyrum

Agalinis Reference: Pennell (1929). 1a. Corolla pink, becoming pale in drying; calyx tube reticulate-veined [Fig. 747]; plants persistently yellow-green, even in drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. acuta 1b. Corolla purple to pink-purple (rarely white); calyx tube scarcely, if at all, veiny [Fig. 748]; plants generally dark green and often purple-tinged in life, darkening in drying 2a. Lobes of the calyx oblong or semiorbicular, obtuse to rounded at the apex [Fig. 748]; anther cells obtuse to subacute at the base; plants fleshy, of coastal saltmarshes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. maritima 2b. Lobes of the calyx triangular to subulate, acute to acuminate at the apex; anther cells mucronate to caudate at the base; plants not fleshy, predominantly not of coastal saltmarshes (except sometimes in A. purpurea) 3a. Lobes of the calyx 1.8–5 mm long, 0.75–2 times as long as the basal, connate portion; pollen sacs ca. 1 mm long; style 4–5 mm long . . . . . . . . . . . . . . . . . . A. neoscotica 3b. Lobes of the calyx 0.3–3.5 mm long, 0.2–1 times as long as the basal, connate portion; pollen sacs 1.4–2.2 mm long; style 8–14 mm long 4a. Pedicels 10–20 mm long at anthesis; corolla glabrous on the adaxial (i.e., inside) surface, the upper lip arched forward over the stamens . . . . . . . . . . . . . . . A. tenuifolia 4b. Pedicels 1–8 mm long at anthesis; corolla pubescent on the adaxial surface (i.e., inside), the lobes of the upper lip reflexed-spreading 5a. Corolla 10–20 (–23) mm long; calyx lobes 2–3.5 mm long, 0.4–1 times as long as the basal, connate portion; style 6–10 mm long . . . . . . . . . . . . . . . . A. paupercula 5b. Corolla 20–38 mm long; calyx lobes 0.5–2 mm long, up to 0.5 times as long as the basal, connate portion; style 15–20 mm long . . . . . . . . . . . . . . . . . . . . A. purpurea 1. Agalinis acuta Pennell

N C Fig. 747

sandplain agalinis. Gerardia acuta (Pennell) Pennell • CT, MA, RI. Sandy fields, roadsides, grasslands, and barrens. The report of Agalinis obtusifolia Raf. in CT by Graves et al. (1910; as Gerardia parvifolia) was referable to this species.

Fig. 747  Flower of Agalinis acuta showing reticulate-veined calyx.

69 0   tricolpate s

2. Agalinis maritima (Raf.) Raf. ssp. maritima N Fig. 748 saltmarsh agalinis. Gerardia maritima Raf. • CT, MA, ME, NH, RI. Saline marshes. 3. Agalinis neoscotica (Greene) Fern.

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Nova Scotia agalinis. Agalinis paupercula (Gray) Britt. var. neoscotica (Greene) Pennell & St. John; Agalinis purpurea L. var. neoscotica (Greene) Boivin; Gerardia neoscotica Greene; Gerardia purpurea L. var. neoscotica (Greene) Gleason • ME; southeastern portion of state. Sandy and peaty soils of headlands and near-coastal areas, edges of tidal marshes and boggy areas. Fig. 748  Flower of Agalinis maritima showing blunt calyx lobes.

4. Agalinis paupercula (Gray) Britt. N small-flowered agalinis.  4a. Agalinis purpurea (L.) Pennell var. parviflora (Benth.) Boivin; Gerardia paupercula (Gray) Britt. ssp. borealis (Pennell) Pennell; G. paupercula (Gray) Britt. var. borealis (Pennell) Deam; G. purpurea L. var. parviflora Benth.; 4b. Gerardia paupercula (Gray) Britt.; G. paupercula (Gray) Britt. var. typica Pennell; G. purpurea L. var. paupercula Gray • CT, MA, ME, NH, RI, VT. Fields, roadsides, open rights-of-way, clearings, trail edges, meadows, shorelines, edges of temporary pools. 1a. Anthers sparsely pubescent with brown or white hairs; corolla 10–17 mm long, with ascending to ascending-spreading lobes; style 6–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a. A. paupercula var. borealis Pennell 1b. Anthers densely pubescent with white hairs; corolla 15–20 (–23) mm long, with spreading lobes; style 7–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. A. paupercula var. paupercula Variety borealis is known from ME, NH, VT. Variety paupercula is known from CT, MA, ME, NH, RI, VT. 5. Agalinis purpurea (L.) Pennell N purple agalinis. Gerardia purpurea L.; G. purpurea L. ssp. parvula Pennell; G. purpurea L. var. grandiflora Benth. • CT, MA, ME, RI; also reported from NH by Magee and Ahles (1999), but specimens are unknown; mainly along the coastal plain, less common inland. Low fields and meadows near the coast, tidal marsh borders, dune hollows. 6. Agalinis tenuifolia (Vahl) Raf.

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slender-leaved agalinis.  6a. Gerardia tenuifolia Vahl; G. tenuifolia Vahl var. humilis Benth.; G. tenuifolia Vahl var. typica Pennell; 6b. Agalinis besseyana (Britt.) Britt.; Gerardia besseyana Britt.; G. tenuifolia Vahl ssp. macrophylla (Benth.) Pennell; G. tenuifolia Vahl var. macrophylla Benth.; 6c. Gerardia tenuifolia Vahl ssp. parviflora (Nutt.) Pennell; G. tenuifolia Vahl var. parviflora Nutt. • CT, MA, ME, NH, RI, VT. Fields, roadsides, trail edges, woodlands, clearings, open rights-of-way. 1a. Calyx lobes 0.2–1.3 (–1.5) mm long; capsules 3–4 (–5) mm long; seeds 0.6–0.9 mm long, with very fine reticulations, the seed surface relatively smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6a. A. tenuifolia var. tenuifolia 1b. Calyx lobes (0.5–) 1–2 mm long; capsules (4–) 5–7 mm long; seeds 0.7–1.3 mm long, with more pronounced reticulations, causing the seed surface to appear rough 2a. Axillary fascicles not or only slightly developed; filaments pubescent; anthers densely pubescent; leaves and branches spreading; leaf blades slightly scabrous on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6b. A. tenuifolia var. macrophylla (Benth.) Blake 2b. Axillary fascicles usually well-developed; filaments sparsely pubescent to glabrous; anthers sparsely to moderately pubescent; leaves and branches ascending; leaf blades slightly to conspicuously scabrous on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6c. A. tenuifolia var. parviflora (Nutt.) Pennell Variety tenuifolia is known from CT, MA, ME, NH, RI, VT. Variety macrophylla is known from CT. Variety parviflora is known from MA, VT. Both varieties macrophylla and parviflora are of conservation concern.

O ro b an c h ac e a e   6 9 1

Aureolaria Species of Aureolaria are parasitic on the roots of various species of Quercus. Reference: Pennell (1928). 1a. Corolla at least sparingly glandular-pubescent on the abaxial (i.e., outside) surface [Fig. 749]; sepals crenate to pinnately lobed [Fig. 749]; capsules glandular-pubescent; seeds 0.8–1 mm long, wingless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pedicularia 1b. Corolla not glandular on abaxial surface; sepals entire (very rarely lobed in A. virginica); capsules glabrous or pubescent with eglandular hairs; seeds 1.5–2.7 mm long, each with several thin wings; plants perennial 2a. Stem pubescent; capsules densely pubescent with red-brown hairs; pedicels 1–3 mm long; seeds 1.5–1.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. virginica 2b. Stem glabrous and glaucous; capsules glabrous; pedicels 4–10 mm long; seeds 2–2.7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. flava 1. Aureolaria flava (L.) Farw. var. flava N smooth false foxglove. Agalinis flava (L.) Boivin; Aureolaria flava (L.) Farw. ssp. typica Pennell; Dasistoma flava (L.) Wood; Gerardia flava L. • CT, MA, ME, NH, RI, VT. Woodlands, rocky forests, dry banks, sandy roadsides, ridges. 2. Aureolaria pedicularia (L.) Raf.

N C Fig. 749

fern-leaved false foxglove.  2a. Aureolaria pedicularia (L.) Raf. ssp. intercedens (Pennell) Pennell; Gerardia pedicularia L. var. intercedens (Pennell) Fern.; 2b. Agalinis pedicularia (L.) Blake; A. pedicularia (L.) Blake var. caesariensis (Pennell) Blake; Aureolaria pedicularia (L.) Raf. ssp. caesariensis Pennell; A. pedicularia (L.) Raf. ssp. carolinensis Pennell; A. pedicularia (L.) Raf. var. carolinensis (Pennell) Pennell; A. pedicularia ssp. typica Pennell; Dasistoma pedicularia (L.) Benth.; Gerardia pedicularia L. • CT, MA, ME, NH, RI, VT. Woodlands, rocky forests, ridges, balds, pine barrens. 1a. Upper portion of stem and leaf blades with abundant glandular hairs; capsules 11–13 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a. A. pedicularia var. intercedens Pennell 1b. Upper portion of stem and leaf blades with few or no glandular hairs; capsules 10–11 (–12) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. A. pedicularia var. pedicularia Variety intercedens is known from MA, NH. It is of conservation concern. Variety pedicularia is known from CT, MA, ME, NH, RI, VT. 3. Aureolaria virginica (L.) Pennell N downy false foxglove. Agalinis virginica (L.) Blake; Dasistoma virginica (L.) Britt.; Gerardia virginica (L.) B.S.P.; Rhinanthus virginicus L. • CT, MA, NH, RI, VT. Woodlands, ridges.

Castilleja Reference: Chuang and Heckard (1991). 1a. Foliage leaf blades entire [Fig. 750]; bracts entire or 1- to 3-toothed [Fig. 750], white to yellow (rarely purple-tinged); plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . C. septentrionalis 1b. Foliage leaf blades 3- to 9-cleft or infrequently entire; bracts 3- to 9-lobed, red, purple, or yellow (rarely white); plants annual or biennial 2a. Upper lip of corolla (i.e., the galea) more than 3 times as long as the lower lip; lower lip of corolla lacking pouches, tipped with 3 highly reduced teeth that are incurved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. coccinea 2b. Upper lip of corolla much less than 3 times as long as the lower lip; lower lip of corolla evidently pouched, tipped by 3 small, but evident, erect, petaloid lobes . . . . . . . . C. exserta

Fig. 749  Flower of Aureolaria pedicularia.

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1. Castilleja coccinea (L.) Spreng.

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scarlet painted-cup. Bartsia coccinea L. • CT, MA, ME, NH, RI. Wet-mesic to hydric meadows, swamps, boggy areas; slopes with seasonal seepage, open rights-of-way, often on higher pH substrate in CT. 2. Castilleja exserta (Heller) Chuang & Heckard ssp. exserta E purple painted-cup. Orthocarpus exsertus Heller; O. purpurascens Benth.; O. purpurascens Benth. var. pallidus Keck • MA. Wool waste. 3. Castilleja septentrionalis Lindl.

N C Fig. 750

northern painted-cup. Castilleja pallida (L.) Spreng. ssp. septentrionalis (Lindl.) Scoggan; C. pallida (L.) Spreng. var. septentrionalis (Lindl.) Gray • ME, NH, VT; northern portion of states. Alpine ravines, plateaus, and snowbank communities, ice-scoured river shores in regions of high-pH bedrock and/or till.

Conopholis 1. Conopholis americana (L.) Wallr. f. N Fig. 750  Inflorescence of Castilleja septentrionalis.

American squaw-root. Orobanche americana L. • CT, MA, ME, NH, RI, VT. Deciduous forests. This plant is parasitic on several different species of the genus Quercus.

Epifagus 1. Epifagus virginiana (L.) W. Bart. N beech-drops. Leptamnium virginianum (L.) Raf. • CT, MA, ME, NH, RI, VT. Deciduous forests. This species is parasitic on the roots of Fagus grandifolia. The upper flowers of this species, though appearing bisexual, are partially abortive and functionally staminate.

Euphrasia Individuals of Euphrasia are known to vary morphologically according to differences in nutrient supply, length of growing season, and height of surrounding vegetation (Downie and McNeill 1990). Therefore, some characters used to discriminate species of Euphrasia (e.g., internode length, number of nodes to first flower, number of branches), though generally useful, can show tremendous variation and should be confirmed through use of additional characters. Sell and Yeo (1970) noted specimens from the Plantae Exsiccatae ‌aequalis Callen (E. nemorosa × Grayanae (number 62) that may be referable to Euphrasia × E. tetraquetra) from Frenchville, ME. Reference: Sell and Yeo (1970). 1a. Bracts with subulate or bristle-tipped teeth [Figs. 751, 752]; corolla 5–8 mm long 2a. Bracts erect to ascending, cuneate at the base, with ascending teeth [Fig. 752] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. stricta 2b. Bracts ascending to spreading, rounded to truncate at the base, with spreading teeth [Fig. 751] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. nemorosa 1b. Bracts with obtuse to acute teeth; corolla 2.5–5 (–7.5) mm long 3a. Corolla 5–7.5 mm long 4a. Corolla white; stem leaf blades suborbicular to rhombic or obovate; capsules (4–) 5–6.5 mm long; floral leaf blades 5–15 mm long; flowering beginning from the 5th to the 9th node (rarely as low as the 2nd node) . . . . . . . . . . . . . . . . . . . . . . . E. suborbicularis 4b. Corolla light purple; stem leaf blades ovate (rarely elliptic); capsules 4–5.5 mm long; floral leaf blades 4–8 mm long; flowering beginning from the 7th to the 11th node . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. micrantha

O ro b a n ch ac e a e   6 9 3

3b. Corolla 2.5–5 mm long 5a. Calyx lobes obtuse at the apex and falcately curved; stems simple or rarely with 1 or 2 branches from the upper nodes; flowering beginning from the 4th or 5th node . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. oakesii 5b. Calyx lobes acute at the apex and ± straight; stems usually with 1–4 (–5) branches from the middle and upper (rarely the lower) nodes or rarely simple; flowering beginning from the 3rd to the 8th node (rarely as high as the 10th node) 6a. Corolla 4–5 mm long, white, sometimes with a light purple upper lip; stem internodes shorter than to slightly longer than the associated leaves; capsules truncate at apex, not reaching tips of calyx lobes . . . . . . . . . . . . . . . . . . . E. tetraquetra 6b. Corolla 2.5–4.5 mm long, light purple to purple (rarely white); stem internodes 1–3 times as long as the associated leaves; capsules retuse or emarginate at apex, as long as or exserted beyond the tips of the calyx lobes . . . . . . . . . . . . . . . . . . . . E. randii 1. Euphrasia micrantha Reichenb. E northern eyebright. MA; western portion of state. Lawns, roadsides, disturbed soil. See Hermann (1958) for the discovery of this species in North America. 2. Euphrasia nemorosa (Pers.) Wallr. N Fig. 751 common eyebright. Euphrasia americana Wettst.; E. canadensis, sensu Fernald; E. officinalis L. var. nemorosa Pers. • MA, ME, NH, VT; also reported from CT by Dowhan (1979), but specimens are unknown. Fields, roadsides, clearings, coastal headlands and open areas. 3. Euphrasia oakesii Wettst.

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Oakes’ eyebright. Euphrasia williamsii B.L. Robins.; Euphrasia williamsii B.L. Robins. var. vestita Fern. & Wieg. • ME, NH; northern portion of states. Alpine ridges, plateaus, and snowbank communities. Two corolla color forms are found in the region—pale violet with darker purple lines (the “oakesii” form) and brown to purple-red (the “williamsii” form).

Fig. 751  Flowers and bracts of Euphrasia nemorosa.

4. Euphrasia randii B. L. Robins. N Rand’s eyebright. Euphrasia purpurea Reeks; E. randii L. var. farlowii B.L. Robins.; E. randii L. var. reeksii Fern. • ME; coastal regions. Atlantic coast turfs, headlands, fields, and open areas. 5. Euphrasia stricta D. Wolff ex J.F. Lehm. E Fig. 752 strict eyebright. Euphrasia brevipila Burn. & Gremli ex Gremli; E. officinalis, auct. non L.; E. rigidula Jord. • MA, ME, ri, vt; also reported from NH by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, waste areas. 6. Euphrasia suborbicularis Sell & Yeo

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round-leaved eyebright. Euphrasia mollis (Ledeb.) Wettst. var. laurentiana Boivin • ME. Ice-scoured river shores in regions of high-pH bedrock and/or till. Though Sell and Yeo (1970) did not explicitly state that this species occurs in New England, they did note that most eastern North American specimens considered to be Euphrasia disjuncta by Fernald and Wiegand (1915) are this species. Further, Sell and Yeo restricted E. disjucta to Newfoundland and Lake Mistassini (Quebec), while E. suborbicularis extends south to the Gaspé Pennisula (Quebec) and Nova Scotia. Given the proximity of the St. John River, where the plant in question was collected in the 1860s, to the Gaspé, it seems likely our plants are E. suborbicularis. Unfortunately, the specimen was apparently lost in a fire in 1866 (Eastman 1981). This is probably the reason that Sell and Yeo did not attribute E. suborbicularis to Maine. 7. Euphrasia tetraquetra (Brébiss.) Arrond. E seacliff eyebright. Euphrasia canadensis Townsend • ME. Open, grassy and turfy areas.

Fig. 752  Flowers and bracts of Euphrasia stricta.

69 4   tricolpate s

Melampyrum Species of Melampyrum exhibit a type of variation called pseudoseasonal polymorphism (de Soó and Webb 1972). This type of variation is correlated with both the time of germination and flowering, as well as the habitat the plant grows in. These variants demonstrate many different morphologies and create difficulty for recognizing true genetic differences (should any exist). Therefore, no infraspecific taxa are recognized here in our species until further study sorts out phenotypic variation from genetic variation. 1. Melampyrum lineare Desr. N American cow-wheat. Melampyrum latifolium (Bart.) Muhl. ex Britt.; M. lineare Desr. var. americanum (Michx.) Beauverd; M. lineare Desr. var. latifolium Bart.; M. lineare Desr. var. pectinatum (Pennell) Fern. • CT, MA, ME, NH, RI, VT. Deciduous to evergreen woodlands and forests, dry fields, sand plains, barrens, roadsides, balds.

Odontites 1. Odontites vernus (Bellardi) Dumort. ssp. serotinus (Dumort.) Corb. E Fig. 753 red false bartsia. Odontites rubra Gilib.; O. serotinus Dumort. • MA, ME, VT. Fields, roadsides, dry banks. Some collections of this species housed at the Harvard University Herbaria (e.g., GH, NEBC) are not easily assignable to this subspecies and may be O. vernus ssp. vernus, which has bracts 10–15 (–20) mm long that clearly exceed the flowers and ascending to suberect branches (vs. bracts 7–10 mm long that are shorter than the flowers and branches usually spreading in ssp. serotinus). More work is needed to verify if ssp. vernus is present in New England. Fig. 753  Inflorescence of Odontites vernus.

Orobanche 1a. Flowers 1–3 per plant, borne on elongate pedicels 5–25 cm long from near the ground; calyx 5-lobed, without closely subtending bracts; corolla 12–20 mm long, weakly zygomorphic, the 5 lobes subequal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. uniflora 1b. Flowers more than 3 per plant (usually more than 10), arranged in a spike, sessile; calyx divided nearly to the base into 2 lateral lobes, each lobe usually bifurcate (therefore, the calyx usually 4-lobed), subtended by a bract; corolla 10–15 mm long, obviously zygomorphic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. minor 1. Orobanche minor Sm. E lesser broom-rape. VT. Gardens. Parasitic on many different genera, including Trifolium. 2. Orobanche uniflora L. N one-flowered broom-rape. Aphyllon uniflorum (L.) Torr. & Gray; Orobanche uniflora L. var. purpurea (Heller) Achey; Thalesia uniflora (L.) Britt. • CT, MA, ME, NH, RI, VT. Forests, woodlands, fields, clearings. Parasitic on many different genera, including Solidago.

Pedicularis 1a. Stem leaves mostly opposite; apical portion of stem and axis of inflorescence glabrous or nearly so; calyx with 2 lateral lobes, each lobe tipped by a foliaceous appendage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lanceolata 1b. Stem leaves mostly alternate; apical portion of stem and axis of inflorescence villous; calyx either with 2 lateral lobes that lack foliaceous appendages or with 5 subequal lobes 2a. Calyx with 2 lateral lobes separated by a shallow apical and deep basal sinus; plants 15–40 cm tall; stems typically clustered, simple; capsules lanceoloid-obloid, ca. 2 times as long as the persistent calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. canadensis

Oro b a n c h ac e a e   6 9 5

2b. Calyx with 5 subequal lobes; plants 40–90 cm tall; stems solitary or sometimes paired, simple or with arching branches; capsules ovoid, barely exceeding the persistent calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. furbishiae 1. Pedicularis canadensis L. ssp. canadensis N forest lousewort. Pedicularis canadensis L. var. dobbsii Fern.; P. gladiata Michx. • CT, MA, ME, NH, RI, VT. Fields, clearings, forest openings, trail edges, woodlands, banks. 2. Pedicularis furbishiae S. Wats.

NC

Furbish’s lousewort. ME; northern portion of state. Ice-scoured river shores, meadows, and shrub thickets in high-pH bedrock and/or till regions. Endemic to the St. John River of ME and New Brunswick. 3. Pedicularis lanceolata Michx.

NC

swamp lousewort. Pedicularis pallida Nutt.; P. virginica Poir. • CT, MA. Swamps, stream banks, hydric meadows, fresh-tidal marshes, wet ditches.

Rhinanthus Rhinanthus serotinus (Schönh) Oborney was reported from ME by Hultén and Fries (1986), but specimens are unknown. 1a. Corolla (15–) 18-24 mm long, the upper lip with small lateral lobes 1.5–2.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. alectorolophus 1b. Corolla 9–15 mm long, the upper lip with small lateral lobes up to 1 mm long . . . . R. minor 1. Rhinanthus alectorolophus (Scop.) Pollich E European yellow-rattle. Rhinanthus major Ehrh. • MA. Fields. 2. Rhinanthus minor L.

n C Fig. 754

little yellow-rattle.  2a. Rhinanthus borealis (Sterneck) Druce; R. groenlandicus (Ostenf.) Chabert; R. minor L. ssp. borealis (Sterneck) A. Löve; R. oblongifolius Fern.; 2b. Rhinanthus crista-galli L., pro parte; R. crista-galli L. var. fallax (Wimmer & Grab.) Druce; R. stenophyllus (Schur) Schinz & Thellung • CT, MA, ME, NH, VT. Fields, roadsides, alpine ravines and slopes. 1a. Leaf blades narrow-oblong to oblong, obtuse to acute at the apex, 2.1–3.7 (–4.4) times as long as wide [Fig. 754]; lower wing margin of seed usually only somewhat corky-thickened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a. R. minor ssp. groenlandicus (Ostenf.) L. Neum. 1b. Leaf blades narrow-lanceolate to lanceolate, acute to acuminate at the apex, (3.5–) 3.8–8.3 times as long as wide; lower wing margin of seed usually conspicuously corky-thickened, the thickened portion clearly defined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. R. minor ssp. minor Subspecies groenlandicus is native and known from NH; also reported from CT, MA, ME, RI, VT by Kartesz (1999), but these reports were in error. It occurs above treeline in open, alpine situations and is of conservation concern. Subspecies minor is non-native and known from CT, MA, ME, NH, VT. It occurs below treeline, generally in human-disturbed or human-maintained places. This variety was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI and was unaware of any collections.

Schwalbea 1. Schwalbea americana L. chaff-seed. Schwalbea americana L. var. australis (Pennell) Reveal & Broome; S. australis Pennell • CT, MA. Woodlands, sandy fields, pastures, roadsides, and waste areas.

NC

Fig. 754  Inflorescence and leaves of Rhinanthus minor ssp. groenlandicus.

69 6   tricolpate s

Oxalidaceae Oxalis The caulescent, yellow-flowered species of Oxalis have a confusing taxonomic history due to misapplication of names in past works (see species discussions for further details). Further, hybridization has been noted between some species in North America (e.g., between O. corniculata and O. dillenii, between O. dillenii and O. florida). References: Eiten (1963), Nesom (2009b). 1a. Plants lacking leafy, aerial stems, the petioles and peduncles arising directly from rhizomes or a bulbous base; corolla largely or entirely white or pink-purple to purple, 10–20 mm long 2a. Inflorescence consisting of a solitary, peduncled flower; petals white with pink veins (rarely pink-purple throughout) [Fig. 756]; sepals without a callus tip; leaves and peduncles arising from slender rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. montana 2b. Inflorescence an umbel with (1–) 2–12 (–18) flowers; petals pink-purple to purple (rarely pink or white); sepals with a pair of orange tubercles at the apex (these confluent and appearing as 1 tubercle in O. violacea); leaves and peduncles arising from one or more bulbs 3a. Leaflets obcordate to obreniform, (5–) 8–15 (–20) mm long, with two marginal oxalate deposits, appearing as small dots on on each side of the base of the apical leaflet notch; plants usually arising from a single bulb, the bulb scales 3-nerved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. violacea 3b. Leaflets obtriangular to broad-obtriangular, 20–50 mm long, lacking marginal oxalate deposits; plants usually arising from a cluster of bulbs (rarely a single bulb), the bulb scales (3–) 5- to 7-nerved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. intermedia 1b. Plants with leafy stems, the petioles and peduncles arising from nodes on the stem; petals yellow, 4–10 mm long 4a. Plants uniformly and moderately to densely pubescent with short, appressed hairs (i.e., strigose) on the stems, branches, peduncles, and pedicels (rarely some hairs ascending); seeds brown with gray to white lines on the transverse ridges . . . . . . O. dillenii 4b. Plants glabrous or sparsely to densely pubescent with appressed, spreading, and/or retrorse hairs, if uniformly strigose then only on peduncles and pedicels; seeds brown or rarely with gray to white lines on the transverse ridges 5a. Stipules absent [Fig. 755, R image]; pubescence of the stem septate [Fig. 755, R]; stems solitary (rarely in pairs or trios) from a rhizome; cymes with (3–) 5–7 (–15) flowers; primary root usually not thicker than stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. stricta 5b. Stipules present, though small and inconspicuous [Fig. 755, L]; pubescence of the stem without septa [Fig. 755, L]; stems either clustered from a crown or taproot or solitary, without horizontal stems or with above-ground horizontal stems (i.e., stolons); cymes with 1–3 (–6) flowers; primary root usually thicker than stem 6a. Stems trailing and rooting at the nodes; stipules relatively well developed (though still small), with distinct margins, rectangular; capsule commonly retrosely pubescent throughout, sometimes with intermixed, spreading septate hairs (varying to glabrous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. corniculata 6b. Stems erect to decumbent, but not rooting at the nodes; stipules very reduced, without distinct margins, appearing as a small, semicircular swelling in the axil of the petiole; capsule glabrous or sparsely pubescent apically or along ridges with appressed hairs, lacking septate hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. florida

Ox a l i dac e a e   6 97

1. Oxalis corniculata L. E Fig. 755L creeping yellow wood sorrel. Acetosella corniculata (L.) Kuntze; Oxalis corniculata L. var. langloisii (Small) Wieg.; O. langloisii (Small) Fedde; O. repens Thunb.; Xanthoxalis corniculata (L.) Small; X. langloisii Small; X. repens (Thunb.) Moldenke • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Lawns, fields, gardens, nurseries, railroads, areas of habitation. 2. Oxalis dillenii Jacq. N slender yellow wood sorrel. Oxalis corniculata L. var. dillenii (Jacq.) Trel.; Xanthoxalis dillenii (Jacq.) Holub • CT, MA, ME, NH, RI, VT. Fields, lawns, roadsides, waste areas. Oxalis dillenii is variable in its growth habit. Extreme forms can approach O. corniculata in that the aerial stems can be horizontal and root at some nodes. However, O. dillenii has a different stem pubescence, different seed color, and has the reduced stipules of O. florida (see identification key), features that don’t vary ecologically. Oxalis dillenii was called O. stricta by Fernald (1950b).

Fig. 755  Pubescence and stipules of Oxalis corniculata (left) and O. stricta (right).

3. Oxalis florida Salisb. N flowering yellow wood sorrel. Oxalis brittoniae Small; O. filipes Small; O. florida Salisb. var. filipes (Small) Ahles; Xanthoxalis brittoniae (Small) Small; Xanthoxalis filipes (Small) Small; X. florida (Salisb.) Moldenke • CT, MA, ME, VT. Fields, lawns, roadsides, waste areas. Oxalis florida has been treated as an infraspecific taxon of O. dillenii; however, the two species differ in stem pubescence and seed color (see identification key), as well as habit and presence/ absence of pedicel bracteoles. Oxalis florida is usually solitary stemmed and commonly possesses bracteoles on the pedicels whereas O. dillenii is usually multistemmed (2–8 stems together) and lacks bracteoles on the pedicels. These differences are sufficient for specieslevel recognition, as followed by Nesom (2009). 4. Oxalis intermedia A. Rich. E West Indian wood sorrel. Ionoxalis intermedia (A. Rich.) Small • MA. Fields, gardens, lawns, roadsides, waste areas. 5. Oxalis montana Raf. N Fig. 756 northern wood sorrel. Oxalis acetosella, sensu Gleason & Cronquist (1991); Oxalis acetosella L. ssp. montana (Raf.) Hultén ex D. Löve • CT, MA, ME, NH, VT. North-temperate to boreal evergreen to mixed evergreen-deciduous forests, sometimes with extensive bryophyte ground cover, often associated Abies and Picea in northern New England and Tsuga in southern New England. 6. Oxalis stricta L. N Fig. 755R common yellow wood sorrel. Ceratoxalis cymosa (Small) Lunell; Oxalis cymosa Small; O. europaea Jord.; O. europaea Jord. var. bushii (Small) Wieg.; O. interior (Small) Fedde; O. stricta L. var. villicaulis (Wieg.) Farw.; Xanthoxalis bushii Small; X. cymosa (Small) Small; X. interior Small; X. stricta (L.) Small • CT, MA, ME, NH, RI, VT. Fields, lawns, roadsides, gardens, railroads, waste areas. 7. Oxalis violacea L.

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violet wood sorrel. Ionoxalis violacea (L.) Small; Oxalis violacea L. var. trichophora Fassett • CT, MA, RI, VT. Ridges and rocky slopes, usually associated with trap rock or marble bedrock under a canopy of Carya, Fraxinus, and/or Quercus.

Paeoniaceae Paeonia 1a. Margins of leaf blades minutely serrulate with cartilaginous papillae; carpels and follicles usually glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lactiflora

Fig. 756  Flower and leaves of Oxalis montana.

69 8   tricolpate s

1b. Leaf margins smooth, not minutely serrulate; carpels and follicles pubescent with white tomentum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. officinalis 1. Paeonia lactiflora Pallas E Chinese peony. Paeonia albiflora Pallas • VT. Fields, roadsides, gardens. 2. Paeonia officinalis L. E common peony. MA, VT. Fields, roadsides, gardens, abandoned homesteads.

Papaveraceae Reference: Mitchell (1983). 1a. Perianth zygomorphic, spurred or saccate at the base [Figs. 757, 758, 759]; plants with a watery latex; petals not crumpled in bud; stamens 6 per flower, the filaments diadelphous; leaf blades finely divided into narrow segments [Fig. 757] 2a. The 2 outer petals dissimilar, only 1 of which is spurred; flowers borne laterally on the pedicels [Fig. 757] 3a. Ovary narrow-cylindric; flowers 7–25 (–30) mm long; fruit dehiscent, (14–) 16– 45 (–50) mm long, with more than 1 seed, with a persistent style; seed with an aril 4a. Corolla pink to pink-purple with a yellow apex; inflorescence a panicle-like cyme, with minute, pale green bracts 1–2 mm long; capsules erect to ascending, 25–45 (–50) mm long [Fig. 757] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Capnoides 4b. Corolla pale yellow, bright yellow, or purple (rarely white, pink, red, or bluepurple); inflorescence a raceme, with conspicuous, usually foliaceous bracts 3–8 mm long; capsules spreading to drooping, (14–) 16–25 (–30) mm long . . . . . . . . . Corydalis 3b. Ovary subglobose; flowers 6–9 mm long; fruit indehiscent, 2.5–3 mm long, 1-seeded, with a deciduous style; seed without an aril . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fumaria 2b. The 2 outer petals similar and spurred (but the spurs short in Adlumia) [Figs. 758, 759]; flowers borne on pendulous pedicels [Figs. 758, 759] 5a. Plants climbing by means of tendril-like, reduced leaflets and their petiolules; corolla connate, except at the very apex, persistent, becoming spongy, enclosing the capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adlumia 5b. Plants not climbing; corolla connate only at the base, deciduous 6a. Stems scapose (i.e., without leaves); corolla 10–25 mm long, the inner petals of ± similar color as the outer petals; stems arising from bulbs or from a scaly rhizome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dicentra 6b. Stems leafy; corolla 30–50 mm long, the inner petals of different color from the outer petals (except in white-flowered forms); stems arising from a rhizome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lamprocapnos 1b. Perianth actinomorphic, without a nectary spur [Figs. 760, 761]; plants with a milky or colored latex (watery in Eschscholzia); petals often crumpled in bud (therefore, wrinkled on expansion); stamens ca. 16–100 or more per flower, the filaments distinct; leaf blades lobed to 1.5 times divided (finely divided in Eschscholzia) 7a. Leaves 1, basal, the blade ± orbicular in outline [Fig. 761]; corolla with 8–12 petals [Fig. 761]; latex red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sanguinaria 7b. Leaves several to many, borne on a stem, the blade not orbicular in outline [Fig. 760]; corolla with 0 or 4–6 petals [Fig. 760]; latex yellow or white

Pa pav e r ac e a e   6 9 9

8a. Foliage spiny; capsules prickly; flowers with (2–) 3 sepals and (4–) 6 petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argemone 8b. Foliage without spines; capsules unarmed; flowers with 2 sepals and 0 or 4 petals 9a. Sepals connate, shed together in flower; gynoecium with 4–8 distinct, linear stigmas; leaf blades finely pinnately or palmately divided into narrow segments; latex watery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eschscholzia 9b. Sepals distinct, shed separately in flower; stigmas either 2-lobed or connate and forming a 4- to 18-lobed disk; leaf blades lobed to 1.5 times divided; latex white or colored 10a. Petals variously colored, but never yellow; stigmas forming a 4- to 18-lobed disk; capsules dehiscing by 4–18 subapical pores; latex white . . . . . . . . . . . Papaver 10b. Petals absent or present and yellow to orange-yellow; stigmas 2-lobed; capsules with 2–4 valves that dehisce to the base; latex yellow or orange-yellow 11a. Flowers solitary on short peduncles, with petals 20–30 mm long; fruit 15–30 cm long at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glaucium 11b. Flowers arranged in umbels or panicles, without petals or with petals 4–12 mm long; fruit 0.8–5 cm long at maturity 12a. Corolla present, of 4 yellow petals; sepals sepaloid, green; leaf blades pinnately divided, the leaflets again lobed; flowers few, in an umbel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelidonium 12b. Corolla absent; sepals petaloid, ± white; leaf blades pinnately lobed; flowers numerous, in a large panicle . . . . . . . . . . . . . . . . . . . . . . . . . . . Macleaya

Adlumia 1. Adlumia fungosa (Ait.) Greene ex B.S.P. N Allegheny-vine. Fumaria fungosa Ait. • CT, MA, ME, NH, RI, VT. Rocky forests, cliff bases, gardens.

Argemone 1a. Petals white (rarely tinged with pink); androecium with 100 or more stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. albiflora 1b. Petals yellow or sometimes pale yellow; androecium with 30–50 stamens . . . . A. mexicana 1. Argemone albiflora Hornem. ssp. albiflora E white prickly-poppy. Argemone alba Lestib.; A. intermedia Sweet, pro parte; A. mexicana L. var. albiflora DC. • CT, MA. Fields, roadsides, lawns. 2. Argemone mexicana L. E Mexican prickly-poppy. Argemone leiocarpa Greene; A. spinosa Moench; A. vulgaris Spach; Ectrus mexicanus (L.) Nieuwl. • CT, MA, VT. Fields, roadsides, waste areas, about buildings.

Capnoides 1. Capnoides sempervirens (L.) Borkh. N Fig. 757 pink-corydalis. Capnoides glauca (Curt.) Moench; Corydalis glauca (Curt.) Pursh; C. sempervirens (L.) Pers.; Fumaria glauca Curt.; Fumaria sempervirens L. • CT, MA, ME, NH, RI, VT. Ledges, balds, woodlands, dry, open clearings.

Fig. 757  Inflorescence and leaves of Capnoides sempervirens.

700  tricolpates

Chelidonium 1. Chelidonium majus L. E greater celandine. Chelidonium majus L. var. laciniatum (P. Mill.) Syme; C. majus L. var. plenum Wehrhahn • CT, MA, ME, NH, RI, VT. Mesic, often shaded, soil of forest edges, roadsides, and waste areas.

Corydalis Reference: Ownbey (1947). 1a. Corolla purple (rarely white, pink, red, or purple-blue), 15–25 (–30) mm long; plants perennial; stem arising from a tuber . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. solida 1b. Corolla pale yellow or bright yellow, 7–16 mm long; plants annual or biennial; stem arising from a taproot 2a. Corolla pale yellow, 7–9 mm long; upper outer petal (i.e., the spurred petal) with a conspicuous undulate to toothed crest toward the tip; spur 1–1.5 (–2) mm long; sepals ca. 0.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. flavula 2b. Corolla bright yellow, 12–16 mm long; upper outer petal with merely a keel, lacking a toothed crest; spur 4–5 mm long; sepals 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . C. aurea 1. Corydalis aurea Willd.

nC

golden corydalis. Capnoides aureum (Willd.) Kuntze; Corydalis washingtoniana Fedde • MA, NH, VT. Lakeshore headlands and cliffs, rocky woodlands. This species is non-native in MA and native elsewhere. 2. Corydalis flavula (Raf.) DC.

NC

short-spurred corydalis. Capnoides flavulum (Raf.) Kuntze • CT. Woodlands, rocky ridges, balds, usually on trap rock. 3. Corydalis solida (L.) Clairv. E spring corydalis. Corydalis bulbosa (L.) DC. • CT, MA, VT. Lawns, gardens, disturbed areas, railroads, forest edges.

Dicentra 1a. Stems arising from bulbs; inflorescence simple (i.e., the pedicels arising from an unbranched axis); corolla largely white or cream (sometimes portions of the corolla pink or yellow-tinged) 2a. Spurs of the corolla parallel, rounded at the apex [Fig. 758], 0.5–1 mm long; bulblets of the rhizome yellow; apical crest of the inner 2 petals conspicuous and projecting; leaf blades typically blue-green; flowers fragrant . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. canadensis 2b. Spurs of the corolla divergent, subacute at the apex [Fig. 759], mostly 2–3 mm long; bulblets of the rhizome white or pink; apical crest of the inner 2 petals inconspicuous; leaf blades typically yellow-green to green; flowers not fragrant . . . . . . . . . . . . . . . . D. cucullaria 1b. Stems arising from a scaly rhizome; inflorescence compound; corolla largely pink to redpurple (rarely white) 3a. Apex of inner petals conspicuous, usually shortly exserted beyond the outer petals (i.e., the spurred petals); plants not colonial (i.e., new plants formed by seed) . . . . . . . . D. eximia 3b. Apex of inner petals visible, but often partly concealed and not exserted beyond the outer petals; plants forming interconnected colonies by rhizome growth . . . . . . D. formosa

Pa pav e r ac e a e   70 1

1. Dicentra canadensis (Goldie) Walp. N Fig. 758 squirrel-corn. Bicuculla canadensis (Goldie) Millsp.; Capnorchis canadensis (Goldie) Kuntze; Corydalis canadensis Goldie • CT, MA, ME, NH, VT; rare in eastern New England. Rich, mesic, deciduous forests. This species often occurs sympatrically with the more common Dicentra cucullaria. When both are present at the same site, D. canadensis usually flowers 7–10 days later than D. cucullaria. Report of this species in RI by George (1997) is based on a vegetative collection taken from a residential setting—15 May 1985, Champlin s.n. (Champlin Herb.). 2. Dicentra cucullaria (L.) Bernh. N Fig. 759 Dutchman’s-breeches. Bicuculla cucullaria (L.) Millsp.; Capnorchis cucullaria (L.) Planch.; Dicentra occidentalis Rydb.; Fumaria cucullaria L. • CT, MA, ME, NH, VT; also reported from RI by George (1997), but specimens are unknown. Rich, mesic, deciduous forests, including upland and riparian types.

Fig. 758  Flowers of Dicentra canadensis.

3. Dicentra eximia (Ker-Gawl.) Torr. E wild bleeding-heart. Bicuculla eximia (Ker-Gawl.) Millsp.; Capnorchis eximia (Ker-Gawl.) Planch.; Fumaria eximia Ker-Gawl. • MA, RI, VT. Fields, roadsides, gardens, banks. 4. Dicentra formosa (Haw.) Walp. ssp. formosa E western bleeding-heart. Capnorchis formosa (Haw.) Planch.; Fumaria formosa Haw. • MA. Fields, roadsides, gardens, areas of habitation.

Eschscholzia 1. Eschscholzia californica Cham. ex Nees ssp. californica E California-poppy. Eschscholzia procera Greene • CT, MA, NH. Waste areas, gardens, dumps.

Fumaria 1. Fumaria officinalis L. ssp. officinalis E common fumitory. CT, MA, ME, NH, VT. Gardens, waste areas, yards, fields, roadsides.

Glaucium 1. Glaucium flavum Crantz E yellow horn-poppy. CT, MA, RI. Sandy sea beaches and waste areas near the coast.

Lamprocapnos 1. Lamprocapnos spectabilis (L.) Fukuhara E common bleeding-heart. Dicentra spectabilis (L.) Lem. • CT, ME. Waste areas, gardens.

Macleaya 1. Macleaya cordata (Willd.) R. Br. E plume-poppy. Bocconia cordata Willd. • CT, MA, ME, NH, RI. Waste areas, areas of habitation.

Papaver Papaver argemone L. was reported from CT by Hultén and Fries (1986), but specimens are unknown. 1a. Leaves of the stem cordate-clasping [Fig. 760]; stems glabrous; flower buds 20–40 mm long; capsules 25–60 mm long; seeds 1–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . P. somniferum

Fig. 759  Flowers of Dicentra cucullaria.

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1b. Leaves of the stem not clasping; stems hispid-bristly; flower buds 5–20 mm long; capsules 10–22 mm long; seeds 0.6–1 mm long 2a. Gynoecium with (5–) 8–15 (–18) stigmas; capsules ovoid to subglobose, 1–1.5 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. rhoeas 2b. Gynoecium with 5–9 stigmas; capsule obconic to obovoid, 2–3 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. dubium 1. Papaver dubium L. E long-podded poppy. Papaver obtusifolium Desf.; P. modestum Jord. • CT, MA, RI. Fields, waste areas. 2. Papaver rhoeas L. E corn poppy. Papaver strigosum (Boenn.) Schur • CT, MA, ME, NH, VT. Abandoned homesteads, fields, forest clearings, dumps. 3. Papaver somniferum L. E Fig. 760 opium poppy. Papaver setigerum, auct. non DC.; P. somniferum L. var. hortense (Huss.) Corb. • CT, MA, ME, NH, RI, VT. Gardens, waste areas. Fig. 760  Flower and leaves of Papaver somniferum.

Sanguinaria 1. Sanguinaria canadensis L. N Fig. 761 blood-root. Sanguinaria canadensis L. var. rotundifolia (Greene) Fedde • CT, MA, ME, NH, RI, VT. Mesic, deciduous forests, including upland and riparian types.

Parnassiaceae Parnassia 1. Parnassia glauca Raf. N fen grass-of-Parnassus. Parnassia americana Muhl. • CT, MA, ME, NH, RI, VT. Fens, river shore seeps, wet meadows, rarely also found on wet lawns and in ditches, usually in high-pH bedrock regions. Fig. 761  Flowering habit of Sanguinaria canadensis.

Paulowniaceae Paulownia 1. Paulownia tomentosa (Thunb.) Sieb. & Zucc. ex Steud. E empress-tree. Bignonia tomentosa Thunb. • CT, MA, RI. Roadsides, forest edges and fragments, waste areas, dumps.

P e da l i ac e a e   703

Pedaliaceae 1a. Flowers in mostly 8- to 20-flowered racemes; corolla 35–55 mm long; fruit 10–20 cm long, tipped by long, arching, horn-like beak that is ultimately split into two halves . . . . . Proboscidea 1b. Flowers solitary in the axils of leaves; corolla 15–33 mm long; fruit 2–3 cm long, tipped by a short, broad, straight beak that is ultimately split into two halves . . . . . . . . . . . . . . . . . Sesamum

Proboscidea Probscidea louisianica (P. Mill.) Thellung ssp. fragrans (Lindl.) Bretting was reported from Essex County, MA, by Weatherby (1932). However, the supporting specimen is P. louisianica ssp. louisianica (Sorrie and Somers 1999). 1. Proboscidea louisianica (P. Mill.) Thellung ssp. louisianica E ram’s-horn. Martynia louisianica P. Mill. • CT, MA, RI, VT; also reported from ME and NH by Kartesz (1999), but specimens are unknown. Fields, areas of cultivation.

Sesamum 1. Sesamum orientale L. E sesame. Sesamum indicum L. • MA. Waste areas.

Penthoraceae Penthorum 1. Penthorum sedoides L. N Fig. 762 ditch-stonecrop. CT, MA, ME, NH, RI, VT. Shorelines, pools, backwater channels, ditches, wetland margins, marshes.

Phrymaceae 1a. Plants aquatic, without aerial stems, forming colonies by slender stolons in submersed populations (annuals in emersed plants); flowers 1–3 mm wide; androecium with 2 stamens; leaf blades narrower than 2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glossostigma 1b. Plants terrestrial or aquatic, with aerial stems, not colonial (forming terrestrial colonies by creeping stems in Mazus miquelii); flowers wider than 3 mm; androecium with 4 stamens; leaf blades much wider than 2 mm 2a. Inflorescence a spike-like raceme, the pedicels very short [Fig. 764]; fruit an achene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phryma 2b. Inflorescence a raceme or of solitary flowers from the axils of foliage leaves, in either case borne on evident pedicels [Fig. 763]; fruit a capsule

Fig. 762  Inflorescence of Penthorum sedoides.

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3a. Bracts subtending the flowers alternate; basal, connate portion of the calyx not ribbed or winged, much shorter than the lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mazus 3b. Bracts subtending the flowers opposite [Fig. 763]; basal, connate portion of the calyx 5-ribbed (the ribs sometimes wing-angled), longer than the lobes . . . . . . . . . . . . Mimulus

Glossostigma Reference: Les et al. (2006). 1. Glossostigma cleistanthum W.R. Barker E mud-mat. CT, RI. Shallow water or wet silt and mud of lakes and rivers, including fresh-tidal reaches. Originally thought to be Glossostigma diandrum (L.) Kunze, New England collections of this species have been confirmed as G. cleistanthum through molecular analysis. This species shows longer leaves than G. diandrum (mostly longer than 10 mm vs. mostly shorter than 8 mm). This new arrival to New England is easily confused with Limosella australis. However, it has a pair of leaves at each node and flowers with 2 stamens (vs. a cluster of leaves at each node and flowers with 4 stamens in L. australis).

Mazus 1a. Plants perennial, glabrous, with horizontal, creeping stems that root at the nodes; corolla 15–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. miquelii 1b. Plants annual, short-pubescent, without creeping stems; corolla 7–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pumilus 1. Mazus miquelii Makino E creeping mazus. Mazus reptans N.E. Br. • MA, ME. Gardens, waste areas. 2. Mazus pumilus (Burm. f.) Stennis E Japanese mazus. Mazus japonicus (Thunb.) Kuntze • CT, MA. Fields, roadsides, lawns.

Mimulus Beardsley and Olmstead (2002) showed that the genus Mimulus is not monophyletic. Mimulus brevipes and M. guttatus (and perhaps M. moschatus, not sampled in their work) need to be removed from Mimulus to create a monophyletic naming system. 1a. Corolla largely blue, lacking hairs on the palate of the lower lip 2a. Leaves sessile; pedicels 20–45 (–60) mm long; calyx lobes 1.5–8 mm long . . . M. ringens 2b. Leaves borne on petioles 10–20 mm long [Fig. 763]; pedicels 2–10 (–15) mm long [Fig. 763]; calyx lobes 1–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. alatus 1b. Corolla largely yellow, pubescent on palate of the lower lip 3a. Pedicels 2–10 mm long; ovary with parietal placentation; plants annual . . . M. brevipes 3b. Pedicels 10–80 mm long; ovary with axile placentation; plants perennial (rarely annual in M. guttatus) 4a. Calyx lobes narrow-triangular, longer than wide, subequal; corolla (14–) 17–22 mm long; stems viscid-villous; leaf blades oblong to ovate, pinnately veined . . . . . . . . . M. moschatus 4b. Calyx lobes broad-triangular, as wide as or wider than long, not equal, the upper one distinctly larger; corolla 25–45 mm long; stems glabrous or pubescent, but not viscid-villous; leaf blades ovate to subrotund, ± palmately veined . . . . . . . . . . M. guttatus 1. Mimulus alatus Ait. N Fig. 763 Fig. 763  Flowers and leaves of Mimulus alatus.

winged monkey-flower. CT, MA. Stream banks, river shores, fresh-tidal marshes, streamside meadows.

Pe da l i ac e a e   70 5

2. Mimulus brevipes Benth. E wide-throated monkey-flower. Eunanus brevipes (Benth.) Greene • MA. Wool waste. 3. Mimulus guttatus DC. E seep monkey-flower. Mimulus guttatus DC. ssp. litoralis Pennell; M. guttatus DC. ssp. micranthus (Heller) Munz; M. langsdorfii Donn ex Greene; M. langsdorfii Donn ex Greene var. guttatus (DC.) Jepson; M. micranthus Heller • CT. Stream shores, wet areas adjacent to streams. 4. Mimulus moschatus Dougl. ex Lindl. var. moschatus n musky monkey-flower. CT, MA, ME, NH, VT. River and stream shores, seeps, stream-side meadows, low roadsides, ditches. This species is native, at least in part, in western counties of MA and NH and in VT. It is considered introduced to other regions of New England. 5. Mimulus ringens L. N Allegheny monkey-flower. Mimulus minthodes Greene; M. ringens var. colpophilus Fern.; M. ringens L. var. minthodes (Greene) A.L. Grant • CT, MA, ME, NH, RI, VT; nearly throughout. Shorelines, marshes, including fresh-tidal marshes, swamps, wetland margins. Mimulus ringens var. colpophilus was considered to be a rare, tidal form of the species that was restricted to ME within New England. Observation of populations along fresh-tidal marshes show continuous variation in supposed diagnostic characters (e.g., calyx lobe length, stem internode length, pedicel length), with more stunted plants showing shorter leaves, stem internodes, calyx lobes, and pedicels found lower on river shores (i.e., experiencing a greater degree of inundation on a daily basis).

Phryma 1. Phryma leptostachya L. N Fig. 764 American lop-seed. Phryma leptostachya L. var. confertifolia Fern. • CT, MA, ME, NH, RI, VT. Rich, mesic, deciduous forests, including upland and riparian types.

Phytolaccaceae Phytolacca 1. Phytolacca americana L. var. americana N American pokeweed. Phytolacca decandra L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings, forest edges, waste areas.

Plantaginaceae 1a. Leaves 6–12 at each node, with entire blades; flowers with a single stamen borne at the summit of the ovary; calyx a minute rim around the apex of the ovary . . . . . . . . . . . . . . Hippuris 1b. Leaves 1 or 2 at each node (but crowded in the apical portion in Callitriche) or all basal, entire or toothed (in whorls of 3–6 in Veronicastrum, but then with serrate blades); flowers with 1–5 stamens, these not borne at the summit of the ovary; calyx absent or present and borne near the base of the ovary

Fig. 764  Inflorescence and leaves of Phryma leptostachya.

706  tricolpates

2a. Perianth absent; plants aquatic, with opposite leaves that are often crowded in the apical portion and forming a floating rosette [Fig. 766] . . . . . . . . . . . . . . . . . . . . . . . Callitriche 2b. Perianth present; plants terrestrial (aquatic in Littorella); leaves not forming a floating rosette near the apex of the stem 3a. Perianth inconspicuous, 3- or 4-merous, with an actinomorphic, scarious corolla [Figs. 771, 774, 775]; leaves all basal (opposite and borne on a stem in P. arenaria) [Fig. 771] 4a. Scape 1- to 3-flowered [Fig. 771]; fruit an achene, 1-locular, 1-seeded; plants aquatic, with slender stolons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Littorella 4b. Scape many-flowered; fruit usually a pyxis, 2- or 3-locular, 2- to many-seeded; plants terrestrial or of tidal wetlands, without stolons . . . . . . . . . . . . . . . . . . . . Plantago 3b. Perianth conspicuous and showy, with a zygomorphic, 5-merous corolla (appearing 4-merous in Veronica and Veronicastrum [Figs. 767, 770, 772]; leaves borne on a stem 5a. Corolla with a slender, basal spur or a saccate swelling protruding between the 2 lower calyx lobes, and with a palate [Figs. 769, 770]; foliage leaves usually alternate (sometimes the lower opposite, especially in Antirrhinum and Chaenorhinum) 6a. Corolla with a saccate (i.e., pouch-like) swelling on the lower side; plants both with flowers arranged in a terminal raceme and with capsules dehiscing by terminal pores 7a. Calyx lobes ± subequal, shorter than the basal, connate portion of the corolla; corolla 25–45 mm long; plants 40–80 (–120) cm tall, perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Antirrhinum 7b. Calyx lobes unequal, all longer than the basal, connate portion of the corolla; corolla 10–15 mm long; plants (5–) 20–50 cm tall, annual . . . Misopates 6b. Corolla with a slender, basal spur [Figs. 769, 770]; plants either with flowers in terminal racemes (but then with capsules opening by terminal slits) or with capsules dehiscing by apical pores (but then with solitary flowers in the axils of leaves) 8a. Inflorescence a terminal raceme; capsules dehiscing by terminal slits; seeds with angular faces 9a. Palate composed of 2 short, white ridges; seeds both ± smooth on the faces and wingless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nuttallanthus 9b. Palate composed of a single, usually yellow or orange, ridge; seeds either with rugose or reticulate patterning or with wings or wing-like angles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Linaria 8b. Inflorescence a solitary flower in the axils of leaves; capsules dehiscing by 2 terminal pores (the pores eventually extending to the capsule base in Cymbalaria); seeds ellipsoid to ovoid 10a. Stems upright; capsules pubescent, asymmetrical, one locule larger than the other . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaenorhinum 10b. Stems prostrate; capsules glabrous, symmetrical, both locules of ± equal size 11a. Leaf blades palmately veined, with 5–9 tooth-like lobes; stems rooting at the nodes in contact with the soil . . . . . . . . . . . . . . . Cymbalaria 11b. Leaf blades pinnately veined, entire (rarely the lower minutely denticulate); stems not rooting at the nodes . . . . . . . . . . . . . . . . . . . Kickxia 5b. Corolla lacking a slender, basal spur, without a palate except in Chelone and 1 species of Penstemon; foliage leaves opposite or whorled (alternate in Digitalis) 12a. Foliage leaves alternate; inflorescence a secund raceme [Fig. 767] . . . . Digitalis

Pl a n tag i n ac e a e   707

12b. Foliage leaves opposite or whorled; inflorescence various, but not a secund raceme 13a. Androecium composed of 5 stamens—4 pollen-bearing stamens and 1 sterile stamen 14a. Corolla subgaleate; sepals subtended by bractlets; sterile stamen shorter than the pollen-bearing stamens; seeds winged . . . . . . . . . . Chelone 14b. Corolla without a galea [Figs. 772, 773]; sepals not subtended by bractlets; sterile stamen nearly equal in length to the pollen-bearing stamens; seeds without wings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Penstemon 13b. Androecium composed of 2 or 4 stamens—all pollen bearing or 2 of the 4 composed of filaments only 15a. Corolla apparently composed of 4 lobes due to fusion of the upper 2 petals [Fig. 778]; calyx composed of 4 sepals (5 sepals in Veronica officinalis); inflorescence a spike or raceme (the bracteal leaves ± similar to the foliage leaves in some Veronica, but then usually alternately arranged) [Figs. 776, 777, 778] 16a. Leaves opposite, sometimes the upper alternate (rarely some whorled in V. longifolia and V. spuria); lobes of the corolla longer than the basal tube; capsules, before dehiscence, emarginate or lobed at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Veronica 16b. Leaves mostly in whorls of 3–7; lobes of the corolla shorter than the basal tube; capsules, before dehiscence, tapering to the apex, not lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Veronicastrum 15b. Corolla composed of 2 lips—the upper lip 2-lobed, the lower lip 3-lobed [Fig. 768]; calyx composed of 5 sepals; inflorescence of solitary or whorled flowers in the axils of oppositely arranged foliage leaves [Fig. 768] 17a. Sepals evidently connate in the basal portion; corolla 4–6 mm long, the middle lobe of the lower corolla lip declined below the lateral lobes and enfolding the 4 pollen-bearing stamens . . . . . . . . . . . . . . . . . . Collinsia 17b. Sepals distinct nearly or quite to the base [Fig. 768]; corolla 5–16 mm long, the middle lobe of the lower lip neither conspicuously declined below the lateral lobes nor enfolding the 2 pollen-bearing stamens [Fig. 768] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gratiola

Antirrhinum 1. Antirrhinum majus L. E garden snapdragon. CT, MA, VT. Waste areas, dumps, roadsides.

Callitriche Reference: Fassett (1951). 1a. Flowers and fruits not subtended by bracts; leaf blades ± uniform, never forming a floating rosette 2a. Schizocarp 0.5–0.7 mm long, borne on a short pedicel 0.1–0.6 mm long, each mericarp with an inconspicuous wing-margin; leaf blades 2–5 mm long, spatulate to oblanceolate, rounded at the apex; plants annual, growing on damp soils . . . . . . . . . . . . . . . . . C. terrestris 2b. Schizocarp 1–2 mm long, subsessile, each mericarp with a conspicuous wing-margin; leaf blades 4–15 (–20) mm long, oblong-linear, rounded to retuse at the apex; plants perennial, typically wholly submersed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. hermaphroditica

708  tricolpate s

1b. Flower and fruits subtended by 2 ± white or translucent bracts mostly 0.5–1.5 mm long; leaf blades often dimorphic—those along the stem linear and relatively distant, those near the stem apex spatulate, crowded, and forming a floating rosette 3a. Schizocarp 1.5–2 mm long, each mericarp with a prominent, thin wing nearly 0.1 mm wide that is ± equally prominent from apex to base of fruit; floating leaf blades 3–8 mm wide, with 5–7 veins by branching of the lateral veins [Fig. 766] . . . . . . . . . . . . . . C. stagnalis 3b. Schizocarp 1–1.4 mm long, each mericarp unwinged or with a thin wing up to 0.05 mm wide that becomes less prominent toward fruit base and is absent in the basal ¼ or more of the fruit; floating leaf blades (when present) up to 5 mm wide, with 3 (–5) veins 4a. Schizocarp 0.2 mm or more longer than wide [Fig. 765], the apical portion of each mericarp with a thin wing, also with an evident commissural groove between the mericarps; pit-like aereolae on surface of fruit tending to be aligned in vertical rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. palustris 4b. Schizocarp ± as wide as long (i.e., not more than 0.1 mm longer than wide), lacking thin wings along the margin of the mericarp (or essentially so), without a commissural groove or with a shallow groove between the mericarps; pit-like aereolae on surface of fruit not at all aligned in vertical rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. heterophylla 1. Callitriche hermaphroditica L.

NC

autumn water-starwort. Callitriche autumnalis L. • VT; also reported from CT by Magee and Ahles (1999), but specimens are unknown; Lake Champlain. Shallow water of lakes. 2. Callitriche heterophylla Pursh var. heterophylla N greater water-starwort. Callitriche anceps Fern. • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving water of rivers, lakes, and pools, ascending to high elevation in NH and VT. Callitriche anceps was originally believed to be a wholly submersed, boreal species allied to C. heterophylla. The characteristics used by Fernald (1950b) to distinguish Callitriche anceps from C. heterophylla (e.g., stem width and length, fruit length, leaf blade morphology) are not diagnostic. Fassett (1951) commented on the lack of utility of these characteristics. However, Fassett maintained C. anceps on the basis of a very minor difference in fruit shape, though he noted that intermediates existed. It is here treated as part of the variation displayed by C. heterophylla. 3. Callitriche palustris L. N Fig. 765 Fig. 765  Fruits and leaves of Callitriche palustris.

vernal water-starwort. Callitriche palustris L. var. verna (L.) Fenley ex Jepson; C. verna L. • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving water of rivers, lakes, and pools, sometimes becoming amphibious as water level declines and then found on shorelines and in wet depressions. 4. Callitriche stagnalis Scop. E Fig. 766 pond water-starwort. CT, MA, ME. Shallow water of lakes and brackish-tidal rivers. Philbrick et al. (1998) have documented the spread of C. stagnalis in the United States. 5. Callitriche terrestris Raf.

NC

terrestrial water-starwort. Callitriche austinii Engelm.; C. deflexa A. Braun; C. deflexa A. Braun var. austinii (Engelm.) Hegelm. • CT, MA, ME. Pools, wet ground, usually associated with puddles, wheel ruts, and low areas in little-used or abandoned roads.

Chaenorhinum 1. Chaenorhinum minus (L.) Lange E Fig. 766  Habit of Callitriche stagnalis showing floating rosette of leaves.

dwarf-snapdragon. Antirrhinum minus L.; Linaria minor (L.) Desf. • CT, MA, ME, NH, VT. Roadsides, railroads, sidewalks, waste areas. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI and was unaware of any collections.

Pl a n tag i n ac e a e   70 9

Chelone 1a. Corolla white, sometimes partly green-yellow or tinged with pink (especially near the apex), the palate pubescent with white hairs; staminode green; leaf blades usually tapering to an ill-defined petiole or subsessile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. glabra 1b. Corolla purple or red-purple throughout, the palate pubescent with yellow (rarely white) hairs; staminode white; leaf blades borne on distinct, slender petioles 5–30 (–40) mm long 2a. Leaf blades ovate, 3–10 cm wide, broadly rounded to truncate at the base; petioles 15–30 (–40) mm long; upper lip of corolla sharply ridged; lower lip of corolla pubescent with yellow hairs and thinly striped with bright purple . . . . . . . . . . . . . . . . . . . . . . . . . C. lyonii 2b. Leaf blades lanceolate to narrow-elliptic, 1.4–4 cm wide, cuneate at the base; petioles 5–15 mm long; upper lip of corolla scarcely ridged; lower lip of corolla pubescent with pale yellow hairs and with or without thin, purple stripes . . . . . . . . . . . . . . . . . . . . . . . . . C. obliqua 1. Chelone glabra L. N white turtlehead. Chelone glabra L. var. chlorantha (Pennell & Wherry) Cooperrider; C. glabra L. var. dilatata Fern. & Wieg.; C. glabra L. var. elatior Raf.; C. glabra L. var. linifolia Coleman • CT, MA, ME, NH, RI, VT; nearly throughout. Shorelines, swamps, wetland margins, wet ditches. 2. Chelone lyonii Pursh E pink turtlehead. CT, MA, ME. Wet soil, stream sides, areas of habitation. 3. Chelone obliqua L. var. obliqua E red turtlehead. MA. Forests.

Collinsia 1. Collinsia parviflora Lindl.

NC

small-flowered blue-eyed Mary. MA, VT. Cliff bases, talus, woodlands, dry roadsides.

Cymbalaria 1. Cymbalaria muralis P.G. Gaertn., B. Mey., & Scherb. E Kenilworth-ivy. Linaria cymbalaria (L.) P. Mill. • CT, MA, VT; also reported from RI by George (1997), but specimens are unknown. Roadsides, waste areas, dumps.

Digitalis 1a. Lobes of the calyx broad-lanceolate to obovate, mostly 6–9 mm wide; corolla purple, pale pink, or white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. purpurea 1b. Lobes of the calyx linear to lanceolate, mostly 1–2.5 mm wide; corolla yellow to white 2a. Corolla 40–50 mm long; leaf blades sparsely pubescent on the abaxial suface; stem pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. grandiflora 2b. Corolla (9–) 15–30 mm long; leaf blades usually glabrous on the abaxial surface; stem usually glabrous 3a. Corolla 20–30 mm long, the connate portion cupuliform, the middle lobe of lower lip much surpassing the lateral lobes and 8–13 mm long; sepals glandular-villous (i.e., glandular hairs found on the surfaces of the sepals) . . . . . . . . . . . . . . . . . . . . . . . . D. lanata 3b. Corolla (9–) 15–22 (–25) mm long, the connate portion cylindrical to cylindricalurceolate, the middle lobe of lower lip slightly exceeding the lateral lobes and 5–6 mm long; sepals glandular-ciliate (i.e., glandular hairs found only on the margins of the sepals) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. lutea

710 tricolpates

1. Digitalis grandiflora P. Mill. E yellow foxglove. Digitalis ambigua Murr. • CT, ME, NH, VT. Fields, clearings. 2. Digitalis lanata Ehrh. E Grecian foxglove. CT, MA, NH, VT. Roadsides, dumps, gardens, ledges near habitation. 3. Digitalis lutea L. E straw foxglove. CT, NH; also reported from VT by Kartesz (1999) and Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, waste areas. Reports of this species from MA (e.g., Seymour 1982) are based on a garden collection that was transplanted from NH—2 Jul 1944, Bean s.n. (NEBC!). 4. Digitalis purpurea L. var. purpurea E Fig. 767 purple foxglove. CT, MA, ME, NH, VT. Fields, roadsides, yards, dumps.

Gratiola 1a. Leaf blades lanceolate to ovate, with a broad base [Fig. 768], glandular-punctate, (8–) 10–25 (–30) mm long; corolla 12–16 mm long, yellow throughout (rarely white); capsule 2–3 mm long; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. aurea Fig. 767  Secund inflorescence of Digitalis purpurea.

1b. Leaf blades lanceolate to oblong or oblanceolate, with a narrow base, not glandularpunctate, (10–) 20–50 (–70) mm long; corolla 8–12 mm long, largely white (commonly with yellow lobes in 1 species); capsules 3–9 mm long; plants annual 2a. Pedicels relatively stout, (0.1–) 1–4 (–14) mm long in fruit, ascending to erect; corolla of chasmogamous flowers pubescent within with filiform hairs; capsules 5–9 mm long; seeds slender-cylindric, 0.7–0.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. virginiana 2b. Pedicels relatively slender, 10–20 (–25) mm long in fruit, spreading; corolla of chasmogamous flowers pubescent within with clavate-tipped hairs; capsules 3–5 mm long; seeds thick-cylindric, 0.4–0.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. neglecta 1. Gratiola aurea Pursh N Fig. 768 golden hedge-hyssop. Gratiola aurea Pursh var. obtusa Pennell; G. lutea Raf. • CT, MA, ME, NH, RI, VT. Lake shores, river shores and banks, including tidal stretches, frequently (but not confined to) sandy substrate. 2. Gratiola neglecta Torr. N

Fig. 768  Flowers and leaves of Gratiola aurea.

clammy hedge-hyssop. Gratiola neglecta Torr. var. glaberrima Fern. • CT, MA, ME, NH, RI, VT. Shorelines, margins of pools, meadows, ditches, low areas in disturbed soils. 3. Gratiola virginiana L. var. virginiana

NC

round-fruited hedge-hyssop. Gratiola sphaerocarpa Ell. • RI. Shorelines, margins of pools, ditches.

Hippuris 1. Hippuris vulgaris L. N common mare’s-tail. MA, ME, NH, VT. Muddy or peaty shores and shallow, still or slow-moving water of streams, ponds, backwater sloughs, and pools.

Kickxia 1a. Leaf blades broad-ovate, rounded to cordate at the base; pedicels pubescent throughout their length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. spuria 1b. Leaf blades ovate to triangular-ovate, sagittate or hastate at the base; pedicels usually glabrous except near the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. elatine

Pl a n tag i nac e a e   71 1

1. Kickxia elatine (L.) Dumort. E sharp-leaved cancerwort. Antirrhinum elatine L.; Linaria elatine (L.) P. Mill. • CT, MA, RI. Fields, roadsides, waste areas, gardens. 2. Kickxia spuria (L.) Dumort. E round-leaved cancerwort. Antirrhinum spurium L. • RI. Fields, roadsides, waste areas.

Linaria Linaria aeruginea (Gouan) Cav. was reported from CT by Kartesz (1999), based on Harger et al. (1922). Harger et al. did not report L. aeruginea but rather reported L. reticulata Willd., a taxon that may belong in the synonymy of the European L. aeruginea (according to Chater et al. 1972). However, the original collection was determined as L. reticulata Desf. by Fernald (note discrepant authorities), a different species from northern Africa now referred to as Linaria pinifolia (Poir.) Thellung. Therefore, the report of L. aeruginea is here considered erroneous (i.e., it should be excluded from the regional flora). Linaria angustissima (Loisel.) Borbás was reported from Worcester County, MA, by several sources (e.g., Seymour 1982), based on Gates 28177 (NEBC!). This specimen is L. vulgaris, as evidenced by the pubescent raceme and spur longer than 10 mm. I have seen two specimens determined as Linaria bipartita (Vent.) Willd., one from Fairfield County, CT (Eames 9511 p.p. GH!), which was actually L. maroccana and one from Middlesex County, MA (17 Jul 1924, Kidder s.n. AMES!), which was taken from a cultivated plant. Therefore, this species, which is now treated as L. incarnata (Vent.) Spreng., is also excluded from New England. Linaria dalmatica (L.) P. Mill. ssp. macedonica (Griseb.) D.A. Sutton was reported from MA and NH by Kartesz (1999), but specimens are unknown. Linaria supina (L.) Chaz. was reported from MA by Knowton and Deane (1923c). However, the specimen (10 Jul 1863, Boott s.n. GH!) was collected from a cultivated plant (i.e., this species is not naturalized in New England; Sorrie and Somers 1999). Reference: Chater et al. (1972). 1a. Corolla white to purple or red-purple and usually faintly to prominently striped with purple, with a white, yellow, or orange palate; seeds ovoid-trigonous, transversely rugose, unwinged or with very narrow, flange-like wings 2a. Spur 2–5 mm long; corolla white to pale purple, therefore, the purple stripes very prominent; stigma capitate; pedicels and sepals glabrous; plants perennial by creeping rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. repens 2b. Spur 5–15 mm long; corolla purple or red-purple, therefore, the purple stripes less conspicuous; stigma bifid; pedicels and often the sepals pubescent with septate hairs; plants annual 3a. Corolla 20–38 mm long, with a largely white palate; spur 8–15 mm long, longer than the remaining portion of the corolla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. maroccana 3b. Corolla 15–18 mm long, with a conspicuous yellow palate; spur 5–7 mm long, shorter than the remaining portion of the corolla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. pinifolia 1b. Corolla yellow, with a yellow or an orange palate; seeds discoid, not rugose, with a marginal wing or compressed-pyrimidal, rugulose, and with flange-like wing angles (ovoidtrigonous, transversely rugose, and unwinged in L. spartea) 4a. Leaf blades narrow-lanceolate to ovate, 3–40 mm wide [Fig. 769], rigid; seeds compressed-pyramidal, rugulose, with a very narrow, often whitish, flange on the angles 5a. Corolla 20–50 mm long; leaf blades 10–40 mm wide, broad-lanceolate to ovate, 2–4 (–5) times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. dalmatica 5b. Corolla 13–22 mm long; leaf blades 3–15 mm wide, narrow-lanceolate to broadlanceolate, (3.5–) 6–12 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . L. genistifolia 4b. Leaf blades filiform to narrow-elliptic, 0.5–5 (–15) mm wide [Fig. 770], herbaceous; seeds discoid, not rugulose, with a narrow, marginal wing (ovoid-pyramidal, transversely rugose, and unwinged in L. spartea)

71 2   tricolpate s

6a. Stigma bifid; seeds ovoid-pyramidal, unwinged, 0.6–0.7 mm long; leaf blades 0.5–1 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. spartea 6b. Stigma capitate; seeds discoid, with a narrow, marginal wing, 1.5–3 (–3.5) mm long; leaf blades 1–5 (–15) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. vulgaris 1. Linaria dalmatica (L.) P. Mill. ssp. dalmatica E Fig. 769 Dalmatian toadflax. Antirrhinum dalmaticum L.; L. genistifolia (L.) P. Mill. ssp. dalmatica (L.) Maire & Petitm. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 2. Linaria genistifolia (L.) P. Mill. E broom-leaved toadflax. Antirrhinum genistifolium L. • MA. Fields, roadsides, waste areas, railroads. 3. Linaria maroccana Hook. f. E Moroccan toadflax. CT, MA, ME, NH. Fields, roadsides, waste areas, gardens. 4. Linaria pinifolia (Poir.) Thellung E Fig. 769  Flowers and leaves of Linaria dalmatica.

pine-needle toadflax. Antirrhinum pinifolium Poir.; A. reticulatum Sm.; Linaria reticulata (Sm.) Desf. • CT. Fields, roadsides, waste areas. 5. Linaria repens (L.) P. Mill. E striped toadflax. Antirrhinum repens L.; Linaria striata DC. • MA, ME; also reported from CT by Hultén and Fries (1986), but specimens are unknown. Fields, roadsides, waste areas. 6. Linaria spartea (L.) Willd. E ballast toadflax. Antirrhinum sparteum L. • CT. Fields, roadsides, waste areas. This species very rarely possesses a purple corolla (Chater et al. 1972). Purple forms would be very difficult to separate from Linaria maroccana. In New England, collections of L. spartea have all had yellow corollas and tend to have relatively straight spurs, whereas collections of L. maroccana show purple to red-purple corollas and somewhat curved spurs. 7. Linaria vulgaris P. Mill. E Fig. 770 butter-and-eggs toadflax. Linaria linaria (L.) Karst. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads.

Fig. 770  Flowers and leaves of Linaria vulgaris.

Littorella Reference: Fernald (1918). 1. Littorella americana Fern. N Fig. 771 American shoreweed. Littorella uniflora (L.) Aschers. var. americana (Fern.) Gleason; Plantago americana (Fern.) K. Rahn • ME, VT. Shallow water of lakes, sometimes emersed as water levels decline later in season. The genus Littorella was combined with Plantago by Rahn 1996). However, Hoggard et al. (2003) provided convincing data that shows Littorella as a genus distinct from Plantago.

Misopates 1. Misopates orontium (L.) Raf. E lesser-snapdragon. Antirrhinum orontium L. • CT, ME. Waste areas, ditches.

Nuttallanthus Fig. 771  Habit of Littorella americana.

1. Nuttallanthus canadensis (L.) D.A. Sutton N oldfield-toadflax. Antirrhinum canadense L.; Linaria canadensis (L.) Chaz. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads.

P l a n tag i nac e a e   7 13

Penstemon Penstemon canescens (Britt.) Britt. was reported from ME by Pennell (1935). However, this species is relatively southern, reaching as far north as PA (Clements et al. 1998). The collection, if indeed determined accurately, would certainly represent an introduction. Reference: Clements et al. (1998). 1a. Corolla red, strongly zygomorphic, the upper lip subgaleate, the lower lip with reflexed lobes; sterile stamen glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. barbatus 1b. Corolla white to pale purple or blue-purple to purple, weakly to moderately zygomorphic, the upper lip not resembling a galea, the lower lip with ascending to spreading lobes; sterile stamen pubescent, at least minutely near the apex 2a. Corolla (35–) 40–50 mm long; calyx at anthesis 7–15 mm long; upper stem leaf blades broad-ovate to orbicular, subcordate to strongly clasping at the base, strongly glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grandiflorus 2b. Corolla 15–35 mm long; calyx at anthesis 2.5–8 mm long; upper stem leaf blades lanceolate or narrow-ovate to elliptic, oblong, or narrow-oblong, rounded to truncate at the base, not or only thinly glaucous 3a. Corolla weakly zygomorphic, the upper and lower lips not well-developed [Fig. 773], pubescent on both surfaces with minute, glandular hairs . . . . . . . . . . . . . . . . P. tubiflorus 3b. Corolla zygomorphic, with conspicuous upper and lower lips, [Fig. 772] pubescent on the inner surface with eglandular hairs 4a. Lower lip with an upward swelling, forming a palate that closes the orifice of the tubular corolla; corolla without purple lines on the inside of the tube; sterile filament densely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. hirsutus 4b. Lower lip without a palate, the orifice of the corolla open; corolla marked with fine purple lines on the inside of the tube; sterile filament sparsely to moderately pubescent or densely pubescent in P. pallidus 5a. Corolla gradually widened distally, strongly ridged within; leaf blades pubescent on both surfaces; stems pubescent . . . . . . . . . . . . . . . . . . . . . . . P. pallidus 5b. Corolla with a slender, basal tube and an evident, abrupt, dilated, distal portion, only moderately ridged within the tube; leaf blades glabrous or sparsely pubescent especially along the abaxial midvein; stems glabrous or pubescent only in lines of decurrence from the leaf bases 6a. Anthers pubescent with stiff, white hairs; corolla white on the abaxial (i.e., outer) surface (sometimes faintly tinged with purple); capsules 9–12 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. digitalis 6b. Anthers glabrous (rarely with a few hairs on one or both locules); corolla pale purple to purple on the abaxial surface; capsules 5–8 mm long 7a. Corolla 20–35 mm long; calyx lobes narrow-lanceolate to linear-subulate, 5–12 mm long; leaf blades often finely serrate . . . . . . . . . . . . . . . . P. calycosus 7b. Corolla 15–20 (–22) mm long; calyx lobes lanceolate to narrow-ovate, 3–6 mm long; leaf blades often obscurely serrate . . . . . . . . . . . . . P. laevigatus 1. Penstemon barbatus (Cav.) Roth ssp. barbatus E beard-lip beardtongue. MA. Waste areas. 2. Penstemon calycosus Small E long-sepaled beardtongue. Penstemon laevigatus Ait. ssp. calycosus (Small) Bennett • CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings. 3. Penstemon digitalis Nutt. ex Sims N foxglove beardtongue. Penstemon laevigatus Ait. ssp. digitalis (Nutt. ex Sims) Bennett • CT, MA, ME, NH, RI, VT. Fields, roadsides, meadows, banks.

71 4   tricolpate s

4. Penstemon grandiflorus Nutt. E large beardtongue. CT, MA. Fields, roadsides. 5. Penstemon hirsutus (L.) Willd. N Fig. 772 northeastern beardtongue. CT, MA, ME, RI, VT. Fields, roadsides, woodlands, outcrops, usually in dry, well-drained soils. The report of this species from NH by Pease (1964) is based on a collection of Penstemon pallidus—Pease 31942 (NEBC!). Penstemon hirsutus is relatively easy to identify in the field due to the palate on the lower corolla lip (unique for New England Penstemon). However, on herbarium specimens this feature can be distorted, and separating this species from P. pallidus is more difficult. Penstemon hirsutus has a corolla 20–30 mm long, capsules 8–9 mm tall, and essentially glabrous adaxial leaf surfaces. Penstemon pallidus has a corolla 15–22 mm long, capsules 6–7 mm tall, and pubescent adaxial leaf surfaces. 6. Penstemon laevigatus Ait. E Fig. 772  Flowers of Penstemon hirsutus.

eastern smooth beardtongue. CT; also reported from RI by George (1997), but specimens are unknown. Fields, roadsides, waste areas, quarries. Most, if not all, reports of this species in New England can be referred to Penstemon calycosus. I have seen one specimen that may be this species (Weatherby 2463 NEBC!). It has a short corolla, but the sepals are ± intermediate between P. calycosus and P. laevigatus (i.e., the specimen is not unequivocal). 7. Penstemon pallidus Small N eastern white beardtongue. CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings, woodlands, banks, usually in dry, sandy places. Some populations of this species are likely introduced in New England. 8. Penstemon tubiflorus Nutt. E Fig. 773 tube beardtongue. Penstemon tubiflorus Nutt. var. achoreus Fern. • CT, MA, ME, NH; also reported from RI by George (1997), but specimens are unknown. Fields, pastures, roadsides, clearings. Reports of this species in VT are based on collections of Penstemon digitalis.

Plantago References: Bassett (1973), Rahn (1996). Fig. 773  Flowers of Penstemon tubiflorus.

1a. Leaf blades dentate to 1- or 2-times pinnately lobed; corolla pubescent on the abaxial (i.e., outer) surface; spike nodding prior to anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . P. coronopus 1b. Leaf blades usually entire (sometimes with obscure teeth); corolla glabrous on the abaxial surface or pubescent in P. maritima; spike not nodding prior to anthesis 2a. Leaves borne on a stem, opposite, the blades linear and 1–3 mm wide [Fig. 774]; peduncles arising from the axils of leaves [Fig. 774] . . . . . . . . . . . . . . . . . . . . . . . . . . P. arenaria 2b. Leaves all basal, the blades linear to ovate and 1–110 mm wide [Fig. 774]; peduncles arising among basal rosettes of leaves [Fig. 775] 3a. Bracts and/or sepals conspicuously pubescent; plants monocarpic, usually annual (sometimes biennial) 4a. Plants subdioecious; lobes of the corolla of carpel-bearing, chasmogamous flowers erect after anthesis, connivent, closing over the fruit; leaf blades oblanceolate to obovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. virginica 4b. Plants synoecious; lobes of the corolla spreading to reflexed after anthesis; leaf blades linear to narrow-oblong 5a. Bracts conspicuously exceeding the flowers and fruits, the lower ones 8–45 mm long, mostly 4–10 times as long as the sepals [Fig. 775]; leaf blades sparsely villous to nearly glabrous on the adaxial surface; inflorescence sparsely tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aristata

Pl a n tag i nac e a e   715

5b. Bracts inconspicuous, scarcely, if at all, extending beyond the flowers and fruits, 2–5.4 mm long, mostly 1–3 times as long as the sepals; leaf blades evidently villous-tomentose on the adaxial surface; inflorescence evidently tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. patagonica 3b. Bracts and sepals glabrous or ciliate only on the apical portion of the keel; plants polycarpic, perennial (monocarpic and annual in P. pusilla) 6a. Flowers with 2 stamens; corolla lobes 0.25–0.5 mm long; plants annual . . . P. pusilla 6b. Flowers with 4 stamens; corolla lobes 0.7–2.5 mm long; plants perennial 7a. Leaf blades linear, 1–12 mm wide, fleshy; corolla tube pubescent on the abaxial surface; plants of tidal wetlands and Atlantic coast shorelines . . . . . . . P. maritima 7b. Leaf blades narrow-elliptic to broad-ovate or obovate, 6–110 mm wide, herbaceous to somewhat fleshy; corolla tube glabrous on the abaxial surface; plants of upland habitats, often weedy (of saline shores in P. intermedia) 8a. Bracts and sepals conspicuously keeled; lobes of the corolla 0.7–1 mm long; pyxis 4- to 34-seeded; all the roots slender and filamentous; spikes 5–30 cm tall 9a. Pyxis (3–) 4–6 mm tall, circumscissile near the base, containing 4–9 (–10) seeds; bracts narrow-triangular; seeds 1.5–2 (–2.5) mm long, not reticulatepatterned; petiole usually anthocyanic near the base . . . . . . . . . . . . . P. rugelii 9b. Pyxis (2–) 2.5–4 mm tall, circumscissile near the middle, containing 6–22 (–34) seeds; bracts broad-ovate; seeds 1–1.7 mm long, reticulate-patterned; petiole usually green near the base (but sometimes anthocyanic in P. major) 10a. Pyxis rounded to pointed at the apex, containing (4–) 6–11 (–13) seeds; leaf blades membranaceous to fleshy-herbaceous, glabrous or pubescent, broad-elliptic to broad-ovate, borne on relatively more elongate and slender petioles; plants of primarily inland habitats, casually found bordering saline communities . . . . . . . . . . . . . . . . . . . . . . . . . . P. major 10b. Pyxis broadly rounded at the apex, containing 14–34 seeds; leaf blades subcoriaceous-fleshy, usually pubescent, narrow-elliptic to ovate, borne on broad, short petioles; plants of saline shores and near coastal habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. intermedia 8b. Bracts and sepals flat or slightly keeled; lobes of the corolla 1.5–2.5 mm long; pyxis 2- to 4-seeded; primary roots prominently thickened, tuberous near junction with foliage; spikes 1.5–10 cm tall

Fig. 774  Inflorescence and stem leaves of Plantago arenaria.

11a. Calyx apparently composed of 3 sepals, 1 of the sepals with 2 midveins and formed by fusion of 2 sepals; leaf blades lanceolate to narrow-elliptic, ascending; scape prominently furrowed . . . . . . . . . . . . . . . . . . . . P. lanceolata 11b. Calyx composed of 4 distinct sepals; leaf blades elliptic to ovate or obovate, prostrate; scape not furrowed . . . . . . . . . . . . . . . . . . . . . . . . . . P. media 1. Plantago arenaria Waldst. & Kit. E Fig. 774 sand plantain. Plantago indica L., nom. illeg.; P. psyllium L., nom. ambig.; P. ramosa Aschers. • CT, MA, ME, NH, VT. Roadsides, parking lots, railroads, waste areas.

2. Plantago aristata Michx. E Fig. 775 bracted plantain. Plantago aristata Michx. var. nuttallii (Rapin) Morris; P. gnaphalioides Pursh var. aristata (Michx.) Hook.; P. nuttallii Rapin; P. patagonica Jacq. var. aristata (Michx.) Gray • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 3. Plantago coronopus L. E buck’s-horn plantain. Asterogeum laciniatum S.F. Gray; Coronopus vulgaris Fourr. • MA. Dumps, waste areas.

Fig. 775  Inflorescence and basal leaves of Plantago aristata.

716 tricolpates

4. Plantago intermedia DC. N many-seeded plantain. Plantago halophila Bickn.; P. major L. ssp. intermedia (DC.) Arcang.; P. major var. intermedia (DC.) Pilger; P. major L. ssp. pleiosperma Pilger; P. major var. scopulorum Fries & Broberg • CT, MA, ME, NH, RI; coastal plain. Open areas near or adjacent to the Atlantic coast, included rocky or gravelly beaches, headlands, and upper edges of saline to brackish marshes. 5. Plantago lanceolata L. E English plantain. Arnoglossum lanceolatum S.F. Gray; Plantago altissima, auct. non L.; P. lanceolata L. var. sphaerostachya Mert. & Koch • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads, lawns. 6. Plantago major L. E common plantain. Plantago major L. var. pilgeri Domin • CT, MA, ME, NH, RI, VT; throughout. Fields, roadsides, waste areas, lawns, trail edges. 7. Plantago maritima L. ssp. juncoides (Lam.) Hultén N seaside plantain. Plantago juncoides Lam.; P. juncoides Lam. var. decipiens (Barneoud) Fern.; P. juncoides Lam. var. glauca (Hornem.) Fern.; P. juncoides Lam. var. laurentiana Fern.; P. maritima L. ssp. borealis (Lange) Blytt & O. Dahl; P. maritima L. var.. juncoides (Lam.) Gray; P. oliganthos Roemer & J.A. Schultes; P. oliganthos Roemer & J.A. Schultes var. fallax Fern. • CT, MA, ME, NH, RI; coastal. Saline and brackish marshes, coastal cliffs and ledges, very rarely found on fresh-tidal river shore ledges. North American plants are recognized as ssp. juncoides on the basis that they are self-compatible, have broader bracts, and usually have 2- to 4-seeded capsules (European plants are usually self-incompatible, have narrower bracts, and 1- to 2-seeded capsules). 8. Plantago media L. E hoary plantain. Arnoglossum incanum S.F. Gray; Plantago incana Stokes • CT, MA, ME, NH, RI. Fields, waste areas, lawns. 9. Plantago patagonica Jacq. E Pursh’s plantain. Plantago gnaphalioides Nutt.; P. patagonica Jacq. var. breviscapa (Shinners) Shinners; P. patagonica Jacq. var. gnaphalioides (Nutt.) Gray; P. purshii Roemer & J.A. Schultes; P. purshii Roemer & J.A. Schultes var. breviscapa Shinners • MA, ME, VT. Fields, waste areas, railroads, wool waste. 10. Plantago pusilla Nutt. E dwarf plantain. Plantago hybrida W. Bart.; P. pusilla Nutt. var. major Engelm. • CT, MA, RI. Sandy roadsides, thin soil near ledges. 11. Plantago rugelii Dcne. N Rugel’s plantain. Plantago rugelii Dcne. var. asperula Farw. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, lawns, trail edges. 12. Plantago virginica L. n pale-seeded plantain. Plantago caroliniana Walt.; P. virginica L. var. viridescens Fern. • CT, MA, ME, RI. Fields, roadsides, waste areas, grasslands, pond shores (for one non-native occurrence). This species is considered native to CT and RI, and non-native elsewhere.

Veronica Reference: Pennell (1921). 1a. Racemes borne in the axils of leaves [Figs. 776, 777]; individual flowers subtended by very small, narrow bracts that are much smaller than the foliage leaves [Figs. 776, 777]; leaves opposite throughout; plants perennial 2a. Stems and usually the leaf blades conspicuously pubescent; styles 2.5–8 mm long; plants of mesic to dry-mesic habitats

P l a n tag i nac e a e   717

3a. Flowering stems creeping near the base, the tip ascending; leaf blades thick, subcoriaceous, permanently pubescent; corolla 4–8 mm wide; capsules distinctly exceeding the calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. officinalis 3b. Flowering stems ascending to erect; leaf blades thin, membranaceous, becoming glabrate; corolla 8–13 mm wide; capsules not, or only slightly, exceeding the calyx 4a. Leaf blades ovate, 1–2 times as long as wide; style 3–5 mm long; sepals subequal in length; stems 10–40 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. chamaedrys 4b. Leaf blades lanceolate to ovate, 2–4 times as long as wide; style 6–8 mm long; upper sepals much shorter than the lower sepals; stems 30–80 cm tall . V. austriaca 2b. Stems and leaf blades essentially glabrous; styles 1.5–4 mm long; plants of hydric habitats 5a. Leaf blades linear to lanceolate, mostly 4–20 times as long as wide; rachis of raceme flexuous; seeds 1.2–1.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. scutellata 5b. Leaf blades lanceolate or narrow-oblong to narrow-ovate or oblong-obovate, 1.5–5 (–8) times as long as wide; rachis of raceme straight; seeds up to 0.5 mm long 6a. Leaf blades borne on short, but evident, petioles [Fig. 776]; racemes with (4–) 10–25 (–30) flowers 7a. Leaf blades lanceolate to narrow-ovate or subtriangular (the lower leaf blades often more elliptic), broadest near the base, acute to subacute at the apex [Fig. 776]; style 2.5–3.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. americana 7b. Leaf blades oval or oblong to oblong-obovate, broadest near or distal to the middle, rounded at the apex; style 1.8–2.2 mm long . . . . . . . . . . . . . . V. beccabunga 6b. Leaf blades of the flowering stems sessile [Fig. 777]; racemes with (15–) 20– 54 (–60) flowers 8a. Leaf blades elliptic to elliptic-ovate, 1.5–3 times as long as wide [Fig. 777]; sepals acute to acuminate; corolla 5–8 mm wide; mature pedicels ascending or upcurved; capsules nearly or fully as wide as tall, scarcely notched at the apex; sterile autumnal shoots formed, with rounder and short-petiolate leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. anagallis‑aquatica 8b. Leaf blades lanceolate to narrow-oblong, (2.5–) 3–5 (–8) times as long as wide; sepals obtuse to subacute; corolla 3–5 mm wide; mature pedicels wide spreading; capsules wider than tall, usually with a conspicuous apical notch; sterile autumnal shoots not formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. catenata 1b. Racemes terminating the main stem [Figs. 778, 779]; flowers subtended by foliaceous bracts or by very small, narrow bracts in V. longifolia and V. spuria [Fig. 778]; upper leaves (i.e., the foliaceous ones that subtend flowers) alternate in most species [Fig. 778]; plants annual or perennial 9a. Plants perennial, from rhizomes, sometimes also rooting at the lower nodes of the stem 10a. Leaf blades 30–100 mm long, the principal ones sharply and conspicuously serrate, often some whorled on a given plant; style 4–10 mm long, ca. 2 times as long as the capsule; capsules pubescent with eglandular hairs; stems 3–15 dm tall 11a. Racemes several to many, arranged in a panicle; corolla 7–14 mm in diameter; stamens not exserted beyond corolla; pedicels (2–) 3–5 mm long, approximately as long as the calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. spuria 11b. Racemes solitary or a few together, not arranged in a panicle; corolla 6–8 mm in diameter; stamens exserted beyond corolla; pedicels 1–2 mm long, shorter than the calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. longifolia 10b. Leaf blades 5–40 mm long, entire or with inconspicuous and/or blunt teeth, none whorled [Fig. 779]; style 1–3.5 mm long, shorter than to about as long as the capsule; capsules pubescent, at least in part, with glandular hairs; stems 0.5–3 dm tall

7 18 tricolpates

12a. Corolla 10–15 mm wide; leaf blades cordate-orbicular to reniform, 5–10 mm long; bracteal leaves gradually reduced in size from foliage leaves (i.e., the flowers appearing axillary) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. filiformis 12b. Corolla 4–10 mm wide; leaf blades lanceolate or elliptic to ovate, 10–40 mm long; bracteal leaves abruptly reduced in size from foliage leaves (i.e., the flowers appearing in racemes) 13a. Capsule 4–7 mm tall, taller than wide; stems erect, pilose in the apical portion; plants of alpine communities . . . . . . . . . . . . . . . . . . . . . . . . . V. wormskjoldii 13b. Capsule 3–4 mm tall, wider than tall; stems creeping at the base or producing prostrate lower branches, minutely puberulent; plants predominantly of humandisturbed communities, sometimes in low-elevation forests . . . . . . V. serpyllifolia 9b. Plants annual, from fibrous roots 14a. Pedicels very short, up to 2 mm long; seeds 0.4–1.2 mm long; bracteal leaves abruptly reduced in size from the foliage leaves (i.e., the flowers appearing in racemes) 15a. Corolla largely or entirely white; leaves linear-oblong to oblanceolate, 3–10 times as long as wide; style 0.1–0.3 mm long; seeds 0.4–0.8 mm long . . . . . . . . . V. peregrina 15b. Corolla blue; leaves ovate to broad-elliptic, 1–2 times as long as wide; style 0.4–1.0 mm long; seeds 0.8–1.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. arvensis 14b. Pedicels longer, 6–40 mm long [Fig. 778]; seeds 1.2–3 mm long; bracteal leaves gradually reduced in size from foliage leaves (i.e., the flowers appearing axillary) [Fig. 778] 16a. Many of the leaf blades with 3–5 palmately arranged, tooth-like lobes; capsules without an apical notch or the notch inconspicuous; style 0.3–1.1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. hederaefolia 16b. Leaf blades toothed, lacking lobes [Fig. 778]; capsules with an evident, apical notch; style 1–3 mm long 17a. Pedicels 15–40 mm long in fruit [Fig. 778]; corolla (5–) 8–11 mm wide; styles 1.8–3 mm long; capsules 5–9 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. persica 17b. Pedicels 6–15 mm long in fruit; corolla 4–8 mm wide; styles up to 1.5 mm long; capsules 3–6 mm wide Fig. 776  Inflorescence and leaves of Veronica americana.

18a. Corolla blue; styles up to 1.5 mm long, extending beyond the apical notch of the capsule; capsules 2.5–4 × 3.5–6 mm, with 9–12 seeds per locule . . . . V. polita 18b. Corolla blue or white; styles up to 1 mm long, not extending beyond the apical notch of the capsules; capsule 4–6 × 3–4 mm, with 3–8 seeds per locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. agrestis 1. Veronica agrestis L. E green field speedwell. CT, MA, ME, NH, VT. Fields, roadsides, waste areas. 2. Veronica americana Schwein. ex Benth. N Fig. 776 American speedwell. Veronica beccabunga L. var. americana Raf. • CT, MA, ME, NH, VT. Stream shores, seeps, swamps, margins of pools. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable occurrence in RI and was unaware of any collections. 3. Veronica anagallis-aquatica L. E Fig. 777 blue water speedwell. Veronica anagallis L.; V. glandifera Pennell • CT, MA, ME, RI, VT. Stream shores, seeps, swamps, wet ditches. 4. Veronica arvensis L. E

Fig. 777  Inflorescence and leaves of Veronica anagallis-aquatica.

corn speedwell. CT, MA, ME, NH, RI, VT. Fields, lawns, roadsides, waste areas.

Pl a n tag i n ac e a e   719

5. Veronica austriaca L. ssp. teucrium (L.) D.A. Webb E broad-leaved speedwell. Veronica latifolia L.; V. teucrium L. • CT, MA, ME, NH, VT. Fields, roadsides, abandoned homesteads. 6. Veronica beccabunga L. E European speedwell. CT, MA, ME. River shores, stream banks, wet ditches, springs. 7. Veronica catenata Pennell

NC

water speedwell. Veronica catenata Pennell var. glandulosa (Farw.) Pennell; V. comosa Richter var. glaberrima (Pennell) Boivin; V. comosa Richter var. glandulosa (Farw.) Boivin; V. connata Raf. ssp. glaberrima Pennell; V. connata Raf. var. glaberrima (Pennell) Fern.; V. connata Raf. var. typica Pennell • MA, VT. Muddy shores and shallow water of streams. 8. Veronica chamaedrys L. E germander speedwell. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 9. Veronica filiformis L. E thread-stalked speedwell. CT, VT. Lawns, fields, gardens. 10. Veronica hederaefolia L. E ivy-leaved speedwell. CT; also reported from RI by George (1997), but specimens are unknown. Fields, roadsides, waste areas. 11. Veronica longifolia L. E long-leaved speedwell. Pseudolysimachion longifolium (L.) Opiz • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, abandoned homesteads, gardens. 12. Veronica officinalis L. E common speedwell. Veronica officinalis L. var. tournefortii (Vill.) Reichenb.; V. tournefortii Vill. • CT, MA, ME, NH, RI, VT. Forests, clearings, trail edges, fields, lawns, roadsides.

13. Veronica peregrina L. n purslane speedwell. 13a. Veronica peregrina L. var. typica Pennell; 13b. Veronica peregrina L. var. xalapensis (Kunth) Pennell; V. xalapensis Kunth • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, waste areas. 1a. Stem, sepals, and capsules glabrous . . . . . . . . . . . . . . . . . . 13a. V. peregrina ssp. peregrina 1b. Stems and commonly the sepals and capsules glandular-pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13b. V. peregrina ssp. xalapensis (Kunth) Pennell Subspecies peregrina is native and known from CT, MA, ME, NH, RI, VT. Subspecies xalapensis is non-native and known from CT, MA, ME, NH, RI, VT. The ranges of these two subspecies now show tremendous overlap, and subspecies xalapensis has become much more common in the East.

Fig. 778  Inflorescence and leaves of Veronica persica.

14. Veronica persica Poir. E Fig. 778 bird’s-eye speedwell. Pocilla persica (Poir.) Fourr. • CT, MA, ME, NH, RI, VT. Fields, roadsides, gardens, waste areas. 15. Veronica polita Fries E gray field speedwell. Pocilla polita (Fries) Fourr. • CT, ME. Fields, roadsides, waste areas. 16. Veronica scutellata L. N narrow-leaved speedwell. Veronica scutellata L. var. villosa Schumacher • CT, MA, ME, NH, RI, VT. Shorelines, swamps, marshes, peatlands. 17. Veronica serpyllifolia L. n Fig. 779 thyme-leaved speedwell.  17a. Veronica humifusa Dickson; V. serpyllifolia L. var. humifusa (Dickson) Vahl; V. tenella All.; Veronicastrum serpyllifolium (L.) Fourr. ssp. humifusum (Dickson) W.A. Weber; 17b. Veronicastrum serpyllifolium (L.) Fourr. • CT, MA, ME, NH, RI, VT. Fields, roadsides, lawns, gardens, waste areas, forests, shorelines, springs, logging roads, clearings.

Fig. 779  Inflorescence and leaves of Veronica serpyllifolia ssp. serpyllifolia.

72 0   tricolpate s

1a. Corolla blue, 5–8 (–10) mm wide; pubescence of pedicels, in part, of viscid or glandular hairs; filaments 2–4 mm long; racemes with 8–15 flowers; larger capsules 4–6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17a. V. serpyllifolia ssp. humifusa (Dickson) Syme 1b. Corolla pale blue with blue lines, 2–4 (–6) mm wide; pedicels puberulent with eglandular hairs; filaments 1–2.5 mm long; racemes with (12–) 20–40 flowers; capsules 3–4 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17b. V. serpyllifolia ssp. serpyllifolia Subspecies humifusa is native and known from ME, NH, VT; also reported from MA by Sorrie and Somers (1999), but specimens are unknown. Subspecies serpyllifolia is non-native and known from CT, MA, ME, NH, RI, VT. Subspecies humifusa is relatively northern and sometimes ascends to moderate elevation. Though it normally occurs in cleared areas (e.g., logging roads, pastures), it sometimes occurs within intact forests. 18. Veronica spuria L. E bastard speedwell. Pseudolysimachion spurium (L.) Rauschert; Veronica paniculata L. • NH, VT. Fields, roadsides, waste areas. 19. Veronica wormskjoldii Roemer & J.A. Schultes var. wormskjoldii

NC

American alpine speedwell. Veronica alpina L. var. terrae-novae Fern.; V. alpina L. var. unalaschcensis Cham. & Schlecht. • ME, NH. Alpine ravines, gullies, and snowbank communities.

Veronicastrum 1. Veronicastrum virginicum (L.) Farw. n Culver’s-root. Leptandra virginica (L.) Nutt.; Veronica virginica L. • CT, MA, ME, VT. Rich, deciduous forests, fields, roadsides, clearings, forest fragments. This species is native in CT, MA, and VT (though individual populations in those states may have originated from cultivation), and is non-native in ME.

Platanaceae Platanus 1a. Carpellate inflorescences paired (rarely solitary or in trios); middle lobe of leaf blade as long as or longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. hybrida 1b. Carpellate inflorescences solitary (rarely paired); lobes of leaf blade shorter than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. occidentalis 1. Platanus hybrida Brot. E London sycamore. Platanus acerifolia (Ait.) Willd. • MA. Railroads, waste areas. The exact origin of this European tree is unknown (Tutin 1993a). Some authorities suggest it is the hybrid product of Platanus occidentalis and P. orientalis L., having arisen in the 17th century. Others suggest it is a cultivar of P. orientalis. 2. Platanus occidentalis L. n American sycamore. CT, MA, ME, NH, RI, VT. River shores, riparian forests, swamps. Records from northern counties in ME represent cultivated or escaped populations. It is only (and formerly) native to the southern portion of the state.

Plu mb ag inac e a e   72 1

Plumbaginaceae Limonium 1. Limonium carolinianum (Walt.) Britt. N Carolina sea-lavender. Limonium angustatum (Gray) Small; L. carolinianum (Walt.) Britt. var. angustatum (Gray) Blake; L. carolinianum (Walt.) Britt. var. nashii (Small) Boivin; L. carolinianum (Walt.) Britt. var. trichogonum (Blake) Boivin; L. nashii Small; L. nashii Small var. albiflorum (Raf.) House; L. nashii Small var. angustatum (Gray) Ahles; L. nashii Small var. trichogonum (Blake) Blake; L. trichogonum Blake • CT, MA, ME, NH, RI; coastal region. Saline marshes, coastal strands and beaches.

Podostemaceae Podostemum 1. Podostemum ceratophyllum Michx. N horn-leaved riverweed. Podostemum abrotanoides Nutt. • CT, MA, ME, NH, RI, VT. Rivers with medium to fast current, frequently on submersed ledge, but also found underwater on cobble, gravel, and sand substrate. More frequent than realized, some populations extending ± continuously for a km or more.

Polemoniaceae Leptosiphon androsaceus Benth. was reported from MA by Collins (1901; as Gilia androsacea). The voucher specimen was later determined to be a plant allied to G. tricolor and became the type for G. tricolor var. longipedicellata (this taxon now considered to be a taxonomic synonym of G. tricolor ssp. diffusa). See Deane (1904) for discussion of this record. 1a. Bracts and calyx lobes spine-tipped; stigmas 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Navarretia 1b. Bracts and calyx lobes not spine-tipped (though sometimes awn-tipped); stigmas usually 3 2a. Leaf blades simple and entire; stamens attached at different heights on the corolla 3a. Most leaves opposite; corolla 12–30 mm wide; calyx with scarious or hyaline intervals between the green, ± herbaceous ribs, ruptured by developing capsule; seeds unchanged when wet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phlox 3b. Leaves alternate; corolla narrower than 10 mm; calyx of nearly uniform chartaceous texture, not ruptured by developing capsule; seeds becoming mucilaginous when wet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Collomia 2b. Leaf blades compound or dissected; stamens attached at the same height on the corolla 4a. Ultimate segments of leaf blades lanceolate to ovate; corolla campanulate [Fig. 782]; calyx herbaceous, somewhat accrescent in fruit . . . . . . . . . . . . . . . . . . . . . . . . Polemonium

72 2   tricolpate s

4b. Ultimate segments of leaf blades narrow-linear to narrow-oblanceolate; corolla salverform; calyx firm, not accrescent 5a. Corolla red on the abaxial surface, 20–40 mm long; inflorescence a slender, elongate thyrse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ipomopsis 5b. Corolla light purple to purple or white on the abaxial surface, 5–13 mm long; inflorescence of cymose clusters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gilia

Collomia 1. Collomia linearis Nutt. E narrow-leaved mountain-trumpet. Gilia linearis (Nutt.) Gray • CT, MA, ME, NH, VT. Yards, fields, railroads, wool waste, around dwellings.

Gilia Gilia capitata Sims ssp. capitata was reported from VT by Atwood et al. (1973). However, the voucher specimen was collected from cultivated material (i.e., this species is not known to be naturalized in VT; specimen at SJFM). 1a. Arachnoid hairs present in the leaf axils and often on the stem; capsule with 2–4 seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. interior 1b. Arachnoid hairs not present on vegetative organs (though other types of hairs may be present); capsule with 10–36 seeds 2a. Corolla ca. 2–3 times as long as calyx, the adaxial surface of connate portion (i.e., inside the basal tube) yellow with purple spots; axis of inflorescence with minute, black stipitate glands; capsule with 16–36 seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. tricolor 2b. Corolla ca. 1–2 times as long as calyx, the adaxial surface of connate portion light purple or white; axis of inflorescence eglandular; capsule with 10–18 seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. achilleifolia 1. Gilia achilleifolia Benth. ssp. multicaulis (Benth.) V. & A. Grant E blue gilia. Gilia multicaulis Benth.; Gilia multicaulis Benth. var. alba Milliken • MA, ME. Dumps, waste areas. 2. Gilia interior (Mason & A. Grant) A. Grant E inland gilia. Gilia tenuiflora Benth. ssp. interior Mason & A. Grant • MA. Wool waste. Reports of Gilia inconspicua (Sm.) Sweet in MA are based on a collection of this species. 3. Gilia tricolor Benth. ssp. diffusa (Congd.) Mason & A. Grant E bird’s-eye gilia. Gilia diffusa Congd.; G. tricolor Benth. var. longipedicellata Greenman in Deane • MA. Wool waste. This species is frequently cultivated and occasionally is found in compost heaps.

Ipomopsis 1. Ipomopsis rubra (L.) Wherry E standing-cypress skyrocket. Gilia rubra (L.) Heller; Polemonium rubrum L. • MA. Waste areas.

Navarretia 1. Navarretia leucocephala Benth. ssp. minima (Nutt.) Day E white-flowered pincushion-plant. Navarretia minima Nutt. • MA. Woolwaste. Reported as Gilia leucocephala Gray by Knowlton and Deane (1923b).

p o l e m o n i ac e a e   723

Phlox References: Wherry (1929, 1930, 1931). 1a. Petals notched at apex [Fig. 781]; leaf blades stiff, with a single primary vein; stems suffrutescent 2a. Leaf blades (15–) 20–40 (–60) mm long; axillary fascicles mostly lacking; apical sinus of petal 0.25–0.5 times as long as entire petal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bifida 2b. Leaf blades (5–) 8–20 mm long; axillary fascicles usually present and conspicuous [Fig. 781]; apical sinus of petal 0.12–0.25 times as long as entire petal . . . . . . . . . . . . . P. subulata 1b. Petals entire at the apex or sometimes emarginate to shallowly notched in P. divaricata; leaf blades herbaceous, with a midvein and several primary lateral veins (though the lateral veins often faint; with only a midvein in P. pilosa); stems herbaceous 3a. Style short, 1–4 mm long, scarcely (if at all) longer than the stigmas or the ovary; stamens and style at most reaching the midpoint of the basal, connate portion of corolla 4a. Upper leaves alternate; corolla commonly red, white, or variegated; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. drummondii 4b. Upper leaves opposite; corolla usually pink, blue, light blue-purple, or light redpurple ; plants perennial 5a. Leaf blades linear to lanceolate or narrow-ovate, tapering to a sharp, indurate awn-tip 0.5–3 mm long; basal, connate portion of corolla usually pubescent; both the reproductive stem, as well as the sterile basal offshoots erect to ascending . . . . P. pilosa 5b. Leaf blades elliptic or oblong to lanceolate or ovate, obtuse to acute at the apex, sometimes tapering to a short awn-tip up to 0.5 mm; basal, connate portion of the corolla glabrous; reproductive stem erect or decumbent at base, whereas the sterile, basal offshoots are prostrate and rooting . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. divaricata 3b. Style elongate, (12–) 14–26 mm long, much longer than the stigmas or the ovary; stamens and style reaching at least the orifice of the basal, connate portion of corolla 6a. Long-creeping, sterile, basal offshoots with spatulate leaf blades present at anthesis; flowering stems 10–40 (–50) cm tall; inflorescence and calyx stipitate-glandular; corolla 25–30 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. stolonifera 6b. Sterile, basal offshoots absent at anthesis or present and short, upcurving, and without spatulate leaf blades; flowering stems 30–200 cm tall; inflorescence and calyx glabrous or pubescent, but not glandular; corolla 12–22 mm wide 7a. Leaf blades ciliolate on the margin, with conspicuous lateral veins that connect near the edge to form a submarginal vein around the perimeter of the blade [Fig. 780]; anthers pale yellow to white . . . . . . . . . . . . . . . . . . . . . . . P. paniculata 7b. Leaf blades eciliate on the margin, without a submarginal, connecting vein; anthers yellow 8a. Calyx 6–8 (–10) mm long; stems streaked or spotted with red (very rarely green throughout), arising from rhizomes, with 7–15 (or more) nodes below the inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. maculata 8b. Calyx (8.5–) 10–13 mm long; stems without red streaks or spots, arising from decumbent stems, with 3–5 nodes below the inflorescence . . . . . . . . . . P. latifolia 1. Phlox bifida Beck var. bifida E sand phlox. VT. Forest edges, fields, lawns. 2. Phlox divaricata L. ssp. divaricata E wild blue phlox. CT, VT. Forested stream banks, forest fragments, waste areas.

72 4   tricolpate s

3. Phlox drummondii Hook. ssp. drummondii. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E annual phlox. CT. Waste areas. Reports of this species for VT (e.g., Atwood et al. 1973, Kartesz 1999) were based on a collection from a cultivated plant (i.e., this species is not known to be naturalized in VT; specimen originally at HNH). 4. Phlox latifolia Michx. E mountain phlox. Phlox ovata, auct. non L. • MA; western portion of state. Fields, gardens. 5. Phlox maculata L. ssp. maculata E spotted phlox. Phlox maculata L. var. odorata (Sweet) Wherry • CT, ME, VT. Fields, roadsides, waste areas, lawns, near dwellings. Previous reports of this species in MA were based on a cultivated specimen (fide Sorrie and Somers 1999). 6. Phlox paniculata L. E Fig. 780 fall phlox. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, gardens. 7. Phlox pilosa L. ssp. pilosa

NC

downy phlox. Phlox pilosa L. var. virens (Michx.) Wherry • CT; southern portion of state. Grassy banks, dry knolls. 8. Phlox stolonifera Sims E Fig. 780  Leaf blade of Phlox paniculata showing lateral veins connecting to form a submarginal vein.

creeping phlox. ME, VT. Fields, roadsides, lawns, gardens. ‌8 × 9. Phlox ×procumbens Lehm. is a very rare phlox hybrid known from ME. It is identified by its petals with a shallow apical notch, leaf blades with a single midvein (but the blades are not stiff as in P. subulata), and inflorescence with glandular hairs mixed with the eglandular, septate ones. 9. Phlox subulata L. ssp. subulata E Fig. 781 moss phlox. CT, MA, ME, NH, RI, VT. Gardens, roadsides, lawns, frequent about old cemeteries.

Polemonium Fig. 781  Habit of Phlox subulata.

Polemonium acutiflorum Willd. was reported from ME and NH by Magee and Ahles (1999), P. cuspidatum was reported from ME by Campbell et al. (1995), and P. occidentale Greene was reported from MA by Knowlton and Deane (1923b), but specimens are unknown. 1a. Corolla shorter than 5 mm; leaves with 5–15 leaflets; plants annual; stems with abundant stipitate-glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. micranthum 1b. Corolla 10–20 mm long; primary leaves with 11–29 leaflets; plants perennial; stems with few or no stipitate-glands (though eglandular hairs may be present) 2a. Leaflets shortly petiolulate, the petiolules mostly 0.1–1.5 mm long; stamens and style well exserted from corolla [Fig. 782] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. vanbruntiae 2b. Leaflets sessile; stamens and style not or shortly exserted from corolla 3a. Many or all of the pedicels as long as or longer than the calyx at anthesis; larger leaves with 11–17 leaflets; lowest bracteal leaves with 9–11 leaflets, nearly as large as the stem leaves; pedicels puberulent with eglandular hairs; corolla 12–16 mm long . . . . . . P. reptans 3b. Most of the pedicels shorter than the calyx at anthesis; larger leaves with 19–29 leaflets; lowest bracteal leaves with 3–7 reduced and crowded leaflets, much reduced compared with the stem leaves; pedicels pubescent with glandular hairs; corolla 15–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. caeruleum 1. Polemonium caeruleum L. E blue Jacob’s-ladder. ME, NH, VT. Fields, roadsides. 2. Polemonium micranthum Benth. E annual Jacob’s-ladder. Polemoniella micrantha (Benth.) Heller • MA. Waste areas, gardens.

p o l e m o n i ac e a e   7 25

3. Polemonium reptans L. var. reptans E spreading Jacob’s-ladder. CT, MA, NH, RI, VT. Fields, roadsides, abandoned homesteads, gardens. 4. Polemonium vanbruntiae Britt.

N C Fig. 782

bog Jacob’s-ladder. Polemonium caeruleum L. ssp. vanbruntiae (Britt.) J.F. Davids. • ME, VT. Swamps, pond shores, fens, stream-side meadows, rarely in seeps within rich, deciduous forests and along mesic roadsides.

Polygalaceae Polygala References: Gillett (1968), Wheelock (1891). 1a. Leaves few, clustered near the tip of the stem, the blades elliptic to oval; inflorescence with 1–4 flowers that are 13–19 mm long; androecium with 6 stamens; plants perennial from creeping rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. paucifolia 1b. Leaves usually many and spaced throughout the stem, the blades linear to narrow-elliptic or narrow-obovate; inflorescence with many flowers that are 3–6 mm long; androecium with 7 or 8 stamens; plants annual, biennial, or perennial, but without creeping rhizomes 2a. Chasmogamous (i.e., petaliferous) flowers borne in loose racemes, on spreading or recurving pedicels 1.5–3.5 mm long [Fig. 784]; cleistogamous flowers produced after the chasmogamous flowers, borne on secund, prostrate or shallowly subterranean racemes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. polygama 2b. Chasmogamous flowers borne in relatively more dense, spike-like racemes, the pedicels up to 1.5 mm long; cleistogamous flowers not produced 3a. Plants perennial, with clustered stems arising from a hard, knotty root; leaf blades (13–) 30–80 × 3–15 (–30) mm, irregularly serrulate . . . . . . . . . . . . . . . . . . . . . . . . . . P. senega 3b. Plants annual, the solitary stems arising from a taproot; leaf blades 10–40 × 1–5 mm, entire 4a. Wing sepals acuminate to awn-tipped at the apex [Fig. 783], minutely roughened on the adaxial surface; raceme 10–15 (–20) mm thick [Fig. 783] . . . . . . . . . . P. cruciata 4b. Wing sepals rounded to obtuse at the apex, not roughened on the adaxial surface; raceme 2–10 (–14) mm thick [Fig. 785] 5a. Leaves 1 at each node; wing sepals 2–5 (–6.3) mm long; racemes blunt at the apex (sometimes apiculate in P. nuttallii due to the flower buds); bracts persistent, each appearing as a minute, upcurved hook, remaining on the raceme axis after the falling of the fruits 6a. Wing sepals 3–5 (–6.3) mm long; ca. 2 times as long as the corolla; racemes 6–10 (–14) mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sanguinea 6b. Wing sepals 2–2.5 mm long, ± as long as the corolla; racemes 4–6 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. nuttallii 5b. Leaves 2–5 at each of the lower nodes (sometimes the upper nodes with only 1 leaf) [Fig. 785]; wing sepals 0.9–2.6 mm long; racemes tapering to a slender point; bracts deciduous from the base of the fruiting raceme, the fallen fruits leaving only a pedicel scar

Fig. 782  Flowers of Polemonium vanbruntiae showing exserted stamens and style.

72 6   tricolpate s

7a. Wing sepals usually ½ to ⅔ as long as fruit [Fig. 785]; most nodes (except sometimes the upper) bearing 2–5 leaves; flowers white to pink to purple; axis of raceme not or scarcely elongating in fruit, the lowest internode 0.3–1.7 (–3.3) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. verticillata 7b. Wing sepals ± as long as the fruit; most nodes (except the lower ones) bearing a single leaf; flowers predominantly green-white to white; axis of raceme elongating in fruit, the lowest internode (1.2–) 2.3–8.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. ambigua 1. Polygala ambigua Nutt.

NC

alternate milkwort. Polygala verticillata L. var. ambigua (Nutt.) Wood; P. verticillata L. var. dolichoptera Fern. • CT, MA, ME, NH, RI, VT. Fields, forest edges, roadsides, railroads, woodlands, forest clearings. This species utilizes anthropogenic habitats over most of New England but has been collected from relatively pristine sites in CT. Fig. 783  Inflorescence of Polygala cruciata.

2. Polygala cruciata L. ssp. aquilonia (Fern. & Schub.) A. Haines N Fig. 783 drum-heads milkwort. Polygala cruciata L. var. aquilonia Fern. & Schub. • CT, MA, ME, NH, RI; primarily along the coastal plain and restricted to the southern portion of northern New England states. Sandy and/or peaty soils of meadows, pond shores, swamps, and bogs. 3. Polygala nuttallii Torr. & Gray N Nuttall’s milkwort. CT, MA, RI. Dry, sandy fields and barrens, sand plains, woodlands. 4. Polygala paucifolia Willd. N fringed milkwort. Triclisperma paucifolia (Willd.) Nieuwl. • CT, MA, ME, NH, RI, VT. Forests, woodlands, field edges. This species shows three color forms in New England: pink to pink-red perianth (the most common), white, and light blue (the rarest). 5. Polygala polygama Walt. N Fig. 784 racemed milkwort. Polygala polygama Walt. var. obtusata Chod.; P. polygama Walt. var. ramulosa Farw. • CT, MA, ME, NH, RI, VT. Dry, sandy fields and barrens, sand plains, forest clearings. This species shows two color forms in New England: pink-red perianth (the most common) and white perianth (relatively rare). 6. Polygala sanguinea L. N

Fig. 784  Inflorescence of Polygala polygama.

blood milkwort. Polygala viridescens L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest clearings, trail and logging road edges. This species shows three color forms in New England: pink-red perianth (the most common), green perianth, and white perianth. 7. Polygala senega L. N Seneca milkwort. Polygala senega L. var. latifolia Torr. & Gray • CT, MA, ME, VT; western and northern New England. River banks and woodlands in limestone and marble bedrock regions, limestone headlands, also on northern, ice-scoured rivers, railroad embankments, roadside clearings in high-pH till regions. 8. Polygala verticillata L. N Fig. 785 whorled milkwort. Polygala verticillata L. var. isocycla Fern.; P. pretzii Pennell; Sexilia verticillata (L.) Raf. • CT, MA, ME, NH, RI, VT. Fields, clearings, ledges, roadsides, open woodlands.

Polygonaceae Fig. 785  Inflorescence and upper leaves of Polygala verticillata.

References: Hong et al. (1998), Mitchell and Dean (1978), Ronse Decraene and Akeroyd (1988), Ronse Decraene et al. (2000). 1a. Achenes trigonous and with 3 evident wing-angles; leaf blades 12–40 (–57) × 10–30 (–61) cm, cordate-ovate to deltate, borne on stout petioles; androecium with 9 stamens . . . . . . . . . Rheum

Po lyg onac e a e   7 27

1b. Achenes lenticular or trigonous, but without wings or with an partially encircling, marginal wing on a lenticular achene in Oxyria; leaf blades shorter and/or narrower, with relatively thinner petioles; androecium with 3–8 stamens (9 in Chorizanthe) 2a. Achenes with a partially encircling, marginal wing; perianth with 4 tepals . . . . . . Oxyria 2b. Achenes without wings; perianth with (4–) 5 or 6 tepals 3a. Outer 3 tepals of carpellate flowers each with a conspicuous spine (note: this most evident in fruit); plants clearly monoecious—the carpellate flowers with 6 tepals positioned below the staminate ones with 5 tepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Emex 3b. None of the tepals bearing spines (though with a spiny involucre in Chorizanthe); plants predominantly synoecious or dioecious (sometimes polygamous), the two types of flowers (when present) with the same number of tepals 4a. Flowers subtended by a 6-parted involucre, the involucre with 3 longer members alternating with 3 shorter members, each member tipped by an awn; androecium with 9 stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chorizanthe 4b. Flowers not subtended by an awn-tipped involucre; androecium with 3–8 stamens 5a. Tepals 6, sepaloid or herbaceous, in 2 series of 3—the outer series spreading to reflexed, the inner series erect and distinctly enlarged in fruit (not accrescent in R. acetosella) [Figs. 801, 803] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rumex 5b. Tepals (4–) 5, often petaloid, usually in a single series, all erect to ascending, of ± equal size relative to one another in fruit or the outer tepals slightly larger than the inner [Figs. 792, 794, 797] 6a. Tepals nearly distinct, remaining small in fruit; achene exserted beyond the remnant tepals for much of its length; filaments winged; nectaries stalked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fagopyrum 6b. Tepals connate at base for a distance up to 57% of their total length [Figs. 796, 797], at least somewhat accrescent in fruit; achene enclosed in remnant tepals or the tip exserted; filaments unwinged; nectaries absent or present and then not stalked 7a. Tepals with trifid veins (i.e., with three main branches at the base); flowers with nectaries, these appearing as mamillae around the stamens; stamens monomorphic, not enlarged at base 8a. Tepals connate at the base for ⅓ or less of their length; gynoecium with 3 carpels and 3 styles or the flowers sterile or replaced by bulbils; androecium with 8 stamens; plants perennial, with basally disposed leaves, unbranched stems, and a solitary, terminal inflorescence [Fig. 786] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bistorta 8b. Tepals connate for ⅓ or more their length into a cup-like structure (except in P. wallichii); gynoecium with 2 or 3 carpels and 1 style that is apically bifid or trifid (with 2 styles in P. virginiana); androecium with 5–8 stamens; plants annual or perennial, with chiefly cauline leaves, unbranched or branched stems, and commonly 2 or more terminal or terminal and axillary inflorescences (commonly solitary and terminal in P. amphibian and P. coccinea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Persicaria 7b. Tepals with dendritic veins (i.e., with a single main vein that smaller veins branch off from) or sometimes the vein unbranched [Fig. 797]; flowers without nectaries; stamens dimorphic, the inner larger, often enlarged at the base 9a. Erect to prostrate herbs up to 7 (–10) dm tall; stipules bilobed (usually not in P. articulatum); papillae absent from base of filaments; outer tepals keeled, but not winged, in fruit; flowers with 3–8 stamens . . . Polygonum

72 8   tricolpate s

9b. Robust, upright herbs 10–40 dm tall or vines; stipules truncate to obliquely pointed at apex (i.e., not bilobed); papillae present at the base of the filaments; outer tepals with a conspicuous keel or wing in fruit; flowers with 8 stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fallopia

Bistorta The genus Bistorta is here recognized separate from Persicaria following the phylogenetic work of Kim and Donoghue (2008). 1a. Inflorescence 3–6 × 0.5–1 cm, bearing sterile flowers in the apical half and bulbils in the basal half (or sometimes bulbils nearly throughout) [Fig. 786]; leaf blades 2–10 cm long, narrowed at the base; plants native, of alpine habitats, at elevations exceeding 1000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. vivipara 1b. Inflorescence 5–9 × 1–2 cm, bearing only fertile flowers; leaf blades 6–20 cm long, rounded to cordate at the base; plants escaped from cultivation . . . . . . . . . . . . . . . B. officinalis 1. Bistorta officinalis Delarbre E meadow bistort. Persicaria bistorta (L.) Sampaio; Polygonum bistortum L. • MA, ME; also reported from VT by Freeman (2005), but specimens are unknown. Gardens, waste areas, forest fragments. 2. Bistorta vivipara (L.) S.F. Gray

N C Fig. 786

alpine bistort. Persicaria vivipara (L.) R. Decr.; Polygonum viviparum L. • ME, NH, VT. Mesic to hydric situations in alpine areas, such as ravines, slides, ledges, plateaus, and snowbank communities.

Chorizanthe 1. Chorizanthe pungens Benth. var. pungens E Monterey spineflower. MA. Wool waste. Fig. 786  Inflorescence of Bistorta vivipara showing bulbils in basal half.

Emex 1. Emex spinosa (L.) Campd. E spiny threecorner-Jack. Rumex spinosus L. • MA. Waste areas.

Fagopyrum 1a. Tepals white to cream, 3–4 mm long; achenes 5–7 mm long; inflorescence corymb-like, clustered near the apex of the plant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. esculentum 1b. Tepals green, 2–3 mm long; achenes 5–5.5 mm long; inflorescence raceme-like, scattered along upper portion of stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. tataricum 1. Fagopyrum esculentum Moench E garden buckwheat. Fagopyrum fagopyrum (L.) Karst.; F. sagittatum Gilib.; F. vulgare Hill; Polygonum fagopyrum L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, most frequent in areas of cultivation. 2. Fagopyrum tataricum (L.) Gaertn. E green buckwheat. Polygonum tataricum L. • MA, ME, NH, VT. Fields, roadsides, waste areas, most frequent in areas of cultivation.

P o lyg o nac e a e   72 9

Fallopia Tepal measurements for the vining species do not include the narrow, decurrent portion that is adnate to the pedicel. Measurements and shape assessment of the leaf blades for non-vining species should be performed on midbranch leaves, as those near the apex of the branches are not typical (usually both smaller and relatively narrower). References: Kim et al. (2000), Freeman and Hinds (2005). 1a. Stems erect, stout, with hollow internodes; stigmas fimbriate; perianth accrescent 2a. Leaf blades broad-cuneate to truncate at the base, abruptly tapering to the apex, 5–15 × 2–10 cm, glabrous on the abaxial surface (though usually with blunt scabrules along the major veins) [Fig. 788, R]; midbranch inflorescences about as long as to longer than the subtending leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. japonica 2b. Leaf blades cordate at the base, gradually tapering to the apex, 10–30 × 7–20 cm, sparsely pubescent with long, multicelled hairs on the abaxial surface [Fig. 788, L]; midbranch inflorescences much shorter than the subtending leaves . . . . . F. sachalinensis 1b. Stems trailing or climbing, slender, solid; stigmas capitate; perianth usually not accrescent 3a. Stipules reflexed-bristly at the base; tepals margins white in life (often drying yellow-white) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. cilinodis 3b. Stipules glabrous; tepals margins green, green-white, yellow-white, or tinged with red in life (white in the introduced F. baldschuanica) 4a. Outer 3 tepals merely keeled or with a very narrow wing in fruit; achene striatepapillose, dull except for the lustrous angles [Fig. 787]; plants annual from a wiry taproot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. convolvulus 4b. Outer 3 tepals conspicuously winged in fruit; achenes smooth, lustrous [Fig. 789]; plants perennial from a rootstock or annual from a taproot 5a. Plants woody near the base (i.e., lianas); flowers white, borne in profuse, branched panicle-like inflorescences; axis of inflorescence brown-white to very pale brown; bracteoles subtending the pedicels (i.e., ocreolae) cream-colored, caudate-tipped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. baldschuanica 5b. Plants herbaceous throughout (i.e., vines); flowers green to green-white (infrequently brown-green or tinged with red), borne in raceme-like inflorescences or small, axillary fascicles; axis of inflorescence green or red (rarely pale); bracteoles subtending the pedicels green to brown, truncate or oblique at apex 6a. Fruiting tepals 3.4–5.1 mm long, with crenate or irregularly jagged and often crisped wings, the wings green to green-white, relatively poorly developed especially near the apex and only 0.25–1 mm wide; achenes 2.4–2.9 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. cristata 6b. Fruiting tepals 4.2–9.4 mm long, with entire to weakly undulate and plane wings, the wings cream to pale brown or tinged with red, well-developed even near the apex and 1–3 mm wide (rarely poorly developed in F. dumetorum); achenes 2.8–5.1 mm long 7a. Fruiting tepals 4.2–6.5 (–7) mm long, often abruptly contracted near the base and only shortly decurrent on the pedicel; achenes 2.8–3.5 (–3.7) mm long; perianth suborbicular to broad-oval or broad-oblong in side view; basal lobes of leaf blades mostly 3–8 mm long when measured parallel to petiole . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. dumetorum 7b. Fruiting tepals (6.2–) 6.5–9.4 mm long, usually gradually narrowed and long-decurrent on the pedicel; achenes (3.5–) 3.7–5.1 mm long; perianth oval to oval-obovate or narrow-obovate in side view; basal lobes of leaf blades mostly 9–19 mm long when measured parallel to the petiole . . . . . . . . . . . . F. scandens

73 0   tricolpate s

1. Fallopia baldschuanica (Regel) Holub E Chinese bindweed. Fallopia aubertii (Henry) Holub; Bilderdykia aubertii (Henry) Moldenke; Bilderdykia baldschuanica (Regel) D.A. Webber; Polygonum aubertii Henry; P. baldschuanica Regel; Reynoutria baldschuanica (Regel) Shinners • MA. Roadsides, waste areas, forest edges. 2. Fallopia cilinodis (Michx.) Holub N fringed bindweed. Bilderdykia cilinodis (Michx.) Greene; Polygonum cilinode Michx. var. laevigatum Fern.; Reynoutria cilinodis (Michx.) Shinners; Tiniaria cilinodis (Michx.) Small • CT, MA, ME, NH, RI, VT. Woodlands, forest borders, clearings, ledges, roadsides, waste areas, rocky banks. 3. Fallopia convolvulus (L.) A. Löve E Fig. 787 black bindweed. Bilderdykia convolvulus (L.) Dumort.; Polygonum convolvulus L. var. subulatum Lej. & Court.; Reynoutria convolvulus (L.) Shinners; Tiniaria convolvulus (L.) Webb & Moq. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, gardens, forest borders, yards. 4. Fallopia cristata (Engelm. & Gray) Holub N Fig. 787  Striate-papillose achene of Fallopia convolvulus with a dull surface.

crested bindweed. Bilderdykia cristata (Engelm. & Gray) Greene; B. scandens (L.) Greene var. cristata (Engelm. & Gray) C.F. Reed; Polygonum cristatum Engelm. & Gray; P. scandens L. var. cristatum (Engelm. & Gray) Gleason; Reynoutria scandens (L.) Shinners var. cristata (Engelm. & Gray) Shinners; Tiniaria cristata (Engelm. & Gray) Small • CT, MA, RI, VT. Woodlands, ridges, balds, rocky slopes. Some older reports of Fallopia dumetorum (e.g., Graves et al. 1910) actually refer to F. cristata. Freeman and Hinds (2005) chose to subsume F. cristata in F. scandens due to apparent intergrading morphology. Review of specimens shows that these two species are distinct. Work by Kim et al. (2000) supports this stance. The main problem is the presence of F. dumetorum, which sits in an intermediate position between these two species in regard to size of reproductive structures. Though it is true that F. cristata overlaps slightly with F. dumetorum and F. dumetorum overlaps slightly with F. scandens, there is no morphological overlap between F. cristata and F. scandens (i.e., confusion with F. dumetorum may be the cause for reporting morphological overlap). Therefore, F. cristata is recognized here as a native element of vining Fallopia with a narrower ecological amplitude than F. scandens. 5. Fallopia dumetorum (L.) Holub E thicket bindweed. Bilderdykia dumetorum (L.) Dumort.; Polygonum dumetorum L.; P. scandens L. var. dumetorum (L.) Gleason; Reynoutria scandens (L.) Shinners var. dumetorum (L.) Shinners; Tiniaria dumetorum (L.) Opiz • CT, MA, RI. Waste areas, roadsides, back dunes, pond shores. Further study is needed to determine the full distribution of this plant in New England; it may occur in additional states.

Fig. 788  Hairs on abaxial leaf blade surface of Fallopia sachalinensis (left), F. ×bohemica (middle), and F. japonica (right).

6. Fallopia japonica (Houtt.) R. Decr. var. japonica E Fig. 788R Japanese knotweed. Pleuropterus cuspidatus (Sieb. & Zucc.) Moldenke; Polygonum cuspidatum Sieb. & Zucc.; Reynoutria japonica Houtt. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, forest edges, river shores and banks. ‌ × 7. Fallopia ×bohemica (Chrtek & Chrtková) J.P. Bailey is an uncommon knotweed 6 hybrid known from CT, MA, ME, RI, VT. It is becoming increasingly more frequent on the landscape. Its leaf morphology is intermediate between F. japonica and F. sachalinensis (i.e., they tend to be large and are truncate to cordate at the base). The best discriminating character is the minute, but firm, stoutly conical, 1- or 2-celled hairs found on the abaxial leaf surface (this type of hair not found in the parental taxa; view at 20× or higher) [Fig. 788, M]. 7. Fallopia sachalinensis (F.S. Petrop. ex Maxim.) R. Decr. E Fig. 788L giant knotweed. Polygonum sachalinense F. Schmidt ex Maxim.; Reynoutria sachalinensis (F. Schmidt ex Maxim.) Nakai • CT, MA, ME, RI, VT. Fields, roadsides, waste areas, open rights-ofway, river banks.

Po lyg o nac e a e   73 1

8. Fallopia scandens (L.) Holub N Fig. 789 climbing bindweed. Bilderdykia scandens (L.) Greene; Polygonum dumetorum L. var. scandens (L.) Gray; P. scandens L.; Reynoutria scandens (L.) Shinners; Tiniaria scandens (L.) Small • CT, MA, ME, NH, RI, VT. Riparian forests, river banks, ledges. See discussion under Fallopia cristata.

Oxyria 1. Oxyria digyna (L.) Hill

N C Fig. 790

mountain-sorrel. Rumex digyna L. • NH; northern portion of state. Alpine gullies, ravines, and brooksides.

Persicaria Reference: Hinds and Freeman (2005).

Fig. 789  Relatively smooth achene of Fallopia scandens with a lustrous surface.

1a. Inflorescence panicle-like; perianth subrotate; tepals connate less than 20% of their total length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. wallichii 1b. Inflorescence raceme-like (sometimes consisting of few flowers and ± capitate or sometimes appearing panicle-like when the upper plant is highly branched); perianth campanulate to narrow-campanulate; tepals connate 25–66% of their total length 2a. Leaf blades extending down the petioles as a wing of tissue that clasps the stem (petiole sometimes only distally winged in P. chinensis); achenes conspicuously cellularreticulate with longitudinal rows of aereolae 3a. Flowers with 2 styles; achenes biconvex; abaxial surface of leaf blade glandularpunctate; peduncle stipitate-glandular near apex just below flowers . . . . . P. nepalensis 3b. Flowers with 3 styles; achenes trigonous; abaxial surface of leaf blade without glandular dots; peduncle stipitate-glandular throughout . . . . . . . . . . . . . . . . . . P. chinensis 2b. Leaf blades without winged petioles, not clasping the stem; achenes lacking a conspicuous cellular-reticulate pattern 4a. Stems armed with reflexed prickles [Fig. 791]; principal leaf blades hastate or sagittate at base (sometimes truncate in P. perfoliata) [Fig. 791]; flowers comparatively few, in a short to capitate raceme-like inflorescence 0.4–2 (–3) cm long 5a. Sheathing stipules apically foliaceous and expanded; leaf blades subpeltate; perianth persistent in fruit, forming a thickened, blue covering over the achene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. perfoliata 5b. Sheathing stipules scarious or hyaline, not expanded upward; leaf blades basifixed; perianth persistent in fruit, but neither thickened nor blue 6a. Leaf blades lanceolate to elliptic, 5–31 mm wide, sagittate at the base, the basal lobes directed backward; peduncles glabrous; styles trifid, 1–1.5 mm long; achenes trigonous, 3–3.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sagittata 6b. Leaf blades deltate, 10–150 mm wide, hastate at the base, the basal lobes divergent [Fig. 791]; peduncles hispid, often stipitate-glandular as well; styles bifid, 0.5–0.6 mm long; achenes biconvex, 3.5–5 mm long . . . . . . . . . . . . . . . . . . P. arifolia 4b. Stems unarmed; leaf blades cuneate to cordate at base, lacking prolonged basal lobes; flowers usually many, in an elongate raceme-like inflorescence 1–50 cm long 7a. Styles elongate, 2–4 mm long, indurate and persistent in fruit, hooked at the tip [Fig. 795]; perianth 4-merous [Fig. 795], not accrescent; flowers conspicuously deflexed in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. virginiana 7b. Styles wanting or up to 2 mm long (up to 4 mm long in short-stamened flowers of P. amphibia), deciduous in fruit, not hooked at tip; perianth 5-merous, accrescent in fruit; flowers ascending to spreading (rarely the short pedicels drooped)

Fig. 790  Habit of Oxyria digyna.

73 2   tricolpate s

8a. Plants perennial with rhizomes or stolons 9a. Inflorescence terminal, solitary or paired, densely flowered, mostly 10–20 mm wide; tepals deep pink to red 10a. Widest leaf blades 17–29 (–32) mm wide; principal emersed leaves with petioles 1–9 mm long, commonly with small, red-brown spots due to infection by Puccinia polygoni-amphibii, usually green in drying; primary inflorescences 13–28 (–30) mm long; floating leaf blades cuneate to rounded (infrequently truncate) at the base; rhizome 1.5–3.5 mm thick between leafless nodes, bearing few-branched to moderately branched roots; stipules of emersed shoots usually developing sheathing stipules with a foliaceous, outward-flange at the summit, these pubescent with wide-ascending to spreading hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. amphibia 10b. Widest leaf blades (23–) 32­–63 mm wide; principal emersed leaves with petioles 5–28 (–38) mm long, lacking the mentioned fungal infection, usually tinged with red or brown in drying; primary inflorescences (25–) 29– 90 mm long; floating leaf blades cordate to rounded at the base; rhizome (2.8–) 3–6.5 mm thick between leafless nodes, bearing highly branched roots; sheathing stipules of emersed shoots without a green, outward flange, glabrous or pubescent with appressed hairs . . . . . . . . . . . . . . . . . . P. coccinea 9b. Inflorescence terminal and often also axillary, often remotely flowered at the base, mostly 2–5 mm wide; tepals pink, white, and/or green 11a. Tepals spotted with numerous yellow to brown or red-brown glandular dots [Fig. 794] 12a. Inflorescence relatively sparsely flowered, usually interrupted in the basal portion with non-overlapping bracteoles; bracteoles subtending the pedicels (i.e., ocreolae) long-ciliate with hairs (0.2–) 0.3–1.9 (–2.2) mm long; leaf blades 6–24 mm wide . . . . . . . . . . . . . . . . . . . . . . . . (in part) P. punctata 12b. Inflorescence relatively densely flowered, usually continuous throughout with contiguous or even overlapping bracteoles (rarely the lowest nodes separate); bracteoles subtending the pedicels usually eciliate or sometimes the lower bracteoles with short cilia up to 0.4 (–1) mm long; leaf blades 20–45 mm wide . . . . . . . . . . . . . . . P. robustior 11b. Tepals not spotted with glandular dots (rarely with a few, pale, scale-like glands in P. hydropiperoides, but these not providing an even dotting over the tepals) 13a. Hairs of sheathing stipules (i.e., ocreae) gradually enlarged at base, adnate to stipule for 28–66% of their total length; perianth usually pink to red or cream-white (rarely purple or green-white to green) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. hydropiperoides 13b. Hairs of the sheathing stipules conspicuously enlarged at base, adnate to the stipule for (0–) 13–28% of their total length; perianth greenwhite to green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. setacea 8b. Plants annual from fibrous roots and/or a taproot 14a. Sheathing stipules (i.e., ocreae) lacking a prominent ring of cilia around the summit, sometimes lacerate into segments in age 15a. Perianth consisting of 4 (–5) tepals; tepals conspicuously 3-nerved, each nerve forking near the end into 2 recurved branches [Fig. 792]; inflorescence relatively slender, 5–11 mm wide, the axis often arched or drooping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lapathifolia

P o lyg o nac e a e   73 3

15b. Perianth consisting of 5 tepals; tepals obscurely nerved; inflorescence relatively thick, 10–15 mm wide, the axis usually straight or with a slight arch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pensylvanica 14b. Sheathing stipules ciliate with a prominent ring of bristles around the summit [Fig. 793] 16a. Tepals spotted with numerous yellow to brown glandular dots [Fig. 794] 17a. Perianth consisting of 4 (–5) tepals; inflorescences terminating the main stem and often axillary, commonly with overlapping bracteoles (i.e., ocreolae), interrupted by small leaves; achenes dull . . . . . . . P. hydropiper 17b. Perianth consisting of 5 tepals; inflorescences terminating the main stem or branches, commonly with non-overlapping bracteoles (at least in the basal portion), without small leaves; achenes lustrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) P. punctata 16b. Tepals not spotted with glandular dots [Fig. 792] 18a. Leaf blades ovate, 3–16 cm wide; sheathing stipules horizontally divergent at the summit creating a conspicuous flange . . . . . P. orientalis 18b. Leaf blades narrow-lanceolate to elliptic or oblanceolate, 1–3 (–4) cm wide; sheathing stipules without a horizontally spreading summit 19a. Upper portion of the stem and peduncles stipitate-glandular; inflorescence 3–10 cm long, lax or drooping . . . . . . . . . . . . . . . . P. careyi 19b. Upper portion of the stem and peduncles without stipitate glands; inflorescence 1–4.5 (–6.5) cm long, commonly erect to ascending 20a. Sheathing stipules long-ciliate with marginal hairs (3–) 4–12 mm long; bracteoles of the flowers long-ciliate with marginal hairs (0.5–) 1–4 (–6) mm long; achenes trigonous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. longiseta 20b. Sheathing stipules ciliate with marginal hairs (0.3–) 1–3.5 (–5) mm long [Fig. 793]; bracteoles of the flowers short-ciliate with marginal hairs (0.1–) 0.2–2 (–2.7) mm long; achenes biconvex or both biconvex and trigonous in the same inflorescence 21a. Inflorescence loose and usually interrupted in the basal portion, 2–4 mm wide; leaf blades linear to narrow-lanceolate, (2–) 4–10 (–23) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. minor 21b. Inflorescence dense, usually uninterrupted, 4–12 mm wide; leaf blades lanceolate to narrow-ovate, (2–) 10–25 (–40) mm wide 22a. Inflorescence 6–12 mm wide, obloid to short-cylindrical; tepals 2–3 mm long at anthesis, enlarging in fruit and then up to 4 mm long, prominently, but minutely, patterned with a raised reticulum near the base; leaf blades commonly with a purple blotch on the adaxial surface [Fig. 793] . . P. maculosa 22b. Inflorescence 4–7 mm wide, cylindrical to subconical; tepals 1.8–2 mm long at anthesis, enlarging in fruit and then up to 2.6 mm long, faintly reticulate-patterned, if at all, near the base; leaf blades usually unblotched . . . . . P. puritanorum 1. Persicaria amphibia (L.) S.F. Gray ssp. laevimarginata (Hultén) Soják N water smartweed. Persicaria amphibia (L.) S.F. Gray var. stipulacea (Coleman) Hara; Polygonum amphibium L. ssp. laevimarginatum Hultén; P. amphibium L. var. natans Michx.; P. amphibium L. var. stipulaceum Coleman; P. coccineum Muhl. ex Willd. var. rigidulum (Sheldon) Stanford; P. natans (Michx.) Eat. • CT, MA, ME, NH, RI, VT. Shallow, still or slow-moving water of lakes and

73 4   tricolpate s

rivers, marshes, shorelines, swamps, ditches. Numerous attempts have been made to subdivide the Persicaria amphibia complex, some with a priori assumptions of the morphological variation. Though some intermediate material does exist, two taxa can be recognized in North America—an aquatic species with an amphibious form when water levels decline (P. amphibia) and a wetland species that can form floating leaves when submerged (P. coccinea). It is known that these taxa have a genetic basis; however, several characters that have been deemed critically important are known to be almost completely controlled by environment (specifically, depth of inundation). These include pubescence of the leaf blades, presence/absence of stipitate-glands on the peduncle, and apex shape of leaf blades. Preliminary surveys suggest about 10% of our material shows substantial intermediacy. These plants are probably best interpreted as F₁ hybrids. European P. amphibia (ssp. amphibia) is very similar to North American P. amphibia (ssp. laevimarginata), except that it usually has truncate- to cordate-based floating leaf blades that frequently show stout, forward-pointing cilia along the margin and never produces foliaceous, outward-flanged stipules on emersed forms. 2. Persicaria arifolia (L.) Haroldson N Fig. 791 halberd-leaved tearthumb. Polygonum arifolium L.; P. arifolium L. var. lentiforme Fern. & Grisc.; P. arifolium L. var. pubescens (Keller) Fern.; Tracaulon arifolium (L.) Raf. • CT, MA, ME, NH, RI, VT. Swamps, marshes, pool margins, stream banks, ditches. 3. Persicaria careyi (Olney) Greene N Carey’s smartweed. Polygonum careyi Olney • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, clearings, shorelines, logging roads. Fig. 791  Leaves and inflorescences of Persicaria arifolia.

4. Persicaria chinensis (L.) H. Gross E Chinese smartweed. Polygonum chinense L. • MA. Fields, roadsides, waste areas. 5. Persicaria coccinea (Muhl. ex Willd.) Greene N scarlet smartweed. Persicaria amphibia (L.) S.F. Gray var. emersa (Michx.) Hickman; P. muehlenbergii (S. Wats.) Small; Polygonum amphibium L. var. coccineum (Muhl. ex Willd.) Farw.; P. amphibium L. var. emersum Michx.; P. coccineum Muhl. ex Willd.; P. coccineum Muhl. ex Willd. var. terrestre Willd.; P. muehlenbergii S. Wats. • CT, MA, ME, NH, RI, VT. Shorelines, marshes, swamps, ditches. See Persicaria amphibia for discussion of this complex. 6. Persicaria hydropiper (L.) Opiz E water-pepper smartweed. Polygonum hydropiper L. • CT, MA, ME, NH, RI, VT. Mesic to hydric soil of fields, ditches, clearings, shorelines, river banks, waste areas, and swamps. 7. Persicaria hydropiperoides (Michx.) Small N false water-pepper smartweed. Persicaria hydropiperoides (Michx.) Small var. breviciliata (Fern.) C.F. Reed; P. hydropiperoides (Michx.) Small var. euronotora (Fern.) C.F. Reed; P. opelousana (Ridd. ex Small) Small; Polygonum hydropiperoides Michx.; P. hydropiperoides Michx. var. adenocalyx (Stanford) Gleason; P. hydropiperoides Michx. var. breviciliatum Fern.; P. hydropiperoides Michx. var. bushianum Stanford; P. hydropiperoides Michx. var. digitatum Fern.; P. hydropiperoides Michx. var. euronotorum Fern.; P. hydropiperoides Michx. var. opelousanum (Riddell ex Small) Riddell ex W. Stone; P. opelousanum Riddell ex Small; P. opelousanum Riddell ex Small var. adenocalyx Stanford • CT, MA, ME, NH, RI, VT. Wet soil or shallow water of rivers, lakes, and swamps. 8. Persicaria lapathifolia (L.) S.F. Gray N Fig. 792 dock-leaved smartweed. Persicaria tomentosa (Schrank) Bickn.; Polygonum lapathifolium L.; P. lapathifolium L. var. ovatum A. Braun; P. lapathifolium L. var. prostratum C.F.H. Wimmer; P. lapathifolium L. var. salicifolium Sibthorp; P. scabrum Moench • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, shorelines, ditches, swamps, river banks. This species is a widespread diploid native to the New and Old World. 9. Persicaria longiseta (Bruijn) Kitagawa E

Fig. 792  Tepals of Persicaria lapathifolia showing veins with recurved branches.

Oriental lady’s-thumb smartweed. Persicaria caespitosa (Blume) Nakai var. longiseta (Bruijn) Reed; Polygonum caespitosum Blume var. longisetum (Bruijn) Steward; P. longisetum Bruijn • CT, MA, ME, NH, VT; also reported from RI by George (1997), but specimens are unknown. Fields, roadsides, waste areas, riparian forests.

Po lyg onac e a e   73 5

10. Persicaria maculosa S.F. Gray E Fig. 793 lady’s-thumb smartweed. Persicaria persicaria (L.) Small; P. vulgaris Webb & Moq.; Polygonum dubium Stein; P. persicaria L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, gardens, shorelines, ditches, yards. 11. Persicaria minor (Huds.) Opiz E small smartweed. Polygonum minus Huds.; P. minus Huds. var. subcontinuum (Meisn.) Fern. • CT, MA; also reported from VT by Hinds and Freeman (2005), but specimens are unknown. Roadsides, waste areas. 12. Persicaria nepalensis (Meisn.) H. Gross E Nepalese smartweed. Polygonum alatum Hamilton ex D. Don; P. nepalense Meisn. • CT, MA. River banks, gravel river beaches.

13. Persicaria orientalis (L.) Spach E prince’s-feather smartweed. Polygonum orientale L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, dumps, gardens.

Fig. 793  Ciliate stipule and blotched leaf blade of Persicaria maculosa.

14. Persicaria pensylvanica (L.) G. Maza N Pennsylvania smartweed. Persicaria bicornis (Raf.) Nieuwl.; P. longistyla (Small) Small; Polygonum bicorne Raf.; P. longistylum Small; P. pensylvanicum L.; P. pensylvanicum L. var. eglandulosum J.C. Myers; P. pensylvanicum L. var. genuinum Fern.; P. pensylvanicum L. var. laevigatum Fern.; P. pensylvanicum L. var. nesophilum Fern. • CT, MA, ME, NH, RI, VT. Shorelines, swamps, marshes, fields, roadsides, ditches, waste areas. 15. Persicaria perfoliata (L.) H. Gross E Asiatic tearthumb. Ampelygonum perfoliatum (L.) Roberty & Vautier; Polygonum perfoliatum L. • CT, MA, RI. Fields, roadsides, waste areas.

16. Persicaria punctata (Ell.) Small N Fig. 794 dotted smartweed. Persicaria punctata (Ell.) Small var. confertiflora (Meisn.) D. Löve & J.-P. Bernard; P. punctata (Ell.) Small var. leptostachya (Meisn.) Small; Polygonum acre Kunth; P. punctatum Ell.; P. punctatum Ell. var. confertiflorum (Meisn.) Fassett; P. punctatum Ell. var. leptostachyum (Meisn.) Small; P. punctatum Ell. var. parviflorum Fassett • CT, MA, ME, NH, RI, VT. Swamps, marshes, shorelines, including brackish to fresh-tidal river shores, shallow water of pools and ponds. 17. Persicaria puritanorum (Fern.) Soják NC Puritan smartweed. Polygonum puritanorum Fern. • MA, me, rI. Coastal plain pond shores. This species has often been synonymized with Persicaria maculosa. However, recent study showed it is derived from hybridization between P. hydropiperoides and P. lapathifolia (Kim et al. 2008). 18. Persicaria robustior (Small) Bickn. N stout dotted smartweed. Persicaria puncta (Ell.) Small var. robustior Small; Polygonum punctatum Ell. var. majus (Meisn.) Fassett; P. punctatum Ell. var. robustius Small; P. robustius (Small) Fern. • CT, MA, ME, NH, RI; primarily on the coastal plain. Shorelines, marshes, swamps, wetland margins. 19. Persicaria sagittata (L.) H. Gross N arrow-leaved tearthumb. Polygonum sagittatum L.; P. sagittatum L. var. gracilentum Fern.; Tracaulon sagittatum (L.) Small; T. sagittatum (L.) Small var. gracilentum (Fern.) C.F. Reed; Truellum sagittatum (L.) Soják • CT, MA, ME, NH, RI, VT; throughout. Marshes, stream banks, swamps, shorelines, ditches, low fields. 20. Persicaria setacea (Baldw.) Small

NC

bristly smartweed. Persicaria setacea (Baldw.) Small var. interjecta (Fern.) C.F. Reed; P. setacea (Baldw.) Small var. tonsa (Fern.) C.F. Reed; Polygonum hydropiperoides Michx. var. setaceum (Baldw.) Gleason; P. setaceum Michx. var. interjectum Fern.; P. setaceum Michx. var. tonsum Fern. • MA, RI; primarily on the coastal plain, disjunct in Hamden County, MA. Pond

Fig. 794  Gland-dotted tepals of Persicaria punctata.

73 6   tricolpate s

shores, streams, marshes. Our specimens in New England are much closer to Persicaria hydropiperoides than many treatments present (as evidenced by the characters they use in their identification keys). 21. Persicaria virginiana (L.) Gaertn. N Fig. 795 jumpseed. Antenoron virginianum (L.) Roberty & Vautier; Polygonum virginianum L.; P. virginianum L. var. glaberrimum (Fern.) Steyermark; Tovara virginiana (L.) Raf.; T. virginiana (L.) Raf. var. glaberrima Fern. • CT, MA, NH, RI, VT. Rich, deciduous, upland and riparian forests. 22. Persicaria wallichii Greuter & Burdet var. wallichii E garden smartweed. Aconogonum polystachyum (Wallich ex Meisn.) Haraldson; Polygonum polystachyum Wallich ex Meisn. • MA. Saline marshes.

Polygonum Fig. 795  Inflorescence of Persicaria virginiana showing deflexed fruits and persistent styles.

Most species of New England Polygonum produce two types of fruits—summer achenes that are brown to black (rarely otherwise) and frequently with surface ornamentation and late-season achenes that are ± olive, smooth, and significantly larger than summer achenes. Late-season achenes are not provided for in the following key. Some species have leaves along the main axes with considerably larger blades than those on the branches (often 3 or more times as long). Such plants are referred to as heterophyllous. Other species have leaves of approximately equal size (or the leaves of the main axes are only slightly larger). These plants are referred to as homophyllous. In the following key, measurements of stipule length must be made on intact stipules (not on those that have disintegrated into fibers). Also, as the perianth is accrescent, it should be measured in fruit. Note also that the leaves are often shed later in the season, leaving only the bracts, which can be substantially smaller and differ in length to width ratios from the leaves. Polygonum argyrocoleon Stued. ex Kunze was reported from MA by Magee and Ahles (1999) and from VT by Kartesz (1999). The MA record was based on a specimen of P. aviculare s.l.—Ahles 77772 (MASS!). The VT report was erroneous. References: Löve and Löve (1956), Costea and Tardif (2003b). 1a. Pedicels articulated near the base; tepal midveins unbranched; flowers with 8 stamens; sheathing stipules truncate to scarcely bilobed at apex . . . . . . . . . . . . . . . . . . . . . P. articulatum 1b. Pedicels articulated at or near the summit; tepal midveins usually dendritically branched; flowers with (3–) 5 (–8) stamens; sheathing stipules bilobed 2a. Stems and branches sharply quadrangular, erect; flowers and fruits in the axils of tiny leaves, giving the appearance of a terminal inflorescence [Fig. 798]; achenes black-brown to black, minutely granular-roughened except on the angles (and often with a variably sized smooth region on the interior of each face prior to maturity) 3a. Pedicels straight, the flowers and fruits erect or ascending; leaf blades pleated with 2 longitudinal folds, minutely serrate on the margin . . . . . . . . . . . . . . . . . . . . . . . . . . P. tenue 3b. Pedicels abruptly recurved, the flowers and fruits soon nodding [Fig. 798]; leaf blades without longitudinal folds, with entire margins . . . . . . . . . . . . . . . . . . . . P. douglasii 2b. Stems and branches terete or 8- to 16-angled, prostrate to erect; flowers and fruits in most species in the axils of normal or slightly reduced leaves, giving the appearance of axillary inflorescences; achenes yellow-green, olive, light brown or brown to black, papillose, smooth, or minutely roughened over each face 4a. Tepals with yellow-green to yellow margins 5a. Tepals basally connate for 40–55% of their length [Fig. 796]; achenes uniformly papillose; leaf blades blue-green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. achoreum 5b. Tepals basally connate for 20–38% of their length; achenes striate-papillose, smooth, or irregularly roughened; leaf blades usually bright green to yellow-green (note: color ± retained in drying)

Po lyg onac e a e   7 37

6a. Leaf blades elliptic to ovate, (8–) 10–30 mm wide; achenes striate-papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. erectum 6b. Leaf blades linear to narrow-elliptic, 1–8 (–10) mm wide; achenes smooth or irregularly roughened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) P. ramosissimum 4b. Tepals with white to pink to red margins 7a. Inflorescence with reduced bracteal leaves usually shorter than to as long as the flowers they subtend (except sometimes the lower bracts, which may be foliaceous) 8a. Achenes striate-papillose, 1.7–2.8 mm long; tepals 1.9–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. arenarium 8b. Achenes smooth or irregularly roughened, 3–5 mm long; tepals 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. bellardii 7b. Inflorescence with conspicuous bracteal leaves that are often several times longer than the flowers they subtend 9a. Outer 3 tepals flat or folded, of approximately equal width and length to the inner tepals and not or scarcely concealing them [Fig. 797]; plants usually of inland, non-saline habitats such as roadsides, sidewalks, and disturbed lots (rarely coastal) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aviculare 9b. Outer 3 tepals cucullate, much wider and often longer the inner tepals, partially or completely concealing them [Fig. 799]; plants usually of brackish and saline habitats such as coastal marshes and dunes (sometimes inland in P. buxiforme and P. ramosissimum) 10a. Leaf blades pale green to white-green, somewhat to strongly glaucous; tepals loosely ascending and not investing apical portion of achene 11a. Leaf blades pale green, only somewhat glaucous; sheathing stipules (i.e., ocreae) of lower nodes 4–7 (–8) mm long, with 3–7 nerves; tepals obovate; achenes 3.5–5 mm long (up to 6.5 mm long in late-season collections) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. oxyspermum 11b. Leaf blades white-green, distinctly glaucous; sheathing stipules of lower nodes 7–10 mm long, with 8–16 nerves; tepals oval; achenes 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. glaucum 10b. Leaf blades green, blue-green, or yellow-green, sometimes tinged with red; tepals ± erect and rather closely investing achene (though achene may be exserted beyond perianth) 12a. Leaf blades linear to narrow-elliptic, mostly 4–12 times as long as wide; achenes smooth or irregularly roughened; plants heterophyllous or nearly homophyllous; androecium with 3–6 stamens . . . . (in part) P. ramosissimum 12b. Leaf blades oblanceolate to oblong or elliptic-oblong, mostly 2–4 times as long as wide; achenes striate-papillose, uniformly granular-roughened, or irregularly roughened; plants homophyllous; androecium with 6–8 stamens 13a. Achenes uniformly granular-roughened, abruptly tapered to a slender beak, 2.8–4 mm long (up to 5 mm long in late-season collections); tepals 3–5 mm long, without a pouch-like swelling near the base; plants prostrate to, more commonly, ascending . . . . . . . . . . . . . . . . . . . . . P. fowleri 13b. Achenes striate-papillose or irregularly roughened, gradually tapering to apex, 2–2.8 mm long; tepals 2–3 mm long, often with a pouchlike protrusion near the base; plants usually prostrate and mat-forming . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. buxiforme

73 8   tricolpate s

1. Polygonum achoreum Blake N Fig. 796 leathery knotweed. Polygonum erectum L. ssp. achoreum (Blake) A. & D. Löve • ME, VT. Fields, roadsides, waste areas, railroads, lawns. The report of this species in CT by Magee and Ahles (1999) is based on a specimen of Polygonum buxiforme—Ahles 85330 (MASS!). 2. Polygonum arenarium Waldst. & Kit. E European knotweed. RI. Fields, roadsides, waste areas. 3. Polygonum articulatum L. N

Fig. 796  Fruiting tepals of Polygonum achoreum.

coastal jointed knotweed. Delopyrum articulatum (L.) Small; Polygonella articulata (L.) Meisn. • CT, MA, ME, NH, RI, VT. Sand soils of fields, roadsides, waste areas, grasslands, railroads, and dunes. 4. Polygonum aviculare L. E Fig. 797 dooryard knotweed. 4a. Polygonum aequale Lindm.; P. arenastrum Jord. ex Boreau; P. aviculare L. var. arenastrum (Jord. ex Boreau) Rovy; P. aviculare L. var. depressum Meisn.; P. microspermum Jord. ex Boreau; 4b. Polygonum aequale Lindm. ssp. oedocarpum Lindm.; P. neglectum Bess.; 4c. Polygonum aviculare L. var. vegetum Ledeb.; P. heterophyllum Lindm.; P. monspeliense Pers.; 4d. Polygonum aviculare L. var. angustissimum Meisn.; P. heterophyllum Aschers. & Graebn. var. angustissimum (Meisn.) Lindm.; P. heterophyllum Aschers. & Graebn. ssp. rurivagum (Jord. ex Boreau) Lindm.; P. rurivagum Jord. ex Boreau • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads, yards, shorelines, cracks in pavement. 1a. Tepals basally connate 40–57% of their length; leaves usually homophyllous or nearly so 2a. Tepals green or red-brown with white margins, basally connate 40–57% of their length, the outer ones with unbranched midveins or with 1 or 2 lateral veins; leaf blades 2.8–5.7 (–6.5) times as long as wide . . . 4a. P. aviculare ssp. depressum (Meisn.) Arcang.

Fig. 797  Fruiting tepals of Polygonum aviculare ssp. aviculare showing subequal tepals.

2b. Tepals green with pink to red (rarely white) margins, basally connate 28–48% of their length, the outer ones with midveins usually showing 2 or more lateral veins; leaf blades (3.4–) 4.2–9.2 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) 4b. P. aviculare ssp. neglectum (Bess.) Arcang. 1b. Tepals basally connate (15–) 20–40 (–42)% of their length [Fig. 797]; leaves homophyllous or, more commonly, heterophyllous 3a. Leaf blades 6–20 mm wide, 2–4.5 times as long as wide; flowers often aggregated at tips of stem and branches, in (1–) 3- to 8-flowered clusters; achenes enclosed in or barely exserted from tepals at maturity . . . . . . . . . . . . . . . . . . . . . . . 4c. P. aviculare ssp. aviculare 3b. Leaf blades 0.5–6.8 (–8) mm wide, (3.4–) 4.2–15 (–19) times as long as wide; flowers usually uniformly distributed along stems and branches, in 1- to 3 (–5)-flowered clusters; achenes often exserted from tepals at maturity 4a. Sheathing stipules 7–12 mm long, many and closely veined, the gaps between the veins mostly 1–3 times the width of the veins . . . . . . . . . . . . . . . . . . . . . . . . . . . 4d. P. aviculare ssp. rurivagum (Jord. ex Boreau) Berher 4b. Sheathing stipules 4–8 mm long, with relatively fewer and more distant veins, the gaps between the veins mostly 3–5 times the width of the veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) 4b. P. aviculare ssp. neglectum (Bess.) Arcang. Subspecies depressum is known from CT, MA, ME, NH, RI, VT. Subspecies neglectum is known from CT, MA, ME, NH, RI, VT. Subspecies aviculare is known from CT, MA, ME, NH, RI, VT. Subspecies rurivagum is known from CT, MA, ME, RI; also reported from NH by Seymour (1982), but specimens are unknown. Subspecies aviculare is far over-reported, most collection are in fact ssp. depressum. In addition to characters used in the key, perianth length and achene length often help to differentiate these two subspecies—ssp. aviculare has tepals mostly 2.4–4.8 mm and achenes mostly mostly 2.4–3.7 mm and ssp. depressum has tepals mostly 2–2.7 (–2.9) mm and achenes mostly 1.8–2.8 mm.

P o lyg o nac e a e   73 9

5. Polygonum bellardii All. E Bellard’s knotweed. MA. Waste areas, dumps. 6. Polygonum buxiforme Small N prairie knotweed. Polygonum aviculare L. var. littorale (Link) Mert.; P. littorale Link • CT, MA, ME, NH, RI. Atlantic coast beaches and shorelines, saline to brackish marshes, rarely inland on shorelines. Costea and Tardiff (2003b) chose to ally Polygonum buxiforme to P. aviculare because they noted that the outer tepals of members of the P. aviculare complex are not always flat. Though that is correct, real differences in the tepal morphology still exist between P. aviculare and P. buxiforme (i.e., simply because the tepals of P. aviculare have been poorly described in the literature does not mean they are similar to P. buxiforme). Polygonum buxiforme has outer tepals that are saccate at the base and the perianth is 0.9–1.3 times as long as wide (vs. non-saccate tepals and perianth 1.5–2.9 times as long as wide in P. aviculare). Given that P. buxiforme also differs in distribution (it is native to North America), it is not here considered part of the largely Eurasian P. aviculare. Polygonum buxiforme has been attributed to VT based on two vegetative collections—Goshen, Dutton s.n. (VT!) and Burlington, Dutton s.n. (VT!). They likely represent collections from the P. aviculare complex but are equivocal and cannot stand as vouchers for P. buxiforme. 7. Polygonum douglasii Greene N Fig. 798 Douglas’ knotweed. Polygonum douglasii Greene var. latifolium (Engelm.) Greene • ME, NH, VT; limited in ME to western portion of the state. Thin soil of ledges, cliff bases, and rocky woodlands. 8. Polygonum erectum L.

NC

upright knotweed. Polygonum aviculare L. var. erectum (L.) Roth ex Meisn. • CT, MA, ME, NH, RI, VT. Waste areas, roadsides, yards, river shores, railroads. 9. Polygonum fowleri B.L. Robins. ssp. fowleri N Fowler’s knotweed. Polygonum allocarpum Blake; P. buxifolium Nutt. ex Bong. • ME. Atlantic coast beaches and shorelines, saline to brackish marshes. The concept of Polygonum fowleri followed here includes P. fowleri s.s. (2n=40) with fleshy, scarcely ridged stems, red-tinged leaves, and nearly veinless stipules that are weakly dissected and P. allocarpum (2n=60) with less-fleshy, ridged stems, ± green leaves, and somewhat veined stipules that, in maturity, are dissected into numerous filaments. The correlations are not perfect and the character states are found intermixed.

Fig. 798  Inflorescence of Polygonum douglasii.

10. Polygonum glaucum Nutt. N seaside knotweed. Polygonum maritimum, auct. non L. • CT, MA, RI. Sandy, Atlantic coast beaches. 11. Polygonum oxyspermum C.A. Mey. & Bunge ex C.F. Ledeb. ssp. raii (Bab.) Webb & Chater

NC

Ray’s knotweed. Polygonum raii Bab. • ME; eastern portion of state. Atlantic coast beaches and dunes hollows. 12. Polygonum ramosissimum Michx. n Fig. 799 yellow-flowered knotweed. 12a. Polygonum atlanticum (B.L. Robins.) Bickn.; P. exsertum Small; P. triangulum Bickn.; 12b. Polygonum prolificum (Small) B.L. Robins.; P. ramosissimum Michx. var. prolificum Small • CT, MA, ME, NH, RI, VT. Saline and brackish marshes, roadsides, waste areas, dumps. 1a. Flowers evidently exserted from their bracteoles (i.e., ocreolae), on pedicels (2–) 2.5–6 mm long; plants heterophyllous; leaf blades obtuse to acuminate at apex, often bright-green to yellow-green, even in drying; tepals with yellow to yellow-green margins, only rarely with white to pink margins . . . . . . . . . . . . . . . . . . . . . . . . . 12a. P. ramosissimum ssp. ramosissimum 1b. Flowers barely or not exserted from their bracteoles, on pedicels 1–2 mm long; plants nearly homophyllous; leaf blades rounded to obtuse at apex, usually blue-green when fresh, drying brown to brown-black; tepals with white to pink margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12b. P. ramosissimum ssp. prolificum (Small) Costea & Tardif

Fig. 799  Fruiting tepals of Polygonum ramosissimum showing outer tepals larger and cucullate.

740 tricolpates

Subspecies ramosissimum is known from CT, MA, ME, NH, RI. It is native along the coast and has been introduced to anthropogenic sites on the interior of New England (e.g., CT, MA, ME, RI). Report of this subspecies in VT by Seymour (1982) was based on a specimen of Polygonum aviculare ssp. aviculare—Day 150 (VT!). Subspecies prolificum is known from CT, MA, ME, NH, RI. It is native along the coast. 13. Polygonum tenue Michx.

NC

slender knotweed. Polygonum tenue Michx. var. protrusum Fern. • CT, MA, ME, NH, RI, VT. Dry soil of sandy fields, roadsides, woodlands, ridges, balds, and ledges.

Rheum 1. Rheum rhabarbarum L. E garden rhubarb. Rheum rhaponticum, auct. non L. • CT, MA, ME, NH, VT. Fields, roadsides, waste areas, abandoned homesteads.

Rumex Pedicel measurements in the identification key include the actual pedicel (the proximal portion below the articulation point) and the pseudopedicel (the distal portion above the articulation point that is formed by the stipe-like hypanthium base). Complete specimens with mature fruit are crucial for confident identification of most species in our area. Rumex conglomeratus Murr. was reported from MA by Mosyakin (2005), but specimens are unknown. Rumex sanguineus L. was reported from CT by Hultén and Fries (1986), but specimens are unknown. Reference: Rechinger (1937). 1a. Leaf blades sagittate or hastate; plants dioecious 2a. Leaf blades predominantly sagittate; inner tepals of carpellate flowers 4–6 mm long in fruit, at least 1 of the them developing a tubercle; tepals of staminate flowers 2–3 mm long; achenes 2–2.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. acetosa 2b. Leaf blades predominantly hastate; inner tepals of carpellate flowers 1.2–3.5 mm long in fruit, none of them developing tubercles; tepals of staminate flowers 1.5–2 mm long; achenes 0.9–1.5 mm long 3a. Inner tepals of carpellate flowers 1.2–1.7 (–2) mm long in fruit, about as long as the achene, unexpanded, not reticulate-veiny; pedicels articulated near summit just below origin of tepals; stems arising from thin, creeping rhizomes . . . . . . . . . . . . . R. acetosella 3b. Inner tepals of carpellate flowers 2.5–3.5 mm long in fruit, much surpassing the achene, accrescent and becoming reticulate-veiny; pedicels articulated in the basal portion; stems arising from a taproot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. hastatulus 1b. Leaf blades without basal lobes; plants synoecious, monoecious, or polygamous 4a. Inner tepals (i.e., valves) with prominent teeth or spines on the margins [Fig. 801] 5a. Margins of the inner tepals with 2 or 3 (–4) slender spines 0.5–2.7 mm long; achenes 0.9–1.75 mm long; basal leaf blades narrow-lanceolate to lanceolate or oblonglanceolate, 4–7 times as long as wide; all 3 of the inner tepals with an enlarged tubercle on the midrib; plants fibrous-rooted annuals 6a. Abaxial leaf blade surface and branches of inflorescence glabrous or inconspicuously papillose; leaf blades narrow-cuneate at base (rarely to broadcuneate); tubercle of inner tepals usually smooth (infrequently finely striate or obscurely pitted) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. maritimus 6b. Abaxial leaf blade surface and branches of inflorescence papillose-pubescent; leaf blades truncate to subcordate at base (rarely broad-cuneate); tubercule of inner tepals distinctly reticulate-pitted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. persicarioides

P o lyg o nac e a e   741

5b. Margins of the inner tepals with 2–6 teeth (0.2–) 0.8–1.5 mm long [Fig. 801]; achenes (1.8–) 2–2.8 mm long; basal leaf blades oblong or oblong-lanceolate to ovate-oblong or obovate-elliptic, up to 4 times as long as wide; only 1 of the inner tepals with an enlarged tubercle on the midrib (all 3 with tubercles in R. violascens); plants taprooted perennials (usually annual or biennial in R. violascens) 7a. Inner tepals all bearing an enlarged tubercle on the midrib, with marginal teeth 0.2–0.5 mm long; basal leaf blades cuneate to rounded at the base (rarely truncate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. violascens 7b. Only 1 of the inner tepals with an enlarged tubercle on the midrib, with marginal teeth 0.8–1.5 mm long; basal leaf blades truncate to cordate at the base (rarely broad-rounded) 8a. Basal leaf blades 20–40 cm long; tubercles smooth [Fig. 801]; branches of inflorescence ascending, usually diverging from primary axis at an angle of 30–45 degrees; stems 60–120 (–150) cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . R. obtusifolius 8b. Basal leaf blades 4–10 (–15) cm long; tubercles usually verrucose; branches of inflorescence spreading, usually diverging from primary axis at an angle of 60–90 degrees; stems 20–60 (–100) cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. pulcher 4b. Inner tepals with entire or obscurely erose margins 9a. Stems with lateral branches and/or axillary leaf tufts, lacking a basal rosette of leaves [Fig. 802]; leaf blades usually pale-green or glaucous, with flat, entire to undulate margins 10a. Pedicels 10–17 mm long, 3–5 times as long as associated inner tepals, articulated near base; tubercle extending below the base of perianth [Fig. 803] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. verticillatus 10b. Pedicels (2–) 3–8 mm long, 1–2.5 times as long as associated inner tepals, articulated in the proximal ⅓ (sometimes near base); tubercle not extending below base of perianth 11a. Inner tepals with a broad tubercle, the tubercle more than half as wide and nearly as long as its associated inner tepal; leaf blades mostly 7–10 times as long as wide; plants predominantly of coastal marshes and shorelines . . . . . R. pallidus 11b. Inner tepals with a narrow tubercle, the tubercle less than half as wide and much shorter than its associated tepal; leaf blades mostly 2.5–6 times as long as wide; plants predominantly of freshwater wetlands and inland disturbed habitats 12a. Inner tepals 4.5–6 × 3–4.5 (–5) mm, 1–3 of them developing tubercles; leaf blades lanceolate to elliptic-lanceolate or broad-lanceolate, the larger 30–55 mm wide, mostly 2.5–4 times as long as wide; achenes 2.5–3.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. altissimus 12b. Inner tepals (2–) 2.5–3.5 (–3.8) × (2–) 2.5–3 (–3.5) mm, usually all 3 of them developing tubercles; leaf blades narrow-lanceolate, the larger 10–40 (–50) mm wide, mostly 5–6 times as long as wide [Fig. 802]; achenes 1.7–2.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. triangulivalvis 9b. Stems lacking both lateral branches and prominent axillary leaf tufts, with a usually conspicuous rosette of basal leaves [Fig. 800]; leaf blades dark green to red-green, with crenate, undulate, and/or crisped margins 13a. Plants with rhizomes; lower leaf blades broad-ovate to orbicular, ± as long as wide; inner tepals without tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. alpinus 13b. Plants with vertical taproots, lacking rhizomes; lower leaf blades narrowlanceolate or lanceolate to oblong-lanceolate or narrow-ovate-triangular, usually more than 3 times as long as wide; inner tepals with or without tubercles

742  tricolpates

14a. Inner tepals lacking well-developed tubercles (rarely the midrib of 1 tepal slightly thickened in R. longifolius) 15a. Pedicels with a distinct articulation point that is swollen and thicker the remaining pedicel; inner tepals broad-orbicular to reniform, (4.5–) 5–6 (–7) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. longifolius 15b. Pedicels with an indistinct articulation point that is scarcely, if at all, thicker than the remaining pedicel; inner tepals ovate or broad-ovate-triangular to suborbicular, 5–10 (–12) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . R. occidentalis 14b. At least 1 of the inner tepals with a well-developed tubercle 16a. All 3 of the inner tepals with a tubercle, the tubercles equal in size or nearly so; pedicels with an indistinct articulation point that is scarcely, if at all, thicker than the remaining pedicel; native plants of wetlands and shores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. britannica 16b. One to 3 of the inner tepals with a tubercle, when 3, then 1 of the tubercles distinctly larger or smaller than the others; pedicels with a distinct articulation point that is swollen and thicker than the remaining pedicel; non-native plants primarily of disturbed habitats, occasionally in wet places such as ditches and shorelines 17a. Inner tepals 5–8 (–10) × 5–9 (–10) mm, usually only 1 of the tepals with a tubercle (rarely as many as 3 tepals with tubercles); tubercle (or the largest one) up to 33% as long as associated inner tepal measuring from the base of the tubercle; achenes 3–3.5 mm long; leaf blades broadlanceolate to oblong-lanceolate, with plane or slightly crisped margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. patientia 17b. Inner tepals 3.5–6 × 3–5 mm, usually all of the tepals with a tubercle (rarely only 1); largest tubercle 40–60% as long as associated inner tepal measuring from the base of the tubercle; achenes 2–3 mm long; leaf blades lanceolate to narrow-lanceolate, with prominently crisped margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. crispus 1. Rumex acetosa L. E common dock. Acetosa pratensis P. Mill.; Rumex acetosa L. ssp. pratensis (P. Mill.) Blytt & Dahl • CT, MA, ME, NH, VT. Fields, roadsides. The report of Rumex arifolius All. (as R. montanus Desf.) in VT by Dole (1937) was based on a specimen of R. acetosa. 2. Rumex acetosella L. ssp. pyrenaicus (Pourret ex Lapeyr.) Akeroyd E sheep dock. Acetosella vulgaris (Koch) Fourr. ssp. pyrenaica (Pourret ex Lapeyr.) A. Löve; Rumex acetosella L. var. pyrenaicus (Pourret ex Lapeyr.) Timbal-Lagrave; R. pyrenaicus Pourret ex Lapeyr. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, shorelines, clearings. Our material of Rumex acetosella (i.e., ssp. pyrenaicus) is characterized by simple basal lobes and inner tepals becoming adnate to the fruit. Other subspecies of R. acetosella are generally considered not to occur in North America (e.g., forms with inner tepals easily separated from fruit; forms with multifid basal lobes), but their distributions are poorly known and they are probably overlooked. 3. Rumex alpinus L. E alpine dock. ME, VT. Fields, roadsides. 4. Rumex altissimus Wood E pale dock. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 5. Rumex britannica L. N greater water dock. Rumex orbiculatus Gray; R. orbiculatus Gray var. borealis Rech. f. • CT, MA, ME, NH, RI, VT. Swamps, marshes, including fresh to brackish types, shorelines, ditches.

Po lyg on ac e a e   743

‌ × 6. Rumex ×dissimilis Rech. f. is a very rare, partially sterile, hybrid dock known 5 from MA, RI. It most closely resembles R. britannica, especially in the long pedicels, relatively broad fruiting inner tepals (in side view), and relatively large tubercles. All three (rarely only 2) inner tepals of a flower produce an enlarged tubercle, 1 of which is larger than the other 2 (all equal or subequal in R. brittanica), the largest tubercle 65–75% as long as the associated inner tepal (measured from the base of the tubercle; the largest tubercle 40–60% as long as the associated inner tepal in R. crispus). The leaf blades show ± crisped margins, and the pedicels show an evident articulation point and average shorter than is usual for R. britannica (5–13 mm long in R. britannica and (3–) 4–8 mm long in R. crispus). 6. Rumex crispus L. ssp. crispus E Fig. 800 curly dock. Lapathum crispum (L.) Scop. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads, shorelines, wetland margins, upper edge of coastal beaches. 6 × 8. This very rare, dock hybrid is known from ME. It has ovate to orbicular inner tepals 4–6.4 mm long, only one of which usually with a tubercle, and achenes 2.6–3 mm long (usually 3 mm or longer in R. longifolius). ‌6 × 10. Rumex ×pratensis Mert. & Koch is a rare, partially sterile, hybrid dock known from MA, ME, NH, RI, VT. Its sterility manifests as fruiting perianths of different shapes and sizes on the same plant. The hybrid has inner tepals with very small teeth along the margins mostly shorter than 0.8 mm tall and leaf blades that are mostly 2–4 times as long as wide, which is intermediate between the parents (mostly 3.2–5.8 times as long as wide in R. crispus; mostly 1.2–2.8 times as long as wide in R. obtusifolius). Length to width measurements should be performed on the larger leaf blades.

Fig. 800  Habit of Rumex crispus with a basal rosette of leaves and leaf blades with crisped margins.

7. Rumex hastatulus Baldw. E wild dock. MA. Atlantic coast beaches. 8. Rumex longifolius DC. E yard dock. Rumex domesticus Hartman • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, lawns, yards, shorelines. 8 × 10. This very rare dock hybrid is known from VT. It shows very small teeth on the margins of the inner tepals mostly shorter than 0.8 mm tall. It was found growing with both parents. 9. Rumex maritimus L. E golden dock. Lapathum minus Lam.; Rumex aureus P. Mill. • MA. Ballast. 10. Rumex obtusifolius L. ssp. obtusifolius E Fig. 801 bitter dock. Acetosa oblongifolia (L.) A. & D. Löve; Rumex obtusifolius L. ssp. agrestis (Fries) Danser • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, shorelines. Two subspecies of this plant are known to occur in North America. Subspecies obtusifolius is marked by a perianth with a single tubercle and prominent, marginal teeth. Subspecies sylvestris Čelak is marked by a perianth with 3 tubercles and short marginal teeth (sometimes the teeth essentially absent). The latter subspecies is rare in North America and is not known to occur in New England. However, North American authors have traditionally not recognized these different forms and, therefore, the distribution of ssp. sylvestris is poorly known. 11. Rumex occidentalis S. Wats.

nC

western dock. Rumex aquaticus L. ssp. fenestratus (Greene) Hultén; R. aquaticus L. var. fenestratus (Greene) Dorn; R. aquaticus L. ssp. occidentalis (S. Wats.) Hultén; R. fenestratus Greene; R. occidentalis S. Wats. var. fenestratus (Greene) Lepage • MA, ME, VT. Swamps, marshes, shorelines. This species appears to be native in ME (Aroostook and Washington Counties) and VT, and introduced elsewhere.

Fig. 801  Fruiting tepals of Rumex obtusifolius.

744 tricolpate s

12. Rumex pallidus Bigelow N seabeach dock. MA, ME, NH, VT; coastal region, disjunct on Lake Champlain in VT. Atlantic coast beaches and marshes, rarely on inland lake shores. 13. Rumex patientia L. E patience dock. Lapathum hortense Lam. • CT, MA, ME, NH, VT. Fields, roadsides, waste areas. 14. Rumex persicarioides L. n American golden dock.  14a. Rumex fueginus Phil.; R. maritimus L. var. athrix St. John; R. maritimus L. ssp. fueginus (Phil.) Hultén; R. maritimus L. var. fueginus (Phil.) Dusen; 14b. Rumex maritimus L. var. persicarioides (L.) R.S. Mitchell • CT, MA, ME, NH, RI, VT. Saline and brackish marshes, Atlantic coast shores, fields, gardens. 1a. Tubercles ± narrow-lanceolate, 0.3–0.4 mm wide, less than ½ as wide as the associated inner tepals exluding the marginal spines, acute to subacute at the apex, brown to red-brown in life; marginal spines of inner tepals 1–3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14a. R. persicaroides var. fueginus (Phil.) A. Haines 1b. Tubercles ± elliptic, 0.4–0.6 mm wide, almost as wide as the inner tepals excluding the marginal spines, obtuse at the apex, cream to white-yellow in life; marginal spines of inner tepals 1–1.5 (–1.7) mm long . . . . . . . . . . . . . . . . . . . . . 14b. R. persicarioides var. persicarioides Variety fueginus is known from CT, MA, ME, NH, RI, VT. It occurs along the coast as a native halophyte and is introduced inland in fields and disturbed habitats. Variety persicarioides is known from CT, MA, RI. It is restricted to coastal habitats where it is native. 15. Rumex pulcher L. E fiddle dock. MA. Roadsides, waste areas. 16. Rumex triangulivalvis (Danser) Rech. f. n Fig. 802 Fig. 802  Habit of Rumex triangulivalvus showing lateral branches and axillary leaf tufts.

white dock. Rumex salicifolius Weinm. ssp. triangulivalvis Danser; R. salicifolius Weinm. var. triangulivalvis (Danser) C.L. Hitchc. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads, shorelines. This species appears to be native to VT (where it is found on shorelines) and introduced elsewhere. 17. Rumex verticillatus L. N Fig. 803 swamp dock. CT, MA, RI, VT; southern and western New England. Swamps, low riparian forests, shorelines. 18. Rumex violascens Rech. f. E violet dock. MA. Waste areas.

Portulacaceae Fig. 803  Fruiting tepals of Rumex verticillatus showing the tubercle extending below base of perianth.

Reference: Mitchell (1993). 1a. Leaves opposite [Figs. 804, 805]; flowers with 3 or 5 stamens 2a. Plants perennial, from a corm (annual from a minute tuber-like body in 1 species); stems with 2 leaves; leaf blades 30–200 mm long; petals 9–15 mm long; androecium with 5 stamens; capsules with 3–6 seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Claytonia 2b. Plants annual; stems with many leaves; leaf blades 3–20 mm long; petals 1–2 mm long; androecium with 3 stamens; capsules with 1–3 seeds . . . . . . . . . . . . . . . . . . . . (in part) Montia 1b. Leaves alternate; flowers with 3–15 stamens 3a. Flowers sessile or subsessile; gynoecium with 4–8 stigmas; fruit a pyxis; ovary partly inferior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Portulaca

P o rt u l ac ac e a e   745

3b. Flowers pedicellate; gynoecium with 3 stigmas; fruit a capsule; ovary superior 4a. Petals red, 4–15 mm long; androecium with 3–15 stamens; capsules with 10–20 seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calandrinia 4b. Petals white to pale pink, 4–6.5 mm long; androecium with 3 stamens; capsules with 1–3 seeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Montia

Calandrinia 1. Calandrinia ciliata (Ruiz & Pavón) DC. E fringed redmaids. Calandrinia ciliata (Ruiz & Pavón) DC. var. menziesii (Hook.) J.F. Macbr.; Talinum ciliatum Ruiz & Pavón • MA. Fields, waste areas.

Claytonia Claytonia perfoliata Donn ex Willd. ssp. perfoliata was attributed to NH based on the following specimen—7 Aug 1915, Wellman s.n. (NEBC!). The label states it was a garden weed and is not here taken as naturalized. 1a. Plants annual or biennial from a minute, tuber-like structure 1–5 mm in diameter or perennial from rhizomes; inflorescence multibracteate . . . . . . . . . . . . . . . . . . . . . . . . . . . C. sibirica 1b. Plants perennial from globose tubers 10–50 mm in diameter; inflorescence unibracteate (rarely with an additional bract) 2a. Leaf blades linear to narrow-oblanceolate, 2–15 mm wide [Fig. 804]; bract subtending lowest pedicel of inflorescence herbaceous, firm . . . . . . . . . . . . . . . . . . . . . . . . . . . C. virginica 2b. Leaf blades broad-oblanceolate to spatulate-oblong or oval, (5–) 10–30 (–33) mm wide; bract subtending lowest pedicel of inflorescence scarious, weak . . . . . . . . . . C. caroliniana 1. Claytonia caroliniana Michx. N Carolina spring-beauty. Claytonia caroliniana Michx. var. lewisii McNeill; C. caroliniana Michx. var. spathulifolia (Salisb.) W.H. Lewis; C. spathulifolia Salisb. • CT, MA, ME, NH, VT. Mesic, deciduous forests and forest edges in upland and riparian settings. 2. Claytonia sibirica L. E Siberian spring-beauty. Claytonia sibirica L. var. bulbifera Gray; Montia sibirica (L.) T.J. Howell; M. sibirica (L.) T.J. Howell var. bulbifera (Gray) B.L. Robins. • MA. Waste areas, near buildings. 3. Claytonia virginica L. N Fig. 804

Fig. 804  Flowers and leaves of Claytonia virginica.

Virginia spring-beauty. Claytonia virginica L. var. hammondiae (Kalmbacher) Doyle, Lewis, & Snyder; C. robusta (Somes) Rydb. • CT, MA, RI, VT; southern and western New England. Mesic, deciduous forests, forest edges, meadows.

Montia 1a. Sepals 1–2 mm long; petals 1–2 mm long; seeds 0.5–1.2 mm long; leaves opposite [Fig. 805], the blades 3–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. fontana 1b. Sepals 3–7 mm long; petals 4–6.5 mm long; seeds 1.2–2.6 mm long; leaves alternate, the blades 10–60 (–100) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. linearis 1. Montia fontana L.

N C Fig. 805

water montia. Claytonia fontana (L.) R.J. Davis; Montia fontana L. var. lamprosperma (Cham.) Fenzl; M. lamprosperma Cham. • ME; coastal. Rock crevices, peaty turf, stream margins, tidal marshes, pools, and forests near the Atlantic coast. The report of this species in NH and VT by Miller (2003) was erroneous.

Fig. 805  Flowers and leaves of Montia fontana.

746  tricolpates

2. Montia linearis (Dougl. ex Hook.) Greene E narrow-leaved montia. Claytonia linearis Dougl. ex Hook.; Montiastrum lineare (Dougl. ex Hook.) Rydb. • MA. Fields, waste areas.

Portulaca 1a. Stems densely pubescent at the nodes; petals 15–25 mm long, white or various shades of yellow or red; leaf blades linear, subterete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grandiflora 1b. Stems glabrous; petals 3–4.6 mm long, yellow; leaf blades spatulate to obovate, flat (but fleshy) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. oleracea 1. Portulaca grandiflora Hook. E cultivated purslane. CT, MA, RI, VT. Waste areas, gardens, dumps. This species has escaped from cultivation and often demonstrates flore pleno (i.e. “double corolla”). 2. Portulaca oleracea L. E common purslane. Portulaca neglecta Mackenzie & Bush; P. retusa Engelm. • CT, MA, ME, NH, RI, VT. Gardens, fields, waste areas, coastal beaches.

Primulaceae Phylogenetic investigation based on DNA sequences and morphology has shown the Primulaceae, as traditionally defined, to be polyphyletic (Källersjö et al. 2000). In order that the taxonomic names refect the evolutionary history of this group, some genera (specifically the terrestrial, non-basal-rosette species) have been moved to other families (Anagallis, Centunculus, Lysimachia, and Trientalis to the Myrsinaceae and Samolus to the Theophrastaceae). 1a. Plants aquatic; leaf blades narrowly dissected [Fig. 806]; peduncles and axes of the terminal racemes conspicuously inflated, the flowers whorled at the constricted nodes [Fig. 806] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hottonia 1b. Plants terrestrial; leaf blades simple; peduncles and axes of inflorescence not or scarcely inflated, the flowers in a solitary, terminal umbel or in racemes with whorls of flowers 2a. Corolla white, the basal, connate portion of the corolla ± equal in lenth to the calyx, constricted at the opening; style very short; plants annual . . . . . . . . . . . . . . . . . . . . Androsace 2b. Corolla pink to purple, red, yellow, or rarely white, the basal, connate portion of the corolla longer than the calyx (+/- equal to the calyx in the yellow-flowered P. veris), not or scarcely constricted at opening; style elongate; plants perennial . . . . . . . . . . . . . . . . . Primula

Androsace 1. Androsace occidentalis Pursh E western androsace. Androsace arizonica (Gray) Derganc; A. occidentalis Pursh var. arizonica (Gray) St. John • MA. Waste areas.

Hottonia Fig. 806  Habit of Hottonia inflata showing inflated inflorescence axes.

1. Hottonia inflata Ell. N Fig. 806 American featherfoil. CT, MA, ME, NH, RI; primarily on the coastal plain. Ponds, pools in swamps, wet ditches.

Pr i m u l ac e a e   747

Primula Flowers must be fresh and fully open or well pressed flat with no wrinkles or folds to supply proper corolla width measurements. 1a. Inflorescence a raceme with (2–) 3–5 whorls of flowers; corolla 19–25 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. japonica 1b. Inflorescence consisting of a solitary, terminal umbel; corolla 8–16 mm wide 2a. Corolla yellow, the basal, connate portion scarcely (if at all) exceeding the calyx; calyx 9–15 mm long, angled in cross-section; leaf blades pubescent . . . . . . . . . . . . . . . . . . . P. veris 2b. Corolla commonly pink to purple with a yellow base, the basal, connate portion longer than the calyx; calyx 3–11 mm long, ± round in cross-section (i.e., without wing-angles); leaf blades glabrous (though sometime farinose) 3a. Sepals 3–6 mm long; scape 3–21 cm tall; largest leaf blades 10–40 (–60) mm long; bracts of umbel 3–6 mm long, barely (if at all) saccate at the base; capsules 2–3 mm in diameter; seeds rounded, their surface nearly smooth; flowers heterostylous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. mistassinica 3b. Sepals 6–9 mm long; scape 10–40 cm tall; largest leaf blades 40–80 mm long; bracts of umbel 6–14 mm long, definitely saccate at the base; capsules 2.5–5 mm in diameter; seeds angled, their surface evidently reticulate; flowers homostylous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. laurentiana 1. Primula japonica Gray E Japanese primrose. MA, VT. Gardens, forest fragments, brook sides, waste areas. The reports of Primula anisodora Balf. f. & G. Forrest (e.g., Weatherbee 1996, Sorrie and Somers 1999) are based on a collection of P. japonica, as evidenced by the relatively more slender and adaxially farinose sepals and larger corollas—Sorrie 2374 (NEBC!). 2. Primula laurentiana Fern.

NC

bird’s-eye primrose. Primula farinosa L. var. macropoda Fern.; P. mistassinica Michx. var. macropoda (Fern.) Boivin • ME. High-pH cliffs and ledges, coastal ledges and turf. Primula laurentiana mainly along the downeast coast of ME (Washington County) on thin, often peaty, soil overlying bedrock usually in close proximity to the Atlantic Ocean. It also occurs at one site in central ME on lake shore cliffs and (formerly) in northern ME on river shore cliffs. 3. Primula mistassinica Michx. N Lake Mistassini primrose. Primula intercedens Fern.; P. mistassinica Michx. var. intercedens (Fern.) Boivin; P. mistassinica Michx. var. noveboracensis Fern. • ME, VT; northern portions of states. River shore outcrops and ledges in high-pH bedrock and/or till regions. 4. Primula veris L. E cowslip primrose. CT, MA, ME, VT. Gardens, lawns, forests.

Ranunculaceae Reference: Mitchell and Dean (1982). 1a. Plants woody, with yellow inner bark; fruit a sparsely pubescent follicle . . . . . Xanthorhiza 1b. Plants herbaceous; fruit an achene, berry, or glabrous follicle 2a. Fruit an achene [Figs. 809, 812, 816]; ovaries with 1 ovule

748 tricolpates

3a. Calyx composed of 5 spurred sepals; receptacle continuing to elongate through and after anthesis, the cluster of achenes becoming slender-conical and 16–50 × 1–3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myosurus 3b. Calyx composed of 4–20 sepals that are not spurred; receptacle not continuing to elongate after anthesis, the cluster of achenes relatively broader compared with length 4a. Petals present, yellow or both white and yellow; sepals (3–) 5, sepaloid 5a. Flowers with 5 sepals [Fig. 818]; seeds both beaked (though the beak sometimes tiny) and lacking a prominent, corky appendage distal to the seed [Fig. 816] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ranunculus 5b. Flowers with 3 (–4) sepals; achenes either beakless or with a prominent, corky appendage distal to the seed 6a. Flowers with 5–8 petals; leaf blades prominently lobed with deep sinuses [Fig. 813]; roots all slender; achenes with both a terminal beak and a prominent, corky appendage prolonged distal to the seed . . . . . . . . . . . . . . . . . . . . Coptidium 6b. Flowers with 7–11 petals; leaf blades entire to crenate; some roots tuberousthickened; achenes lacking both a terminal beak and a corky appendage distal to the seed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ficaria 4b. Petals absent or inconspicuous; sepals 4–20, petaloid or sepaloid in some species of Thalictrum 7a. Plants trailing or climbing vines; sepals valvate in bud; style plumose [Fig. 812]; stem leaves opposite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clematis 7b. Plants upright, not vining; sepals imbricate in bud; style not plumose; stem leaves alternate or lacking (note: the opposite or whorled leaves found on the stems of some Thalictrum and Anemone are involucral leaves associated with the inflorescence) 8a. Stem leaves alternate; sepals 1.5–5 mm long, often inconspicuous and sometimes caducous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Thalictrum 8b. Stem leaves lacking, but appearing opposite or whorled in most species (these leaves actually involucral bracts associated with the inflorescence); sepals 5–25 mm long, conspicuous, usually persisting through anthesis 9a. Leaf blades palmately lobed or palmately compound (subternately divided in A. multifida); style present; achenes lacking evident veins on lateral surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anemone 9b. Leaf blades pinnately compound in ternate fashion [Fig. 821]; style absent (i.e., the stigma sessile); achenes with evident veins or ribs on lateral surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Thalictrum 2b. Fruit a follicle, capsule, or berry [Fig. 807]; ovaries with 2 or more ovules 10a. Flowers zygomorphic 11a. Upper sepal arched or hooded; petals not spurred . . . . . . . . . . . . . . . . . . Aconitum 11b. Upper sepal neither arched nor hooded, prolonged into a spur; upper petal or all the petals spurred 12a. Flowers with 4 distinct petals and 3 carpels; ultimate lobes of leaf blades 5–25 mm wide; plants perennial, from fasciculate roots . . . . . . . . . . . . . . Delphinium 12b. Flowers with 2 connate petals and 1 carpel; ultimate lobes of leaf blades narrower than 1.5 mm; plants annual, from a slender taproot . . . . . . . . . . Consolida 10b. Flowers actinomorphic

R a nun c ul ac e a e   749

13a. Leaf blades pinnately dissected into filiform segments; gynoecium of 5–10 connate carpels; fruit a capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nigella 13b. Leaf blades toothed to divided, but the divisions not threadlike; gynoecium of 1–15 distinct carpels; fruit a follicle or berry 14a. Leaf blades palmately lobed [Fig. 814]; perianth consisting of only 3 petaloid, caducous sepals; inflorescence a solitary flower borne at the top of a leafy stem [Fig. 814]; mature fruit a red berry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrastis 14b. Leaf blades simple or compound, but not palmately lobed; perianth either with both sepals and petals present or with 4–9 petaloid sepals; inflorescence either with 2 or more flowers or a solitary flower borne on a scape (solitary and terminal in Trollius and also rarely in Caltha); mature fruit a follicle or berry (a red berry in some Actaea) 15a. Leaf blades not pinnately compound, either simple or palmately divided; calyx composed of 5–9 petaloid, pale yellow to bright yellow or orange-yellow sepals 16a. Leaf blades palmately divided to the base; flowers with 10–15 (–25) yellow staminodes 3–6 mm long; inflorescence with a solitary, terminal flower. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trollius 16b. Leaf blades simple; flowers lacking staminodes; inflorescence terminal or both terminal and axillary, with 1–7 flowers . . . . . . . . . . . . . . . . . . . . . . Caltha 15b. Leaf blades pinnately compound, often in ternate fashion; perianth not yellow, the calyx with 5 or fewer sepals 17a. Perianth inconspicuous and often partly or entirely caducous, the sepals 2.5–4 (–6) mm long; inflorescence a raceme or panicle [Fig. 807]; fruit a berry or a follicle in A. racemosa [Fig. 807]; ultimate segments of leaf blades sharply cleft and toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Actaea 17b. Perianth conspicous, not caducous, the petaloid sepals (4–) 5–25 mm long; inflorescence a solitary flower or a 1- to 10-flowered cyme; fruit a follicle; ultimate segments of leaves crenately toothed or crenately lobed 18a. Sepals red or blue to people (rarely white); petals conspicuous, prolonged into long spurs [Fig. 811], 15–31 mm long including the spur; leaves borne on an aerial, leafy stem, the blades 2- or 3-times compound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aquilegia 18b. Sepals white; petals represented by small, clavate-spatulate staminodia, without spurs, 2.5–3 mm long; leaves all basal, the blades once-compound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coptis

Aconitum 1a. Leaf blades compound or lobed with very deep sinuses, usually with less than 2 (–4) mm of tissue from the base of the sinus to the base of the blade, the central leaflet or lobe lobed again with sinuses extending more than ½ the distance to the midrib of the lobe; stems erect; seeds winged on the angles and rugose on the faces, but without thin plates and ridge segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. napellus 1b. Leaf blades lobed with moderately deep sinuses, usually with more than 2 mm of tissue from the base of the sinus to the base of the blade, the central lobe toothed or lobed with sinuses extending less than ½ the distance to the midrib of the lobe; stems erect, reclining, or climbing; seeds ornamented with thin plates and ridge segments . . . . . . . . . . . . A. uncinatum

75 0   tricolpate s

1. Aconitum napellus L. E garden monkshood. CT, Ma, Me, NH, VT. Fields, roadsides, waste areas, yards, forest fragments. This species was reported from RI by Kartesz (1999), based on George (1997); however, George (1999) stated it had questionable naturalization in RI. 2. Aconitum uncinatum L. E wild monkshood. NH. Fields, roadsides, waste areas.

Actaea Actaea is circumscribed broadly here to include Cimicifuga, following the phylogenetic work of Compton et al. (1998). 1a. Axis of inflorescence usually branched (i.e., the inflorescence a panicle); stigma borne on a short, stout style; fruit a follicle; petals represented by linear staminodes that are bifid at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. racemosa 1b. Axis of inflorescence simple (i.e., the inflorescence a raceme) [Fig. 807]; stigma sessile or subsessile; fruit a berry [Fig. 807]; petals entire at apex 2a. Pedicels of infructescence slender, 0.3–0.7 mm in diameter [Fig. 807]; berries red (rarey white); stigma narrower than ovary; leaflets commonly pubescent on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. rubra 2b. Pedicels of infructescence thick, 1–2.5 mm in diameter; berries white (rarely red); stigma wider than ovary; leaflets glabrous or nearly so on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pachypoda 1. Actaea pachypoda Ell. N white baneberry. Actaea alba, auct. non (L.) P. Mill. • CT, MA, ME, NH, RI, VT. Forests, usually mesic, deciduous types. Red-fruited plants (forma rubrocarpa), though generally rare, can be locally frequent. ‌1 × 3. Actaea ×ludovici Boivin is a very rare baneberry hybrid known from MA, VT; also reported from RI by George (1997), but specimens are unknown. The hybrids are variable and show different combinations of character states (e.g., white berries on plants that appear close to A. rubra but also show more prominent stigmas and less pubescence on the abaxial leaf blades, red or pink berries on plants that appear close to A. pachypoda but also show more pubescence on the abaxial leaf surface). These plants frequently show some sterile fruits. 2. Actaea racemosa L. n black bugbane. Cimicifuga racemosa (L.) Nutt.; C. serpentaria Pursh; Macrotrys racemosa (L.) Sweet; Thalictrodes racemosum (L.) Kuntze • CT, MA, ME. Rich, mesic, deciduous forests. This species is introduced in ME and, in part, to the other states. 3. Actaea rubra (Ait.) Willd. N Fig. 807 Fig. 807  Infructescence of Actaea rubra.

red baneberry. Actaea arguta Nutt.; A. spicata L. var. arguta (Nutt.) Torr.; A. spicata L. ssp. rubra (Ait.) Hultén • CT, MA, ME, NH, RI, VT. Forests, usually mesic, deciduous types, often associated with rich soils. White-fruited plants (forma neglecta) are much less common than red-fruited plants.

Anemone Anemone is circumscribed broadly to include Hepatica based on the phylogenetic work of Hoot et al. (1994). Reference: Dutton et al. (1997). 1a. Leaves dimorphic—the basal ones with 3 (–5)-lobed blades [Fig. 808], the stem ones subtending the flower with simple, lanceolate blades; stem leaves closely subtending the flower 2a. Apex of leaf lobes rounded to bluntly pointed; sinuses, at most, extending to middle of leaf blade; stem leaves (i.e., bracts) rounded at the apex; achenes 1–1.4 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. americana

R a nun c u l ac e a e   75 1

2b. Apex of leaf lobes acute to acuminate [Fig. 808]; sinuses indented beyond middle of leaf blade [Fig. 808]; bracts acute to obtuse at the apex; achenes 1.4–1.8 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. acutiloba 1b. Leaf blades compound or lobed with deep sinuses in A. canadensis, the basal and cauline ones ± similar; stem leaves remotely subtending the flower 3a. Carpels and achenes glabrous or thinly pubescent, but not tomentose [Fig. 809]; achenes in capitate to spherical clusters; plants with rhizomes (these tuberous in A. blanda) 4a. Sepals blue (rarely white), numbering 8–14 (–18) per flower; rhizomes tuberous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. blanda 4b. Sepals white to pink (rarely blue or purple), numbering (4–) 5–7 (–12) per flower; rhizomes without tubers (but with caudices in A. canadensis) 5a. Leaves subtending flowers without petioles or these obscure; stems branched, with 1–6 flowers; sepals white; achenes broadly wing-margined; above-ground stems produced from caudices on rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. canadensis 5b. Leaves subtending flowers with evident petioles; stems simple, with 1 flower; sepals pink to white (rarely blue or purple in A. nemorosa); achenes merely with a sharp margin; above-ground stems produced directly from rhizomes 6a. Flowers with 5 sepals; rhizomes white or brown to black, 1–3 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. quinquefolia 6b. Flowers with (5–) 6 or 7 (–12) sepals; rhizomes brown to black, 3–5 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. nemorosa 3b. Carpels and achenes densely pubescent, the achenes with long, tomentose hairs [Fig. 810]; achenes in oblong to cylindric clusters; plants commonly without rhizomes 7a. Leaf blades dissected into numerous, narrow segments (1.5–) 2–3.5 (–5) mm wide, those subtending the inflorescence sessile or nearly so; styles deciduous from achenes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. multifida 7b. Leaf blades divided into 3–5 broad segments (4–) 6–30 (–40) mm wide, those subtending the inflorescence distinctly petioled; styles persistent on achenes 8a. Inflorescence subtended by (3–) 5–9 leaves (i.e., involucral bracts); peduncles naked; cluster of achenes 2–4.5 cm tall, 2–5 times as tall as broad; styles crimson; beak of the achene 0.3–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. cylindrica 8b. Inflorescence subtended by 2–3 (–5) leaves; peduncles with 2 additional subtending leaves; cluster of achenes 1.5–3 cm tall, mostly 1–1.5 times as tall as broad; styles pale or crimson-tipped; beak of the achene 1–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. virginiana 1. Anemone acutiloba (DC.) G. Lawson N Fig. 808 sharp-lobed hepatica. Hepatica acutiloba DC.; H. nobilis Schreber var. acuta (Pursh) Steyermark • CT, MA, ME, NH, VT. Rich, mesic, often rocky, deciduous forests.

Fig. 808  Flower and leaves of Anemone acutiloba.

2. Anemone americana (DC.) H. Hara N blunt-lobed hepatica. Hepatica americana DC.; H. nobilis Schreber var. obtusa (Pursh) Steyermark • CT, MA, ME, NH, RI, VT. Rich, mesic, often rocky, deciduous forests. 3. Anemone blanda Schott & Kotschy E Greek windflower. MA. Forest fragments, dumps, waste areas. 4. Anemone canadensis L. N Fig. 809 Canada windflower. Anemonidium canadense (L.) A. & D. Löve • CT, MA, ME, NH, VT. River and lake shores, fields, meadows. Reports of this species from RI are based on a specimen consisting of a vegetative fragment (i.e., the collection is equivocal)—Champlin 661 (Champlin Herb.).

Fig. 809  Infructescence of Anemone canadensis showing achenes that lack tomentum.

75 2   tricolpate s

5. Anemone cylindrica Gray N long-headed windflower. CT, MA, ME, NH, RI, VT. Sandy and/or rocky soil of fields, hillsides, and woodlands. 6. Anemone multifida Poir. var. multifida N cut-leaved windflower. Anemone globosa (Torr. & Gray) Nutt. ex Pritz.; A. hudsoniana (DC.) Richards.; A. multifida Poir. var. hudsoniana DC.; A. multifida Poir. var. globosa Torr. & Gray • ME, VT. High-pH river shore outcrops. 7. Anemone nemorosa L. E European windflower. Anemonidium nemorosum (L.) Holub • MA. Forests, forest fragments. 8. Anemone quinquefolia L. var. quinquefolia N wood windflower. Anemone nemorosa L. var. bifolia (Farw.) Boivin; A. quinquefolia L. var. bifolia Farw.; Anemone quinquefolia L. var. interior Fern. • CT, MA, ME, NH, RI, VT. Deciduous or mixed evergreen-deciduous forests, forest edges, riparian forests, river banks, fields. 9. Anemone virginiana L. N Fig. 810 tall windflower. 9a. Anemone cylindrica Gray var. alba Oakes; A. riparia Fern.; A. virginiana L. var. riparia (Fern.) Boivin • CT, MA, ME, NH, RI, VT. River shores, banks, and shoreline outcrops, forests, forest edges, woodlands, roadsides, fields, ledges.

Fig. 810  Infructescence of Anemone virginiana var. virginiana showing achenes that have tomentum.

1a. Clusters of fruits 8–10 mm wide; anthers 0.7–1.2 mm long; achenes with spreadingascending to ascending styles; involucral bracts usually broad-cuneate to truncate at the base, the terminal leaflet with concave- or straight-tapered margins in the basal portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9a. A. virginiana var. alba (Oakes) Wood 1b. Clusters of fruits 11–14 mm wide; anthers 1.2–1.6 mm long; achenes with divergent styles; involucral bracts usually cordate to truncate at the base, the terminal leaflet with convex- to straight-tapered margins in the basal portion . . . . . . . . . . . . . 9b. A. virginiana var. virginiana Variety alba is known from CT, MA, ME, NH, VT. It is commonly associated with riparian communities, though it also occurs in open and forested upland communities. Variety virginiana is known from CT, MA, ME, NH, RI, VT. It is commonly associated with upland communities, though it is also known from riparian habitats.

Aquilegia 1a. Perianth red (the sepals sometimes green at the apex); spurs straight to arching [Fig. 811]; follicle with a persistent style beak 15–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . A. canadensis 1b. Perianth blue to purple (rarely white); spurs strongly curved inward; follicle with a persistent style beak 5–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. vulgaris 1. Aquilegia canadensis L. N Fig. 811 red columbine. Aquilegia australis Small; A. canadensis L. var. australis (Small) Munz; A. canadensis L. var. coccinea (Small) Munz; A. canadensis L. var. eminens (Greene) Boivin; A. canadensis L. var. latiuscula (Greene) Munz; A. coccinea Small • CT, MA, ME, NH, RI, VT. Rich, deciduous forests and woodlands, cliffs, river shore outcrops, river banks, rocky slopes. Rare forms of this plant have spurless petals. Fig. 811  Flower of Aquilegia canadensis.

2. Aquilegia vulgaris L. E European columbine. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, abandoned homesteads, forest edges, gardens.

Caltha 1. Caltha palustris L. N marsh-marigold. Caltha asarifolia DC.; C. palustris L. ssp. asarifolia (DC.) Hultén; C. palustris L. var. asarifolia (DC.) Rothrock; C. palustris L. var. flabellifolia (Pursh) Torr. & Gray • CT, MA, ME, NH, RI, VT. Swamps, stream margins, low riparian forests.

R a nun c ul ac e a e   75 3

Clematis 1a. Flowers and fruits solitary in axils of leaves [Fig. 812], with petaloid staminodes between the sepals and stamens; sepals 20–60 mm long 2a. Sepals 4 per flower, light red-purple to blue-purple, 25–60 mm long; style pubescent ± throughout its length in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. occidentalis 2b. Sepals 6 per flower, usually white, 20–30 mm long; style apically glabrous in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. florida 1b. Flowers and fruits in many-flowered cymes or panicles; sepals 5–22 mm long, white to cream 3a. Stems ascending to erect, not vining; leaves with 5–9 leaflets; achene beak 12–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. recta 3b. Stems vining (i.e., trailing or climbing on other vegetation); leaves with 3 or 5 leaflets; achene beak 20–60 mm long 4a. Leaves with 3 leaflets; flowers unisexual, the carpellate ones with 40–70 carpels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. virginiana 4b. Leaves with 3 or 5 leaflets; flowers bisexual (sometimes some staminate flowers mixed in the same inflorescence), with 5–10 carpels (20 or more in C. vitalba) 5a. Sepals adaxially glabrous and abaxially tomentose along margins; leaflet margins entire; flowers with 10 or fewer carpels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. terniflora 5b. Sepals adaxially and abaxially tomentose; leaflet margins entire to crenate or dentate; flowers with 20 or more carpels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. vitalba 1. Clematis florida Thunb. E Asian virgin’s-bower. Atragene florida Pers.; Clematis japonica Thunb. var. simsii Makino • CT. Roadsides, waste areas. 2. Clematis occidentalis (Hornem.) DC. ssp. occidentalis N Fig. 812 purple virgin’s-bower. Atragene americana Sims; A. occidentalis Hornem.; Clematis verticillaris DC.; C. verticillaris DC. var. grandiflora Boivin • CT, MA, ME, NH, RI, VT. Rich, rocky, deciduous forests, ledges, talus, river banks. 3. Clematis recta L. E ground virgin’s-bower. NH. Fields, forest edges and fragments, about dwellings. 4. Clematis terniflora DC. E yam-leaved virgin’s-bower. Clematis dioscoreifolia Levl. & Vaniot; C. dioscoreifolia Levl. & Vaniot var. robusta (Carr.) Rehd.; C. paniculata Thunb. • CT, MA, NH, RI, VT. Roadsides, fields, forest fragments, near dwellings. 5. Clematis virginiana L. N Virginia virgin’s-bower. Clematis canadensis P. Mill.; C. missouriensis Rydb.; C. virginiana L. var. missouriensis (Rydb.) Palmer & Steyermark • CT, MA, ME, NH, RI, VT. Roadsides, forest edges, shrub thickets, river banks, pond shores, swamps. 6. Clematis vitalba L. E white virgin’s-bower. ME. Roadsides, abandoned homesteads.

Consolida 1a. All the bracts of the inflorescence simple, sometimes the lowermost one dissected; sepals dark blue (rarely pink or white); upper lobe of petals 3.5–5 mm long . . . . . . . . . . . . . . C. regalis 1b. Lower bracts of the inflorescence (at least the lower 2, and usually more) dissected into 3 or more lobes; sepals blue, purple, red-purple, or white; upper lobe of petals 5–8 mm long

Fig. 812  Infructescence and leaves of Clematis occidentalis.

75 4   tricolpate s

2a. Bracteoles on pedicels not touching the sepals, positioned 4–20 mm from the flower; spur 12–20 mm long; seeds dark brown to black; upper lobe of petals apically cleft for 0.2–1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ajacis 2b. Bracteoles on pedicels touching the sepals, positioned 1–5 mm from the flower; spur 8–12 mm long; seeds red-brown; upper lobe of petals apically cleft for 0.5–1.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. orientalis 1. Consolida ajacis (L.) Schur E doubtful knight’s-spur. Consolida ambigua (L.) P.W. Ball & Heywood; Delphinium ajacis L.; D. ambiguum L. • CT, MA, RI, VT; also reported from ME and NH by Kartesz (1999), but specimens are unknown. Fields, roadsides, waste areas, dumps. 2. Consolida orientalis (J. Gay) Schroedinger E Oriental knight’s-spur. Delphinium orientale J. Gay • VT. Fields, roadsides. 3. Consolida regalis S.F. Gray E royal knight’s-spur. Delphinium consolida L. • CT. Fields, roadsides, waste areas.

Coptidium 1. Coptidium lapponicum (L.) Gandog.

N C Fig. 813

Lapland-crowfoot. Ranunculus lapponicus L. • ME; northern portion of state. Swamps, usually dominated by Thuja occidentalis.

Coptis 1. Coptis trifolia (L.) Salisb. N Fig. 813  Habit of Coptidium lapponicum.

three-leaved goldthread. Anemone groenlandica Oeder; Coptis groenlandica (Oeder) Fern.; C. trifolia (L.) Salisb. ssp. groenlandica (Oeder) Hultén; C. trifolia (L.) Salisb. var. groenlandica (Oeder) Fassett; Helleborus trifolius L. • CT, MA, ME, NH, RI, VT. Forests and swamps, commonly associated with evergreen trees and ascending high into the mountains.

Delphinium The report of Delphinium grandiflorum L. for VT by Seymour (1982) was based on a cultivated specimen. 1. Delphinium exaltatum Ait. E tall larkspur. MA; also reported from ME by Campbell et al. (1995), but specimens are unknown. Fields, roadsides.

Ficaria 1. Ficaria verna Huds. ssp. bulbifera A. & D. Löve E fig-crowfoot. Ranunculus ficaria L. ssp. bulbilifer Lambinon; R. ficaria L. ssp. bulbifera (Marsden-Jones) Lawalree, nom. illeg.; R. ficaria L. var. bulbifera Marsden-Jones • CT, MA, NH, RI. Riparian forests, river banks, waste areas, gardens.

Hydrastis 1. Hydrastis canadensis L. Fig. 814  Flower and leaf of Hydrastis canadensis.

n C Fig. 814

goldenseal. CT, MA, VT; found mainly in western New England. Rich, mesic, often rocky forests, usually associated with limestone and trap bedrock. Some MA populations (e.g., Franklin County, at least in part) represent introductions.

R a nun c u l ac e a e   75 5

Myosurus 1. Myosurus minimus L. E tiny mousetail. MA. Fields, roadsides, waste areas.

Nigella 1a. Inflorescence, a solitary flower, subtended by an involucre composed of a whorl of bracts that are pinnately dissected into filiform segments; capsule smooth on outer surface, with 10 locules; ultimate segments of upper leaves linear . . . . . . . . . . . . . . . . . . . . . . . . . . N. damascena 1b. Inflorescence not subtended by an involucre; capsule tuberculate on outer surface, with 5 locules; ultimate segments of upper leaves broad-linear to narrow-lanceolate . . . . . . N. sativa 1. Nigella damascena L. E devil-in-the-bush. CT, MA, VT; also reported from ME by Knowlton et al. (1918), but specimens are unknown. Waste areas, gardens, near dwellings. Reports of Nigella sativa L. (e.g., Graves et al. 1910) are based on collections of N. damascena. Nigella damascena is often cultivated for its attractive flowers. Escaped forms sometimes have supernumerary flower parts (i.e., they are double-flowered). While single-flowered plants usually possess both sepals and petals, double-flowered plants usually possess only the petaloid sepals. 2. Nigella sativa L. E black-cumin. CT. Waste areas.

Ranunculus Phylogeny and morphology support a division of a broadly defined Ranunculus (Emadzade et al. 2010). In New England, the genera Coptidium and Ficaria are segregated from the remainder of Ranunculus. Reports of the European Ranunculus circinatus Sibth. (e.g., Graves et al. 1910) were based on collections of R. aquatilis var. diffusus. Ranunculus laxicaulis (Torr. & Gray) Darby was reported from VT by Magee and Ahles (1999), but specimens are unknown. References: Duncan (1980), Whittemore (1997). 1a. Plants aquatic, with submersed leaf blades that are finely divided into narrow segments less than 1 mm wide (the emersed leaves, if present, with broader segments) [Fig. 817] 2a. Petals white with yellow bases [Fig. 817]; leaf segments capillary; achenes coarsely wrinkled on the surface, with a beak 0.2–0.5 mm long . . . . . . . . . . . . . . . . . . . . . . R. aquatilis 2b. Petals yellow; leaf segments flat; achenes smooth on the surface, with a beak 0.4–1.8 mm long 3a. Petals 7–12 mm long; achenes 1.8–2.2 mm long, the lower portion thickened and spongy; achene beak 1–1.8 mm long; styles 0.8–1.2 mm long . . . . . (in part) R. flabellaris 3b. Petals 3–7 mm long; achenes 1–1.6 mm long, not thickened in the lower half; achene beak 0.4–0.8 mm long; styles 0.2–0.4 mm long . . . . . . . . . . . . . . . . . . . (in part) R. gmelinii 1b. Plants terrestrial, of wetlands, or sometimes aquatic, with leaf blades that are simple or divided with relatively broad segments [Figs. 815, 819, 820] 4a. All the leaf blades simple and unlobed [Fig. 820] 5a. Leaf blades cordate to reniform [Fig. 820]; achenes with longitudinal nerves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. cymbalaria 5b. Leaf blades linear to lanceolate; achenes without longitudinal nerves 6a. Leaf blades 0.4–8 mm wide; plants with filiform, arching stolons; petals 1–5 mm long; achenes essentially smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. flammula

75 6   tricolpate s

6b. Lower leaf blades (6–) 8–24 mm wide; plants lacking stolons, though rooting at the base of the stem; petals 5–8 mm long; achenes reticulate-patterned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. ambigens 4b. Either the lower leaf blades or those borne on the stem or both conspicuously lobed to compound [Figs. 815, 819] 7a. Leaves dimorphic—the basal ones simple with ovate to reniform blades and borne on elongate petioles, the stem ones becoming with conspicuously lobed blades and becoming sessile upward; nectary scale fused to petals on 3 sides, forming a pocket that encloses the nectary 8a. Petals 6–8 mm long; achenes 1.8–2.2 mm long . . . . . . . . . . . . . . . . R. rhomboideus 8b. Petals 1–3.5 mm long; achenes 1.1–2 mm long 9a. Stems villous, the hairs usually discernable to the naked eye; base of some roots tuberous-thickened and 1–2 mm thick; basal leaf blades usually broadcuneate to truncate at base; receptacle glabrous . . . . . . . . . . . . . . . . . R. micranthus 9b. Stems glabrous (infrequently with some tiny, sparse hairs that can be seen with magnification); none of the roots tuberous-thickened, the base 0.2–1.5 mm thick; basal leaf blades usually cordate to subcordate at base; receptacle sparsely pubescent 10a. Beak of achene 0.6–1 mm long, strongly curved [Fig. 816]; sepals pubescent on abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. allegheniensis 10b. Beak of achene 0.1–0.2 mm long, straight to barely curved; sepals glabrous on abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. abortivus 7b. Leaves ± monomorphic, relatively similar in general pattern, gradually becoming smaller upward, with shorter sinuses, and/or with shorter petioles; nectary scale separate from petal for at least 50% of its length and forming a flap over the nectary or the scale poorly developed and forming a crescent-shaped to circular ridge that surrounds but does not cover the nectary 11a. At least the larger leaves with definite, unwinged petiolules (i.e., these leaves compound) [Fig. 819] 12a. Petals 2–5 mm long, as long as or shorter than the sepals; anthers up to 1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. pensylvanicus 12b. Petals 6–15 mm long, usually longer than the sepals; anthers 1.2 mm long or longer 13a. Sepals reflexed along a defined, transverse fold 2–3 mm above the base [Fig. 818]; plants either perennial with stems bulbous-thickened at base or annual 14a. Achene faces smooth; petals 9–13 mm long; plants perennial, with stems bulbous-thickened at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. bulbosus 14b. Achene faces usually sparsely papillose (rarely smooth); petals 7–10 mm long; plants annual, without bulbous-thickened stems . . (in part) R. sardous 13b. Sepals spreading (sometimes reflexed from the very base in age); plants perennial, the stems not thickened at the base 15a. Some roots tuberous-thickened, the others slender; petals usually widest at or below middle; stems erect to ascending, not rooting at the nodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. fascicularis 15b. Roots all slender; petals usually widest distal to the middle; stems decumbent to creeping and often rooting at the nodes (or upright and not rooting in R. hispidus) 16a. Styles short, curved, the persistent stigma spread out over upper side of style; achene beak (0.7–) 0.8–1.2 (–1.4) mm long . . . . . . . . . . . . R. repens

R a nun c u l ac e a e   757

16b. Styles elongate, with a terminal, deciduous stigma; achene beak (0.8–) 1.5–2.6 (–3) mm long 17a. Stems spreading to procumbent, 50–80 (–91) cm long at anthesis, sometimes rooting at the nodes; plants of hydric soils (e.g., swamps, marshes, ditches) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. caricetorum 17b. Stems upright, 14–45 (–60) cm tall at anthesis, never rooting at the nodes; plants of mesic to dry-mesic, upland woodlands and forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. hispidus 11b. Leaves with flattened, winged, sometimes ill-defined petiolules (i.e., the leaves lobed but not compound) [Fig. 815] 18a. Stems floating or prostrate and rooting at the nodes; achenes plump, without a sharp or wing-like keel 19a. Petals 7–12 mm long; achenes 1.8–2.2 mm long, the lower portion thickened and spongy; achene beak 1–1.8 mm long; styles 0.8–1.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. flabellaris 19b. Petals 3–7 mm long; achenes 1–1.6 mm long, not thickened in the lower half; achene beak 0.4–0.8 mm long; styles 0.2–0.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. gmelinii 18b. Stems erect, rooting at only the base; achenes flattened, with a sharp or wing-like keel between the faces 20a. Achene faces usually sparsely papillose (rarely smooth); sepals reflexed along a defined, transverse fold 2–3 mm above the base; plants annual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. sardous 20b. Achene faces smooth; sepals spreading or reflexed from the very base; plants perennial from rhizomes, caudices, or bulbous bases or annual in R. sceleratus 21a. Petals 7–11 (–15) mm long, longer than the sepals; sepals spreading; receptacle glabrous; achenes 2–3 × 1.8–2.4 mm . . . . . . . . . . . . . . . . . . . R. acris 21b. Petals 2–5 mm long, equal to or shorter than sepals; sepals reflexed from near the base (sometimes spreading in R. sceleratus); receptacle pubescent (sometimes glabrous in R. sceleratus); achenes 1–2.2 × 0.8–1.8 mm 22a. Anthers up to 1 mm long; style absent; achenes in a cylindric cluster, each with a very short, straight beak ca. 0.1 mm long; stems not thickened at the base; nectary scale forming a crescent-shaped to circular ridge that surrounds but does not cover the nectary . . . . . . . . . . . . . . . . R. sceleratus 22b. Anthers longer than 1 mm; style present; achenes in a spherical cluster, each with a hooked beak 1–1.4 mm long; stems bulbous-thickened at the base; nectary scale forming a flap over the nectar . . . . R. recurvatus

Fig. 815  Lobed leaf blade of Ranunculus acris.

1. Ranunculus abortivus L. N kidney-leaved crowfoot. Ranunculus abortivus L. ssp. acrolasius (Fern.) Kapoor & A. & D. Löve; R. abortivus L. var. acrolasius Fern.; R. abortivus L. var. eucyclus Fern.; R. abortivus L. var. indivisus Fern. • CT, MA, ME, NH, RI, VT. Mesic, deciduous forests and woodlands, cliff bases, and rocky slopes, often associated with rich soils. 2. Ranunculus acris L. E Fig. 815 tall crowfoot. Ranunculus acris L. var. latisectus G. Beck • CT, MA, ME, NH, RI, VT; nearly throughout. Mesic to hydric soil of fields, roadsides, waste areas, river banks, shorelines. 3. Ranunculus allegheniensis Britt. N Fig. 816 Allegheny crowfoot. CT, MA, RI, VT. Mesic forests, woodlands, forest edges, fields, and roadsides, usually in regions of high-pH bedrock.

Fig. 816  Infructescence of Ranunculus allegheniensis showing achenes with curved style beak.

75 8   tricolpate s

4. Ranunculus ambigens S. Wats.

NC

water-plantain crowfoot. Ranunculus obtusiusculus Raf. • CT, MA, ME, NH, RI. Pond shores, stream shores, ditches, pools, hydric fields, marshes, swamps. 5. Ranunculus aquatilis L. var. diffusus Withering N Fig. 817 white water crowfoot. Batrachium aquatile (L.) Dumort.; B. longirostre (Godr.) F.W. Schultz; B. trichophyllum (Chaix) F.W. Schultz; Ranunculus amphibius James; R. aquatilis L. var. capillaceus (Thuill.) DC.; R. aquatilis L. var. longirostris (Godr.) Lawson; R. aquatilis L. var. subrigidus (W. Drew) Breitung; R. longirostris Godr.; R. subrigidus W. Drew; R. trichophyllus Chaix; R. trichophyllus Chaix var. confervoides (Fries) Rikli; R. trichophyllus Chaix ssp. eradicatus (Laestad.) C.D.K. Cook; R. trichophyllus Chaix var. eradicatus (Laestad.) W. Drew; R. trichophyllus Chaix ssp. lutulentus (Perrier & Song.) Vierh. • CT, MA, ME, NH, RI, VT. Still or slow-moving water of lakes and rivers, commonly in circumneutral to basic water. 6. Ranunculus bulbosus L. E Fig. 818 bulbous crowfoot. Ranunculus bulbosus L. var. dissectus Barbey • CT, MA, ME, NH, RI, VT. Fields, roadsides, forests. Fig. 817  Habit of Ranunculus aquatilis.

7. Ranunculus caricetorum Greene N Fig. 819 swamp crowfoot. Ranunculus hispidus Michx. var. caricetorum (Greene) T. Duncan; R. intermedius Eat.; R. septentrionalis Poir. in Lam. var. caricetorum (Greene) Fern. • CT, MA, ME, NH, VT; also reported from RI by George (1997), but specimens are unknown. Swamps, hydric fields and meadows, marshes, wetland edges, riparian forests, stream banks. Ranunculus caricetorum was included within R. hispidus as a variety by Duncan (1980). However, it is distinct in its morphology, ecology, and chromosome number. In light of such, it is treated here as a species. 8. Ranunculus cymbalaria Pursh N Fig. 820

Fig. 818  Flower of Ranunculus bulbosus showing sepals reflexed on a fold above the base of the sepal.

seaside crowfoot. Cyrtorhyncha cymbalaria (Pursh) Britt.; Halerpestes cymbalaria (Pursh) Greene; H. cymbalaria (Pursh) Greene ssp. saximontana (Fern.) Moldenke; Ranunculus cymbalaria Pursh var. alpinus Hook.; R. cymbalaria Pursh ssp. saximontanus (Fern.) Thorne; R. cymbalaria Pursh var. saximontanus Fern. • CT, MA, ME, NH, RI; coastal region but disjunct in Berkshire County, MA, and restricted in CT to the eastern portion of the state. Saline to brackish shorelines and marshes of the Atlantic Ocean and tidal rivers. 9. Ranunculus fascicularis Muhl. ex Bigelow N early crowfoot. Ranunculus fascicularis Muhl. ex Bigelow var. apricus (Greene) Fern. • CT, MA, ME, NH, VT. Rocky, commonly dry-mesic, forests and woodlands, ledges, often associated with circumneutral to high-pH bedrock. 10. Ranunculus flabellaris Raf. N greater yellow water crowfoot. Ranunculus delphiniifolius Torr. ex Eat. • CT, MA, ME, NH, RI, VT. Swamps, ponds, pools, slow streams. 11. Ranunculus flammula L. N

Fig. 819  Compound leaf blade of Ranunculus caricetorum.

creeping crowfoot.  11a. Ranunculus filiformis Michx. var. ovalis Bigelow; R. flammula L. var. samolifolius (Greene) L. Benson; R. reptans L. var. ovalis (Bigelow) Torr. & Gray;  11b. Ranunculus filiformis Michx.; R. reptans L.; R. reptans L. var. filiformis (Michx.) DC. • CT, MA, ME, NH, VT. River shores, lakes shores, still or slow-moving, shallow water, marshes, pools, low riparian forests. 1a. Leaf blades narrow-elliptic to lanceolate or linear, 2–8 mm wide; stems 0.5–2 mm thick; sepals 2–3 mm long . . . . . . . . . . . . . . . . . . . . . 11a. R. flammula var. ovalis (Bigelow) L. Benson 1b. Leaf blades linear to filiform, 0.4–1.5 mm wide; stems 0.2–1 mm thick; sepals 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11b. R. flammula var. reptans (L.) E. Meyer Variety ovalis is known from CT, MA, ME, NH, VT. Variety reptans is known from CT, MA, ME, NH, VT. This variety was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it could be in RI and was unaware of any collections. Though both varieties

R a nun c u l ac e a e   75 9

are widespread in New England, var. reptans is more frequent and extends further north in New England. 12. Ranunculus gmelinii DC.

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lesser yellow water crowfoot. Ranunculus gmelinii DC. var. hookeri (D. Don) L. Benson; R. gmelinii DC. var. prolificus (Fern.) Hara; R. gmelinii DC. ssp. purshii (Richards.) Hultén; R. gmelinii DC. var. purshii (Richards.) Hara; R. gmelinii DC. var. terrestris (Ledeb.) L. Benson; R. purshii Richards. • ME; found mainly in the northern portion of state. Slow streams, evergreen swamps dominated by Thuja occidentalis, and ditches in regions of high-pH bedrock and/or till. 13. Ranunculus hispidus Michx.

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hispid crowfoot. Ranunculus hispidus Michx. var. eurylobus L. Benson; R. hispidus Michx. var. falsus Fern. • CT, MA, RI, VT. Dry-mesic to mesic soils of deciduous forests and woodlands, cliff bases, clearings, fields, and roadsides, commonly found in areas of high-pH bedrock. 14. Ranunculus micranthus Nutt.

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small-flowered crowfoot. Ranunculus micranthus Nutt. var. cymbalistes (Greene) Fern.; R. micranthus Nutt. var. delitescens (Greene) Fern. • CT, MA, RI. Rich, rocky, deciduous forests and woodlands, ridges. 15. Ranunculus pensylvanicus L. f. N bristly crowfoot. CT, MA, ME, NH, RI, VT; becoming rare to the south. Shorelines, river banks, swamps, ditches, marshes. 16. Ranunculus recurvatus Poir. var. recurvatus N hooked crowfoot. Ranunculus recurvatus Poir. var. adpressipilis Weatherby; R. recurvatus Poir. var. typicus L. Benson • CT, MA, ME, NH, RI, VT. Rich, mesic forests, stream banks, swamps dominated by deciduous trees, forest seeps. 17. Ranunculus repens L. E spot-leaved crowfoot. Ranunculus repens L. var. degeneratus Schur; R. repens L. var. erectus DC.; R. repens L. var. glabratus DC.; R. repens L. var. pleniflorus Fern.; R. repens L. var. villosus Lamotte • CT, MA, ME, NH, RI, VT. Swamps, shorelines, ditches, wet-mesic to hydric fields, lawns. 18. Ranunculus rhomboideus Goldie E prairie crowfoot. MA, RI. Fields, roadsides, waste areas. 19. Ranunculus sardous Crantz E hairy crowfoot. Ranunculus parvulus L.; R. pseudobulbosus Schur • RI. Fields, roadsides. 20. Ranunculus sceleratus L. var. sceleratus N cursed crowfoot. Hecatonia scelerata (L.) Fourr.; Ranunculus sceleratus L. var. typicus L. Benson • CT, MA, ME, NH, RI, VT. Swamps, marshes, ditches, borders of tidal wetlands.

Thalictrum Reference: Park and Festerling (1997). 1a. Leaves subtending the inflorescence opposite (though sometimes appearing whorled when the leaves are divided to the base); sepals 5–18 mm long, conspicuous, usually persisting through anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. thalictroides 1b. Stem leaves alternate; sepals 1.5–5 mm long, often inconspicuous and sometimes caducous 2a. Leaf segments mostly with 3 lobes, each lobe entire and without additional sublobes; filaments usually white, sometimes tinged with purple, somewhat to prominently dilated distally (slender throughout in T. dasycarpum); anthers 0.5–2.7 (–3) mm long (up to 3.6 (–4) mm long in T. dasycarpum)

Fig. 820  Habit of Ranunculus cymbalaria.

76 0   tricolpate s

3a. Filaments ± slender throughout; anthers 1–3.6 (–4) mm long, usually with a prominent apiculus at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. dasycarpum 3b. Filaments somewhat to prominently dilated distally; anthers 0.5–2.7 (–3) mm long, without an apiculus at the apex or a poorly developed one 4a. Abaxial surface of leaflets with sessile or stalked glands or white-papillose; achenes usually stipitate-glandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. revolutum 4b. Abaxial surface of leaflets lacking glands or papillae; achenes glabrous or pubescent with eglandular hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. pubescens 2b. Leaf segments with 4 or more lobes, these often appearing as 3 lobes and each lobe with 1–3 tooth-like sublobes [Fig. 821]; filaments colored, usually red-purple to purple or yellow to green-yellow, ± slender throughout; anthers 2–4.5 mm long 5a. Lowest branch of the inflorescence subtended by a short-petioled leaf, the petiole shorter than 30 mm; achenes inwardly curved; plants with rhizomes, flowering from early June through early August . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. venulosum 5b. Lowest branch of the inflorescence subtended by a long-petioled leaf, the petiole 30–60 mm long; achenes straight; plants arising from a stout caudex, without rhizomes, flowering from early May through early June . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. dioicum 1. Thalictrum dasycarpum Fisch. & Avé-Lall. E purple meadow-rue. Thalictrum dasycarpum Fisch. & Avé-Lall. var. hypoglaucum (Rydb.) Boivin; T. hypoglaucum Rydb. • CT. Roadsides, forest fragments. This species is noted to rarely possess stipitate-glands on petioles, leaf rachises, peduncles, and/or pedicels and rarely possess sessile glands on abaxial leaf surfaces. Such specimens would cause confusion with Thalictrum revolutum. In addition to characters used in the identification key, these two species differ in mean achene body and achene stipe measurements. Thalictrum dasycarpum has achene bodies 2–4.6 mm long and achene stipes 0–1.1 mm long vs. 3.5–5 mm and 0.2–1.7 mm in T. revolutum, respectively. 2. Thalictrum dioicum L. N early meadow-rue. CT, MA, ME, NH, RI, VT. Rich, mesic, deciduous forests, riparian forests, shaded river banks. 3. Thalictrum pubescens Pursh N tall meadow-rue. Thalictrum polygamum Muhl. ex Spreng.; T. polygamum Muhl. ex Spreng. var. hebecarpum Fern.; T. polygamum Muhl. ex Spreng. var. intermedium Boivin; T. pubescens Pursh var. hebecarpum (Fern.) Boivin • CT, MA, ME, NH, RI, VT; throughout. Swamps, marshes, meadows, shorelines, river banks, wet-mesic to hydric fields, ditches, deciduous forests, roadsides. 4. Thalictrum revolutum DC. N waxy-leaved meadow-rue. Thalictrum revolutum DC. var. glandulosior Boivin • CT, MA, NH, RI, VT. Dry-mesic to mesic, rocky forests, woodlands, ridges, and fields. Thalictrum revolutum is noted to rarely lack glands on leaves and achenes. Such forms are extremely rare in New England and would be confused with T. pubescens. In addition to characters used in the identification key, these two species differ in filament flexibility, achene beak length, and flower type/distribution. Thalictrum revolutum has flexible filaments, achene beaks (1–) 1.5–3.3 (–5) mm long, and usually has unisexual flowers on different plants (i.e., the plants are usually dioecious) vs. rigid filaments, achene beaks 0.6–2.5 mm long, and sometimes with some bisexual flowers on a given plant (i.e., the plants are sometimes polygamous) in T. pubescens. Thalictrum revolutum has been attributed to VT by Park and Festerling (1997) based on 13 Jul 1930, Knowlton s.n. (NEBC!). This specimen, which does show stipitateglandular carpels, is atypical in its thin, non-glandular leaf segments. This species possesses a pungent odor in life, this especially obvious after rain.

R a nun c u l ac e a e   76 1

5. Thalictrum thalictroides (L.) Eames & Boivin N anemone meadow-rue. Anemone thalictroides L.; Anemonella thalictroides (L.) Spach; Syndesmon thalictroides (L.) Hoffmgg. ex Britt. • CT, MA, ME, NH, RI, VT; rare in northern New England. Dry-mesic to mesic forests, woodlands, banks, and open areas. 6. Thalictrum venulosum Trel. var. confine (Fern.) Boivin

N C Fig. 821

veiny-leaved meadow-rue. Thalictrum confine Fern.; T. turneri Boivin • ME, VT, northern portion of states. Lake shores and northern, ice-scoured river shores in regions of high-pH bedrock and/or till.

Trollius 1. Trollius laxus Salisb.

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American globe-flower. CT, northwestern portion of state. Meadows and swamps in regions of high-pH bedrock.

Xanthorhiza

Fig. 821  Ultimate segments of leaves of Thalictrum venulosum with 4 or more lobes.

1. Xanthorhiza simplicissima Marsh. E shrub yellowroot. Xanthorhiza apiifolia L’Hér. • CT, MA, ME. Forests, stream banks.

Resedaceae Reseda 1a. Leaf blades entire (rarely with a few, minute, hyaline teeth near the base) 2a. Pedicels 1–3 mm long; corolla with 4 petals, each petal either entire or with 4 short lobes; capsules 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. luteola 2b. Pedicels 4–6 mm long (elongating to 16 mm in fruit); corolla with 6 petals, each petal divided at the apex into 9–15 linear-spatulate segments; capsules 9–11 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. odorata 1b. Leaf blades lobed or dissected 3a. Petals green-white; filaments persistent until maturation of fruit; gynoecium with 4 carpels; capsules with 4 apical teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. alba 3b. Petals green to green-yellow or yellow; filaments deciduous before maturation of fruit; gynoecium with 3 carpels; capsules with 3 apical teeth 4a. Petals apically entire or with 2 or 3 linear-spathulate segments [Fig. 822]; capsules 4.5–5.5 mm wide; seeds lustrous, smooth; most leaf blades pinnatifid with 1 or 2 (–4) pairs of lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. lutea 4b. Petals divided at the apex into 9–15 linear-spathulate segments; capsules 7–11 mm wide; seeds dull, undulate-rugose; leaf blades entire or sometimes the upper ones with 2 or 3 lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. odorata 1. Reseda alba L. E white upright mignonette. CT, MA, ME, NH, VT. Roadsides, waste areas, dumps. 2. Reseda lutea L. E Fig. 822 yellow upright mignonette. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas.

Fig. 822  Flowers of Reseda lutea.

762  tricolpates

3. Reseda luteola L. E dyer’s mignonette. CT, MA, NH, RI. Fields, roadsides, waste areas. 4. Reseda odorata L. E garden mignonette. CT, MA, ME. Fields, roadsides, and waste areas. Previous reports of Reseda odorata in VT (e.g., Seymour 1982, Magee and Ahles 1999) were based on specimens of R. lutea (in part) and on a collection of cultivated material (in part)—Sep 1879, Leland s.n. (NEBC).

Rhamnaceae 1a. Leaf blades with 3 prominent veins from the base [Fig. 823]; petals with an evident claw; fruit a dry, capsule-like drupe; inflorescence a terminal umbel composed of numerous flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceanothus 1b. Leaf blades pinnately veined; petals absent or present and with a short claw; fruit a fleshy drupe; inflorescence axillary, either a solitary flower or a few-flowered umbel 2a. Leaf blades with entire margins; winter buds naked; style undivided; flowers bisexual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Frangula 2b. Leaf blades with toothed margins; winter buds covered by scales; style divided for part of its length; flowers functionally unisexual . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhamnus

Ceanothus 1a. Leaf blades elliptic to oblong or narrow-oblong, 10–25 mm wide; inflorescence on peduncles terminating the branchlets of the current season; fruits 4–5 mm long; lowest pair of lateral veins of the leaf blade emerging 1–3 mm above the base of the blade, the outer margin of the vein within the lateral edge of the blade . . . . . . . . . . . . . . . . . . . . . . C. herbaceus 1b. Leaf blades narrow-ovate to broad-ovate, 20–60 mm wide; inflorescence terminating axillary peduncles; fruits 5–6 mm long; lowest pair of lateral veins of the leaf blade emerging 1–3 mm below the base of the blade, the outer margin of the vein forming the basal lateral edge of the blade [Fig. 823] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. americanus 1. Ceanothus americanus L. N Fig. 823 New Jersey redroot. Ceanothus americanus L. var. intermedius (Pursh) Torr. & Gray; C. americanus L. var. pitcheri Torr. & Gray; C. intermedius Pursh • CT, MA, ME, NH, RI, VT. Woodlands, edges of dry fields and roadsides, rocky slopes. 2. Ceanothus herbaceus L.

Fig. 823  Leaf blade of Ceanothus americanus with lowest pair of lateral veins exposed for a short distance on the basal edge.

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prairie redroot. Ceanothus herbaceus L. var. pubescens (Torr. & Gray ex S. Wats.) Shinners; C. ovatus, auct. non Desf.; C. ovatus Desf. var. pubescens Torr. & Gray ex S. Wats.; C. pubescens (Torr. & Gray ex S. Wats.) Rydb. ex Small • MA, VT. Sandy lake shores, banks, and fields.

Frangula 1. Frangula alnus P. Mill. E glossy false buckthorn. Rhamnus frangula L. • CT, MA, ME, NH, RI, VT. Forests, forest fragments, roadsides, fields.

Rhamnus 1a. Flowers with 5 sepals, 0 petals, and 5 stamens; leaves alternate; branches unarmed; plants 0.15–1 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. alnifolia

Rh a m nac e a e   7 63

1b. Flowers with 4 sepals, petals, and stamens [Fig. 824]; leaves alternate, subopposite, or opposite (often displaying all arrangements on a given plant); plants often armed with a spine-like process near the apex of some branchlets or in the forks of some branches; plants up to 10 m tall 2a. Petioles 5–15 mm long, (0.03–) 0.10–0.16 (–0.23) times as long as the blade; leaf blades with 5 or 6 (–8) pairs of primary lateral veins, abaxially pubescent with gold-yellow hairs on the veins and/or in the leaf axils (at least when young), turning yellow or gold-yellow abaxially when dry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. utilis 2b. Petioles 10–40 mm long, (0.08–) 0.25–0.50 (–0.75) times as long as the blade; leaf blades with 3–5 (–6) pairs of primary lateral veins; usually some branchlets and branches on a plant with a spine-like process; leaf blades glabrous abaxially or with ± white hairs on the veins and/or in the vein axils, remaining green to pale green abaxially when dry 3a. Leaf blades (1.9–) 3–6.6 cm long, 3 or 4 pairs of lateral veins, elliptic to ovate or broad-ovate; drupe with 3 or 4 pyrenes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. cathartica 3b. Leaf blades (4–) 6–13 cm long, with 4 or 5 (–6) pairs of lateral veins, oblong to elliptic or elliptic-obovate; drupe with 2 pyrenes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. davurica 1. Rhamnus alnifolia ĽHér. N alder-leaved buckthorn. CT, MA, ME, NH, VT. Swamps, fens, often associated with Thuja. 2. Rhamnus cathartica L. E Fig. 824 European buckthorn. CT, MA, ME, NH, RI, VT. Forests, forest fragments, roadsides, fields. 3. Rhamnus davurica Pallas ssp. davurica E Dahurian buckthorn. Rhamnus citrifolia (Weston) Hess & Stern • CT, MA, VT. Forest fragments, roadsides, field edges. 4. Rhamnus utilis Dcne. E Chinese buckthorn. CT. Forest edges, waste areas. Rhamnus utilis has drupes with 2 pyrenes (similar to R. davurica).

Rosaceae Dryas integrifolia Vahl. has been persistently reported to occur in New England since at least 1814. This early report by Pursh was based on a specimen collected by Peck in 1781 from the White Hills of Englée Island off the coast of Newfoundland. Fernald (1903) contended that an error occurred during preparation of the early flora where the White Hills of Newfoundland became confused with the White Mountains of New Hampshire. Therefore, D. integrifolia is not considered part of the New England tracheophyte flora. 1a. Leaf blades simple, the margins entire, toothed, or lobed [Figs. 828, 836] 2a. Plants herbaceous, less than 1 m tall 3a. Androecium with 4 stamens; petals absent; bractlets present and alternating with the sepals (i.e., an epicalyx is present); gynoecium with 1 (–3) ovaries, permanently enclosed within the hypanthium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alchemilla 3b. Androecium with many stamens; petals present (absent in apetalous flowers of R. dalibarda); bractlets absent; gynoecium with 5–many ovaries, not enclosed within the hypanthium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Rubus 2b. Plants trees or shrubs (i.e., woody), commonly taller than 1 m 4a. Gynoecium apocarpous, composed of 3–many distinct carpels; fruit an achene, follicle, capsule, or aggregate of drupes (a drupe in Rhodotypos)

Fig. 824  Flowers of Rhamnus cathartica.

76 4   tricolpate s

5a. Leaves opposite; fruit a drupe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhodotypos 5b. Leaves alternate; fruit an achene, capsule, follicle, or aggregate of drupes 6a. Petals yellow; fruit an achene (i.e., both dry and indehiscent) . . . . . . . . . . Kerria 6b. Petals white to pink or red-purple; fruit an aggregate of drupes, follicle, or capsule (i.e., not both dry and indehiscent) 7a. Fruit fleshy, indehiscent; petals 15–30 mm long; leaf blades 10–20 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Rubus 7b. Fruit dry, dehiscent; petals 2–7 mm long; leaf blades 3–10 cm long ‌vanhouttei); 8a. Leaf blades toothed or double-toothed (often 3-lobed in S. × stipules lacking; carpels not inflated in fruit, dehiscing on 1 suture . . . . . Spiraea 8b. Leaf blades usually 3-lobed; stipules present (or at least stipule scars present on the branchlets); carpels prominently inflated in fruit, dehiscing on both sutures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physocarpus 4b. Gynoecium syncarpous and composed of 2–5 carpels or consisting of a single carpel; fruit a pome, drupe, or 5-angled capsule 9a. Leaf blades entire (sometimes serrate near the apex on vigorous shoots in Exochorda) 10a. Flowers solitary at the apex of leafy branchlets, 4–5 cm in diameter; fruit a pome 30–50 mm in diameter; leaf blades 5–10 × 3–5 cm . . . . . . . . . . . . . . . Cydonia 10b. Flowers borne 2–10 together in a raceme or corymb-like cyme, up to 3.5 cm in diameter; fruit a pome 5–7 mm in diameter or a capsule; leaf blades 0.7–6.5 × 0.5–3.5 cm 11a. Inflorescence a raceme of 4–10 flowers; petals white, 12–20 mm long; fruit a 5-angled capsule with deep sinuses containing winged seeds . . . . Exochorda 11b. Inflorescence a corymb or corymb-like cyme with 2–4 flowers; petals pink, 3–4 mm long; fruit a red pome with (1–) 2 (–3) pyrenes . . . . . . . . . . . Cotoneaster 9b. Leaf blades serrate to lobed (often subentire over part of the margin in Prunus pumila and some Amelanchier) 12a. Flowers with 1 style; fruit a drupe; ovary superior . . . . . . . . . . . . . . . . . . . Prunus 12b. Flowers with 2–5 styles; fruit a pome; ovary inferior, or at least partly so in fruit (note: the ovary is superior in flower in the following genera) 13a. Inflorescence a raceme (+/- a fascicle in A. bartramiana); ovary and fruit 10-locular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amelanchier 13b. Inflorescence a cyme, umbel, or fascicle, in any case lacking an elongate, main axis; ovary and fruit 2- to 5-locular 14a. Carpels hard and bony, difficult to open to expose the seeds; styles distinct; plants usually armed with evident thorns; leaf blades often lobed or with lobe-like teeth, especially on long shoots [Figs. 831, 833, 835] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crataegus 14b. Carpels papery or cartilaginous, easily opened to expose the seeds; styles connate at the base (distinct in Pyrus); plants unarmed or with thorns in Chaenomeles (often with thorn-like short shoots in Malus and Pyrus); leaf blades unlobed (usually lobed in Malus sieboldii) 15a. Leaf midrib with a row of glands on the adaxial surface; petals 4–6 mm long; fruit 0.4–1 cm in diameter, astringent at maturity . . . . . . . . . . . . . Aronia 15b. Leaf midrib without a row of glands on the adaxial surface; petals 15– 25 mm long (5–7 mm long in Sorbus alnifolia); fruit 0.6–12 cm in diameter, sweet or sour-sweet at maturity 16a. Leaf blades prominently double toothed; petals 5–7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Sorbus

Ros ac e a e   76 5

16b. Leaf blades single toothed (sometimes lobed in some Malus); petals 15–25 mm long 17a. Branchlets usually with thorns; carpels each with many ovules; stipules large and conspicuous, mostly 5–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaenomeles 17b. Branchlets without thorns (though the short-shoots often thornlike); carpels each with 2 ovules; stipules small and/or caducous 18a. Styles distinct; anthers ± red (i.e., anthocyanic); orifice of hypanthium nearly closed by a ring of tissue; pome pyriform or globose, with abundant sclerenchyma cells (i.e., with a gritty feel); inflorescence a corymb-like raceme; leaves involute in bud, with a thick, waxy cuticle on the adaxial surface . . . . . Pyrus 18b. Styles connate at the base; anthers yellow to white (i.e., nonanthocyanic); orifice of hypanthium open; pome globose to ovoid, without or with few sclerenchyma cells (i.e., without a gritty feel); inflorescence a fascicle; leaves convolute or conduplicate in bud, without a thick, waxy cuticle adaxially . . . . . . . . . . . . . . . . . . . . Malus 1b. Leaf blades compound [Figs. 841, 849, 871] 19a. Leaves pinnately compound with 3 leaflets or palmately compound [Figs. 841, 849] 20a. Gynoecium with 2–6 (–10) carpels 21a. Petals white (sometimes pale pink); inflorescence at the apex of a leafy stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gillenia 21b. Petals yellow; inflorescence on a ± naked peduncle originating from near the ground level, or sometimes with a small leaf or 2 immediately below the cyme 22a. Leaflets toothed along the margins and at the apex; petals 5–10 mm long; androecium with many stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Geum 22b. Leaflets 3 (–5) toothed at the very apex, the margins entire for much of their length; petals ca. 1 mm long; androecium with 5 stamens . . . . . . Sibbaldia 20b. Gynoecium with many carpels [Figs. 842, 869] 23a. Styles elongate, jointed near or above the middle, the basal portion with a hooked tip, becoming indurate and persistent in fruit, the apical portion deciduous [Fig. 844]; lower and upper stem leaves distinctly different in size, shape, and division . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Geum 23b. Styles short, not jointed near middle, deciduous in fruit [Fig. 839]; lower and upper stem leaves relatively similar in outline, though the upper somewhat reduced in size 24a. Flowers without bractlets alternating with the sepals; carpels maturing as small drupes that are coherent in fruit; plants often armed with bristles or prickles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Rubus 24b. Flowers with bractlets that resemble the sepals and alternate with them (i.e., the sepals appear to number (8–) 10 per flower); carpels maturing as achenes; plants unarmed 25a. Bractlets definitely larger than the sepals, with 3 conspicous teeth at the apex; receptacle enlarged in fruit but rather dry and insipid tasting; petals yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Potentilla 25b. Bractlets of ± similar size as the sepals or smaller, lacking conspicuous apical teeth (though the margins may have small teeth); receptacle either enlarged in fruit and sweet-tasting or not enlarged and remaining dry; petals white to pink or ochroleucous to yellow

76 6   tricolpate s

26a. Receptacle enlarged in fruit and becoming fleshy, sweet-tasting at maturity; petals white or tinged with pink; bractlets ± subequal in size to the sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fragaria 26b. Receptacle neither enlarged nor fleshy in fruit; petals yellow or white; bractlets in most species narrower and/or shorter than the sepals 27a. Style originating from the side of the carpel (i.e., lateral) [Fig. 839, L]; petals white; surface of ovary and achenes pubescent; leaflets entire for most of the margin and with 3 (–5) teeth at the very apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sibbaldiopsis 27b. Style originating from or very near the apex of the carpel (i.e., terminal) [Fig. 839, R]; petals yellow (white in P. alba); surface of ovary and achenes glabrous; leaflets with teeth or lobes usually distributed over more of the margin, restricted to the very apex in a few species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Potentilla 19b. Leaves once- or twice-pinnately compound, always with more than 3 leaflets [Fig. 871] 28a. Plants trees or shrubs (i.e., woody) 29a. Flowers with 2–5 carpels; inflorescence a many-flowered cyme or panicle, usually with 25 or more flowers 30a. Rachis of leaves adaxially with clusters of dark glands at the nodes; flowers with 2–4 carpels, partly epigynous; leaflet margins serrate to obscurely double-serrate [Fig. 871]; inflorescence a flat- to round-topped cyme; fruit a pome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Sorbus 30b. Rachis of leaves lacking dark glands; flowers with 5 carpels, perigynous; leaflet margins double-serrate; inflorescence a tall panicle; fruit a follicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sorbaria 29b. Flowers with many carpels; inflorescence a solitary flower, a corymb, a cyme, or a raceme, most species with an inflorescence with fewer than 25 flowers 31a. Plants unarmed; leaflets entire; fruit and hypanthium not fleshy; petals yellow (often white or ochroleucous in planted populations) . . . . . . . Dasiphora 31b. Plants usually armed with bristles or prickles; leaflets toothed; fruit fleshy or enclosed in a fleshy hypanthium; petals white to pink to red-purple or ochroleucous 32a. Fruit an aggregate of achenes, surrounded and concealed by a fleshy hypanthium; stipules connate to the petiole for at least half their length [Fig. 862]; plants perennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rosa 32b. Fruit an aggregate of drupes, not surrounded by a hypanthium and, therefore, clearly visible [Fig. 869]; stipules distinct from the petiole; plants biennial—producing vegetative stems the first year and reproductive the second . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Rubus 28b. Plants herbaceous (sometimes woody at the very base) 33a. Leaves twice-pinnately compound; fruit dehiscent . . . . . . . . . . . . . . . . Aruncus 33b. Leaves once-pinnately compound; fruit indehiscent 34a. Flowers with 4 petaloid sepals, 0 petals, and 0–2 carpels . . . . Sanguisorba 34b. Flowers with 5 sepaloid sepals, 5 petals, and 2–many carpels 35a. Flowers with 2 carpels; hypanthium armed with hooked bristles [Figs. 825, 826]; inflorescence an elongate raceme . . . . . . . . . . . . . Agrimonia

Rosac e a e   767

35b. Flowers with 5–many carpels; hypanthium not armed; inflorescence a solitary flower, a corymb, or a cyme 36a. Styles elongate, jointed near or above the middle, the basal portion with a hooked tip, becoming indurate and persistent in fruit, the apical portion deciduous [Fig. 844]; lower and upper stem leaves distinctly different in size, shape, and division . . . . . . . . . . . . . . . . . . . . . (in part) Geum 36b. Styles short, not jointed near the middle, deciduous in fruit; lower and upper stem leaves relatively similar in outline, though the upper somewhat reduced in size 37a. Flowers without bractlets alternating with the sepals; petals white or pink to pink-red; carpels 5–15 per flower, arranged in a single whorl . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Filipendula 37b. Flowers with bractlets that resemble the sepals and alternate with them (i.e., the sepals appearing to number (8–) 10 per flower); petals yellow, white, or red; carpels many per flower, spirally arranged in several cycles 38a. Flowers solitary at the tips of leafless peduncles produced from the nodes of an eventually prostrate to spreading stem 39a. Leaves pinnately compound with 5–31 principal leaflets, conspicuously pubescent with gray-white tomentum on the abaxial surface (this sometimes partially concealed by lustrous, sericeous hairs); style lateral, emerging from the side of the carpel [Fig. 839, L] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argentina 39b. Leaves palmately compound with 3–5 leaflets, glabrous or moderately pubescent with villous or sericeous hairs on the abaxial surface; style terminal, emerging from near the apex of the carpel [Fig. 839, R] . . . . . . . . . . . . . . . . . . . . . (in part) Potentilla 38b. Flowers few to numerous, arranged in cymes borne at the summit of a leafy stem 40a. Petals ochroleucous; style subbasal, emerging from near the base of the carpel; plants viscid-pubescent, especially above, the hairs with brown septa; stamens mostly 25–30 per flower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drymocallis 40b. Petals red or yellow; style terminal or lateral; plants not viscid-pubescent; stamens mostly 20 per flower 41a. Petals red; style originating from the side of the carpel (i.e., lateral) [Fig. 839, L]; principal leaves with 5–7 pinnately arranged leaflets; wetland species . . . . . . . . . . . . . . . . Comarum 41b. Petals yellow; style originating from near the apex of the carpel (i.e., terminal) [Fig. 839, R]; leaves with 3–7 palmately or subpalmately arranged leaflets; primarily upland plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) Potentilla

Agrimonia 1a. Hypanthium densely and evidently hirsute in and between the furrows; petals 4–6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. eupatoria 1b. Hypanthium glabrous or stipitate-glandular, sometimes with minute, eglandular hairs in the furrows; petals 2–5 mm long

76 8   tricolpate s

2a. Axis of raceme with abundant stipitate glands [Fig. 825]; leaflets with small, glandular dots on the abaxial surface 3a. Larger leaf blades with 11–23 leaflets excluding the smaller leaflets distributed along the leaf rachis; axis of raceme with dense, short eglandular hairs in addition to the glandular ones and occasional long, eglandular ones . . . . . . . . . . . . . . . . . . . A. parviflora 3b. Larger leaf blades with 3–9 leaflets excluding the smaller leaflets distributed along the leaf rachis; axis of raceme either lacking eglandular hairs or with longer, ascending to spreading, eglandular hairs 4a. Axis of raceme with scattered, divaricately spreading eglandular hairs 0.5–2 mm long [Fig. 825]; hypanthium 3–5 mm tall; outer series of hooked bristles on hypanthium wide-spreading [Fig. 825]; roots all fibrous . . . . . . . . . A. gryposepala 4b. Axis of raceme with sparse (sometimes very sparse), ascending eglandular hairs 0.5–1 mm long; hypanthium 2–2.5 mm long; outer series of hooked bristles on hypanthium ascending to erect [Fig. 826]; some roots tuberous thickened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. rostellata 2b. Axis of raceme without stipitate glands [Fig. 826]; leaflets without glandular dots on the abaxial surface or with them in A. striata 5a. Leaflets with small, glandular dots on the abaxial surface; hypanthium 4–5 mm long; roots all fibrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. striata 5b. Leaflets without glandular dots on the abaxial surface (or these obscure); hypanthium 2–3 mm long; some roots tuberous-thickened 6a. Leaves with 3 or 5 leaflets excluding the smaller leaflets distributed along the leaf rachis; leaflets rounded at apex; hypanthium 2–2.5 mm long; pubescence of stem wide-spreading, 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. microcarpa 6b. Larger leaves with 5–13 leaflets excluding the smaller leaflets distributed along the leaf rachis; leaflets acute to obtuse at the apex; hypanthium 2.5–3 mm long; pubescence of stem loosely ascending or incurved, 2–3 mm long . . . . . A. pubescens 1. Agrimonia eupatoria L. E medicinal agrimony. MA. Wool waste. 2. Agrimonia gryposepala Wallr. N Fig. 825 common agrimony. CT, MA, ME, NH, RI, VT. Forests, forest margins, roadsides. Fig. 825  Fruits of Agrimonia gryposepala with outer bristles of hypanthium reflexed, borne along a stipitate-glandular axis.

3. Agrimonia microcarpa Wallr. E low agrimony. Agrimonia platycarpa Wallr.; A. pubescens Wallr. var. microcarpa (Wallr.) Ahles; A. pumila Muhl. ex Bickn. • CT. Forests, woodlands. Two collections were reported to occur at NEBC by Mehrhoff (1995), although only one was cited. Searches of that herbarium have failed to locate the voucher specimens. 4. Agrimonia parviflora Ait.

NC

southern agrimony. CT, MA; western portion of states. Marshes, marsh edges, stream banks, roadsides. 5. Agrimonia pubescens Wallr. N downy agrimony. Agrimonia bicknellii (Kearney) Rydb.; A. mollis (Torr. & Gray) Britt. • CT, MA, RI, VT; also reported from ME by McGregor (1986), but specimens are unknown. Woodlands, often in regions of high-pH bedrock. 6. Agrimonia rostellata Wallr. N Fig. 826  Fruits of Agrimonia striata with outer bristles of hypanthium ascending, borne along a pubescent axis that is eglandular.

woodland agrimony. CT, MA. Woodlands, ridges, rocky slopes, often in dry-mesic soil. 7. Agrimonia striata Michx. N Fig. 826 roadside agrimony. CT, MA, ME, NH, RI, VT. Mesic to wet-mesic forests, woodlands, fields, forest edges, roadsides.

Rosac e a e   76 9

Alchemilla 1. Alchemilla monticola Opiz E hairy lady’s-mantle. Alchemilla pratensis, auct. non F.W. Schmidt; A. vulgaris, auct. non L.; A. xanthochlora, auct. non Rothm. • CT, MA, ME, NH, VT. Fields, roadsides, waste areas, yards.

Amelanchier Despite its small size, Amelanchier is a difficult genus. Frequent hybridization creates local, novel morphologies that can be very confusing, especially on herbarium sheets where the plants are removed from the context of sympatric species. Several species that exist in multiple races based on ploidy level sometimes differ subtly from one another. There also exist undescribed morphologies in New England, particularly in the eastern half of ME. Use caution when assessing leaf teeth/vein characters, as the character states used in the key are for normal growth. Stump sprouts possess misleading leaves. Given that determinations are significantly easier to perform on flowering material, every effort should be made to collect during this brief time in the spring. References: Wiegand (1912), Jones (1946). 1a. Flowers borne in fascicles of 1–4; leaves imbricate in bud, with the outer ones enfolding the inner ones in bud, at maturity with petioles 2–10 mm long, cuneate at the base; bark often brown; ovary summit conical and pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bartramiana 1b. Flowers borne in racemes of 4–10 (–15); leaves conduplicate and positioned beside one another in bud, at maturity with petioles (8–) 10–30 mm long, rounded to cordate at the base; bark often gray; ovary summit rounded, pubescent or glabrous 2a. Petals 3–6 (–7) mm long, sometimes pollen-bearing [Fig. 827]; plants colonial, with many close stems up to 2 (–3) m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. nantucketensis 2b. Petals (6–) 7–22 mm long, not bearing pollen; plants colonial or not, 0.3–13 m tall 3a. Summit of the ovary pubescent (note: character state visible in flower and fruit); sepals reflexed from near the middle in fruit 4a. Lateral veins of leaf blade visible but not prominent, curved forward, forking 3 or more times, anastomosing and evanescent near the margin, not extending into the teeth [Fig. 829]; margin of leaf blade usually serrulate with 6–10 teeth per cm, the teeth more than twice as many as primary lateral veins (caution: leaves on stump sprouts can show fewer, larger teeth); fruit 7–12 mm in diameter . . . . . . . . . . A. spicata 4b. Lateral veins of leaf blade prominent, nearly straight, forking 1 or more times, extending into the teeth or not; margin of leaf blade coarsely serrate-dentate with 0–5 teeth per cm, sometimes entire near the base (or nearly throughout), the teeth less than twice as many as primary lateral veins; fruit 5–8 mm in diameter (up to 10 mm in diameter in A. gaspensis, but that species with leaves sparsely pubescent during flowering) 5a. Flowering racemes compact, 20–40 (–50) mm long, the rachis and pedicels densely pubescent; lowest pedicel 9–13 (–15) mm long in flower; expanding leaf blades densely tomentose abaxially, retaining the tomentum long after flowering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. humilis 5b. Flowering racemes somewhat open, 40–80 mm long, the rachis and pedicels sparsely pubescent; lowest pedicel (7–) 10–30 mm long in flower; expanding leaf blades soon losing much of the abaxial tomentum and becoming glabrescent post flowering 6a. Low, colonial plants up to 1 m tall; petals 6–9 mm long; leaf blades with 6–13 pairs of primary veins that mostly anastomose near the margin; hypanthium scutelliform, 3–4 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. gaspensis

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6b. Taller shrubs with solitary or clumped stems 1–3 m tall; petals (10–) 11–15 (–20) mm long; leaf blades with 12–15 pairs of primary veins that mostly enter the teeth [Fig. 828]; hypanthium open and shallow, (3.5–) 4–6 (–8) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sanguinea 3b. Summit of the ovary glabrous; sepals ascending to recurved in fruit 7a. Petals 7–12 mm long; racemes straight, erect or ascending, the axis 2.5–6 cm long; sepals ascending to spreading (infrequently recurving) in fruit; stems several to many, close together, forming clumps; mature leaf blades rounded to subcordate at base 8a. Leaf blades sparsely pubescent and often tinged with purple at anthesis, at maturity truncate to subcordate at the base and acute to short-acuminate at the apex, with 5–7 teeth per cm; petals 9–12 mm long; flowering racemes sparsely pubescent on the rachis and pedicels; lower pedicels 20–35 mm long in fruit; sepals ascending to somewhat recurving in fruit . . . . . . . . . . . . . . . . . . A. intermedia 8b. Leaf blades densely pubescent and green at anthesis (the blade color obscured by the hairs), at maturity rounded at the base and rounded or mucronate at the apex, with 6–11 teeth per cm; petals 7–10 mm long; flowering racemes densely pubescent on the rachis and pedicels; lower pedicels 5–10 mm long in fruit; sepals erect to loosely ascending in fruit . . . . . . . . . . . . . . . . . . . A. canadensis 7b. Petals 12–22 mm long; racemes lax or arching, the axis 4–12 cm long; sepals recurved in fruit; stems solitary or few together; mature leaf blades cordate at the base 9a. Leaf blades at anthesis 50–75% expanded, nearly glabrous, strongly tinged with red-purple or purple, essentially glabrous at maturity; lower pedicels 20–50 mm long in fruit; pomes fleshy and sweet . . . . . . . . . . . . . . . . . . . . . . A. laevis 9b. Leaf blades at anthesis less than 50% expanded, densely tomentose on the abaxial surface, green, retaining some pubescence in maturity especially along the midvein and in the axils of the primary lateral veins; lower pedicels 10–25 mm long in fruit; pomes drier and bland tasting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. arborea 1. Amelanchier arborea (Michx. f.) Fern. N downy shadbush. Mespilus arborea Michx. f.; Pyrus wangenheimiana Tausch • CT, MA, ME, NH, RI, VT. Forests, woodlands, ridges, field edges, roadsides, most frequent in well drained soils. 1 × 2. This very rare shadbush hybrid is known from VT. It resembles Amelanchier × ‌neglecta in many features but has relatively more pubescence on the expanding leaf blades and relatively shorter pedicels. 1 × 7. This rare shadbush hybrid is known from MA, VT. It closely resembles Amelanchier arborea but is slightly earlier flowering (i.e., will have dropped most of its petals when other A. arborea in the local area are flowering) and shows relatively more-expanded leaves during anthesis (i.e., 50% or more developed and not tightly folded during anthesis). From A. laevis it is separated by the moderate pubescence on the flowering leaf blades and raceme. 2. Amelanchier bartramiana (Tausch) M. Roemer N mountain shadbush. Amelanchier canadensis (L.) Medik. var. pauciflora Farw.; A. oligocarpa (Torr. & Gray) M. Roemer; A. canadensis (L.) Medik. var. oligocarpa Torr. & Gray.; Pyrus bartramiana Tausch • MA, ME, NH, VT; most frequent in north-temperate to boreal portions of states. North temperate to subalpine forests and forest openings, field edges, ridges, roadsides. ‌ × 7. Amelanchier ×neglecta Egglest. ex G.N. Jones is a relatively common shadbush 2 hybrid in the northern, cooler locations of New England where A. bartramiana is frequent. It is known from MA, ME, NH, VT. It shows short racemes (the axis 2–4 cm long) bearing mostly 4–7 flowers, pedicels 15–25 mm long, petals 8–10 mm long, and ovaries with a pubescent summit. The leaf blades are intermediate between the parental taxa and show sparse pubescence at anthesis. This nothospecies usually flowers prior to either of its parental species at a given site.

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3. Amelanchier canadensis (L.) Medik. N eastern shadbush. Amelanchier canadensis (L.) Medik. var. oblongifolia Torr. & Gray; A. lucida Fern.; A. oblongifolia (Torr. & Gray) M. Roemer; Crataegus racemosa Lam.; Mespilus canadensis L. • CT, MA, ME, NH, RI, VT. Fields, forest borders and openings, roadsides, open rights-of-way. Very rare in VT, many collections determined to be this species were misidentified (specimens at NEBC!). 4. Amelanchier gaspensis (Wieg.) Fern. & Weatherby

NC

Gaspé shadbush. Amelanchier sanguinea (Pursh) DC. var. gaspensis Wieg. • ME; northern portion of state. Ice-scoured river shores, river shore outcrops. This species could easily be confused with Amelanchier spicata given the similarity in habit (e.g., dwarf, colonial); however, it has sparse pubescence on the inflorescences and leaf blades during flowering, whereas A. spicata shows dense pubescence. 5. Amelanchier humilis Wieg. N low shadbush. Amelanchier humilis Wieg. var. compacta Nielsen; A. humilis Wieg. var. exserrata Nielson • VT; mainly in the Lake Champlain Valley but also found in the lower Connecticut River valley. Limestone headlands, woodlands, and rocky lake shores in high-pH bedrock regions. Amelanchier humilis can be difficult to separate from A. spicata, especially those forms of the latter with coarsely toothed leaf blades. The details of the veins (e.g., curvature, degree of branching) on the leaf blades are critical features. 6. Amelanchier intermedia Spach N intermediate shadbush. ME, NH, VT; also reported from CT by Seymour (1982), but specimens are unknown. Fields, forest borders and openings, roadsides, open rights-of-way, swamps, wetland margins. Very likely of hybrid origin, intermediate in many respects between Amelanchier canadensis and A. laevis. 7. Amelanchier laevis Wieg. N smooth shadbush. Amelanchier canadensis (L.) Medik. var. botryapium Gray; A. arborea (Michx. f.) Fern. var. laevis (Wieg.) Ahles • CT, MA, ME, NH, RI, VT. Fields, forest borders and openings, roadsides, open rights-of-way. Among our most common and recognizable species. 7 ‌ × 9. Amelanchier ×wiegandii Nielson is a relatively rare shadbush hybrid in New England known from me. Its leaf blades show the coarsely serrate margin of A. sanguinea with mostly 4 or 5 teeth per cm; however, the lateral veins fork 1 or more times near the margin, and the leaf apex is apiculate to short-acuminate (rather than rounded to obtusely pointed as in A. sanguinea). This putative hybrid is further characterized by sparsely pubescent, bronze- or purple-tinged leaf blades at anthesis with mostly 9–12 pairs of primary lateral veins at maturity and lower pedicels 15–40 mm long at anthesis. 8. Amelanchier nantucketensis Bickn. N Fig. 827 Nantucket shadbush. Amelanchier micropetala (B.L. Robins.) Ashe; A. oblongifolia (Torr. & Gray) M. Roam. var. micropetala B.L. Robins.; A. stolonifera Wieg. forma micropetala (B.L. Robins.) Rehd. • CT, MA, ME, NH, ri. Fields, roadsides, sand plains, ledges, river shore outcrops, ridges. Two forms of this species are found in New England—one with a glabrous ovary summit (nantucketensis s.s.) that is found on the coastal islands of MA (rarely elsewhere) and another with a pubescent ovary summit (micropetala) that is found on the mainland of MA and in ME and NH. This species is very difficult to distinguish from Amelanchier spicata in the absence of flowers. Some forms of A. nantucketensis are readily distinguished from A. spicata—those that grow taller than 2 m and those that produce arrow-straight shoots (A. spicata never exceeds 2 m in the wild and never produces arrow-straight shoots). Unfortunately, these forms of A. nantucketensis are rare. Amelanchier nantucketensis commonly has leaf blades that are subentire in the basal third, whereas A. spicata has leaf blades that are commonly toothed to the base. Unfortunately, variation in these species makes applying this characters difficult.

Fig. 827  Andropetalous flower of Amelanchier nantucketensis.

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9. Amelanchier sanguinea (Pursh) DC. N Fig. 828

Fig. 828  Leaf blade of Amelanchier sanguinea with prominent, relatively straight, and few-branched veins.

round-leaved shadbush. Amelanchier canadensis (L.) Medik. var. rotundifolia (Michx.) Torr. & Gray; Mespilus canadensis L. var. rotundifolia Michx.; Pyrus sanguinea Pursh • MA, ME, NH, VT. Forest edges, fields, cliffs, river shore outcrops and ledges. Tremendous confusion exists in regional herbaria concerning this species. It appears that collectors who observed oval to orbicular leaf blades that were coarsely dentate assumed they had Amelanchier sanguinea in hand; however, many of these collections represent forms of A. spicata, detectable due to the curving, forking, and anastomosing veins. Additionally, several features of the inflorescence and flowers make it relatively easy to distinguish these two species. Amelanchier spicata has a shorter inflorescence that is relatively compact (lowest pedicels 6–18 mm long) and densely pubescent during flowering, with a cup-shaped hypanthium, and petals 7–10 mm long, whereas A. sanguinea has a longer inflorescence that is more open (lowest pedicels (7–) 10–30 mm long) and sparsely pubescent during flowering, with an open, scutelliform hypanthium, and petals (10–) 11–15 (–20) mm long. Many reports of A. sanguinea from MA and all reports from CT are based on A. spicata. 10. Amelanchier spicata (Lam.) K. Koch N Fig. 829 dwarf shadbush. Amelanchier stolonifera Wieg.; Crataegus spicata Lam.; Pyrus ovalis Willd. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest edges, woodlands, sand plains, open rightsof-way, balds. The correct name for this low, colonial shadbush has long been debated. Wiegand (1912) regarded the name Amelanchier spicata as applying to a hybrid, this based solely on height measurements of plants growing in a garden setting originally taken from Canada (Willdenow reported 2–2.5 m tall). Therefore, he named our eastern North American species A. stolonifera. Plants growing in a garden setting should not be used as evidence for inapplicability of a name, especially considering many plants grown in a garden setting are more robust than their wild counterparts (in fact, I have observed this species in a garden in ME where it does grow taller than in the wild). Jones (1946) examined the type specimen and concluded that A. spicata and A. stolonifera were conspecific (observations of type specimens are directly applicable to choice of names). Wiegand also conceded that the type specimen of A. spicata agreed well with A. stolonifera. Therefore, I follow Jones in using Lamarck’s epithet for this well-known shadbush (A. spicata).

Fig. 829  Leaf blade of Amelanchier spicata with less prominent, curving, and branched veins.

Argentina 1a. Leaflets densely sericeous over the abaxial surface with lustrous hairs that overlay and conceal the duller tomentum; stolons, peduncles, and petioles usually villous; achenes as thick as wide, with a deep groove on the abaxial ridge . . . . . . . . . . . . . . . . . . . . . . . . . A. anserina 1b. Leaflets sericeous on the veins of the abaxial side, tomentose on the remainder of the surface; stolons, peduncles, and petioles glabrous to sparsely villous; achenes flattened, not as thick as wide, rounded or weakly grooved on the abaxial ridge . . . . . . . . . . . . . . . . . A. egedii 1. Argentina anserina (L.) Rydb. n common silverweed. Argentina anserina (L.) Rydb. var. concolor (Ser.) Rydb.; Potentilla anserina L.; P. anserina L. var. concolor Ser.; P. anserina L. var. sericea Hayne • MA, ME, NH, VT. River shores, lake shores, roadsides in the vicinity of water bodies. This species is considered introduced in MA and native elsewhere. The epithet “anserina” has been, and continues to be, used too inclusively. Many floras (particular those targeting amateur botanists) consider this to be the species along the Atlantic coast. Reports of this species from CT and RI (e.g., Magee and Ahles 1999) were based on collections of Argentina egedii. 2. Argentina egedii (Wormsk.) Rydb. ssp. groenlandica (Tratt.) A. Löve N Pacific silverweed. Potentilla anserina L. var. groenlandica Tratt.; P. egedii Wormsk. var. groenlandica (Tratt.) Polunin • CT, MA, ME, NH, RI. Saline marshes, Atlantic coast strands.

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Aronia Robertson et al. (1991) placed Aronia (eastern North America) in Photinia (Asia and Central America) based on morphological similarity and phylogenetic evidence. However, Aronia is the older name. Therefore, unless Photinia is formally conserved, it needs to be subsumed under Aronia. 1a. Pome black, generally falling from the plant in the fall and not persistent through the winter; abaxial surface of leaf blades, branchlets, and pedicels glabrous or nearly so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. melanocarpa 1b. Pome red or purple, either long-persistent on the plant or falling in autumn; abaxial surface of leaf blades, branchlets, and pedicels varying from sparsely to densely pubescent (rarely one or more surfaces glabrous in A. floribunda) 2a. Pome red; leaf blades usually varying from moderately to densely pubescent on the abaxial surface, turning bright red in the fall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. arbutifolia 2b. Pome purple; leaf blades usually varying from sparsely to moderately pubescent on the abaxial surface, not or only partially turning red in the fall . . . . . . . . . . . . . . . . . A. floribunda 1. Aronia arbutifolia (L.) Pers. N red chokeberry. Aronia arbutifolia (L.) Pers. var. glabra Ell.; A. pyrifolia Lam.; Mespilus arbutifolia L.; Photinia pyrifolia (Lam.) Robertson & Phipps; Pyrus arbutifolia (L.) L. f.; P. arbutifolia (L.) L. f. var. glabra Cronq.; Sorbus arbutifolia (L.) Heynh. • CT, MA, RI; also reported from ME and NH by Magee and Ahles (1999), but specimens are unknown. Woodlands, fields, roadsides, swamps, peaty soils. 1 × Sorbus aucuparia L. ×Sorbaronia hybrida (Moench) Schneid. is a rare intergeneric hybrid that sometimes escapes cultivation in New England (it is known from MA). The nothogenus is recognized by its leaves, some or all of which are variably pinnate in which the terminal leaflet is substantially larger than the lateral leaflets (often 2 or more times longer), and the leaf is adaxially provided with dark glands both on the rachis clustered at the origins of the leaflets and along the midvein of the terminal leaflet. The hybrid is similar in appearance to Sorbus hybrida L., but that species with larger leaves (7–12 cm long vs. 3–8 cm long), broader flowering inflorescences (6–10 cm wide vs. 2–5 cm wide), and red fruit (vs. purple to black). This nothospecies is pubescent in the inflorescence, along new branchlets, and on abaxial leaf surfaces, and it has a dark purple pome. 2. Aronia floribunda (Lindl.) Spach N purple chokeberry. Aronia arbutifolia (L.) Pers. var. atropurpurea (Britt.) Seymour; A. atropurpurea Britt.; A. prunifolia (Marsh.) Rehd.; Photinia floribunda (Lindl.) Robertson & Phipps; Pyrus floribunda Lindl.; Sorbus arbutifolia (L.) Heynh. var. atropurpurea (Britt.) Schneid. • CT, MA, ME, NH, RI, VT. Swamps, wetland margins, peaty soils, balds, shorelines, fields. This species is frequently treated as the first generation hybrid of A. arbutifolia and A. melanocarpa. However, it is found outside of the range of sympatry of the parental taxa, it produces viable fruit, and it possesses a distinctive morphology. All of these facts suggest this entity should be treated as a species. 3. Aronia melanocarpa (Michx.) Ell. N black chokeberry. Aronia arbutifolia (L.) Ell. var. nigra (Willd.) Seymour; A. nigra (Willd.) Koehne; Mespilus arbutifolia L. var. melanocarpa Michx.; Photinia melanocarpa (Michx.) Robertson & Phipps; Pyrus arbutifolia (L.) L. f. var. nigra Willd.; P. melanocarpa (Michx.) Willd.; Sorbus melanocarpa (Michx.) Heynh. • CT, MA, ME, NH, RI, VT. Woodlands, balds, peatlands, swamps, roadsides. 3 × Sorbus aucuparia L. ×Sorbaronia fallax (Schneid.) Schneid. is a rare intergeneric hybrid that sometimes escapes cultivation in New England (it is known from MA, ME). The nothogenus is recognized by characters described under hybrids of Aronia arbutifolia. This nothospecies is nearly glabrous in the inflorescence, along new branchlets, and on abaxial leaf surfaces and it has a black pome.

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Aruncus 1. Aruncus dioicus (Walt.) Fern. var. dioicus E goat’s beard. Aruncus allegheniensis Rydb. • MA, ME; also reported from RI by George (1997), but specimens are unknown. Fields, roadsides, waste areas.

Chaenomeles 1a. Branchlets scabrous, becoming verrucose when old; pome 30–40 mm in diameter; leaf blade obovate to spatulate, crenate; sepals ovate (rarely suborbicular), 4–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. japonica 1b. Branchlets smooth, not verrucose when old; pome 40–60 mm in diameter; leaf blade elliptic-ovate to elliptic-oblong (rarely obovate), sharply serrate; sepals suborbicular (rarely ovate), 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. speciosa 1. Chaenomeles japonica (Thunb.) Lindl. ex Spach E Japanese flowering-quince. Cydonia japonica (Thunb.) Pers.; Pyrus japonica Thunb. • VT; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Railroads, ditches, roadsides. 2. Chaenomeles speciosa (Sweet) Nakai E common flowering-quince. Chaenomeles lagenaria (Loisel.) Koidzumi; Cydonia japonica (Thunb.) Pers. var. lagenaria (Loisel.) Makino; C. lagenaria Loisel.; C. speciosa Sweet • CT, MA. Roadsides, forest fragments.

Comarum 1. Comarum palustre L. N marsh-cinquefoil. Potentilla palustris (L.) Scop.; P. palustris (L.) Scop. var. parvifolia (Raf.) Fern. & Long; P. palustris (L.) Scop. var. villosa (Pers.) Lehm. • CT, MA, ME, NH, VT. Fens, especially lakeside types, peaty shorelines, wet meadows, marshes.

Cotoneaster 1. Cotoneaster divaricatus Rehd. & Wilson in Sarg. E spreading cotoneaster. MA. Roadsides, yards, forest fragments and edges. This species is responsible for a report of Cotoneaster horizontalis Dcne. from MA (Searcy 2008).

Crataegus Crataegus is taxonomically complex and, as treated here, the second largest genus of tracheophytes in New England. Though the genus does suffer from real phenomena, such as infrequent hybridization, apomixis, and rarely collected (i.e., poorly known) taxa, the major hurdle to learning the genus is the name proliferation by early hawthorn researchers. The following key will work best with flowering material for most species, though some taxa will require mature fruits as well for confident identification. Most measurements were performed on dried specimens, so adjustments may be needed for freshly collected material. Species with moderate to dense pubescence in the inflorescence during flowering usually maintain a few hairs along the peduncle and/or pedicels in fruit. Species with pubescent fruits also showed pubescent hypanthia during flowering. Species with one cycle of stamens have 5–10 (–11) total stamens, whereas those with two cycles of stamens have 12–20 total stamens. When assessing the character states of bracteoles within the inflorescence, those at the very base of the inflorescence should not be used as they are often different from those in the middle and upper portions of the inflorescence (they are often toothed or lobed and

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sometimes of herbaceous texture when the upper are not). Inflorescences/infructescences are always borne on short shoots, but not all short shoots bear flowers/fruits. Leaves from long shoots are usually larger, with more pronounced lobes, and/or broader at the base than short shoot leaves. Erosions on the inner surfaces of the pyrenes are a critical character for the identification of Crataegus; however, these are sometimes caused by insect feeding within the immature pomes. Pomes that contain living insect larvae, have insect exit holes on the exterior of the fruit, have insect feces within the erosions, or have erosions that lead to the outer surface of the pyrene should be considered suspect (i.e., find pomes that do not possess any of these features). The distribution reports for New England Crataegus are hopelessly confused by misidentifications and equivocal specimens. Except for selected species (e.g., those of regional conservation concern), no effort has been made to list all of the erroneous reports. The report of C. magniflora Sarg. from VT by Macklin (2001) is based on a collection C. coccinioides (specimen at VT!). References: Palmer (1952), Phipps and Muniyamma (1980). The following are imperfectly known taxa: Crataegus asperifolia Sarg. This taxon has been considered allied to C. scabrida due to the shallow and/or irregular erosions on the inner surface of some pyrenes. However, the erosions, which are very infrequent, all appear to have been caused by insects. It is known from VT. Crataegus baccata Sarg. This hawthorn is very similar to Crataegus macracantha; however, the pomes are very small and pyrenes are plane on the inner surfaces. It is known from MA. Crataegus bellula Sarg. This taxon has been considered to be an interserial hybrid between series Intricatae (e.g., C. intricata) and series Pruinosae (e.g., C. pruinosa). It is known from MA. Crataegus bristolensis Sarg. This taxon is very close to C. populnea in many important morphological details. Its pyrenes show irregular erosions on the inner surfaces; however, these appear to be caused by insect damage. It is known from MA. Crataegus cyclophylla Sarg. This taxon has been considered allied to C. scabrida due to the shallow and/or irregular erosions on the inner surface of some pyrenes. However, the erosions, which are very infrequent, all appear to be caused by insects. Leaf blade morphology suggests this is an interserial hybrid between series Rotundifoliae (e.g., C. chrysocarpa) and series Tenuifoliae (e.g., C. macrosperma). It is known from VT. Crataegus emersoniana Sarg. This hawthorn appears to be an interserial hybrid derived from series Macracanthae (e.g., C. macracantha), as evidenced by the shallow and/or irregular erosions on the inner surface of some pyrenes. The other possible parent has not been determined with confidence but may involve series Pruinosae (e.g., C. pruinosa). It is known from MA. Crataegus pilosa Sarg. This taxon has been considered to be an interserial hybrid between series Intricatae (e.g., C. intricata) and series Pruinosae (e.g., C. pruinosa). However, key aspects of its morphology contradict this hypothesis, including the absence of bracteoles in the inflorescence. It is known from MA. Crataegus thayeri Sarg. This taxon is related to the C. scabrida complex but appears to be transitional between some taxa in regard to its leaf blade outline. It is known from MA. Crataegus websteri Sarg. This hawthorn is similar to C. egglestonii in many key features and likely has a similar origin—as a hybrid taxon involving series Macracanthae (e.g., C. macracantha). It is known from NH. 1a. Primary lateral veins of the leaf blades running to the sinuses as well as the points of the lobes [Fig. 836] 2a. Flowers with 3–5 styles; leaf blades rounded to subcordate at base, sharply acute at the apex [Fig. 836]; anthers non-anthocyanic; thorns (12–) 23–55 mm long . . . . . C. phaenopyrum 2b. Flowers with 1 style; leaf blades cuneate to broad-cuneate at base, rounded to acute at apex; anthers anthocyanic; thorns 5–17 mm long . . . . . . . . . . . . . . . . . . . . . . . . . C. monogyna

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1b. Primary lateral veins of the leaf blades running only to the points of the lobes or the leaves unlobed and the veins running to the teeth [Fig. 833] 3a. Androecium with 2 cycles of stamens [Fig. 838] 4a. Inflorescence sparsely to densely pubescent, retaining some hairs at least until full maturity of pome 5a. Flowers 20–24 mm wide; leaf blades broad-ovate to broad-triangular or suborbicular and (50–) 61–110 mm long at maturity; pomes usually prominently glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. coccinioides 5b. Flowers 13–20.5 mm wide; leaf blades usually of narrower shape or of shorter length or both; pomes not or only slightly glaucous 6a. Sepals entire or nearly so; leaf blades elliptic to narrow-elliptic-obovate to obovate, usually widest above the middle, barely (if at all) lobed on short shoots, 1.4–2.1 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. punctata 6b. Sepals glandular-serrate to glandular-pinnatifid; leaf blades not combining the above-mentioned characters 7a. Anthers non-anthocyanic; leaf blades near apex of long shoots suborbicular or nearly so, with small, acuminate-tipped lobes . . . . . . . . . . . . . . . . C. oakesiana 7b. Anthers anthocyanic; leaf blades near apex of long shoots not combining the above mentioned characters 8a. Pyrenes with a definite depression occupying most of each half of the inner face (pyrenes plane or with shallow and irregular erosions in the very rare C. pisifera); anthers 0.7–1 (–1.3) mm long; inflorescence relatively large, with (7–) 12–20 flowers; leaf blades mostly rhombic to obovate, those on short shoots often with obscure lobes 9a. Pomes 9–13 mm in diameter; pyrenes with a deep excavation on the inner surfaces; inflorescence villous (glabrous or sparsely villous in very rare forms); flowers 15–20.5 mm in diameter . . . . . . (in part) C. succulenta 9b. Pomes 5–9 mm in diameter; pyrenes plane to shallowly and irregularly eroded on the inner surfaces; inflorescence glabrous to sparsely villous; flowers 13–15 mm in diameter . . . . . . . . . . . . . . . . . . . . . . (in part) C. pisifera 8b. Pyrenes plane on the inner faces; anthers 1.2–1.5 mm long; inflorescence mostly of small to medium size, with 5–12 flowers; leaf blades elliptic or ovate to elliptic-ovate or orbicular, those on short shoots with obscure to evident lobes 10a. Leaf blades adaxially with relatively long and soft appressed hairs mostly 0.3–0.8 mm long; short shoot leaf blades usually broad-cuneate at the base, the basal margins forming an angle of mostly less than 95 degrees; petioles frequently with a narrow, but evident, wing of green tissue, especially near the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. irrasa 10b. Leaf blades adaxially with relatively short and stiff appressed hairs mostly 0.2–0.4 mm long; short shoot leaf blades usually rounded at the base, the basal margins forming an angle of mostly more than 95 degrees; petioles scarcely winged 11a. Leaf blades broad-ovate to suborbicular (occasionally some ovate on short shoots), 1–1.2 times as long as wide [Fig. 835]; pomes globose to, infrequently, short-obloid . . . . . . . . . . . . . . . . . . (in part) C. flabellata 11b. Leaf blades ovate or elliptic-ovate, infrequently broad-ovate, 1.1–1.6 times as long as wide; pomes obloid to obovoid . . . . . . . . . . C. lucorum

Rosac e a e   777

4b. Inflorescence glabrous 12a. Pyrenes with a definite depression occupying most of each half of the inner face (pyrenes plane or with shallow and irregular erosions in the very rare C. pisifera) [Fig. 834, R]; bracts of leaf opening strongly tinged with red to orange-red and conspicuous; inflorescence relatively large, with (7–) 12–20 flowers; sepals glandularserrate to glandular-pinnatifid 13a. Pomes 9–13 mm in diameter; pyrenes with a deep excavation on the inner surfaces; inflorescence villous (glabrous or sparsely villous in very rare forms); flowers 15–20.5 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. succulenta 13b. Pomes 5–9 mm in diameter; pyrenes plane to shallowly and irregularly eroded on the inner surfaces; inflorescence glabrous to sparsely villous; flowers 13–15 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. pisifera 12b. Pyrenes plane on the inner faces or some faces with irregular and shallow erosions [Fig. 834, L]; bracts of leaf opening yellow-brown or weakly tinged with red, less conspicuous; inflorescence mostly of small to medium size, with 4–13 flowers; sepals entire to glandular-serrate 14a. Anthers (1.6–) 1.8–2.3 mm long; flowers 20–23 mm wide; pyrenes plane to shallowly and irregularly eroded on the inner surfaces . . . . . . . . . . . . . C. umbratilis 14b. Anthers 0.7–1.6 mm long; flowers (16–) 17–21.5 mm wide; pyrenes plane on the inner surfaces (shallowly and irregularly eroded on some pyrenes in C. brainerdii, but that species with 3 styles vs. (4–) 5 styles) 15a. Anthers 0.7–1 mm long; pyrenes plane to shallowly and irregularly eroded on the inner surfaces; leaf blades on short shoots mostly cuneate at the base, the basal margins forming an angle of mostly less than 95 degrees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. brainerdii 15b. Anthers 1.1–1.6 mm long; pyrenes plane on the inner surfaces; short shoot leaf blades usually broad-cuneate to rounded, truncate, or subcordate at the base, the basal margins forming an angle of mostly more than 95 degrees 16a. Leaf blades adaxially with abundant strigae during flowering, these persisting at least along the main veins and/or in patches between the veins until early summer (in some taxa persisting nearly until maturation of the fruit); pomes not or only thinly glaucous; pedicels and short shoot branches not glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. schuettei 16b. Leaf blades adaxially without strigae or the hairs sparse to moderately abundant during flowering and absent on mature, fully expanded blades; pomes moderately to prominently glaucous at maturity; pedicels and short shoot branches often thinly glaucous in late season 17a. Leaf blades glabrous adaxially from very early on (rarely with very few hairs); fruiting sepals prominently elevated on a collar 0.5–1.7 mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pruinosa 17b. Leaf blades adaxially with sparse to moderately abundant strigae during flowering; fruiting sepals slightly elevated on a collar 0.3–0.8 mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. forbesae 3b. Androecium with 1 cycle of stamens [Fig. 837] 18a. Bracteoles of inflorescence abundant at flowering, with prominently elevated marginal glands on stipes (0.1–) 0.2–1.4 mm long [Fig. 830]; petioles prominently glandular, sometimes even glandular-toothed; mature pomes yellow-green to bronzegreen or dull red 19a. Foliage, inflorescence, and pome ± glabrous

77 8   tricolpate s

20a. Leaf blades with 4 or 5 pairs of acute to subacuminate lobes, the tips of the lobes spreading or even slightly outcurved, those on long shoots with ovate to oblong-ovate blades; flowers 13–17 mm wide; pome obloid to obovoid, 8–13 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. intricata 20b. Leaf blades with 3 or 4 pairs of obtuse to subacute lobes, the tips of the lobes ascending, those on long shoots with broad-ovate to suborbicular blades; flowers 17–21 mm wide; pome globose to subglobose (rarely short-obloid), 10–14 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. boyntonii 19b. Foliage, inflorescence, and pome (at least when young) pubescent 21a. Anthers non-anthocyanic; pome subglobose to depressed-globose; leaf blades broad-cuneate to rounded, truncate, or subcordate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. biltmoreana 21b. Anthers anthocyanic; pome obovoid to subglobose; leaf blades cuneate at the base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. stonei 18b. Bracteoles of inflorescence absent to moderately abundant at flowering, with sessile to slightly elevated marginal glands on stipes as long as 0.2 mm; petioles glandular or not, but usually not glandular-toothed on short shoot leaves; mature pomes usually orange to red, often brightly so 22a. Inflorescence glabrous 23a. Petioles 2–12 (–15) mm long; leaf blades (1.2–) 1.5–2.5 times as long as wide, usually broadest at or above the middle of the blade, those on short shoots without lobes or with mostly short, obscure lobes 24a. Leaf blades 1.5–2.5 times as long as wide, those on short shoots usually unlobed and widest well above the middle, with faint veins on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. crus‑galli 24b. Leaf blades 1.2–1.8 times as long as wide, those on short shoots often with short lobes and widest near or slightly above the middle, with somewhat evident veins on the adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . C. schizophylla 23b. Petioles 10–40 mm long; leaf blades 1–1.6 times as long as wide, usually broadest at or below the middle of the blade, those on short shoots with obscure to evident lobes 25a. Leaf blades adaxially with abundant strigae during flowering, the hairs persisting at least along the main veins and/or in patches between the veins until early summer (in some taxa persisting nearly until maturation of the fruit); anthers anthocyanic 26a. Short shoot leaves mostly cuneate at the base during flowering, the basal margins forming an angle of less than 90 degrees; pyrenes plane to shallowly and irregularly eroded on the inner surfaces . (in part) C. scabrida 26b. Short shoot leaves mostly broad-cuneate to rounded or truncate (infrequently subcordate) at the base, the basal margins forming an angle of greater than 90 degrees; pyrenes plane on the inner surfaces 27a. Sepals entire or weakly glandular-serrate; flowers 14–16 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. macrosperma 27b. Sepals glandular-serrate, often prominently so; flowers 16–20 mm wide 28a. Short shoot leaves elliptic to elliptic-ovate, 1.2–1.5 times as long as wide [Fig. 833] . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. holmesiana 28b. Short shoot leaves elliptic-ovate to broad-ovate or triangularovate, 1.1–1.3 times as long as wide

Ros ac e a e   779

29a. Pomes obloid to obovoid, 8–10 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. fretalis 29b. Pomes globose to subglobose (rarely short-obloid), 9–12 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. coccinea 25b. Leaf blades adaxially glabrous or with sparse strigae during flowering and then soon glabrous (sometimes with moderately abundant strigae in C. dodgei); anthers anthocyanic or non-anthocyanic 30a. Sepals prominently glandular-serrate to glandular-pinnatifid with elongate calyx teeth, the longest teeth (0.3–) 0.5–1.5 mm long; flowers with 4 or 5 styles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. bicknellii 30b. Sepals entire to glandular-serrate, the longest teeth (when present) 0.1–0.5 (–0.6) mm long; flowers with 2–4 (–5) styles 31a. Short shoot leaves mostly cuneate at the base during flowering, the basal margins forming an angle of less than 90 degrees [Fig. 831]; anthers non-anthocyanic 32a. Leaf blades on short shoots obtuse to subactue at the apex, with acute lobes, borne on glandular (rarely sparsely so) petioles; sepals weakly to somewhat prominently glandular-serrate . . . . . . . . C. flavida 32b. Leaf blades on short-shoots rounded to obtuse at the apex, with rounded to obtuse lobes, borne on eglandular (rarely sparsely glandular) petioles [Fig. 832]; sepals entire to weakly glandular-serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. dodgei 31b. Short shoot leaves mostly broad-cuneate to roundeded or truncate (infrequently subcordate) at the base, the basal margins forming an angle of greater than 90 degrees; anthers anthocyanic (non-anthocyanic in some C. handyae, but that species with erosions on the inner surfaces of some pyrenes) 33a. Inner surface of some pyrenes with shallow and irregular erosions; anthers usually non-anthocyanic . . . . . . (in part) C. handyae 33b. Inner surface of pyrenes plane; anthers anthocyanic 34a. Sepals glandular-serrate; branch thorns very fine, usually 1.6–1.7 mm wide at the midpoint; pomes obloid to obpyriform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. levis 34b. Sepals entire to weakly glandular-serrate; branch thorns thicker, (1.7–) 1.8–2.5 mm thick at the midpoint; pomes globose to subglobose (to obloid or obovoid in some C. populnea, but that species sparsely pubescent adaxially during flowering) 35a. Leaf blades adaxially with sparse to moderately abundant strigae during flowering and then soon glabrous; pomes variable in shape, sometimes elongate and tapering to the base in drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. populnea 35b. Leaf blades adaxially glabrous or with very few hairs during flowering; pomes globose to subglobose, rounded to truncate at the base in drying 36a. Leaf blades with relatively shallow sinuses, the lobes near the middle of the blade on mature leaves 4–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. dissona 36b. Leaf blades with relatively deep sinuses, the lobes near the middle of the blade on mature leaves 7–15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. jesupii

78 0   tricolpate s

22b. Inflorescence sparsely to densely villous 37a. Pyrenes with a definite depression occupying most of each half of the inner face [Fig. 834, R]; bracts of leaf opening strongly tinged with red to orange-red and conspicuous; inflorescence relatively large, with (8–) 10–31 flowers; sepals conspicuously glandular-serrate to glandular-pinnatifid; leaf blades mostly rhombic to obovate, those on short shoots often with inconspicuous lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. macracantha 37b. Pyrenes plane on the inner faces or some faces with irregular and shallow erosions [Fig. 834, L]; bracts of leaf opening yellow-brown or weakly tinged with red, less conspicuous; inflorescence mostly of small to medium size, with 3–16 (–19) flowers; sepals entire or weakly to conspicuously glandular-serrate; leaf blades rhombic or elliptic to ovate, suborbicular, or elliptic-ovate (rarely oblongobovate), those on short shoots with inconspicuous to evident lobes 38a. Leaf blades adaxially glabrous or with very sparse strigae during flowering and then soon glabrous 39a. Leaf blades oblanceolate to broad-oblanceolate-elliptic, 1.5–2.5 times as long as wide, borne on petioles 2–12 (–15) mm long, those on short shoots essentially unlobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. crus‑galli 39b. Leaf blades broad-rhombic or broad-elliptic to ovate, rhombic-ovate broad-ovate, or triangular-ovate, 1–1.3 times as long as wide, borne on petioles 17–35 mm long, those on short shoots with evident lobes 40a. Flowers 19–22 mm wide; pomes globose; some pyrenes with shallow and irregular erosions in the inner surfaces . . . . . . . . . (in part) C. handyae 40b. Flowers 14–16 mm wide; pomes obovoid to obpyriform; pyrenes plane on the inner surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. kennedyi 38b. Leaf blades adaxially with abundant strigae during flowering, the hairs persisting at least along the main veins and/or in patches between the veins until early summer (in some taxa persisting nearly until maturation of the fruit) 41a. Short shoot leaves mostly cuneate at the base during flowering, the basal margins forming an angle of less than 95 degrees 42a. Flowers 19–22 mm wide; sepals entire . . . . . . . . . . . . . . . . C. jonesiae 42b. Flowers 13–19 mm wide; sepals weakly to conspicuously glandular-serrate 43a. Pyrenes plane on the inner surfaces [Fig. 834, L]; anthers nonanthocyanic (anthocyanic in the very rare C. chrysocarpa var. praecox) 44a. Inflorescence sparsely pubescent; hypanthium glabrous or sparsely pubescent near the base; sepals weakly glandular-serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lumaria 44b. Inflorescence moderately to densely pubescent; hypanthium pubescent ± throughout; sepals conspicuously glandular-serrate 45a. Leaf blades broad-rhombic to suborbicular, 36–61 mm long at maturity, 1–1.2 times as long as wide [Fig. 831]; pomes globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. chrysocarpa 45b. Leaf blades rhombic-elliptic to ovate, 48–78 (–87) mm long at maturity, 1.2–1.7 (–1.9) times as long as wide; pomes subglobose to short-obloid . . . . . . . . . . . . . . . . . . . . . . . . . . C. keepii 43b. Some pyrenes with shallow and irregular erosions on the inner surfaces; anthers anthocyanic

Rosac e a e   78 1

46a. Inflorescence moderately villous; hypanthium moderately villous; young branchlets sparsely villous; flowers 18–19 mm wide; pomes obloid to obovoid, sparsely villous; leaf blades elliptic to rhombic or rhombic-ovate, usually widest near the middle, the margins with ± straight taper to the apex . . . . . . . . . . C. laurentiana 46b. Inflorescence sparsely pubescent; hypanthium glabrous or sparsely pubescent; young branchlets glabrous; flowers 14–18 mm wide; pomes globose to subglobose (rarely to short-obloid), glabrous; leaf blades very variable in shape but usually with one or more features: somewhat oblong in outline; broadest above or below the middle; and/or the margins with concave taper to the apex 47a. Branch thorns long, slender, and straight or nearly so, (35–) 45–60 × 1.9–2.1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ideae 47b. Branch thorns variable, usually shorter, more stout, and slightly curved, 22–45 (–65) × (1.9–) 2–2.9 mm 48a. Most leaf blades on short shoots elliptic to ellipticobovate to oblong-obovate, with concave taper to the apex; hypanthium usually glabrous . . . . . . . . . . . (in part) C. scabrida 48b. Most leaf blades on short shoots ovate to rhombic-ovate or broad-ovate, with convex or straight taper to the apex; hypanthium usually sparsely villous near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. egglestonii 41b. Short shoot leaves mostly broad-cuneate to rounded or truncate (infrequently subcordate) at the base, the basal margins forming an angle of greater than 95 degrees 49a. Anthers non-anthocyanic; subherbaceous bracteoles sometimes present in the apical half of the inflorescence; stamens ca. 50% as long as the petals; leaf blades permanently pubescent abaxially on and between the primary lateral veins 50a. Flowers 20–26 mm wide; mature leaf blades 55–111 mm long  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. submollis 50b. Flowers 16–17.5 mm wide; mature leaf blades 36–70 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. faxonii 49b. Anthers anthocyanic; herbaceous bracteoles rarely present in the apical half of the inflorescence; stamens ca. 65–95% as long as the petals; leaf blades abaxially pubescent on only the primary lateral veins at maturity (if at all; sometimes permanently pubescent across the surface in C. holmesiana, but that species usually with obloid to obovoid pomes vs. globose to short-obloid pomes) 51a. Sepals subentire to weakly glandular-serrate (i.e., sepals without marginal glands or these present and then usually sparse and mostly not elevated from the margin more than 0.2 mm), somewhat persistent or deciduous from mature pomes; flowers 15–18 mm wide 52a. Leaf blades broad-ovate to suborbicular (rarely some ovate or broad-triangular-ovate), 1–1.2 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. flabellata 52b. Leaf blades ovate to broad-elliptic or elliptic-ovate, (1.1–) 1.2–1.7 times as long as wide . . . . . . . . . . . . . . . . . . . C. fluviatilis 51b. Sepals glandular-serrate, often conspicuously so (i.e., some of the glands elevated from the margin of the sepal on teeth 0.2–0.6 mm),

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somewhat to firmly persistent on the mature pomes; flowers (16–) 17–21 mm wide 53a. Mature leaf blades (i.e., post-flowering) on short shoots mostly elliptic to elliptic-ovate, 1.2–1.5 times as long as wide [Fig. 833] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. holmesiana 53b. Mature leaf blades on short shoots ovate to suborbicular, 1–1.3 times as long as wide 54a. Pomes obloid to obovoid; hypanthium glabrous; lobes of the leaf blades with relatively more frequent and more prominent outcurved tips [Fig. 835] . . . . . . . . . . . . . . . . . . . . (in part) C. fretalis 54b. Pomes globose to subglobose (rarely to short-obloid); hypanthium villous (sometimes glabrous in C. coccinea); lobes of the leaf blades with relatively fewer and less prominent outcurved tips 55a. Leaf blades ovate to broad-ovate, usually truncate to subcordate at the base, ± plane in life; thorns (20–) 25–60 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) C. coccinea 55b. Leaf blades broad-ovate to suborbicular, usually rounded at the base, often convex in life (i.e., with drooping sides); thorns 15–35 (–42) mm long . . . . . . . . . . . . . . . . . . . . . C. pringlei 1. Crataegus bicknellii Egglest.

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Bicknell’s hawthorn. Crataegus chrysocarpa Ashe var. bicknellii (Egglest.) Palmer; C. rotundifolia Moench var. bicknellii Egglest. • MA; restricted to Nantucket Island. Roadsides, woodlands, field edges. This species has been considered allied to Crataegus chrysocarpa. However, the glabrous inflorescence, shallow erosions on some pyrene inner surfaces, short-acuminate leaf apices, and few, caducous bracteoles ally this plant with the C. scabrida complex. Reports of C. bicknellii from Barnstable County, MA (e.g., Sorrie and Somers 1999), were based on C. macracantha. 2. Crataegus biltmoreana Beadle N Fig. 830 Fig. 830  Bracteole of Crataegus biltmoreana with prominently elevated glands along the margin.

Biltmore hawthorn. Crataegus modesta Sarg.; C. peckii Sarg. • CT, MA, NH, VT. Woodlands, ridges, forest openings. 3. Crataegus boyntonii Beadle N stinking hawthorn. Crataegus baxteri Sarg.; C. bissellii Sarg.; C. foetida Ashe; C. hargeri Sarg. • CT, MA, VT. Woodlands, ridges, fields, forest borders, roadsides. 4. Crataegus brainerdii Sarg.

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Brainerd’s hawthorn. Crataegus seelyana Sarg.; C. shirleyensis Sarg. • MA, VT. Fields, forest borders, roadsides, early successional forests. Crataegus shirleyensis (known from MA) differs in its pubescent inflorescence and hypanthium. It is otherwise very similar to C. brainerdii. It probably deserves recognition at some level. Reports of C. brainerdii from CT, ME, and NH (e.g., Seymour 1982, Kartesz 1999) are erroneous and based on misidentified specimens. 5. Crataegus chrysocarpa Ashe

N C Fig. 831

fireberry hawthorn. 5a. Crataegus faxonii Sarg. var. praecoqua (Sarg.) Kruschke; C. praecoqua Sarg.; C. praecox Sarg. • CT, MA, ME, NH, RI, VT. Fields, forest borders, roadsides, early successional forests, river banks. 1a. Anthers anthocyanic . . . . . . . . . . . . . . . 5a. C. chrysocarpa var. praecox (Sarg.) J.B. Phipps 1b. Anthers non-anthocyanic . . . . . . . . . . . . . . . . . . . . . . . 5b. C. chrysocarpa var. chrysocarpa Fig. 831  Short-shoot leaf blade of Crataegus chrysocarpa.

Variety praecox is known from VT and is of regional conservation concern. Variety chrysocarpa is known from CT, MA, ME, NH, VT.

Ros ac e a e   783

6. Crataegus coccinea L. N scarlet hawthorn. Crataegus lobulata Sarg.; C. pedicellata Sarg,; C. pringlei Sarg. var. lobulata (Sarg.) Egglest.; C. polita Sarg.; C. robesoniana Sarg. • CT, MA, VT. Fields, forest borders, roadsides, early successional forests. 7. Crataegus coccinioides Ashe N Kansas hawthorn. Crataegus coccinioides Ashe var. dilatata (Sarg.) Egglest.; C. dilatata Sarg. • CT, MA,VT. Fields, forest borders, roadsides, early successional forests. 8. Crataegus crus-galli L. n cockspur hawthorn. Crataegus exigua Sarg. • CT, MA, ME, RI, VT. Fields, forest borders, roadsides, early successional forests, lacustrine flood plains. This species appears to be native in the Lake Champlain Valley (in part) of VT and possibly in parts of CT. It is introduced elsewhere. 9. Crataegus dissona Sarg. N northern hawthorn. Crataegus pruinosa (Wendl. f.) K. Koch var. dissona (Sarg.) Egglest. • CT, MA, NH, RI, VT. Fields, forest borders, roadsides, early successional forests. 10. Crataegus dodgei Ashe

N C Fig. 832

Dodge’s hawthorn. Crataegus dodgei Ashe var. rotundata (Sarg.) Kruschke; C. gravesii Sarg.; C. rotundata Sarg. • CT, MA, VT; western New England. Fields, forest borders, roadsides, early successional forests. Reports of this species in ME, NH, and RI (e.g., Seymour 1982, Kartesz 1999) are erroneous and based on misidentified specimens (primarily collections of Crataegus flavida). 11. Crataegus egglestonii Sarg.

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Eggleston’s hawthorn. VT. Fields, forest borders, roadsides, early successional forests. Considered to be closely related to Crataegus scabrida by many authors due to the shared character state of eroded pyrenes but with a very different leaf blade outline (i.e., with a different parentage). See C. scabrida for additional discussion. 12. Crataegus faxonii Sarg.

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Faxon’s hawthorn. Crataegus faxonii Sarg. var. praetermissa (Sarg.) Palmer; C. praetermissa Sarg.; C. rotundifolia Moench var. faxonii (Sarg.) Egglest. • ME, NH, VT; northern New England. Fields, forest borders, roadsides, early successional forests. Reports of this species in CT and MA (e.g., Seymour 1982, Kartesz 1999) are erroneous and based on misidentified specimens. Crataegus faxonii has been relegated to the synonymy of C. chrysocarpa by some authors. However, the broad leaf blade bases, dense indument during flowering, and occasional presence of herbaceous bracteoles suggests this plant may be derived through hybridization of C. chrysocarpa and C. submollis. 13. Crataegus flabellata (Bosc ex Spach) K. Koch N Gray’s hawthorn.  13a. Crataegus grayana Egglest.; 13b. Crataegus crudelis Sarg.; C. lemingtonensis Sarg.; C. propria Sarg. • MA, ME, NH, VT. Forest edges, successional fields, roadsides, early successional forests, forest clearings. 1a. Androecium with 2 cycles of stamens . . . . 13a. C. flabellata var. grayana (Egglest.) Palmer 1b. Androecium with 1 cycle of stamens . . . . . . . . . . . . . . . . . . . 13b. C. flabellata var. flabellata Variety grayana is known from CT, MA, ME, NH, VT. Variety flabellata is known from MA, ME, NH, VT. 14. Crataegus flavida Sarg. N yellow hawthorn. Crataegus chrysocarpa Ashe var. phoenicea Palmer, sensu New England authors; C. dodgei Ashe var. flavida (Sarg.) P.G. Sm. & J.B. Phipps; C. grossa Sarg. • CT, MA, ME, NH, RI, VT. Fields, forest borders, roadsides, early successional forests. Glabrous forms of Crataegus chrysocarpa have long been called C. chrysocarpa var. phoenicea, a problematic name given its absence of a type. Plants labeled as this variety in regional herbaria clearly belong to a separate species (here called C. flavida) as evidenced by glabrous inflorescences and pomes, less prominent teeth on sepals, and differences in leaf blade outline.

Fig. 832  Short-shoot leaf blade of Crataegus dodgei.

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15. Crataegus fluviatilis Sarg. N river hawthorn. Crataegus apiomorpha Sarg. • ME, NH, VT. Fields, forest borders, roadsides, early successional forests. Similar to Crataegus flabellata but with a relatively more elongate leaf blade and always with one cycle of stamens. 16. Crataegus forbesae Sarg.

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Forbes’ hawthorn. CT, MA. Fields, forest borders, roadsides, early successional forests. This species is similar to Crataegus populnea except that it has two cycles of stamens. It is very likely derived from hybridization of a member of series Pruinosae (e.g., C. pruinosa) and a member of series Tenuifoliae (e.g., C. macrosperma). 17. Crataegus fretalis Sarg.

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Long Island Sound hawthorn. Crataegus ampla Sarg. • CT, MA, ME. Fields, forest borders, roadsides, early successional forests. Crataegus fretalis is hypothesized to be of hybrid origin between a member of series Coccineae (e.g., C. coccinea) and a member of series Tenuifoliae (e.g., C. macrosperma; Palmer 1952). Its occurrence in ME, outside the range of any member of series Coccineae, suggests it is acting as a species. 18. Crataegus handyae Sarg.

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Handy’s hawthorn. Crataegus chadsfordiana Sarg.; Crataegus spatiosa Sarg. • CT, MA. Fields, forest borders, roadsides, early successional forests. This rare taxon appears to be derived through hybridization between a member of series Macracanthae (e.g., Crataegus macracantha) and a member of series Pruinosae (e.g., C. pruinosa). Most of the collections shows non-anthocyanic anthers; however, rare collections show anthocyanic anthers. Though these collections were considered synonymous by Sargent (under the name C. chadsfordiana), they are better treated as a new variety but are currently lacking a published name. 19. Crataegus holmesiana Ashe N Fig. 833 Holmes’ hawthorn. Crataegus anomala Sarg., C. eamesii Sarg.; C. holmesiana Ashe var. villipes Ashe; C. tardipes Sarg.; C. villipes (Ashe) Ashe • CT, MA, ME, NH, VT. Fields, forest borders, roadsides, early successional forests.

Fig. 833  Short-shoot leaf blade of Crataegus holmesiana.

20. Crataegus ideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N C Ide’s hawthorn. VT; northern portion of state. Fields, forest borders, roadsides, early successional forests. This poorly known taxon is likely derived from hybridization given the shallow and/or irregular excavations on some of the pyrene inner surfaces, indicating series Macracanthae (e.g., Crataegus macracantha) as a probable parent. 21. Crataegus intricata Lange

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entangled hawthorn. 21a. Crataegus straminea Beadle • CT, MA, NH, RI, VT. Fields, forest borders, roadsides, early successional forests. 1a. Anthers anthocyanic; leaf blades usually broadest near middle; pome yellow to yellow-green (rarely dull orange), 8–10 mm thick . . . . . . . . . . . 21a. C. intricata var. straminea (Beadle) Palmer 1b. Anthers non-anthocyanic; leaf blades usually broadest middle; pome bronze-green to redbrown at maturity, 9–13 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . 21b. C. intricata var. intricata Variety straminea is known from CT and is of regional conservation concern. Variety intricata is known from CT, MA, NH, RI, VT. 22. Crataegus irrasa Sarg. var. blanchardii (Sarg.) Egglest.

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zigzag hawthorn. Crataegus blanchardii Sarg. • ME, VT; mainly northern portion of states. Fields, forest borders, roadsides, early successional forests, rocky balds, ridges. 23. Crataegus jesupii Sarg.

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Jesup’s hawthorn. Crataegus incisa Sarg. • CT, RI, VT; primarily western New England. Fields, forest borders, roadsides, early successional forests. Crataegus incisa is very similar, but some sepals show a few long teeth on the margins. Plants from Berkshire County, MA, are very close to C. jesupii, but show non-anthocyanic anthers and serrate sepals (collections at A!). Their identities have not been determined.

Ros ac e a e   78 5

24. Crataegus jonesiae Sarg.

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Miss Jones’ hawthorn. ME. Fields, forest borders, roadsides, early successional forests. Reports of this species in RI and VT (e.g., Seymour 1982, Kartesz 1999) are erroneous and based on misidentified specimens. 25. Crataegus keepii Sarg. N Keep’s hawthorn. Crataegus brunetiana, auct. non Sarg.; C. searsii Sarg. • MA, ME, NH, VT. Fields, forest borders, roadsides, early successional forests. Crataegus keepii is the proper name for what has passed as C. brunetiana by most New England authors (see C. laurentiana for discussion). 26. Crataegus kennedyi Sarg.

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Kennedy’s hawthorn. VT. Fields, early successional forests. Recent field searches of the type locality have failed to find this hawthorn. It may now be extinct. 27. Crataegus laurentiana Sarg.

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Brunet’s hawthorn. Crataegus brunetiana Sarg.; C. fernaldii Sarg.; C. laurentiana Sarg. var. brunetiana (Sarg.) Kruschke; C. laurentiana Sarg. var. fernaldii (Sarg.) Kruschke • ME. Fields, forest borders, roadsides, early successional forests, river shores and banks. Crataegus brunetiana has been considered to be a member of the C. chrysocarpa compex with more elongate leaf blades. However, the type collections show anthocyanic anthers and shallow erosions on the pyrene inner surfaces, indicating a different relationship. Crataegus brunetiana appears to have a similar evolutionary history as C. laurentiana and is here considered to represent the same taxon. This rare species appears to be derived through hybridization between a member of series Macracanthae (e.g., Crataegus macracantha) and a member of series Rotundifoliae (e.g., C. chrysocarpa). Plants that are actually similar to C. chrysocarpa in indument, anther color, pyrene inner surfaces, etc., but have more elongate leaf blades are properly called C. keepii. 28. Crataegus levis Sarg.

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smooth hawthorn. CT, RI. Fields, forest borders, roadsides, early successional forests. 29. Crataegus lucorum Sarg.

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grove hawthorn. Crataegus insolens Sarg. • VT. Fields, forest borders, roadsides, early successional forests. Crataegus lucorum has been suspected of being an interserial hybrid between a member of series Coccineae (C. coccinea and near relatives) and a member of series Tenuifoliae (C. macrosperma and near relatives). However, it is very similar to C. fluviatilis except for differences in stamen number and subtle differences in leaf blade and pome shape. 30. Crataegus lumaria Ashe

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thorny hawthorn. Crataegus caesariata Sarg.; C. chrysocarpa Ashe var. caesariata (Sarg.) Palmer; C. dodgei Ashe var. lumaria (Ashe) Sarg. • CT, MA. Fields, forest borders, roadsides, early successional forests. 31. Crataegus macracantha Lodd. ex Loud

N C Fig. 834R

long-thorned hawthorn. 31a. Crataegus dumicola Sarg.; C. fertilis Sarg.; C. fulgens Sarg.; C. rhombifolia Sarg.; C. succulenta Schrad. ex Link var. occidentalis Britt.; C. succulenta Schrad. ex Link var. rhombifolia (Sarg.) Egglest.; 31b. Crataegus ferentaria Sarg.; C. pellucidula Sarg.; C. succulenta Schrad. ex Link var. macracantha (Lodd. ex Loud) Egglest.; C. stratfordensis Sarg. • CT, MA, ME, NH, RI, VT. Forest edges, successional fields, roadsides, early successional forests, forest clearings. 1a. Anthers anthocyanic; flowers 15–18 (–19) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31a. C. macracantha var. occidentalis (Britt.) Egglest. 1b. Anthers non-anthocyanic; flowers 17–22 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31b. C. macrcantha var. macracantha Variety occidentalis is known from CT, MA, ME, VT and is of regional conservation concern. Variety macracantha is known from CT, MA, ME, NH, RI, VT

Fig. 834  Inner surface of pyrenes of Crataegus macrosperma (left) and C. macracantha (right).

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32. Crataegus macrosperma Ashe N Fig. 834L, 835 large-seeded hawthorn. Crataegus ascendens Sarg.; C. culta Sarg.; C. florea Sarg.; C. macrosperma Ashe var. acutiloba (Sarg.) Egglest.; C. macrosperma Ashe var. matura (Sarg.) Egglest.; C. macrosperma Ashe var. pentandra (Sarg.) Egglest.; C. macrosperma Ashe var. roanensis (Ashe) Palmer; C. matura Sarg.; C. monstrata Sarg.; C. napaea Sarg.; C. paddockeae Sarg.; C. pentandra Sarg.; C. randiana Sarg.; C. roanensis Ashe; C. serena Sarg.; C. viridimontana Sarg. • CT, MA, ME, NH, RI, VT. Forest edges, successional fields, roadsides, early successional forests, forest clearings. 33. Crataegus monogyna Jacq. E one-seeded hawthorn. CT, MA, ME, NH, RI, VT. Forest edges, fields, lawns, early succesional forests, areas of habitation. This species is often cultivated and frequently shows flore pleno (i.e., double corolla). Corolla color ranges from white to pink-red. Fig. 835  Short shoot leaf blade of Crataegus macrosperma.

34. Crataegus oakesiana Egglest.

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Oakes’ hawthorn. ME, NH, VT. Forest edges, successional fields, roadsides, early successional forests, river banks. 35. Crataegus phaenopyrum (L. f.) Medik. E Fig. 836 Washington hawthorn. Crataegus cordata Ait.; C. populifolia Walt. • MA, ME, RI. Forest edges, fields, early succesional forests, areas of habitation. 36. Crataegus pisifera Sarg.

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pea hawthorn. Crataegus succulenta Schrad. ex Link var. pisifera (Sarg.) Kruschke • VT. Forest edges, successional fields, roadsides, early successional forests. 37. Crataegus populnea Ashe

Fig. 836  Short shoot leaf blade of Crataegus phaenopyrum.

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poplar hawthorn. Crataegus beckwithae Sarg.; C. blandita Sarg.; C. compta Sarg.; C. damei Sarg.; C. demissa Sarg.; C. dissimilis Sarg.; C. fuscosa Sarg.; C. genialis Sarg.; C. glaucophylla Sarg.; C. gravis Ashe; C. iracunda Beadle var. populnea (Ashe) Kruschke; C. iracunda Beadle var. stolonifera (Ashe) Kruschke; C. macrosperma Ashe var. demissa (Sarg.) Egglest.; C. media Sarg., non Bechst.; C. robbinsiana Sarg.; C. stolonifera Sarg. • CT, MA, NH, VT. Forest edges, successional fields, roadsides, early successional forests. Reports of this species in ME and RI by Seymour (1982) and Kartesz (1999, as Crataegus compta and C. iracunda) are erroneous and based misidentified specimens. Though treatments have varied, generally at least three species of this series (Silvicolae) have been attributed to New England—C. brumalis, C. populnea, and C. stolonifera. Collections of the first were often misidentified specimens of C. macrosperma, the remaining were named taxa that belong here in synonymy. Collections of the last are not morphologically distinct from C. populnea when all of the variation on the New England landscape is accounted for (i.e., though the types of C. populnea and C. stolonifera are distinct, there are taxa that Sargent named that completely blur any morphological distinctions). 38. Crataegus pringlei Sarg. N Pringle’s hawthorn. Crataegus exclusa Sarg.; C. pringlei Sarg. var. exclusa (Sarg.) Egglest. • MA, NH, VT. Forest edges, successional fields, roadsides, early successional forests. 39. Crataegus pruinosa (Wendl. f.) K. Koch

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frosted hawthorn.  39a. Crataegus caerulescens Sarg.; C. cognata Sarg.; C. conjuncta Sarg.; C. leiophylla Sarg.; C. porteri Britt.; C. porteri Britt. var. caerulescens (Sarg.) Palmer;  39b. Crataegus festiva Sarg.; C. littoralis Sarg.; C. parvula Sarg.; C. pequatorum Sarg.; C. philadelphica Sarg.; C. pruinosa (Wendl. f.) K. Koch var. latisepala (Ashe) Egglest.; C. pruinosa (Wendl. f.) K. Koch var. leiophylla (Sarg.) J.B. Phipps; C. pruinosa (Wendl. f.) K. Koch var. parvula (Sarg.) J.B. Phipps; C. pruinosa (Wendl. f.) K. Koch var. philadelphica (Ashe) Egglest.; Mespilus pruinosa Wendl. f. • CT, MA, ME, NH, RI, VT; generally restricted to the southern portion of the northern New England states, but extending north in the Lake Champlain Valley of VT. Forest edges, successional fields, roadsides, early successional forests,

Ros ac e a e   7 87

forest clearings. Though this species has been subdivided into numerous varieties (Palmer 1952, Phipps and Muniyamma 1980, etc.), only two groups can be confidently recognized on the New England landscape. 1a. Anthers non-anthocyanic; pomes globose to obovoid to obpyriform, rounded to truncate or tapering to the base; flowers 17–19 mm wide . . . . . 39a. C. pruinosa var. porteri (Britt.) Egglest. 1b. Anthers anthocyanic; pomes globose to obloid, rounded to truncate at the base; flowers 18–22 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39b. C. pruinosa var. pruinosa Variety porteri is known from CT, MA and is of conservation concern. Variety pruinosa is known from CT, MA, ME, NH, RI, VT. 40. Crataegus punctata Jacq. var. punctata N dotted hawthorn. CT, MA, NH, VT. Forest edges, successional fields, riparian and early successional forests. 41. Crataegus scabrida Sarg.

N C Fig. 837

scabrous hawthorn. Crataegus affinis Sarg.; C. brainerdii Sarg. var. scabrida (Sarg.) Egglest.; C. glabrata Sarg.; C. picta Sarg. • NH, VT. Forest edges, successional fields, early successional forests. Crataegus scabrida has been considered to be a complex of taxa derived from hybridization with one parent in common—C. macracantha—due to the shallow and irregular erosions on the pyrene inner surfaces. Four varieties have traditionally been recognized in New England and, in the absence of additional taxa, are quite recognizable—var. asperifolia, var. cyclophylla, var. egglestonii, and var. scabrida. However, there are numerous problems with this approach. First, there are additional named taxa to contend with that blur the distinctions between the aforementioned varietes, including Crataegus thayeri Sarg. and C. websteri Sarg. These two taxa, which have their type locations within New England, have character states (particularly leaf blade shape) that are transitional between the named varities below and suggest that some elements of the C. scabrida complex may be best treated as a complex assesmblage of hybrid derived taxa (i.e., treated without formally recognized varieties). Second, C. scabrida s.s. is quite different from the remaining taxa and probably should be treated separately from them (its leaf blade outline suggests a different parentage). Third, it has been assumed that the erosions on the pyrenes of C. cyclophylla and C. asperifolia were genetically inherited and imply a close relationship to the remaining taxa of this complex. However, their erosions appear to be caused by insect damage, suggesting that taxonomic changes are necessary. 42. Crataegus schizophylla Egglest.

Fig. 837  Flower of Crataegus scabrida with a single cycle of stamens.

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cleft-leaved hawthorn. MA. Forest edges, successional fields, early successional forests. 43. Crataegus schuettei Ashe var. basilica (Beadle) J.B. Phipps N Fig. 838 royal hawthorn. Crataegus alnorum Sarg.; C. basilica Beadle; C. contigua Sarg.; C. declivus Ashe, nomen nudum; C. edsonii Sarg. • CT, MA, ME, NH, VT. Forest edges, successional fields, early successional forests. 44. Crataegus stonei Sarg.

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Stone’s hawthorn. MA. Forest edges, successional fields, early successional forests. 45. Crataegus submollis Sarg. N Quebec hawthorn. Crataegus arnoldiana Sarg.; C. champlainensis Sarg.; Crataegus pennsylvanica Ashe • CT, MA, ME, NH, VT. Forest edges, successional fields, roadsides, early successional forests, forest clearings. 46. Crataegus succulenta Schrad. ex Link var. succulenta

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fleshy hawthorn. Crataegus ambrosia Sarg.; C. ardula Sarg.; C. ferta Sarg.; C. florifera Sarg.; C. gemmosa Sarg.; C. succulenta Schrad. ex Link var. gemmosa (Sarg.) Kruschke; C. venulosa Sarg.; C. virilis Sarg. • MA, VT. Forest edges, successional fields, early successional forests. One of the rarest and most over-reported hawthorns in the region. Most collections determined as this species were in fact C. macracantha.

Fig. 838  Flower of Crataegus schuettei with two cycles of stamens.

78 8   tricolpate s

47. Crataegus umbratilis Sarg.

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shaded hawthorn. CT. Forest edges, successional fields, early successional forests. Likely derived through hybridization with a member of series Macracanthae (e.g., Crataegus macracantha) and a member of series Pruinosae (e.g., C. pruinosa). It is very similar to C. handyae but with two cycles of stamens and very large anthers.

Cydonia 1. Cydonia oblonga P. Mill. E quince. Cydonia vulgaris Pers.; Pyrus cydonia L. • CT, MA. Forest fragments, roadsides, waste areas, areas of habitation. Reports of this species in VT (e.g., Atwood et al. 1973) are based, in part, on collections taken from a cultivated plant—23 Jul 1905, Wheeler s.n. (NEBC!).

Dasiphora 1. Dasiphora floribunda Raf. N shrubby-cinquefoil. Dasiphora fruticosa (L.) Rydb. ssp. floribunda (Pursh) Kartesz; Pentaphylloides floribunda (Pursh) A. Löve; Potentilla floribunda Pursh; P. fruticosa L. ssp. floribunda (Pursh) Elkington • CT, MA, ME, NH, VT. Fens, meadows, ledges, cliffs, and river shore outcrops in regions of high-pH bedrock and/or till.

Drymocallis 1. Drymocallis arguta (Pursh) Rydb. N Fig. 839L tall wood-beauty. Drymocallis agrimonioides (Pursh) Rydb.; Geum agrimonioides Pursh; Potentilla arguta Pursh • CT, MA, ME, NH, VT. Cliffs, river shore ledges, fields, rocky slopes, woodlands. Fig. 839  Achenes and styles of Drymocallis arguta (left) and Potentilla argentea (right).

Exochorda 1. Exochorda racemosa (Lindl.) Rehd. E common pearlbrush. Amelanchier racemosa Lindl.; Exochorda grandiflora (Hook.) Lindl. • CT, MA. Forest fragments, field edges, roadsides, areas of habitation.

Filipendula 1a. Leaves with 7–17 (–25) pairs of well-developed lateral leaflets; carpels pubescent; corolla with (5–) 6–7 (–9) petals [Fig. 840]; roots producing tubers . . . . . . . . . . . . . . . . . . . . . . F. vulgaris 1b. Leaves with 2–5 pairs of well-developed lateral leaflets; carpels glabrous; corolla usually with 5 petals; roots without tubers 2a. Petals pink to red, with irregularly toothed margins; lateral leaflets with 3–5 lobes; terminal leaflet with usually 2 or 3 lobes; fruit stipitate, straight . . . . . . . . . . . . . . . . . . F. rubra 2b. Petals white, with entire margins; lateral leaflets usually without lobes, merely coarsely toothed; terminal leaflet with 3–5 lobes; fruit sessile, twisted . . . . . . . . . . . . . . . . . . F. ulmaria 1. Filipendula rubra (Hill) B.L. Robins. E prairie dropwort. Ulmaria rubra Hill. • CT, MA, ME, NH, VT. Fields, roadsides, trail edges. 2. Filipendula ulmaria (L.) Maxim. E meadow dropwort. Filipendula denudata (J. & K. Presl) Rydb., F. ulmaria (L.) Maxim. ssp. denudata (J. & K. Presl) Hayek; F. ulmaria (L.) Maxim. var. denudata (J. & K. Presl) Hayne ex Maxim., Spiraea ulmaria L. • CT, MA, ME, NH, VT. Fields, roadsides, wastes areas, gardens.

Ros ac e a e   78 9

3. Filipendula vulgaris Moench E Fig. 840 common dropwort. Filipendula hexapetala Gilib. ex Maxim. • CT, ME, VT; also reported from MA by Bean et al. (1967), but specimens are unknown. Fields, roadsides, waste areas.

Fragaria Reference: Staudt (1999). 1a. Terminal tooth of leaflets commonly less than half as wide as adjacent teeth and surpassed by them [Fig. 841]; leaflets usually petiolulate; petals 7–10 (–12) mm long; achenes embedded in the surface of the fruiting receptacle; inflorescence resembling a corymb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. virginiana

Fig. 840  Flower of Filipendula vulgaris.

1b. Terminal tooth of leaflets commonly more than half as wide as adjacent teeth and surpassing them; leaflets usually sessile or nearly so; petals 4–7 mm long; achenes not or only slightly embedded in the surface of the fruiting receptacle; inflorescence resembling a raceme or panicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. vesca 1. Fragaria vesca L. n woodland strawberry.  1a. Fragaria americana (Porter) Britt.; F. vesca L. var. americana Porter; 1b. Fragaria vesca L. var. alba (Ehrh.) Rydb. • CT, MA, ME, NH, RI, VT. Forests, woodlands, outcrops, fields, roadsides, clearings, waste areas. 1a. Flowers 12–14.6 mm wide in life; fruit slender-ovoid to ellipsoid; stolons with appressed hairs in proximal portion . . . . . . . . . . . . . . . . . . . . . . 1a. F. vesca ssp. americana (Porter) Staudt 1b. Flowers 13.2–20.5 mm wide in life; fruit ovoid to subglobose; stolons with spreading hairs in proximal portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. F. vesca ssp. vesca Subspecies americana is native and known from CT, MA, ME, NH, RI, VT. Subspecies vesca is non-native and known from CT, MA, ME, NH, RI, VT. It sometimes produces a white to creamwhite fruit (rather than the normal red color). 2. Fragaria virginiana Duchesne n Fig. 841 common strawberry.  2a. Fragaria glauca (S. Wats.) Rydb.; F. multicipita Fern.; F. virginiana Duchesne var. glauca S. Wats.; F. virginiana Duchesne var. terrae-novae (Rydb.) Fern. & Wieg.; 2b. Fragaria grayana Vilm. ex J. Gay; F. virginiana Duchesne var. illinoensis Gray;  2c. Fragaria canadensis Michx.; F. virginiana Duchesne var. canadensis (Michx.) Farw. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, lawns, clearings, shorelines. 1a. Petioles, peduncles, pedicels, and stolons with appressed to appressed-ascending hairs, often appearing nearly glabrous to the naked eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2a. F. virginiana ssp. glauca (S. Wats.) Staudt 1b. Petioles with spreading hairs [Fig. 841], the hairs visible to the naked eye, the peduncles, pedicels, and stolons either similarly pubescent (especially in the proximal portion) or with appressed to appressed-ascending hairs 2a. Peduncles, pedicels, and stolons with spreading hairs throughout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. F. virginiana ssp. grayana (Vilm. ex J. Gay) Staudt 2b. Peduncles, pedicels, and stolons with appressed to appressed-ascending hairs (sometimes the peduncles and stolons with spreading hairs in the proximal portion or glabrate in the distal portion) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2c. F. virginiana ssp. virginiana Subspecies glauca is native and known from MA, ME, NH, RI, VT. Subspecies grayana is non-native and known from CT, VT. Subspecies virginiana is native and known from CT, MA, ME, NH, RI, VT. 2 ‌ × Fragaria chiloensis (L.) P. Mill. Fragaria ×ananasa Duchesne nssp. ananasa is the commonly planted, cultivated strawberry. It can be recognized by its large fruits (mostly 25–65 mm in diameter vs. 5–20 mm in diameter for F. virginiana), large flowers (25–55 mm in diameter vs. 11.5–25.5 mm in F. virginiana), and thick, often evergreen leaflets. It is known from CT, MA, ME, NH, RI, VT.

Fig. 841  Leaf blade of Fragaria virginiana var. virginiana showing the tiny apical tooth on the terminal leaflet.

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Geum Fresh petal color should be noted on herbarium collections because the color sometimes fades to a dingy yellow-white on old collections, regardless of the color in life. Petal length is an important character for identification. However, the petals do not reach a fully expanded size until the sepals are in a reflexed position for most species (i.e., measuring petal length while the sepals are ascending or merely spreading will result in a shorter measurement than is typical for the species). See Smedmark et al. (2003) for rationale of including Waldsteinia in Geum. 1a. Flowering stems scapose or subscapose, with highly reduced stem leaves or lacking leaves altogether; styles ± straight, no portion conspicuously curved, not articulated or articulated at the base 2a. Basal leaves with 3 leaflets, the lateral leaflets not or only somewhat smaller than the terminal leaflet; gynoecium with 2–6 (–10) carpels; styles deciduous at the base in fruit by an articulation point; petals 5–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . G. fragarioides 2b. Basal leaves with 1 large terminal leaflet and 0–6 pairs of very small lateral leaflets; gynoecium with 30 or more carpels; styles persistent and plumose in fruit; petals 8–15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. peckii 1b. Flowering stems with evident, well-developed leaves; styles articulated near the middle, the basal part hooked at the tip and becoming indurate, the apical part usually deciduous 3a. Epicalyx absent; cluster of achenes stipitate on a stalk 1–2 mm long, elevated above the persistent calyx; petals 1–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. vernum 3b. Epicalyx present; cluster of achenes sessile or nearly so with respect to the calyx; petals 2–10 mm long 4a. Sepals petaloid, purple to red-purple, ascending to erect at anthesis [Fig. 845]; petals erect to ascending; flowers somewhat to conspicuously nodding (becoming erect in fruit) [Fig. 845] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. rivale 4b. Sepals sepaloid, green, reflexed at anthesis [Figs. 842, 843]; petals spreading; flowers erect 5a. Petals yellow to orange-yellow 6a. Basal segment of style minutely glandular [Fig. 844]; terminal segment of basal leaf blades suborbicular to reniform, truncate to cordate at the base, much larger than the lateral segments; pedicels minutely and densely puberulent, sometimes also with scattered, longer hairs; fruiting receptacle glabrous or inconspicuously short-pubescent, the carpel scars plainly visible on the denuded receptacle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. macrophyllum 6b. Basal segment of style eglandular; terminal segment of basal leaf blades oblanceolate to obovate, cuneate at the base, not or scarcely larger than the lateral segments; pedicels both conspicuously hirsute and minutely puberulent [Fig. 842]; fruiting receptacle hirsute, the hairs partially obscuring the carpel scars on the denuded receptacle 7a. Achenes usually numbering 200–250 per cluster, evidently spreadingpubescent near the base of the basal segment of style; petals (5–) 6–10 mm long; pedicels relatively stout, mostly thicker than 1 mm; stipules of stem leaves relatively small, conspicuously reduced compared with the leaflets; leaf teeth usually sharply pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. aleppicum 7b. Achenes usually numbering fewer than 100 per cluster, the basal segment of the style glabrous; petals 4–7 mm long; pedicels relatively slender, mostly thinner than 1 mm; stipules of stem leaves relatively large, ± appearing as a pair of leaflets set close to the stem; leaf teeth usually blunt to obtusely pointed . . . G. urbanum 5b. Petals white to yellow-white

Ros ac e a e   79 1

8a. Pedicels both conspicuously hirsute and minutely puberulent [Fig. 842], relatively stout, mostly thicker than 1 mm; basal leaf blades variable, but commonly with 3–7 principal leaflets (smaller leaflets also often present; sometimes with only 1 principal leaflet); achenes usually numbering more than 160 per cluster [Fig. 842] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. laciniatum 8b. Pedicels minutely and densely puberulent, sometimes also with scattered longer hairs [Fig. 843], relatively slender, mostly thinner than 1 mm; basal leaf blades with usually 3 principals leaflets; achenes usually numbering 30–160 per cluster [Fig. 843] 9a. Petals ochroleucous, 2–4 mm long, much shorter than the sepals; leaf teeth usually blunt to obtusely pointed; stem moderately hirsute (at least in the basal half) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. virginianum 9b. Petals white, (3.5–) 4–9 mm long, nearly as long as or longer than the sepals; leaf teeth sharply pointed; stem glabrous or sparsely hirsute near the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. canadense 1. Geum aleppicum Jacq. ssp. strictum (Ait.) Clausen N Fig. 842

Fig. 842  Fruits of Geum aleppicum borne on hirsute and minutely puberulent pedicels.

yellow avens. Geum aleppicum Jacq. var. strictum (Ait.) Fern.; G. strictum Ait. • ct, ma, me, nh, ri, vt. Forests, woodlands, swamps, fields, roadsides, trail edges, stream banks. North American populations of Geum aleppicum show sparser and shorter pubescence on the faces of the carpels compared with Eurasian populations. On this basis, ssp. strictum is here recognized. Fruiting collections are sometimes difficult to separate from G. laciniatum, especially those forms of the latter with fewer leaflets on basal leaves. Geum aleppicum has evidently pubescent carpels and densely hirsute fruiting receptacles (vs. glabrous carpel bodies or with some hairs restricted to the apex and short-pubescent fruiting receptacles, the carpel scars plainly visible). 2. Geum canadense Jacq. var. canadense N Fig. 843 white avens. Geum canadense Jacq. var. brevipes Fern.; G. canadense Jacq. var. camporum (Rydb.) Fern. & Weatherby; G. canadense Jacq. var. grimesii Fern. & Weatherby • CT, MA, ME, NH, RI, VT. Forests, swamps, clearings, roadsides, waste areas. 3. Geum fragarioides (Michx.) Smedmark

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Appalachian barren-strawberry. Dalibarda fragarioides Michx.; Waldsteinia fragarioides (Michx.) Tratt. • CT, MA, ME, NH, VT. Forests, woodlands, riparian terraces, river banks, fields, clearings, logging roads.

Fig. 843  Fruits of Geum canadense borne on minutely puberulent pedicels.

4. Geum laciniatum Murr. N floodplain avens. CT, MA, ME, NH, RI, VT. Forests, roadsides, fields, forest edges, usually associated with riparian habitats. 1a. Carpel bodies glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a. G. laciniatum var. laciniatum 1b. Carpel bodies bristly pubescent near the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. G. laciniatum var. trichocarpum Fern. Variety laciniatum is known from CT, MA, ME, NH, RI, VT. Variety trichocarpum is known from CT, MA, ME, NH, RI, VT. The morphological variation present in New England populations of this species appears slight (i.e., does it warrant formal taxonomic status?). However, the variation does appear discrete and is reported to be weakly correlated with latitude (var. trichocarpum is more common to the south; Fernald 1950b). See also Geum aleppicum. 5. Geum macrophyllum Willd. N Fig. 844 large-leaved avens. ME, NH, VT; most frequent in the northern portion of states. Forests, swamps, lacustrine forests, fields. ‌5 × 7. Geum ×pulchrum Fern. is a rare hybrid avens in New England known from NH, VT. It shows a purple, spreading sepals 4–5 mm long and gold-yellow, spreading, obovate, clawed petals (the petals of G. macrophyllum are narrowed to the base but not with a conspicuous claw; the petals of G. rivale are clawed but are yellow suffused with purple to entirely purple).

Fig. 844  Achene of Geum macrocarpon showing minute glands at summit of ovary.

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6. Geum peckii Pursh

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White Mountain avens. Acomastylis peckii (Pursh) Bolle; Geum radiatum Michx. var. peckii (Pursh) Gray; Sieversia peckii (Pursh) Rydb. • NH. Alpine ravines, cliffs, and plateaus. Reports of this species in VT are erroneous (Zika 1992). Zinck (1994) found no consistent morphological differences between Geum radiatum Michx., a species of the southern Appalachians, and G. peckii. Paterson and Snyder (1999) performed a RAPD study and found that the genetic divergence between these two taxa is comparable to other species. Based on these results, Paterson and Snyder supported the maintenance of G. peckii as distinct from G. radiatum. This means that in the absence of geographic data, one would be forced to examine the DNA to confidently identify some individuals of these species. 7. Geum rivale L. N Fig. 845

Fig. 845  Nodding flowers of Geum rivale.

water avens. CT, MA, ME, NH, RI, VT. Fens, meadows, swamps, stream banks, low fields, ditches. Plants with green sepals and pale yellow petals has been collected in Kennebec County, ME. It is not yet know whether this occurrence represents a known color morph of Geum rivale (forma virescens) or a hybrid population involving a yellow-flowered species (e.g., G. aleppicum, G. macrophyllum). 8. Geum urbanum L. E town avens. MA, RI, VT. Roadsides, waste areas, gardens. 9. Geum vernum (Raf.) Torr. & Gray

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spring avens. Stylypus vernus Raf. • MA, VT. Rich, deciduous forests. This species is introduced in Berkshire County, MA. 10. Geum virginianum L. N cream-colored avens. Geum flavum (Porter) Bickn.; G. hirsutum Muhl. ex Link • CT, MA, RI. Rich forests, riparian forests, stream banks, meadows. This taxon is much over-reported in New England. Many collections labeled as such are in fact Geum canadense. The report of this species in VT by Magee and Ahles (1999) is based on a collection of G. laciniatum var. laciniatum—1 Jul 1908, Kirk s.n. (MASS!).

Gillenia 1. Gillenia trifoliata (L.) Moench E Bowman’s-root. Porteranthus trifoliatus (L.) Britt. • MA. Forests, forest fragments. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI and was unaware of any collections.

Kerria 1. Kerria japonica (L.) DC. E Japanese-rose. Rubus japonicus • CT, NH, RI. Roadsides, forest borders, about dwellings.

Malus A difficult and frequently cultivated genus. Some plants do not appear to match descriptions of species known to occur in New England, these possibly the result of hybridization or perhaps merely escaped cultivars that have not yet been identified. Malus sargentii Rehd. was reported from ME by Greene et al. (2005). The voucher specimens (at HCOA!) are vegetative and the records are not accepted here. 1a. Leaf blades (at least those on long shoots) usually lobed, conduplicate in bud; pomes with some sclerenchyma cells. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. sieboldii 1b. Leaf blades unlobed, convolute in bud (conduplicate in M. floribunda); pomes lacking sclerenchyma cells

Ros ac e a e   79 3

2a. Calyx lobes deciduous [Fig. 846]; pomes 0.6–1 cm in diameter 3a. Calyx glabrous; young branchlets glabrous; leaves convolute in bud; petiole 20–50 mm long; gynoecium with (4–) 5 styles; mature pome 8–10 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. baccata 3b. Calyx pubescent; young branchlets pubescent; leaves conduplicate in bud; petiole 15–25 mm long; gynoecium with usually 4 styles; mature pome 6–8 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. floribunda 2b. Calyx lobes persistent in fruit; pome 2–12 cm in diameter 4a. Leaf blades sharply serrate; pedicels 20–35 mm long, slender; calyx glabrous or somewhat villous; pomes ovoid, 2–3.5 cm in diameter . . . . . . . . . . . . . . . . . . . M. prunifolia 4b. Leaf blades crenate-serrate; pedicels 10–25 mm long, stout; calyx tomentose; pomes subglobose, (2–) 6–12 cm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pumila 1. Malus baccata (L.) Borkh. E Fig. 846 Siberian crab apple. Pyrus baccata L. • CT, MA, ME, NH, RI, VT. Roadsides, forest borders, successional fields, areas of habitation. 1‌ × 2. Malus ×arnoldiana (Rehd.) Rehd. is a rare apple hybrid known from MA. It is recognized by its ovate leaf blades, yellow pome ca. 10 mm in diameter, and pink petals that gradually pale to white (the petals of M. floribunda begin deep red and pale to nearly white). 2. Malus floribunda Sieb. ex Van Houtte E Japanese flowering crab apple. Malus pulcherrima (Sieb.) Makino • CT, MA. Roadsides, forest borders, successional fields, areas of habitation. 3. Malus prunifolia (Willd.) Borkh. E pear-leaved crab apple. Pyrus prunifolia Willd. • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Roadsides, forest borders, successional fields, areas of habitation. 4. Malus pumila P. Mill. E cultivated apple. Malus domestica (Borkh.) Borkh.; M. sylvestris, auct. non P. Mill.; Pyrus pumila (P. Mill.) K. Koch • CT, MA, ME, NH, RI, VT. Roadsides, forest borders, successional fields, orchards, areas of habitation. ‌4 × M. ioensis (Wood) Britt. Malus ×soulardii (Bailey) Britt. is a very rare apple hybrid known from MA. It is recognized by its broad-ovate to elliptic-ovate leaf blades that are sometimes slightly lobed, have irregularly crenate margins, and often an obtuse apex, short and stout, tomentose pedicels, and a yellow pome that is often red tinged and 50 mm or more in diameter. Malus sieboldii, a naturalized species that has some leaf blades lobed, is different in its sharply serrate leaf blades and small pomes (6–8 mm in diameter). 5. Malus sieboldii (Regel) Rehd. E Toringo crab apple. Pyrus sieboldii Regel • ct, mA. Roadsides, forest borders and fragments, railroads, areas of habitation.

Physocarpus 1. Physocarpus opulifolius (L.) Maxim. n Atlantic ninebark. Opulaster alabamensis Rydb.; O. opulifolius (L.) Kuntze; Spiraea opulifolia L. • CT, MA, ME, NH, RI, VT. Forest fragments, fields, river banks and shores, areas of habitation. Native, in part, to VT (and possibly CT) and introduced in other states. Native populations are usually associated with river banks and shores.

Fig. 846  Pomes of Malus baccata with deciduous calyces.

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Potentilla Phylogenetic study by Erikkson et al. (1998) showed that an inclusive Potentilla is nonmonophyletic and also showed that Duchesnea is nested within it. Therefore, several segregate genera are recognized here (Argentina, Comarum, Dasiphora, and Sibbaldiopsis), and Duchesnea is subsumed under Potentilla. Reference: Rydberg (1898). 1a. Bractlets definitely larger than the sepals, with 3 conspicous teeth at the apex; receptacle enlarged in fruit, somewhat resembling a “strawberry”, but rather dry and insipid tasting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. indica 1b. Bractlets of ± similar size as the sepals or smaller, lacking conspicuous apical teeth (though the margins may have small teeth); receptacle not enlarged and remaining dry 2a. Flowers solitary on axillary peduncles [Fig. 847]; stems slender, herbaceous, prostrate and trailing (though often ascending at anthesis), rooting at some nodes and/or the apex (usually erect to ascending and not rooting in P. erecta); style slender, not thickened near the middle or base (conical at base and thickned near apex in P. erecta and P. reptans, both very rare introductions) 3a. Stems erect to ascending, though sometimes procumbent at anthesis, not rooting at the nodes; stem leaves sessile or subsessile; basal rosette ephemeral, usually absent at anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. erecta 3b. Stems prostrate (though initially ascending), rooting at some nodes and/or the apex; stem leaves petiolate (sometimes sessile or subsessile in P. simplex); basal rosettes persisting through anthesis 4a. Many or all of the flowers with 4 sepals and petals; leaf blades glabrous or glabrate; upper leaves with 3 or 4 leaflets; bractlets lanceolate; stipules narrow-ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. anglica 4b. Flowers with 5 sepals and petals; leaf blades pubescent on the abaxial surface (+/- glabrous in P. reptans); upper leaves with usually 5 leaflets; bractlets linearlanceolate; stipules lanceolate 5a. Petals (6–) 8–12 mm long; anthers (1–) 1.3–2 mm long; epicalyx segments lanceolate to oblong or elliptic; stems from a slender root . . . . . . . . . . . . . P. reptans 5b. Petals 4–7 (–10) mm long; anthers 0.6–1 mm long; epicalyx segments linear to narrow-lanceolate; stems from a thickened rhizome 6a. Flowers from the axil of the first well-developed cauline leaf [Fig. 847]; terminal leaflet usually less than 2 times as long as wide; plants shorter, 5–15 cm at anthesis; rhizome 5–20 mm × 4–8 mm . . . . . . . . . . . . . . P. canadensis 6b. Flowers usually from the axil of the second well-developed cauline leaf; terminal leaflet usually more than 2 times as long as wide; plants taller, 20–40 cm at anthesis; rhizome 20–80 × 5–20 mm . . . . . . . . . . . . . . . . P. simplex 2b. Flowers usually few to many in a terminal cyme (the flowers rarely solitary); stems erect to spreading or sometimes prostrate, but not trailing and rooting (trailing and rooting with woody stems in P. verna), the inflorescence usually with bracts; style thickened near the middle or base 7a. Leaf blades sparsely to densely tomentose on the abaxial surface (long, straight or merely curved hairs also present in some species, especially P. inclinata and P. intermedia) 8a. Styles (1–) 1.5–2.5 (–3) mm long, short-conic near the base and then filiform and prolonged, the longer ones 2–3 times as long as the mature ovary; petals 4–10 (–11) mm long; filaments 1.5–3.5 mm long; anthers 0.7–1.2 (–1.6) mm long

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9a. Leaflets sparsely tomentose abaxially, toothed with elongate, triangularlanceolate teeth mostly 5–15 mm long, the sinuses between the teeth usually extending ⅓ to ½ the distance to the midrib . . . . . . . . . . . . . . . . . . . . . . . P. gracilis 9b. Leaflets densely tomentose abaxially, crenate to crenate-serrate with teeth mostly 2–5 mm long, the sinuses between the teeth usually extending less than ⅓ the distance to the midrib . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pulcherrima 8b. Styles 0.6–1.2 mm long, conic or columnar, shorter than to slightly longer than the mature ovary; petals (2.5–) 4–7 mm long; filaments 0.8–1.5 (–2.5) mm long; anthers 0.3–0.6 (–1.2) mm long 10a. Lower leaves with 5–7 leaflets, these pinnately arranged or the upper 3 confluent and appearing subpalmately arranged [Fig. 848]; styles glandular at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. litoralis 10b. Lower leaves with 4–7 leaflets, these definitely palmately arranged when numbering 5 or more, styles not glandular at the base (though sometimes papillose) 11a. Leaflets revolute, each with 2–4 (–6) tooth-like lobes, densely silvertomentose abaxially; basal leaves much larger than the middle stem leaves; style base papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. argentea 11b. Leaflets with plane or scarcely revolute margins, coarsely serrate to pinnatifid with (3–) 4–10 (–12) teeth or lobes, sparsely to moderately graytomentose abaxially; basal leaves relatively similar in size and outline to the middle stem leaves; style base epapillose 12a. Leaflets pale green abaxially; petals 4–5 (–5.7) mm long; segments of epicalyx oblong-ovate to oblong, obtuse to acute at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) P. intermedia 12b. Leaflets usually gray-green abaxially; petals 5–7 mm long; segments of epicalyx narrow-lanceolate to lanceolate, acute at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. inclinata 7b. Leaf blades glabrous or pubescent with straight or merely curved hairs on the abaxial surface 13a. Lower leaves with 3 leaflets 14a. Plants cespitose or mat-forming, the flowering stems 1–10 (–20) cm tall 15a. Dwarf, cespitose plants without procumbent woody stems, 1–4 cm tall; stipules ovate, blunt to abruptly pointed at the apex; leaflets 5–13 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. robbinsiana 15b. Mat-forming plants with numerous, procumbent woody stems that root at the nodes, the flowering stems up to 10 (–20) cm tall; stipules linear to narrow-lanceolate, sharply pointed at the apex; leaflets (5–) 8–35 (–40) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) P. verna 14b. Plants simple to branched, but neither cespitose nor mat-forming, the flowering stems (10–) 20–120 cm tall 16a. Androecium with 15–20 stamens bearing anthers 0.3–0.5 mm long; stems hirsute; petals 2–5 mm long, slightly shorter than to subequal in length to the sepals; achenes green-brown to brown, striate-ribbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. norvegica 16b. Androecium with (5–) 10 (–15) stamens bearing anthers 0.2–0.3 mm long [Fig. 850]; stems villous, petals 1.3–3 mm long, conspicuously shorter than the sepals [Fig. 850]; achenes yellow-brown, smooth . . . . . (in part) P. rivalis

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13b. Lower leaves with (4–) 5–9 leaflets [Fig. 849] 17a. Petals white; leaflets with a few teeth near the apex; achenes smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. alba 17b. Petals yellow; leaflets with teeth or tooth-like lobes extending from the apex to at least below the middle of the leaflet and often nearly or quite to the base of the leaflet; achenes striate-ribbed or smooth in P. rivalis 18a. Mat-forming plants with numerous procumbent woody stems that root at the nodes, the flowering stems 3–10 (–20) cm tall; style slightly dilated near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) P. verna 18b. Simple or branched, erect to spreading plants, without procumbent, rooting stems, the flowering stems (–15) 25–80 cm tall; style slender at apex 19a. Androecium with (5–) 10 (–15) stamens bearing anthers 0.2–0.3 mm long [Fig. 850]; petals 1.3–3 mm long, conspicuously shorter than the sepals [Fig. 850]; achenes smooth . . . . . . . . . . . . . . . . . . . . (in part) P. rivalis 19b. Androecium with 15–30 stamens bearing anthers 0.6–1.2 (–1.5) mm long; petals subequal to or longer than the sepals; achenes usually striate-ribbed 20a. Petals 4–5 (–5.7) mm long, slightly shorter to slightly longer than the sepals; stems without glandular hairs, often tomentose (at least above, varying to nearly glabrous) . . . . . . . . . . . . (in part) P. intermedia 20b. Petals (6–) 7–12 mm long, slightly to evidently longer than the sepals; stems usually with glandular hairs (in addition to longer, eglandular ones), lacking tomentum 21a. Stems arising centrally from a withered rosette of leaves (i.e., primary leaves borne on the stem); anthers 0.8–1.2 (–1.5) mm long; stipules lobed; achenes 1.2–1.8 mm long . . . . . . . . . . . . . . . . . . P. recta 21b. Stems arising laterally to a persistent basal rosette of leaves (i.e., primary leaves basal); anthers 0.7–0.9 mm long; stipules entire; achenes 1–1.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. thuringiaca 1. Potentilla alba L. E white cinquefoil. CT, MA. Gardens, waste areas. 2. Potentilla anglica Laicharding E English cinquefoil. Potentilla procumbens Sibthorp • ME. Fields, pastures, waste areas. Considered to be a hybrid-derived species with Potentilla erecta and P. reptans as parents. 3. Potentilla argentea L. E Fig. 839R silver-leaved cinquefoil. Potentilla argentea L. var. pseudocalabra T. Wolf • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, clearings, dry lawns. Fig. 847  Habit of Potentilla canadensis showing flower from the axil of the first welldeveloped leaf on the stem.

4. Potentilla canadensis L. N Fig. 847 dwarf cinquefoil. Potentilla canadensis L. var. villosissima Fern.; P. caroliniana Poir.; P. pumila Poir. • CT, MA, ME, NH, RI, VT. Fields, roadsides, clearings, dry lawns. 5. Potentilla erecta (L.) Raeusch. E erect cinquefoil. Tormentilla erecta L. • MA. Waste areas. 6. Potentilla gracilis Doug. ex Hook. var. gracilis E graceful cinquefoil. Potentilla longipedunculata Rydb.; P. macropetala Rydb. • NH. Fields. 7. Potentilla inclinata Vill. E ashy cinquefoil. Potentilla canescens Bess.; P. intermedia L. var. canescens (Bess.) Rupr. • CT. Fields, roadsides, waste areas. This species has been confused with Potentilla intermedia. The

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distributions given here for these two species represent only those collections that have been confirmed (i.e., most literature reports cannot be accepted because of taxonomic confusion); therefore, it should be regarded as tentative. Considered to be a hybrid-derived species with Potentilla argentea and P. recta as parents. 8. Potentilla indica (Andr.) T. Wolf E Indian-strawberry. Duchesnea indica (Andr.) Focke; Fragaria indica Andr. • CT. Lawns, clearings, forest borders, waste areas. 9. Potentilla intermedia L. E downy cinquefoil. CT, MA, NH, RI, VT. Fields, roadsides, waste areas. See Potentilla inclinata for discussion. Considered to be a hybrid-derived species with Potentilla argentea and P. norvegica as parents. 10. Potentilla litoralis Rydb. n Fig. 848 coast cinquefoil. Potentilla pectinata Raf.; P. pensylvanica L. var. litoralis (Rydb.) Boivin; P. pensylvanica L. var. pectinata (Raf.) Boivin; P. pensylvanica L. var. virgulata (A. Nels.) T. Wolf; P. virgulata A. Nels. • ME, NH, VT; primarily near the Atlantic coastline. Fields, open headlands, rock outcrops. Introduced to an open hilltop pasture in VT, but native along the Atlantic coast. This plant belongs to a widespread species complex with a very troubled nomenclatural history. The name Potentilla pectinata Raf., formerly used for this species, is both superfluous and a later homonym.

Fig. 848  Leaf of Potentilla litoralis.

11. Potentilla norvegica L. N Norwegian cinquefoil. Potentilla monspeliensis L.; P. norvegica L. ssp. hirsuta (Michx.) Hyl.; P. norvegica L. var. hirsuta (Michx.) Lehm.; P. norvegica L. var. labradorica (Lehm.) Fern.; P. norvegica L. ssp. monspeliensis (L.) Aschers. & Graebn. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, river shores. Our native North American race (ssp. monspeliensis) generally shows obovate leaflets and hirsute-type hairs on the stem (some high-elevation plants have ± glabrous stem, these recognized by some authors as var. labradorica). The European form (ssp. norvegica), which has been introduced to North America, generally shows oblong leaflets and fine hairs on the stem (Rydberg 1898). However, these differences are not absolute, and many researchers recognize a broadly defined Potentilla norvegica instead of multiple taxa. 12. Potentilla pulcherrima Lehm. E soft cinquefoil. Potentilla camporum Rydb.; P. gracilis Doug. ex Hook. var. pulcherrima (Lehm.) Fern. • NH. Fields. 13. Potentilla recta L. E Fig. 849 sulphur cinquefoil. Potentilla recta L. var. obscura (Nestler) W.D.J. Koch; P. recta L. var. pilosa (Willd.) Ledeb.; P. recta L. var. sulphurea (Lam. & DC.) Peyr. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas.

Fig. 849  Leaf of Potentilla recta.

14. Potentilla reptans L. E creeping cinquefoil. MA. Lawns. 15. Potentilla rivalis Nutt. E Fig. 850 brook cinquefoil. Potentilla millegrana Engelm. ex Lehm.; P. pentandra Engelm.; P. rivalis Nutt. var. millegrana (Engelm. ex Lehm.) S. Wats.; P. rivalis Nutt. var. pentandra (Engelm.) S. Wats. • MA, ME. Waste areas. 16. Potentilla robbinsiana Oakes ex Rydb.

NC

Robbins’ cinquefoil. Potentilla hyparctica Malte ssp. robbinsiana (Oakes ex Rydb.) A. & D. Löve • NH. Alpine plateaus. 17. Potentilla simplex Michx. N old-field cinquefoil. Potentilla simplex Michx. var. argyrisma Fern.; P. simplex Michx. var. calvescens Fern.; P. simplex Michx. var. typica Fern. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, lawns.

Fig. 850  Flower of Potentilla rivalis.

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18. Potentilla thuringiaca Bernh. ex Link E European cinquefoil. RI. Fields, roadsides, waste areas. 19. Potentilla verna L. E spring cinquefoil. Potentilla neumanniana, auct. non Reichenb.; P. tabernaemontani Aschers. • CT. Roadsides, lawns. Report of this species in MA by Kartesz (1999) was based on comments in Seymour (1982) and was unverified (i.e., specimens are unknown).

Prunus 1a. Flowers and fruits sessile or subsessile; ovary and drupe densely pubescent; winter branchlets with 3 axillary buds at each node, the lateral ones inflorescence buds, the central one a leaf bud . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. persica 1b. Flowers and fruits borne on evident pedicels; ovary and drupe glabrous; winter branchlets with a single bud at each node 2a. Drupes grooved, often glaucous; endocarp compressed and ± 2-edged; inflorescence in a fascicle that lacks leafy bracts at the base [Fig. 852]; sepals pubescent on the adaxial surface (at least near the base; glabrous in P. domestica); terminal winter buds absent 3a. Sepals with marginal glands; leaf teeth with a gland (note: the gland sometimes deciduous at maturity of the leaf and its former presence can be detected by observing a small gland scar on the tooth) [Fig. 854] 4a. Flowers solitary or paired (rarely in trios) [Fig. 852]; drupes dark blue to black (red, yellow, or black in P. cerasifera); leaves convolute in bud; branches sometimes tipped by a spine-like process 5a. Young branchlets and bracts densely pubescent; pedicels pubescent 6a. Leaf blades 2–4 × 0.8–1.8 cm; drupes 10–15 mm in diameter; endocarp rugose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. spinosa 6b. Leaf blades 3.5–6 (–8) × 2–4 cm; drupes (10–) 15–25 mm in diameter; endocarp nearly smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. insititia 5b. Young branchlets and bracts glabrous to sparsely pubescent; pedicels glabrous or sparsely pubescent [Fig. 852] 7a. Hypanthium pubescent; leaf blades pubescent on the abaxial surface; petioles 10–20 mm long; flowers 10–20 mm in diameter; drupes 40–75 mm in diameter; branches usually lacking thorns . . . . . . . . . . . . . . . . . . . . . . P. domestica 7b. Hypanthium glabrous; leaf blades glabrous on the abaxial surface or sparsely pubescent along the midvein; petioles 6–12 mm long; flowers 20– 25 mm in diameter; drupes 20–30 mm in diameter; branches sometimes with thorns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cerasifera 4b. Flowers in fascicles of 3–5; drupes red to yellow; leaves conduplicate in bud; branches not tipped by a spine-like process 8a. Petals 8–13 mm long, usually white changing to pink; sepals and pedicels tinged with or completely red; leaf blades coarsely and sometimes doubly serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. nigra 8b. Petals 5–7 mm long, usually remaining white; sepals and pedicels ± green; leaf blades finely crenulate-serrulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. hortulana 3b. Sepals lacking marginal glands (though they may be toothed); leaf teeth without a gland (though sometimes with a callous point) [Fig. 853] 9a. Petals 7–12 mm long; drupes red to yellow, (15–) 20–30 mm in diameter; leaf blades abruptly acuminate at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. americana

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9b. Petals 3–8 mm long; drupes blue-purple to purple-black (rarely yellow), 10–15 (–25) mm in diameter; leaf blades truncate to acute or gradually acuminate at the apex 10a. Leaf blades ovate to elliptic or obovate (suborbicular in var. gravesii), truncate to acute at the apex, pubescent on the abaxial surface; pedicels pubescent; drupes (10–) 13–15 (–25) mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. maritima 10b. Leaf blades lanceolate to oblong-ovate or narrow-obovate, acute to acuminate at the apex, glabrate to sparsely pubescent on the abaxial surface; pedicels glabrous (rarely pubescent); drupes 10–12 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. alleghaniensis 2b. Drupes not grooved, not glaucous; endocarp globose to subglobose (ellipsoid in P. susquehanae); inflorescence a raceme or a fascicle with leafy bracts at the base from the same winter bud (lacking leafy bracts in P. pumila, P. pensylvanica, and P. susquehanae) [Fig. 851]; sepals glabrous on the adaxial surface; terminal winter buds present 11a. Inflorescence a raceme with 18–64 (–90) flowers, the pedicels shorter than the axis of the inflorescence [Fig. 856] 12a. Leaf blades with more than 15 pairs of lateral veins (some of the veins inconspicuous), lanceolate-elliptic to ovate-elliptic, toothed with blunt, callous-tipped teeth, usually pubescent along the abaxial midrib with patches of white, turning red-brown, hairs; sepals oblong to triangular, entire or sparsely glandular-erose, persistent in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. serotina 12b. Leaf blades with mostly 8–11 pairs of lateral veins, oblong to obovate, toothed with sharply pointed teeth, glabrous on the abaxial surface or with some obscure patches of hairs in the axils of the primary veins; sepals broad-triangular to semicircular, conspicuously glandular-serrate, deciduous post anthesis . . . . . P. virginiana 11b. Inflorescence a fascicle with 2–10 flowers, the pedicels much longer than the axis of the inflorescence (if an axis is present) 13a. Leaf blades with teeth that lack a gland [Fig. 853]; low shrubs 0.2–1 (–2.5) m tall 14a. Branchlets glabrous; leaf blades narrow-oblanceolate, 5–18 mm wide, 3–6 times as long as wide [Fig. 855], scarcely glaucous on the abaxial surface; plants depressed, mat-forming, with short, erect branches commonly 1–6 dm tall [Fig. 855]; endocarp subglobse, 6–8 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pumila 14b. Branchlets minutely pubescent; leaf blades oblong to oblong-obovate, 15–30 mm wide, 2–3 times as long as wide, glaucous on the abaxial surface; plants ascending to erect, commonly 3–10 dm tall; endocarp ellipsoid, 5–6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. susquehanae 13b. Leaf blades with teeth that possess a gland (note: the gland sometimes deciduous at maturity of the leaf and its former presence can be detected by observing a small gland scar on the tooth) [Fig. 854]; tall shrubs to trees, 3–20 m tall 15a. Leaf blades broad-ovate to suborbicular, broadly rounded to subcordate at the base; young branchlets glandular-puberulent; inflorescence a 4to 10-flowered raceme with a short axis, the axis present during flowering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. mahaleb 15b. Leaf blades lanceolate or elliptic or elliptic-ovate or elliptic-obovate, cuneate to narrowly rounded at the base; branchlets glabrous or pubescent, but lacking glandular hairs; inflorescence a 2- to 5-flowered fascicle without an axis or sometimes with a very short axis in fruit [Fig. 851] 16a. Petals 4–7 mm long; flowers 12–16 mm in diameter; drupes 5–7 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pensylvanica 16b. Petals 10–15 mm long; flowers 25–40 mm in diameter; drupes (8–) 15–25 mm in diameter

8 00   tricolpate s

17a. Petals emarginate at the apex; sepals ascending to spreading; bracts of inflorescence brown to green-brown, usually deciduous; drupe 8–10 mm in diameter, purple-black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. serrulata 17b. Petals entire at the apex [Fig. 851]; sepals recurved or reflexed (at least post anthesis); bracts of inflorescence green, persistent; drupe 15–25 mm in diameter, red to purple-black (rarely yellow) 18a. Sepals margins glandular-serrate; petiole 10–20 (–25) mm long; proximal margin of leaf teeth much longer than the distal margin, therefore, the gland close to the sinus; leaf blades glabrous or essentially so on the abaxial surface (pubescent in early season) . . . . . . . . P. cerasus 18b. Sepals margins entire; petiole 20–70 mm long; proximal margin of leaf teeth only slightly longer than distal margin, therefore, the gland well removed from the sinus; leaf blades persistently (though sometimes sparsely) pubescent on the abaxial surface . . . . . . . . . . . . . . . . . . . . P. avium 1. Prunus alleghaniensis Porter var. alleghaniensis

nC

Allegheny plum. CT, MA. River banks, sandy river bottoms, fields, roadsides. Native to CT and introduced in MA. Though this species is reported to have glabrous pedicels, most of our New England material shows pubescent pedicels. Further, the sepals show some glands at the apex of the marginal teeth. It is clear that our material of this species, if truly Prunus alleghaneiensis, is not typical for the taxon. 2. Prunus americana Marsh. n American plum. CT, MA, ME, NH, RI, VT. Successional fields, forest borders and fragments, roadsides. Native, in part, to much of CT, the western counties of MA, to RI, and to VT, introduced elsewhere. 3. Prunus avium (L.) L. E Fig. 851

Fig. 851  Inflorescence of Prunus avium with leafy bracts at the base.

sweet cherry. Cerasus avium (L.) Moench • CT, MA, ME, RI; also reported from NH by Hodgdon and Steele (1958), but specimens are unknown. Roadsides, abandoned homesteads, forests, waste areas. Reports of this species in VT are based on a collection of Prunus cerasus— Day 371 (NEBC!). 4. Prunus cerasifera Ehrh. E cherry plum. MA, NH, VT. Roadsides, abandoned homesteads. 5. Prunus cerasus L. E sour cherry. Cerasus vulgaris P. Mill. • CT, MA, ME, NH, RI, VT. Roadsides, forest fragments, abandoned homesteads, forest borders. 6. Prunus domestica L. E Fig. 852

Fig. 852  Inflorescence of Prunus domestica without leafy bracts at the base.

European plum. CT, MA, VT; also reported from NH by Magee and Ahles (1999). Roadsides, abandonded homesteads, forest borders. Prunus domestica is similar to P. insititia. In addition to the characters used in the key, they can be separated by characteristics of their fruit. Prunus domestica has fruits 40–75 mm long with a conspicuously compressed and keeled endocarp that separates from the mesocarp (i.e., flesh). Prunus insititia has fruits (10–) 15– 25 mm long with a somewhat compressed and scarcely keeled endocarp that adheres to the mesocarp. 7. Prunus hortulana Bailey E Hortulan plum. MA. Forest edges. 8. Prunus insititia L. E Damson plum. Prunus domestica L. ssp. insititia (L.) Schneid.; P. domestica L. var. insititia (L.) Boivin • CT, MA, ME, NH, VT Roadsides, abandoned homesteads, forests, waste areas. See Prunus domestica for additional diagnostic characters. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI and was unaware of any collections.

Rosac e a e   8 0 1

9. Prunus mahaleb L. E Mahaleb cherry. CT, MA. Fields, roadsides, abandoned homesteads, forest borders. 10. Prunus maritima Marsh.

N C Fig. 853

beach plum. 10a. Prunus gravesii Small • CT, MA, ME, NH, RI. Fields, roadsides, dunes, forest borders, and sandy openings near the Atlantic coast (rarely inland some distance from the coast). 1a. Leaf blades suborbicular, mostly 1.1–1.3 times as long as wide, 19–38 (–40) mm long; style 4–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . 10a. P. maritima var. gravesii (Small) G.J. Anderson 1b. Leaf blades ovate to oval or obovate, mostly 1.3–2.2 times as long as wide, (27–) 30– 60 (–70) mm long; style 6.1–9.3 mm long . . . . . . . . . . . . . . . . . . . 10b. P. maritima var. maritima Variety gravesii is known from CT and is of conservation concern. Variety maritima is known from CT, MA, ME, NH, RI.

Fig. 853  Leaf blade margin of Prunus maritima showing eglandular teeth.

11. Prunus nigra Ait. n Fig. 854 Canada plum. Prunus americana Marsh. var. nigra (Ait.) Waugh • CT, MA, ME, NH, VT. Fields, roadsides, abandoned homesteads, forest fragments, waste areas. This species is native, in part, to the western counties of MA and in VT and is introduced elsewhere. 12. Prunus pensylvanica L. f. var. pensylvanica N pin cherry. Cerasus pensylvanica (L. f.) Loisel. • CT, MA, ME, NH, RI, VT. Woodlands, forest borders and clearings, logged areas, roadsides, successional fields, waste areas. 13. Prunus persica (L.) Batsch E peach. Amygdalus persica L.; Persica vulgaris P. Mill. • CT, MA, ME, RI, VT. Roadsides, abandoned homesteads, forest borders. 14. Prunus pumila L. var. depressa (Pursh) Bean N Fig. 855 dwarf sand plum. Cerasus depressa (Pursh) Ser.; Prunus depressa Pursh • MA, ME, NH, VT. Sand and gravel river beaches, river shore outcrops, talus slopes, sandy roadsides. Prunus pumila var. pumila was reported from VT by Kartesz (1999); however, the report was erroneous and based on plants of P. susquehanae.

Fig. 854  Leaf blade margin of Prunus nigra showing glandtipped teeth.

15. Prunus serotina Ehrh. var. serotina N black cherry. Padus serotina (Ehrh.) Borkh. • CT, MA, ME, NH, RI, VT; nearly throughout. Deciduous and mixed evergreen-deciduous forests, riparian terraces, roadsides. 16. Prunus serrulata Lindl. E Japanese flowering cherry. Cerasus serrulata (Lindl.) G. Don ex Loud. • MA. Gardens, forest edges. 17. Prunus spinosa L. E

Fig. 855  Inflorescence and depressed stem of Prunus pumila var. depressa.

blackthorn plum. Prunus domestica L. var. spinosa (L.) Kuntze • CT, MA, ME. Fields, roadsides, forest fragments. 18. Prunus susquehanae hort. ex Willd. N Appalachian sand plum. Cerasus susquehanae (hort. ex Willd.) Sweet; Prunus cuneata Raf.; P. pumila L. var. cuneata (Raf.) Bailey; P. pumila L. var. susquehanae (hort. ex Willd.) Jaeger. • CT, MA, ME, NH, RI, VT; absent from much of central and eastern ME and eastern VT. Sandy fields, clearings, roadsides, beaches, and barrens, ridges. 19. Prunus virginiana L. var. virginiana N Fig. 856 choke cherry. Padus virginiana (L.) P. Mill. • CT, MA, ME, NH, RI, VT. Successional fields, forest borders, roadsides, woodlands, openings.

Pyrus 1a. Flowers with 5 styles, 25–30 mm in diamter; hypanthium pubescent; calyx persistent on fruit; pome (5–) 6–16 × 4–12 cm, yellow to green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. communis

Fig. 856  Raceme of Prunus virginiana.

802  tricolpates

1b. Flowers with 2 or 3 styles, 20–25 (–28) mm in diameter; hypanthium glabrous; calyx deciduous from fruit; pome 0.9–1.5 × 0.9–1.5 cm, brown to dark brown and with pale dots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. calleryana 1. Pyrus calleryana Dcne. E Bradford pear. CT, MA, NH. Fields, roadsides, forest borders. 2. Pyrus communis L. E common pear. CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, abandoned homesteads.

Rhodotypos 1. Rhodotypos scandens (Thunb.) Makino . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E black jetbead. Corchorus scandens Thunb. • CT, MA; also reported from NH and VT by Bean et al. (1967), but specimens are unknown. Forest fragments, waste areas.

Rosa Prickle stature and density are important characters for Rosa identification. They must be assessed on normal shoots. Plants sometimes produce vigorous shoots with greatly elongated growth (identifiable by the absence of both abrupt color changes of the bark and annual scars normally found at the junction of year’s growth). These vigorous shoots are usually much more densely prickly than the normal shoots and are misleading for identification. Stipule morphology is also very important and should not be assessed on the apical-most one or two leaves, as these are sometimes different from those near the base and middle of the branchlet. Peduncle and pedicel bracts are frequently caducous and often are shed soon after flowering. They are best observed on peduncles supporting budding flowers. Species noted to have pinnatifid sepals actually bear flowers with 2 entire sepals, 2 sepals bearing slender lobes on both margins, and 1 sepal bearing slender lobes on only 1 margin. Many of the cultivated species from Europe that are naturalized in New England normally produce double corollas (flore pleno; e.g., R. gallica [Fig. 859], R. cinnamomea). References: Lewis (1957), Graham and Primavesi (2005), Joly and Bruneau (2007). 1a. Styles connate (though sometimes becoming distinct, in part, in age), the stigmas conspicuously exserted 3–6 mm from the orifice of the hypanthium and ± equaling the height of the stamens [Fig. 860]; inflorescences (1–) 3- to 30-flowered; stems climbing or scrambling 2a. Leaflets 30–70 (–90) × 10–40 mm, numbering 3 or 5 (–7) per leaf; petals (10–) 20–30 mm long 3a. Petals pink, fading toward white prior to falling; styles glabrous; leaf blades firmherbaceous to subcoriaceous, dull to sublustrous adaxially, deciduous . . . . . . R. setigera 3b. Petals white; styles pubescent; leaf blades coriaceous, lustrous adaxially, evergreen or partly evergreen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. sempervirens 2b. Leaflets 8–30 (–45) × 8–25 mm, numbering (5–) 7 or 9 (–11) per leaf; petals 7–15 (–20) mm long 4a. Stipules fimbriately or pectinately toothed; styles glabrous; leaflets oblong or elliptic to obovate, herbaceous, deciduous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. multiflora 4b. Stipules dentate; styles pubescent; leaflets suborbicular to broad-ovate or broadobovate, firm, sometimes partly evergreen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. luciae 1b. Styles distinct (though often closely connivent), the stigmas barely to slightly exserted beyond the orifice of the hypanthium (i.e., up to 2 mm) and shorter than the stamens; inflorescences 1- to 6 (–15)-flowered; stems erect to arched

Ros ac e a e   8 03

5a. Flowers solitary at the tips of branches (rarely in pairs or trios); peduncle borne from the axil of a leaf but without additional bracts; stems often provided with abundant prickles and bristles (i.e., prickles evidently varying in thickness) 6a. Leaves with 7–11 leaflets 5–20 × 5–10 mm; petals 10–25 mm long, white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. spinosissima 6b. Leaves with 3–7 leaflets 20–60 × 20–30 mm; petals 25–35 mm long, pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. gallica 5b. Flowers solitary or 2–6 and arranged in a corymb; peduncle and/or pedicels bracteates [Fig. 861]; stems unarmed or provided with sparse to abundant prickles, when prickles abundant, then of relatively uniform thickness (except in R. rugosa, but then the prickle bases pubescent) 7a. Branchlets, young prickles, and older prickle bases densely tomentose; fruiting hypanthia 20–30 mm wide, borne on outward-curved to down-curved pedicels; petals mostly (30–) 35–50 mm long; leaflets strongly rugose-veiny . . . . . . . . . . . . . . . . . R. rugosa 7b. Branchlets, young prickles, and older prickle bases glabrous or sparsely pubescent; fruiting hypanthia 7–20 mm wide, borne on straight to weakly curved pedicels (sometimes curved in R. acicularis); petals 9–28 mm long (up to 45 mm long in R. gallica, but that species with pinnatifid sepals); leaflets not or only somewhat rugose-veiny 8a. Orifice of the hypanthium 1–2 mm wide, the stigmas exserted 1–2 mm beyond it (i.e.; the apical portion of the styles visible in intact flowers and fruits); sepals pinnatifid; achenes usually lining the inner wall of the hypanthium as well as the receptacle 9a. Leaflets ± single serrate, lacking marginal stipitate glands and lacking glands on the surfaces; pedicels glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. canina 9b. Leaflets double serrate, stipitate-glandular on the margin and on one or both surfaces (usually lacking glands on the surfaces in R. gallica); pedicels stipitate-glandular 10a. Pedicels (10–) 20–60 mm long, stout; petals 25–35 (–45) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. gallica 10b. Pedicels 6–20 mm long, moderately thick to slender; petals 8–25 mm long 11a. Foliar glands subsessile, when fresh mostly less than 0.05 mm in diameter, brown-red to translucent, and with a resinous odor; leaflets tomentose abaxially (at least along the veins), sparsely tomentose adaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. sherardii 11b. Foliar glands evidently stalked, when fresh mostly 0.05–0.1 mm in diameter, translucent, and with a sweet, fruity odor; leaflets pubescent along the major veins abaxially and glabrous to sparsely pubescent adaxially 12a. Styles pubescent; sepals erect-ascending post anthesis, sometimes persisting in fruit; leaflets evidently glandular on the abaxial surface or on both surfaces, very aromatic; prickles of unequal sizes, ranging from acicular to stout; mature hypanthia 10–15 mm wide . . . . . . . R. rubiginosa 12b. Styles glabrous; sepals reflexed post anthesis, deciduous in fruit; leaflets glandular on only the abaxial surface, weakly aromatic; prickles of ± equal size, none of them acicular; mature hypanthia 7–10 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. micrantha 8b. Orifice of the hypanthium (1.5–) 2–4 mm wide, the stigmas barely exserted beyond it and closing off the opening (i.e., the styles not visible in intact flowers and fruits); sepals entire or with 1–4 linear appendages; achenes confined to the receptacle or also lining the walls of the hypanthium 13a. Pedicels and hypanthium stipitate-glandular [Fig. 861]; sepals spreading or reflexed after anthesis, promptly deciduous in fruit; achenes confined to the receptacle

8 04  tricolpate s

14a. Leaflets with fine teeth, the teeth near the middle of the leaflet (0.3–) 0.4–0.7 mm long; plants of hydric soils 15a. Stems with a pair of stout, broad-based, often curved prickles at most of the nodes, these prickles much larger than the few, slender ones of the internodes; leaflets dull or scarcely lustrous adaxially, the terminal one with (16–) 21–25 (–27) teeth per margin; hypanthium with (40–) 70–98 (–135) stipitate-glands; connate portion of the stipules with ± parallel margins, scarcely widened distally; inflorescence with usually 2 or more flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. palustris 15b. Stems with numerous internodal prickles that are slender, small-based, and straight, similar to the prickles found at the nodes; leaflets lustrous on the adaxial surface, the terminal one with (13–) 14–20 (–23) teeth per margin; hypanthium with (8–) 35–55 (–80) stipitate-glands; connate portion of the stipules without parallel margins, widened distally; inflorescence with usually 1 flower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. nitida 14b. Leaflets with coarse teeth, the teeth near the middle of the leaflet (0.4–) 0.7–0.9 (–1.3) mm long; plants of mesic to xeric soils (infrequently of hydric soils in R. virginiana) 16a. Connate portion of the stipules without parallel margins, widened distally, each wider than 1.1 mm [Fig. 862, L]; nodal prickles usually relatively stout, sometimes curved; internodal prickles usually absent on branchlets; leaflets lustrous adaxially; inflorescence commonly with 3 or more flowers; petioles rarely bristly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. virginiana 16b. Connate portion of the stipules with ± parallel margins, scarcely widened distally, each narrower than 1.1 mm [Fig. 862, R]; nodal prickles usually slender and straight; internodal prickles often present on branchlets; leaflets dull or scarcely lustrous adaxially; inflorescence commonly with 1 or 2 flowers; petioles commonly bristly . . . . . . . . . . . . . . . . . . . . . . . . . . R. carolina 13b. Pedicels and hypanthium glabrous (pedicels or both the pedicels and hypanthia stipitate-glandular in rare forms); sepals erect or connivent after anthesis (reflexed in R. blanda var. glabra), persistent in fruit (frequently deciduous at maturation of fruit in R. glauca); achenes usually lining the inner wall of the hypanthium as well as the receptacle 17a. Stems with a pair of stout, broad-based, often curved prickles at most of the nodes [Fig. 858], these prickles much larger than the few slender ones of the internodes; corolla usually double [Fig. 859] . . . . . . . . . . . . . R. cinnamomea 17b. Stems without prickles or with few to numerous prickles, the prickles of the nodes not strongly differentiated from those of the internodes (i.e., all the prickles of ± similar dimension); corolla usually single 18a. Leaflets softly tomentose on both surfaces, abaxially with tiny, red to red-brown, subsessile glands that present a resiny odor when bruised on living plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. mollis 18b. Leaflets glabrous to pubescent, but not tomentose on both surfaces, lacking aromatic resin glands (but sometimes with subsessile glands in R. acicularis) 19a. Leaflets glaucous and/or stronged tinged with red to brown-purple, single-serrate on the margin; branchlets glaucous, the bloom usually persisting into fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. glauca 19b. Leaflets green, single or double-serrate on the margin; branchlets without bloom or glaucous only when young

Ros ac e a e   8 0 5

20a. Leaves with (7–) 9 or 11 leaflets; leaflets obovate to oblongobovate; inflorescence with (1–) 3 or 4 (–5) flowers, usually borne both on lateral branchlets from branches of the previous year and on terminal branchlets of the current season . . . . . . . . . . R. arkansana 20b. Leaves with 5 or 7 leaflets; leaflets elliptic or oblong to ovate or obovate; inflorescence with 1 or 2 (–3) flowers, usually borne only on lateral branchlets that are borne on branches of the previous year 21a. Branchlets with numerous prickles [Fig. 857]; usually the upper stipules, floral bracts, petioles, and/or leaf rachises stipitateglandular, at least when young; fruiting hypanthium dark blue in drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. acicularis 21b. Branchlets without prickles (rarely with a few prickles); upper stipules, floral bracts, petioles, and leaf rachises without stipitate glands (though often with sessile glands at the tips of the teeth of the stipules and/or floral bracts); fruiting hypanthium usually red to red-brown in drying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. blanda 1. Rosa acicularis Lindl. ssp. sayi (Schwein.) W.H. Lewis

N C Fig. 857

bristly rose. Rosa acicularis Lindl. var. bourgeauiana (Crépin) Crépin; R. bourgeauiana Crépin; R. sayi Schwein. • MA, ME, NH, VT; also reported from RI by Kartesz (1999), but specimens are unknown. Ridges, cliffs, and balds in high-pH bedrock regions, rocky headlands, railroads. This taxon usually has a globose mature hypanthium; however, rare collections in New England show elongate hypanthia—Sorrie et al. 2562 (NEBC!). Specimens from CT considered to be Rosa acicularis—Weatherby 5176 (NEBC!)—are not this species, as evidenced the presence of stipitate-glands on the hypanthia and scarcity of glands on leaves and stipules. 2. Rosa arkansana Porter E prairie rose. Rosa arkansana Porter var. suffulta (Greene) Cockerell; R. rydbergii Greene; R. suffulta Greene • MA, nh. Fields, roadsides. 3. Rosa blanda Ait. N smooth rose.  3a. Rosa blanda Ait. var. glandulosa Schuette; R. subblanda Rydb.;  3b. Rosa johannensis Fern. • CT, MA, ME, NH, VT. River shores and banks, meadows, rocky slopes, roadsides. 1a. Sepals erect in fruit; petiole and leaf rachis pubescent (rarely glabrous); abaxial surface of leaflets pubescent (rarely the surface glabrous) . . . . . . . . . . . . . . . . . 3a. R. blanda var. blanda 1b. Sepals wide-spreading to reflexed in fruit; petiole and leaf rachis glabrous or promptly so; abaxial surface of leaflets glabrous or sparsely pubescent on the veins only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. R. blanda var. glabra Crépin Variety blanda is known from CT, MA, ME, NH, VT. Variety glabra is known from ME and is of conservation concern. It is restricted to northern, ice-scoured rivers. Report of var. glabra in VT by Seymour (1982) was erroneous, the specimen is Rosa blanda var. blanda. 3a ‌ × 14. Rosa ×palustriformis Rydb. is a rare rose hybrid in New England known from ME. It is morphologically intermediate in critical characters that separate the parental taxa. Rosa blanda shows largely unarmed branchlets, distinct portions of the stipules averaging 4.6 mm long, petioles and rachises rarely with prickles (8% of individuals), pedicels rarely with stipitate-glands (1% of individuals), and hypanthia rarely with stipitate-glands (1% of individuals). Rosa palustris normally has nodal prickles on branchlets, distinct portions of the stipules averaging 2.6 mm, petioles and rachises often with prickles (75% of individuals), pedicels usually with stipitate-glands (92% of individuals), and hypanthia always with stipitate-glands.

Fig. 857  Prickles of Rosa acicularis.

8 06   tricolpate s

4. Rosa canina L. E Fig. 858 dog rose. CT, MA, ME, RI, VT. Fields, roadsides, waste areas. 5. Rosa carolina L. n Fig. 862L Carolina rose.  5a. Rosa carolina L. var. deamii (Erlanson) Deam; R. carolina L. var. glandulosa (Crépin) Farw.; R. carolina L. var. villosa (Best) Rehd.; R. lyonii Pursh; 5b. Rosa subserrulata Rydb. • CT, MA, ME, NH, RI, VT. Woodlands, clearings, fields, roadsides. 1a. Reproductive branchlets armed primarily with thin (rarely stout) nodal prickles, the internodal prickles and/or bristles lacking or present and few and scattered; hypanthia and pedicels almost always stipitate-glandular; branches relatively thin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5a. R. carolina ssp. carolina Fig. 858  Prickles of Rosa canina.

1b. Reproductive branchlets armed with stout or, less commonly, thin nodal prickles and abundant internodal prickles and bristles; hypanthia and pedicels stipitate-glandular or not; branches relatively thick . . . . . . . . . . . . . . 5b. R. carolina ssp. subserrulata (Rydb.) W.H. Lewis Subspecies carolina is known from CT, MA, ME, NH, RI, VT. Subspecies subserrulata is known from VT and is of regional conservation concern. ‌5 × 21. Rosa ×novae-angliae W.H. Lewis is a rare rose hybrid known from MA. It is difficult to detect due to the morphological variability of both parents. It is intermediate in many characters that distinguish the parental taxa (see couplet 16 of the identification key). 6. Rosa cinnamomea L. E cinnamon rose. Rosa majalis Herrm. • CT, MA, ME, NH, RI, VT. Fields, roadsides, abandoned homesteads. This species occurs only as a double corolla form in New England.

Fig. 862  Stipules of Rosa carolina (left) and R. virginiana (right).

7. Rosa gallica L. E Fig. 859 French rose. CT, MA, ME, NH, RI, VT. Fields, roadsides, abandoned homesteads. Rosa gallica is sometimes described as having solitary flowers borne on ebracteate peduncles (e.g., Gleason and Cronquist 1991). However, this is only sometimes true. Two morphologies of this species appear to occur on the New England landscape. One has 1- or 2-flowered inflorescences with ebracteate pedicels that bear abundant stipitate-glands and have abundant prickles of various sizes on the branchlets. The other morphology has 2- to 4-flowered inflorescences with bracteate pedicels that bear fewer stipitate-glands and have fewer prickles on the branchlets. 8. Rosa glauca Pourret E red-leaved rose. Rosa rubrifolia Vill. • MA, ME. Fields, roadsides, abandoned homesteads. 9. Rosa luciae Franch. & Rochebr. ex Crépin E memorial rose. Rosa wichuraiana Crépin • CT, MA. Fields, roadsides.

Fig. 859  Flower of Rosa gallica with supernumerary petals.

10. Rosa micrantha Borrer ex Sm. E small-flowered sweet-briar rose. Rosa rubiginosa L. var. nemoralis (Léman) Thory; R. rubiginosa L. var. micrantha (Sm.) Lindl. • CT, MA, ME, RI. Fields, roadsides. 11. Rosa mollis Sm. E soft downy rose. VT. Fields, roadsides. This species is somewhat similar to Rosa sherardii. Both species belong to the Rosa tomentosa Sm. complex of Europe that is identified by its tomentose and glandular leaflets. Rosa mollis has horizontally spreading, straight prickles, fleshy sepal bases, and an orifice to the hypanthium that is ½ or more the diameter of the disk (i.e., the area at the summit of the hypanthium within the sepal bases). Rosa sherardii has slightly curved prickles, thin and chartaceous sepal bases, and an orifice to the hypanthium that is ca. ⅓ the diameter of the disk.

Fig. 860  Flower of Rosa multiflora with connate and exserted styles.

12. Rosa multiflora Thunb. ex Murr. E Fig. 860 rambler rose. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, forest fragments, river banks.

Ros ac e a e   8 07

13. Rosa nitida Willd. N shining rose. CT, MA, ME, NH, VT. Swamps, peatlands, pond shores. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it could be in RI and was unaware of any collections. 14. Rosa palustris Marsh. N Fig. 861 swamp rose. Rosa floridana Rydb. • CT, MA, ME, NH, RI, VT. Swamps, marshes, low meadows, pond shores. 15. Rosa rubiginosa L. E sweet-briar rose. Rosa eglanteria L. • CT, MA, ME, NH, RI, VT. Fields, roadsides. This species sometimes shows double corollas in New England. 16. Rosa rugosa Thunb. E beach rose. CT, MA, ME, NH, RI, VT. Atlantic coast beaches and strands, dunes, roadsides. This species shows single and double corollas in New England. 16 × ?. This very rare rose hybrid is known from VT. It is commonly referred to as Rosa ‘Hollandica’ in horticultural literature. It is similar to Rosa rugosa in having the larger prickles pubescent and numerous small glands on the abaxial leaf surface. It differs from that species in its lighter green, less rugose leaflets, straight or weakly curved fruiting pedicels, ascending fruiting sepals, and hips 8–15 (–20) mm in diameter (vs. dark green and prominently rugose leaflets, curved fruiting pedicels, erect fruiting sepals, and hips 20–25 mm in diameter in R. rugosa). The other parent of this rose is unknown, but study suggests it may be R. davurica Pall., R. luciae, or R. multiflora (Mikanagi et al. 1994).

Fig. 861  Flower of Rosa palustris.

17. Rosa sempervirens L. E evergreen rose. MA. Fields, hedgerows, roadsides. 18. Rosa setigera Michx. E climbing rose. Rosa rubifolia R. Br. in Ait. f.; R. setigera Michx. var. serena Palmer & Steyermark; R. setigera Michx. var. tomentosa Torr. & Gray • CT, MA, ME, NH, VT. Fields, roadsides. 19. Rosa sherardii Davies E Sherard’s downy rose. VT. Fields, roadsides. 20. Rosa spinosissima L. E Scotch rose. Rosa pimpinellifolia L. • CT, MA, ME, NH, VT. Fields, roadsides, abandoned homesteads. This species has white flowers. It is sometimes considered also to have yellowwhite petals (e.g., Gleason and Cronquist 1991; Seymour 1982). However, these latter plants are likely introgressants involving Rosa foetida J. Herrm., a species with yellow flowers. Rosa spinosissima sometimes shows double corollas in New England. 21. Rosa virginiana P. Mill. N Fig. 862R Virginia rose. Rosa carolina L. var. lucida (Ehrh.) Farw.; R. lucida Ehrh.; R. nanella Rydb.; R. virginiana P. Mill. var. lamprophylla (Rydb.) Fern.; R. virginiana P. Mill. forma nanella (Rydb.) Fern. • CT, MA, ME, NH, RI, VT. Fields, roadsides, openings.

Rubus Treatment of Rubus subgenus Rubus (i.e., the blackberries) varies greatly between different authors. At one extreme are treatments that recognize highly variable “megaspecies”, ignoring vast amounts of morphological diversity through the use of broadly defined taxa (e.g., Gleason and Cronquist 1991). At the other extreme are treatments that recognize a multitude of species, these often based on a single difference, such as more pointed leaflets or a more compact inflorescence (e.g., Davis et al. 1967, 1968a, 1968b, 1969a, 1969b, 1970). Followed here is Hodgdon and Steele (1966), a treatment of intermediate nature that provides names to some of the variation seen on the New England landscape. The species are, in most cases, defined on substantial differences in morphology (though some are undoubtedly arbitrary

Fig. 862  Stipules of Rosa carolina (left) and R. virginiana (right).

8 08   tricolpate s

and seek to divide continuous variation). The work of Hodgdon and Steele appears to be the best compromise until a thorough biosystematic study is performed. The synonymy of the blackberries follows Hodgdon and Steele and should be considered tentative (as admonished by the authors). Further, preliminary work examining types suggest that some of these synonyms were incorrectly placed by them. In the key, distinguishing the types of stems is very important. Primocanes are the first-year stems and are green for the first half of the growing season, later forming winter buds at the tip. Floricanes are the second-year stems and are usually red or red-brown to purple or purple-brown and frequently have dead tips (i.e., no buds). Stem habit (e.g., ascending, doming, trailing) must be assessed on primocanes from the summer and fall (i.e., during or after flowering on the floricanes) for two important reasons. First, primocanes that may eventually become prostrate often grow upward during initial growth in the spring and do not arch over to the ground until summer. Second, various items can depress floricanes and make them appear decumbent (e.g., winter snow, fallen tree branches). Prickle stature is an important component of the identification key. The following measurements define terms used in the key: bristles are 0.1–0.4 (–0.5) mm thick at base, small-based prickles are (0.3–) 0.4–1 (–1.2) mm thick at base, and broad-based prickles are (0.8–) 1–4 mm thick at base. Many hybrid combinations between species of blackberries have been noted. Those in the R. flagellaris complex (R. arenicola, R. enslenii, R. flagellaris, R. jaysmithii, R. recurvicaulis) would, in most cases, be so difficult to detect that not all the combinations that likely occur on the landscape are noted in the following text. Rubus parviflorus Nutt. was reported from MA by Kartesz (1999), but specimens are unknown and the record is likely erroneous. References: Hodgdon and Steele (1966), Steele and Hodgdon (1970). 1a. Principal leaves simple; plants unarmed 2a. Leaf blades (6–) 10–20 cm wide; sepals conspicuously stipitate-glandular; petals 15–30 mm long; stems 10–20 dm tall, woody 3a. Petals pink-purple (rarely white); sepals provided with elongate, purple, stipitateglands; aggregate rather dry, scarcely juicy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. odoratus 3b. Petals white (rarely pink); sepals provided with short, yellow-orange, stipitate glands; aggregate juicy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. parviflorus 2b. Leaf blades 3–9 cm wide; sepals without glands; petals 4–18 mm long, or absent; stems up to 3 dm tall, herbaceous 4a. Carpels maturing as dry, achene-like drupes; petals 4–8 mm long or absent; leaf blades unlobed, the margin crenate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. dalibarda 4b. Carpels maturing as white-yellow, fleshy drupes; petals 10–18 mm long; leaf blades with 5–7 lobes as well as toothed on the margin . . . . . . . . . . . . . . . . . . . R. chamaemorus 1b. Principal leaves compound; plants usually armed with prickles or stiff bristles (always unarmed in R. pubescens and sometimes unarmed in R. canadensis) 5a. Stems herbaceous, not clearly differentiated into primocanes and floricanes, unarmed; stipules oblanceolate to obovate; fruit separating from the receptacle only with difficulty . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. pubescens 5b. Stems woody, clearly differentiated into primocanes and floricanes (except R. illecebrosus with annual, above-ground stems), usually armed with bristles or prickles; stipules linear or setaceous; fruit separating from the receptacle or not 6a. Leaves with 3–9 pinnately arranged leaflets (sometimes palmately arranged in R. occidentalis); fruit red or black, separating from the dry receptacle, the receptacle remaining behind on the pedicel (separating from the pedicel with the fleshy receptacle in R. illecebrosus) 7a. Petals large, surpassing the sepals; floricane leaves with 5–9 leaflets; leaflets glabrous on the abaxial surface; fruit red, not separating from the receptacle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. illecebrosus

Ros ac e a e  8 0 9

7b. Petals smaller, equaled or surpassed by the sepals; floricane leaves with 3 leaflets; leaflets densely tomentose with white-gray to gray hairs on the abaxial surface; fruit red or black, separating from the receptacle 8a. Fruit purple-black (rarely yellow), the individual drupes separated by bands of tomentum; pedicels with only moderately stout, often curved, prickles [Fig. 868]; primocanes very glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. occidentalis 8b. Fruit red to orange (rarely yellow), the individual drupes not separated by bands of tomentose pubescence; pedicels with straight bristles or minute prickles and often also glandular hairs; primocanes not, or only slightly, glaucous 9a. Rachis of inflorescence, pedicels, and often also the primocanes with stipitate glands or glandular bristles 10a. Petals white or pink, erect and incurved, appressed to the stamens; pedicels and calyx densely beset with elongate, red to purple, gland-tipped bristles 3–5 mm long [Fig. 869]; primocanes arching to doming, sometimes rooting at the tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. phoenicolasius 10b. Petals white, ascending to erect, but not appressed to the stamens; pedicels and calyx with minute, stipitate glands, these often exceeded by eglandular bristles; primocanes erect to arching, not rooting at the tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. idaeus 9b. Rachis of inflorescence, pedicels, and primocanes eglandular 11a. Petals white, ascending to erect, but not appressed to the stamens; terminal leaflet acute to acuminate at the apex; endocarps shallowly pitted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. idaeus 11b. Petals pink to purple-red, erect and incurved, appressed to the stamens; terminal leaflet obtuse to broad-acute at the apex; endocarps shallowly rugose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. parvifolius 6b. Leaves with 3 pinnately arranged or 5 palmately arranged leaflets; fruit black at maturity, separating from the pedicel with the fleshy receptacle 12a. Inflorescence a panicle, the branches and pedicels armed with stout, flattened prickles [Fig. 864]; coarse, scrambling and climbing plants armed with stout, flattened prickles 13a. Leaflets laciniately toothed or sometimes again cleft, ± green on the abaxial surface; calyx lobes with prickles, terminated by a narrow, elongate appendage; petals apically 3-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. laciniatus 13b. Leaflets toothed, gray-tomentose on the abaxial surface; calyx lobes without prickles, lacking prolonged appendages; petals entire at the apex 14a. Primocanes canescent toward apex; prickles curved; petals white or sometimes tinged with pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. discolor 14b. Primocanes ± glabrous; prickles ± straight; petals pale pink to red  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. bifrons 12b. Inflorescence a raceme, corymb-like raceme, or rarely a solitary flower, the branches and pedicels usually unarmed (never with stout, flattened prickles) [Fig. 863, 866, 867]; upright to trailing plants with weak to strong armature, the prickles usually ± terete above the base 15a. Primocanes doming, prostrate, or trailing, rooting, or with the opportunity to root, at the tip (in some species the depressed forms of the primocane not developing until midsummer when the apex of the stems finally reaches the ground) [Fig. 867]

810  tricolpates

16a. Primocanes armed with hairs, bristles, or slender, small-based prickles [Fig. 867]; petals 5–12 mm long; leaves usually lustrous, tending to be evergreen, with 3 leaflets 3.5–5 (–7) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. hispidus 16b. Primocanes armed, at least in part, with stout, broad-based prickles [Fig. 865]; petals 10–25 mm long; leaves lustrous to, more commonly, dull, usually deciduous, with 3 or 5 leaflets 6–15 (–16) cm long (2–5 cm long in the rare R. cuneifolius) 17a. Inflorescence with 1–4 (–5) flowers [Fig. 866]; pedicels ± erect, 10–60 (–80) mm long, the uppermost one 10–20 mm long (or the peduncle to 50 mm when only 1 flower is present) [Fig. 866] 18a. Stems very slender, 1–2 mm in diameter, with prickles (0.5–) 1–2 mm long; primocane leaves usually with 3 leaflets; inflorescences with 1 (–3) flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. enslenii 18b. Stems thicker, 1.5–4 mm in diameter, with prickles 1–3 (–4) mm long; primocane leaves with 3 or 5 leaflets; inflorescences with 1–4 (–5) flowers 19a. Leaf blades ± glabrous abaxially, the hairs, when present, essentially confined to the leaflet midribs and primary lateral veins, usually not noticeable to the touch . . . . . . . . . . . . . . . . . . . . R. flagellaris 19b. Leaf blades ± evidently pubescent abaxially, with hairs present across the surface (i.e., between the primary lateral veins), usually noticeable to the touch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. jaysmithii 17b. Inflorescence with (1–) 2–8 (–12) flowers; pedicels ascending to spreading, 5–40 mm long, the uppermost one 3–11 (–15) mm long 20a. Leaf blades ± glabrous abaxially, the hairs, when present, essentially confined to the leaflet midribs and primary lateral veins, usually not noticeable to the touch; inflorescence rachis and pedicels sparsely to moderately pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. recurvicaulis 20b. Leaf blades ± evidently pubescent abaxially, with hairs present across the surface (i.e., between the primary lateral veins), usually noticeable to the touch; inflorescence rachis and pedicels moderately to densely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. arenicola 15b. Primocanes arching or erect, not rooting at the tip (sometimes doming in R. vermontanus) 21a. Primocanes armed with hairs, bristles, or slender, small-based prickles [Fig. 870]; stems 0.3–1 m tall; axis of inflorescence commonly with stipitateglands (always lacking in R. elegantulus and often lacking in R. semisetosus) 22a. Leaf blades ± evidently pubescent abaxially, with hairs present across the surface (i.e., between the primary lateral veins), usually noticeable to the touch; primocanes armed, at least in part, with slender, small-based prickles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. semisetosus 22b. Leaf blades ± glabrous abaxially, the hairs, when present, essentially confined to the leaflet midribs and primary lateral veins, usually not noticeable to the touch; primocane armature various 23a. Primocanes armed with hairs or stiff bristles only, the hairs/bristles numbering 600–5000 per 10 cm [Fig. 870] . . . . . . . . . . . . . . . . . . R. setosus 23b. Primocanes armed, at least in part, by slender, small-based prickles, numbering 10–500 per 10 cm 24a. Central petiolule of primocane leaves (8–) 14–20 (–30) mm long; inflorescence rachis frequently with stipitate glands; stems 2–4 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. vermontanus

Ros ac e a e   8 1 1

24b. Central petiolule of primocane leaves (15–) 25–40 (–50) mm long; inflorescence rachis never with stipitate glands; stems 3–6 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. elegantulus 21b. Primocanes armed, at least in part, with stout, broad-based prickles; stems 0.5–3 m tall; axis of inflorescence lacking stipitate-glands except in R. allegheniensis and hybrids with that species 25a. Inflorescence, and often also the primocane, pubescent with stipitate glands [Fig. 863], the inflorescence (3–) 8–20 cm long and with (3–) 9–22 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. allegheniensis 25b. Inflorescence and primocane ± lacking stipitate glands, the inflorescence 3–11 (–17) cm long and with (2–) 3–15 (–21) flowers 26a. Leaf blades ± glabrous abaxially, the hairs, when present, essentially confined to the leaflet midribs and primary lateral veins, usually not noticeable to the touch 27a. Prickles of the stem absent or few, numbering 0–10 per 10 cm; inflorescence rachis pubescent to subglabrous, usually without prickles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. canadensis 27b. Prickles of the stem more numerous, numbering 10–60 (–100) per 10 cm [Fig. 865]; inflorescence rachis pubescent, often with thin prickles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) R. elegantulus 26b. Leaf blades ± evidently pubescent abaxially, with hairs present across the surface (i.e., between the primary lateral veins), usually noticeable to the touch 28a. Leaflets oblanceolate to obovate, definitely widest above the middle, densely tomentose abaxially with white to gray hairs (sometimes becoming partly glabrate in late season) . . . . . . . . . . . . . . . . R. cuneifolius 28b. Leaflets elliptic or elliptic-oblanceolate to broad-ovate, widest near or below the middle, pubescent abaxially, but not densely tomentose 29a. Terminal leaflet of primocane leaves elliptic-oblanceolate to elliptic, cuneate to narrow-rounded at the base, 2 or more times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. argutus 29b. Terminal leaflet of primocane leaves ovate to broad-ovate, broad-cuneate to cordate at the base, less than 2 times as long as wide 30a. Prickles of the stem mostly 2–4 mm long, numbering 0–20 per 10 cm; leaf blades herbaceous; inflorescence with (0–) 1 or 2 (–6) leafy bracts (i.e., most of the pedicels subtended by small, stipule-like bracts) . . . . . . . . . . . . . . . . . R. pensilvanicus 30b. Prickles of the stem mostly 4–6 mm long, numbering 4–60 per 10 cm; leaf blades tending to be more chartaceous; inflorescence with (0–) 1–7 leafy bracts (i.e., many of the pedicels subtended by foliaceous bracts) . . . . . . . . . . . . . . . . . . R. frondosus 1. Rubus allegheniensis Porter N Fig. 863 common blackberry. Rubus allegheniensis Porter var. gravesii Fern.; R. auroralis Bailey; R. fernaldianus Bailey; R. longissimus Bailey; R. nigrobaccus Bailey; R. pugnax Bailey; R. saltuensis Bailey; R. sativus Brainerd • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, open rights-of-way, logged areas.

Fig. 863  Raceme of Rubus allegheniensis.

8 12   tricolpate s

1 × 5. This rare blackberry hybrid is known from ME, NH, VT. It is marked by stems with relatively thin and short prickles (more like Rubus canadensis than R. allegheniensis). The leaves are pubescent abaxially and tend to be much like R. allegheniensis. The leaflets of the primocane leaves are with elongate, acuminate apices. The axis of the inflorescence has occasional stipitate-glands. 1 × 10. This very rare blackberry hybrid is known from NH. It has leaves that generally resemble Rubus allegheniensis and are pubescent on the abaxial surface, with stems bearing thinner prickles than found in that species (but as in R. elegantulus), and an inflorescence axis bearing stipitate-glands. 1 × 12. This uncommon blackberry hybrid is known from Ma, Me, NH. It is marked by primocanes with low-doming to nearly trailing habit that show occasional stipitateglands, primocane leaflets mostly 7–11 cm long that are sparsely pubescent abaxially, armature of straight to hooked prickles, and inflorescences to 11 cm long that bear frequent glands on the rachis and pedicels. 1 × 13. This very rare blackberry hybrid is known from MA, RI. It is marked by stems armed with stout prickles, leaves that are pubescent abaxially, and an inflorescence with scattered stipitate-glands. Tentative synonym: Rubus rosa Bailey. 1 × 14. This rare blackberry hybrid is known from MA, ME, NH, VT. It is marked by variably oriented primocanes (erect, doming, or trailing) armed with both short, stiff, curved prickles (but usually thinner than in typical R. allegheniensis) and stipitate-glands, subcoriaceous and lustrous, ovate to elliptic leaflets that are pubescent abaxially, and a small- to medium-sized inflorescence. The petals are relatively small (close to R. hispidus). Tentative synonyms: Rubus biformispinus Blanch.; R. electus Bailey; R. jactus Bailey; R. laevior (Bailey) Fern.; R. permixtus Blanch.; R. sanfordii Bailey. 1 × 22. This rare blackberry hybrid is known from CT, MA, ME, NH, RI. It is marked by stems with scattered, relatively short and thin prickles (compared with Rubus allegheniensis) and an inflorescence with occasional stipitate-glands. Tentative synonyms: Rubus alumnus Bailey; R. licitus Bailey; R. paulus Bailey. 1 × 25. This very rare blackberry hybrid is known from ME. It is intermediate between its parental species. 1 × 27. This uncommon blackberry hybrid is known from MA, ME, NH, VT. It is marked by arching to erect habit, stems 0.6–1.2 (–1.5) m tall armed with prickles of variable size, slender bristles, and stipitate-glands (small-based prickles and bristles are usually numerous), leaflets that usually resemble Rubus allegheniensis with sparse to dense pubescence abaxially, and an inflorescence that is smaller than usual for R. allegheniensis that has stipitate-glands along its axis. Tentative synonyms: Rubus abbrevians Blanch.; R. aculiferus Fern.; R. glandicaulis Blanch.; R. montpelierensis Blanch.; R. sceleratus Brainerd ex Fern. 1 × 28. This rare blackberry hybrid is known from ME, NH, VT. It is marked by arching habit, stems 3–5 mm in diameter armed with slender prickles that are shorter and thinner than is typical for Rubus allegheniensis and often stipitate-glands as well. The leaves are pubescent abaxially and the inflorescence is shorter than normal for R. allegheniensis with stipitate-glands along the axis. The fruits are also smaller than typical for R. allegheniensis. Tentative synonym: Rubus ravus Bailey. 2. Rubus arenicola Blanch. N sandy field blackberry. Rubus alius Bailey; R. brainerdii Fern.; R. curtipes Bailey; R. janssonii Bailey; R. obsessus Bailey; R. pauper Bailey; R. perpauper Bailey; R. prosper Bailey • MA, ME, NH, RI. Fields, roadsides, sand plains, forest borders, woodlands, balds, open rights-of-way. Steele and Hodgdon (1970) tentatively reported the hybrid Rubus arenicola × R. semisetosus from NH. 3. Rubus argutus Link N southern blackberry. Rubus jugosus Bailey; R. paludivagus Fern. • CT, MA, RI. Fields, roadsides, forest borders, woodlands, balds, open rights-of-way. This species is similar in many ways to Rubus frondosus, especially in terms of the abundant, large armature on the stems.

Ros ac e a e   8 13

4. Rubus bifrons Vest ex Tratt. E Fig. 864 twice-leaved blackberry. MA, RI. Fields, roadsides, forest borders, waste areas. 5. Rubus canadensis L. N smooth blackberry. Rubus argutus Link var. randii (Bailey) Bailey; R. millspaughii Bailey; R. randii (Bailey) Rydb. • CT, MA, ME, NH, VT. Fields, roadsides, forest borders and clearings, wetland margins, seeps, stream banks, bases of ledges, open rights-of-way, sometimes ascending to boreal settings. 5 × 10. This very rare blackberry hybrid is likely more common than is currently vouchered. It is known from NH. Due to the close morphological similarity of the parental species, knowledge of both species present at a given site is very useful information for detecting most occurrences.

Fig. 864  Panicle of Rubus bifrons.

5 × 22. This rare blackberry hybrid is known from ME, NH. It is marked by stems with sparse, thin prickles that number 0–7 per 10 cm. The leaves closely resemble those of Rubus canadensis but have sparse to moderate pubescence abaxially. The raceme axis is densely (rarely only moderately) pubescent. 6. Rubus chamaemorus L. N baked-apple-berry. ME, NH. Mossy, alpine heaths, coastal peatlands. 7. Rubus cuneifolius Pursh

NC

sand blackberry. Rubus chapmanii Bailey; R. cuneifolius Pursh var. angustior Bailey; R. cuneifolius Pursh var. subellipticus Fern. • CT, MA, NH. Fields, rocky pastures and hillsides, clearings. 8. Rubus dalibarda L. N dewdrop. Dalibarda repens L.; Rubus repens (L.) Kuntze • CT, MA, ME, NH, RI, VT. Deciduous to evergreen forests, swamps. 9. Rubus discolor Weihe & Nees E Himalayan blackberry. Rubus procerus, auct. non P.J. Muell. ex Genev • MA. Forest edges, waste areas. 10. Rubus elegantulus Blanch. N Fig. 865 showy blackberry. Rubus amicalis Blanch.; R. kennedyanus Fern.; R. multiformis Blanch. • CT, MA, ME, NH, VT. Fields, roadsides, forest borders and clearings, wetland margins, graminoid marshes, seeps, stream banks, bases of ledges, open rights-of-way, sometimes ascending to boreal settings. 10 × 17. This very rare blackberry hybrid is known from VT. It resembles hybrids between Rubus elegantulus and R. recurvicaulis (10 × 25) except that the leaf blades are pubescent abaxially. 10 × 25. This rare blackberry hybrid is known from ME, VT. It is marked by stout primocanes that arch, dome, or trail, large leaflets that are glabrous on the abaxial surface, relatively thin prickles, and inflorescences similar to Rubus elegantulus but without stipitate-glands along the axis. 10 × 27. This rare blackberry hybrid is known from ME, NH. It is marked by doming to arching habit, stems armed with sparse to abundant thin prickles and bristles (stipitateglands are usually also present), and leaves with ovate to lanceolate leaflets. It is very similar to Rubus setosus × R. vermontanus and is best separated from that hybrid by examining the parental species at the site. 10 × 28. This very rare blackberry hybrid is known from ME, NH. It is very difficult to detect due to the close morphological similarity of the two parental taxa and is likely more common than is currently vouchered. Knowledge of the two species occurring at the same site is important information for detecting most of the occurrences. Tentative synonym: Rhus multilicius Bailey.

Fig. 865  Armature of Rubus elegantulus.

8 14   tricolpate s

11. Rubus enslenii Tratt. N Enslen’s blackberry. Rubus sempervirens Bigelow • CT, MA, ME, NH, RI, VT. Forests and woodlands, often in association with Quercus and/or Carya, balds, ledges. 11 × 22. This very rare blackberry hybrid is known from NH. It is marked by primocanes with arched to trailing habit, primocane leaflets numbering 3–5 and sparsely pubescent abaxially, stems with sparse, thin, slender prickles, 1- to 5-flowered inflorescences up to 8 cm long, and pedicels 2–6 cm long that lack stipitate-glands. 11 × 25. This rare blackberry hybrid is known from ME, NH. It is marked by primocanes with low-doming to arching habit, primocane leaflets that are ± glabrous abaxially, and 1- to 4-flowered inflorescences 2–3.5 cm long with short, slender pedicels. 12. Rubus flagellaris Willd. N Fig. 866 northern blackberry. Rubus felix Bailey; R. geophilus Blanch.; R. maniseesensis Bailey • CT, MA, ME, NH, RI, VT. Fields, roadsides, sand plains, forest borders, woodlands, balds, open rights-of-way. 12 × 13. This uncommon blackberry hybrid is known from CT, MA, NH. It is marked by primocanes with arching to doming or trailing habit, primocane leaflets pubescent abaxially, armature of hooked prickles, and 2- to 6-flowered inflorescences with a pubescent rachis (but lacking stipitate-glands). Fig. 866  Inflorescence of Rubus flagellaris.

12 × 14. This rare blackberry hybrid is known from ME. It has primocane leaves with thick leaflets, the leaflets usually numbering 3 but often the lateral leaflets with a lobe on the outer, basal margin (these lobes variably developed, sometimes the leaf with 5 total leaflets). The primocane stems are armed with both prickles and bristles. Tentative synonym: Rubus mainensis Bailey. 12 × 25. This rare blackberry hybrid is known from MA. It has the habit of Rubus flagellaris and R. recurvicaulis (e.g., trailing stems). The inflorescence has 3–7 flowers on ascending to erect pedicels (i.e., the aspect is of R. flagellaris but with too many flowers). The lower pedicels commonly exceed 40 mm in length. 13. Rubus frondosus Bigelow N leafy-flowered blackberry. Rubus bellobatus Bailey; R. heterogeneous Bailey; R. insulanus Bailey; R. multispinus Blanch.; R. recurvans Blanch. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, open rights-of-way. 13 × 14. This very rare blackberry hybrid is known from MA. It has primocanes with a mixture of small-based and broad-based prickles that are collectively abundant and abaxially pubescent leaves. The inflorescences have few, relatively small flowers. 13 × 17. This very rare blackberry hybrid is known from MA. It is marked by primocanes with arching to trailing habit and relatively stout prickles. The primocane leaves are abaxially moderately pubescent across the surface. The inflorescences are variable and have 1–6 flowers. This hybrid is very similar to Rubus flagellaris × R. frondosus but has a greater degree of pubescence on the leaf blades. 13 × 25. This rare blackberry hybrid is known from Ma, Me. It is very similar to Rubus flagellaris × R. frondosus hybrids, but occurs with R. recurvicaulis as the low-growing parent rather than R. flagellaris. 13 × 26. This very rare blackberry hybrid is known from MA. It is marked by arching habit, stems 3–5 mm in diameter armed with thin, numerous prickles 2–5 mm long that number up to 30 per 10 cm (the prickles are usually more slender than in typical Rubus frondosus). The leaves that are pubescent abaxially. The inflorescence resembles most closely R. semisetosus and has a pubescent, eglandular axis. 14. Rubus hispidus L. N Fig. 867

Fig. 867  Habit of Rubus hispidus showing bristly floricane.

bristly blackberry. Rubus cubitans Blanch.; R. pervarius (Bailey) Bailey; R. spiculosus Fern. • CT, MA, ME, NH, RI, VT. Fields, roadsides, woodlands, swamps, wetland margins, forest borders, open-rights-of-way, borrow pits. This species rarely has stipitate-glands along the axis of the inflorescence. When present, the stipitate-glands are usually sparse and of uniform length.

Rosac e a e   8 15

14 × 17. This very rare blackberry hybrid is known from RI. It is intermediate between its parental species. 14 × 25. This rare blackberry hybrid is known from MA, NH. It is marked by primocanes with trailing habit, primocane leaves with 3 or 5 leaflets that are chartaceous and not lustrous, stems armed with short prickles, bristles, and stipitate-glands, and short, compact, 6- to 10-flowered inflorescences up to 4 cm long. 14 × 26. This very rare blackberry hybrid is known from MA. The primocanes are armed with mostly 100–300 bristles and small-based prickles per 10 cm and bear leaves with 3–5 oblong-ovate to ovate leaflets that are not lustrous (as in Rubus hispidus) and are abaxially pubescent on the midrib and primary lateral veins and sparsely so between them. It is very similar to R. hispidus × R. vermontanus but has a greater degree of pubescence on the leaf blades. 14 × 27. This uncommon blackberry hybrid is known from MA, ME, NH, VT. It is marked by doming or trailing primocanes armed with abundant prickles, bristles, and stipitateglands (mostly 2000–3000 per 10 cm), leaves with 3 or 5 sublustrous leaflets, and inflorescences with some stipitate-glands along the axis. Considered by Steele and Hodgdon (1970) to be one of the most common blackberry hybrids in New England. Tentative synonyms: Rubus adjacens Fern.; R. alter Bailey; R. blanchardianus Bailey; R. harmonicus Bailey; R. jacens Blanch.; R. pudens Bailey; R. rixosus Bailey; R. tholiformis Fern.; R. trifrons Blanch.; R. vigoratus Bailey. 14 × 28. This uncommon blackberry hybrid is known from MA, ME, NH, VT. It closely resembles Rubus hispidus × R. setosus but has some prickles that are slightly more stout and curved (in addition to the numerous bristles and stipitate-glands). Tentative synonym: Rubus viridifrons Bailey. 15. Rubus idaeus L. n red raspberry. 15b. Batidea strigosa (Michx.) Greene; Rubus idaeus L. var. aculeatissimus Regel & Tiling; R. idaeus L. var. canadensis Richards.; R. idaeus L. var. caudatus (Robins. & Schrenk) Fern.; R. idaeus L. var. egglestonii (Blanch.) Fern.; R. idaeus L. var. eucyclus Fern. & Weatherby; R. idaeus L. var. heterolasius Fern.; R. idaeus L. ssp. melanolasius (Dieck) Focke; R. idaeus L. ssp. sachalinensis (Levl.) Focke; R. melanolasius Dieck; R. strigosus Michx.; R. strigosus Michx. var. canadensis (Richards.) House • CT, MA, ME, NH, RI, VT. Fields, roadsides, forests, fields, swamps, clearings, logged forests, shorelines, gardens, banks, forest fragments, abandoned homesteads. 1a. Plants lacking stipitate glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15a. R. idaeus ssp. idaeus 1b. Inflorescence and often the primocanes with stipitate glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15b. R. idaeus ssp. strigosus (Michx.) Focke Subspecies idaeus is known from CT, MA, ME, VT. It is non-native and more commonly found in human-disturbed and human-modified habitats. Subspecies strigosus is known from CT, MA, ME, NH, RI, VT. It is native and found in many different habitats, including pristine and human-disturbed. ‌ 15b × 19. Rubus ×neglectus Peck is a somewhat rare raspberry hybrid known from CT, MA, NH, RI, VT. It is intermediate between the two parental species and can be distinguished by its fruit color and pedicel morphology. The mature fruit is red-purple to purple (vs. red in R. idaeus and purple-black in R. occidentalis). The pedicels are armed with very thin prickles 0.7–1.5 (–2) mm long and have stipitate-glands (those of R. idaeus have thin bristles and are stipitate-glandular whereas those of R. occidentalis have stouter prickles (1.1–) 1.5–2.7 mm long and lack stipitate-glands). Additionally, the stipitateglands of R. × tend to be sparser and/or shorter than those of R. idaeus. ‌neglectus 16. Rubus illecebrosus Focke E strawberry raspberry. CT, MA, ME. Roadsides, forest fragments, abandoned homesteads. 17. Rubus jaysmithii Bailey N Smith’s blackberry. Rubus scambens Bailey; R. tetricus Bailey • MA, NH, RI, VT. Fields, roadsides, sand plains, woodlands, forest borders, open rights-of-way.

8 16  tricolpates

18. Rubus laciniatus Willd. E cut-leaved blackberry. CT, MA; also reported from RI by George (1992), but specimens are unknown. Fields, roadsides, waste areas. The reports of this species in VT are based on a cultivated specimen collected by Brainerd (ex horto, specimen at VT). 19. Rubus occidentalis L. N Fig. 868 black raspberry. Rubus occidentalis L. var. pallidus Bailey • CT, MA, ME, NH, RI, VT. Forests, forest borders, clearings, fields, roadsides, banks. 20. Rubus odoratus L. N flowering raspberry. Rubacer columbianum (Millsp.) Rydb.; R. odoratum (L.) Rydb.; Rubus odoratus L. var. columbianus Millsp.; R. odoratus L. var. malachophyllus Fern. • CT, MA, ME, NH, RI, VT. Forests, woodlands, talus slopes, bases of ledges, roadsides, railroads. Fig. 868  Inflorescence of Rubus occidentalis showing prickles on pedicels.

21. Rubus parvifolius L. E Japanese raspberry. Rubus parviflorus L. var. triphyllus (Thunb.) Nakai; R. triphyllus Thunb. • MA. Waste places.

22. Rubus pensilvanicus Poir. N Pennsylvania blackberry. Rubus abactus Bailey; R. amnicola Blanch.; R. andrewsianus Blanch.; R. avipes Bailey; R. barbarus Bailey; R. conanicutensis Bailey; R. facetus Bailey; R. honorus Bailey; R. insons Bailey; R. orarius Blanch.; R. ostryifolius Rydb.; R. pergratus Blanch.; R. philadelphicus Blanch. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, swamps, open rights-of-way. 22 × 25. This rare blackberry hybrid is known from ME, NH. It is marked by low-doming primocanes that are armed with relatively slender, straight to slightly curved prickles, leaflets that are sparsely pubescent abaxially, and a variable inflorescence (but always without stipitate-glands). 22 × 28. This rare blackberry hybrid is known from ME. It is marked by erect to arching primocanes with numerous small-based prickles (ca. 200 per 10 dm), and lacking stipitate-glands with relatively glabrous abaxial leaf blade surface, and with short racemes (2.5–5 cm long) that lack stipitate-glands. Tentative synonyms: Rubus miscix Bailey; R. peculiaris Blanch. 23. Rubus phoenicolasius Maxim. E Fig. 869 wine raspberry. CT, MA, RI, VT. Fields, roadsides, forest fragments and borders, talus slopes. 24. Rubus pubescens Raf.

Fig. 869  Inflorescence of Rubus occidentalis showing stipitateglands on pedicels and sepals.

NC

dwarf raspberry. Cylactis pubescens (Raf.) W.A. Weber; Rubus pubescens Raf. var. pilosifolius A.F. Hill; R. triflorus Richards. • CT, MA, ME, NH, RI, VT. Forests, swamps, wetland margins. A variety with abaxially closely pubescent leaflets has traditionally been recognized (var. pilosifolius). However, there are specimens with intermediate hair density that suggest this is not a distinct taxon; e.g., 25 Jun 1918, Bean s.n. (GH!) 25. Rubus recurvicaulis Blanch. N arching blackberry. Rubus arundelanus Blanch.; R. brevipedalis Bailey; R. coloniatus Bailey; R. plicatifolius Blanch.; R. positivus Bailey; R. rhodinsulanus Bailey; R. semierectus Blanch.; R. severus Brainerd; R. usus Bailey • MA, ME, NH, VT. Fields, roadsides, sand plains, forest borders, woodlands, balds, open rights-of-way, railroads. 25 × 27. This rare blackberry hybrid is known from MA, ME, NH. It is marked by primocanes with doming or trailing habit, primocane leaves with (3–) 5 leaflets, stems armed with slender prickles and few to many intermixed bristles, and an inflorescence up to 9 cm long with stipitate-glands on the rachis and pedicels (the inflorescence mainly resembling Rubus setosus). Tentative synonyms: Rubus arcuans Fern. & St. John; R. bicknellii Bailey.

Rosac e a e   8 17

25 × 28. This rare blackberry hybrid is known from MA, ME, NH. It is marked by primocanes with arching or doming habit, glabrous and chartaceous leaflets, stems armed with numerous sharp, slender, straight or hooked prickles. Tentative synonym: Rubus severus Brainerd ex Fern. 26. Rubus semisetosus Blanch. N northeastern blackberry. Rubus ascendens Blanch.; R. bigelovianus Bailey; R. hispidoides Bailey; R. ortivus Bailey; R. perinvisus Bailey • CT, MA, ME, NH, RI; also reported from VT by Atwood et al. (1973), but specimens are unknown. Fields, roadsides, forest borders, graminoid marshes, open rights-of-way. 26 × 27. This very rare blackberry hybrid is known from MA. It resembles Rubus setosus, but the leaves abaxially have sparse pubescence and small prickles along the leaflet midveins. 26 × 28. This rare blackberry hybrid is known from ct, mA. It is similar to Rubus semisetosus, but the pubescence is much less dense and mainly present on the principal veins. 27. Rubus setosus Bigelow N Fig. 870 setose blackberry. Rubus boottianus Bailey; R. frondisentus Blanch.; R. junior Bailey; R. lawrencei Bailey; R. nigricans Rydb.; R. notatus Bailey; R. significans Bailey; R. udus Bailey • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, open rights-of-way, marshes edges, ascending to boreal habitats. 27 × 28. This uncommon blackberry hybrid is known from CT, MA, ME, NH, VT. It is similar to both Rubus setosus and R. vermontanus. The stems are armed with stipitate-glands, bristles of various lengths, and some slender-based prickles that are relatively rigid, collectively numbering 1000–3000 per 10 cm. Very difficult to separate from R. elegantulus × R. setosus without knowledge of parental species present at the site (though the latter hybrid usually has fewer prickles and bristles). Tentative synonyms: Rubus groutianus Blanch.; R. parlinii Bailey; R. univocus Bailey. 28. Rubus vermontanus Blanch. N Vermont blackberry. Rubus deaneanus Bailey; R. junceus Bailey; R. singulus Bailey; R. tardatus Blanch.; R. vermontanus Blanch. var. viridiflorus Blanch. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, balds, open rights-of-way.

Sanguisorba 1a. Leaflets 5–20 mm long; flowers with 12 or more stamens and 2 carpels . . . . . . . . . S. minor 1b. Leaflets mostly 25–80 (–100) mm long; flowers with 4 stamens and 1 carpel 2a. Sepals white, much shorter than the stamens; spikes 3–12 (–20) cm tall; filaments distally widened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. canadensis 2b. Sepals purple to red-purple, equaling or slightly longer than the stamens; spikes 1–3 cm tall; filaments slender throughout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. officinalis 1. Sanguisorba canadensis L. N Canada burnet. CT, MA, ME, NH, RI, VT. Swamps, borders of tidal marshes, low riparian forests, stream banks, meadows, river shores. 2. Sanguisorba minor Scop. ssp. balearica (Bourg. ex Nyman) M. Garm. & C. Navarro E salad burnet. Poterium polygamum Waldst. & Kit.; P. sanguisorba L.; Sanguisorba minor Scop. ssp. muricata (Spach) Briq.; S. muricata (Spach) Gremli • CT, MA, ME, RI, VT. Fields, roadsides, waste areas. 3. Sanguisorba officinalis L. E great burnet. Poterium officinale (L.) Gray; Sanguisorba officinalis L. var. polygama (F. Nyl.) Mela & A. Caj.; S. polygama F. Nyl. • ME. Fields, roadsides.

Fig. 870  Armature of Rubus setosus.

8 18  tricolpates

Sibbaldia 1. Sibbaldia procumbens L.

NC

sibbaldia. Potentilla sibbaldii Haller f. • NH. Alpine ravines.

Sibbaldiopsis 1. Sibbaldiopsis tridentata (Ait.) Rydb. N three-toothed-cinquefoil. Potentilla tridentata Ait. • CT, MA, ME, NH, RI, VT. Open, often sterile, sandy and rocky areas such as fields, roadsides, sand plains, balds, ledges, mountain tops, and alpine plateaus.

Sorbaria 1. Sorbaria sorbifolia (L.) A. Braun E false spiraea. Schizonotus sorbifolius (L.) Lindl.; Spiraea sorbifolia L. • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, abandoned homesteads.

Sorbus See the genus Aronia for discussion of intergeneric hybrids placed in the nothogenus ×Sorbaronia. 1a. Leaf blades simple; calyx lobes deciduous in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . S. alnifolia 1b. Leaf blades compound; calyx lobes persistent in fruit 2a. Leaf blades with 3–7 pairs of leaflets; terminal leaflet much larger than the lateral leaflets, often lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hybrida 2b. Leaf blades with 11–17 pairs of leaflets; terminal leaflet ± similar to lateral leaflets, serrate 3a. Branchlets, inflorescence, adaxial surface of leaflets, and hypanthium villous; bud scales villous, not glutinous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. aucuparia 3b. Branchlets, inflorescence, adaxial surface of leaflets, and hypanthium glabrous or some surfaces sparsely villous; bud scales glabrous or ciliate on the margin, glutinous 4a. Leaflets acuminate at the apex, 3–5 times as long as wide [Fig. 871, R]; winter bud scales eciliate or sparsely ciliate; petals 3–4 mm long; pomes 4–7 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. americana 4b. Leaflets acute to short-acuminate at the apex, 2–3.5 times as long as wide [Fig. 871, L]; winter bud scales ciliate; petals 4–5 mm long; pomes 7–10 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. decora 1. Sorbus alnifolia (Sieb. & Zucc.) K. Koch E Korean mountain-ash. Crataegus alnifolia Sieb. & Zucc.; Micromeles alnifolia Koehne • MA. Forest fragments. 2. Sorbus americana Marsh. N Fig. 871R American mountain-ash. Pyrus americana (Marsh.) DC. • CT, MA, ME, NH, VT. Temperate, boreal, and subalpine forests, ridge tops, swamps. George (1999) reported this species from RI; however, all specimens seen by me identified as this species from RI either are Sorbus aucuparia or were collected from residential settings. Fig. 871  Leaflets of Sorbus decora (left side of rachis) and S. americana (right side of rachis).

3. Sorbus aucuparia L. E European mountain-ash. Pyrus aucuparia (L.) Gaertn. • CT, MA, ME, NH, RI, VT. Fields, forest borders and fragments, areas of habitation.

Rosac e a e   8 19

4. Sorbus decora (Sarg.) Schneid. n Fig. 871L showy mountain-ash. Pyrus americana (Marsh.) DC. var. decora Sarg.; Pyrus decora (Sarg.) Hyl.; Pyrus decora (Sarg.) Hyl. var. groenlandica (Schneid.) Fern.; Sorbus decora (Sarg.) Schneid. var. groenlandica (Schneid.) G.N. Jones; S. groenlandica (Schneid.) A. & D. Löve • CT, MA, ME, NH, RI, VT. Temperate, boreal, and subalpine forests, ridge tops, swamps. This species in nonnative in RI and native elsewhere in New England. Sorbus decora flowers ca. 7–10 earlier than S. americana when both occur at the same site. Sorbus groenlandica is a frequently recognized taxon that supposedly differs from S. decora in its more pointed leaflets that are green abaxially (rather than glaucous). Review of herbarium specimens revealed that these characters are not always correlated. Further, one can observe leaves with acutely tipped leaflets and leaves with acuminately tipped leaflets on a single plant in New England. Löve and Löve (1966) supported recognition of S. groenlandica by its different chromosome number. However, lacking morphological distinction, it makes sense to subsume this taxon in S. decora (i.e., treat S. decora as a species that includes multiple chromosome races). 5. Sorbus hybrida L. E oak-leaved mountain-ash. Pyrus pinnatifida Ehrh.; Sorbus fennica (Kalm) Fries • ME, NH. Fields, roadsides. Reports of this species from VT (e.g., Kartesz 1999) are based on a cultivated specimen with no evidence of naturalization noted on the herbarium label—Kittridge s.n. (VT).

Spiraea The name Spiraea salicifolia L. was at one time used for our species of western New England now called S. alba var. alba (e.g., Wiegand 1900). Recent reports of S. salicifolia in New England (e.g., Magee and Ahles 1999) are erroneous, as they were based on older publications that were referring to S. alba var. alba. 1a. Inflorescence a simple umbel or corymb, terminating a short-shoot [Fig. 873] 2a. Inflorescences (except sometimes the uppermost) borne on a peduncle, with 5–12 flowers; petioles 4–7 mm long; follicles pubescent with appressed hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chamaedryfolia 2b. Inflorescences sessile, with 3–7 flowers [Fig. 873]; petioles 1–4 mm long; follicles glabrous or sometimes pubescent along the adaxial suture in S. prunifolia 3a. Leaf blades ovate to oblong-lanceolate, pubescent on the abaxial surface; pedicels 10–24 mm long, pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. prunifolia 3b. Leaf blades narrow-lanceolate, glabrous on the abaxial surface; pedicels 6–10 mm long, glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. thunbergii 1b. Inflorescence a panicle or compound corymb (i.e., with branches), terminating a longshoot [Fig. 872] 4a. Inflorescence a compound corymb, as wide as or wider than tall; leaf blades (5–) 8–15 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. japonica 4b. Inflorescence a panicle, mostly wider than tall (except in high-elevation plants) [Fig. 872]; leaf blades 3–7 (–8) cm long 5a. Mature leaf blades with white or red-brown tomentum abaxially; branchlets tomentose, especially about the nodes; flowers without a nectar disk; sepals reflexed in fruit; follicles pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tomentosa 5b. Mature leaf blades ± glabrous abaxially; branchlets ± glabrous; flowers with a nectar disk; sepals erect in fruit; follicles glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. alba 1. Spiraea alba Du Roi N Fig. 872 white meadowsweet. 1b. Spiraea alba Du Roi var. septentrionalis (Fern.) Seymour; S. latifolia (Ait.) Borkh.; S. latifolia (Ait.) Borkh. var. septentrionalis Fern.; S. septentrionalis (Fern.) A. & D. Löve • CT, MA, ME, NH, RI, VT. Fields, roadsides, swamps, marshes, wetland margins, shorelines, clearings, boreal and alpine ravines.

Fig. 872  Inflorescence of Spiraea alba var. latifolia.

8 20   tricolpate s

1a. Rachis of panicle and hypanthium tomentulose; leaf blades oblanceolate, 3–4 (or more) times as long as wide; branchlets brown to yellow-brown; sepals obtuse at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. S. alba var. alba 1b. Rachis of panicle and hypanthium ± glabrous; leaf blades broad-oblanceolate to oblong or obovate, 2–3 times as long as wide; branchlets purple-brown to red-brown; sepals acute at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. S. alba var. latifolia (Ait.) Dippel Variety alba is known from MA, VT. Variety latifolia is known from CT, MA, ME, NH, RI, VT. Spiraea septentrionalis is a frequently recognized taxon that, in New England, occurs in the higher Appalachian mountains. It is said to differ from S. alba var. latifolia in its larger flowers and diminutive panicles. Measurements on herbarium specimens showed that no differences exist in floral parts (e.g., sepals, petals, anthers). The small panicle with short, lower branches is not unique to S. septentrionalis. This character state can be seen on depauperate plants of S. alba at low elevation. On peaks such as Mount Washington (NH) and Katahdin (ME), one can observe a morphological cline with elevation (with progressively smaller panicles produced by higher elevation plants). Löve and Löve (1966) supported the recognition of S. septentrionalis by its different chromosome number. However, given the lack of morphological distinction, it makes sense to subsume S. septentrionalis in S. alba var. latifolia (i.e., treat this variety as one that includes multiple chromosome races). 2. Spiraea chamaedryfolia L. E germander meadowsweet. Spiraea chamaedryfolia L. var. ulmifolia (Scop.) Maxim.; S. ulmifolia Scop. • CT. Fields, roadsides, waste areas. 3. Spiraea japonica L. f. var. fortunei (Planch.) Rehd. E Japanese meadowsweet. Spiraea fortunei Planch. • CT, MA, ME, NH, RI. Fields, roadsides, waste areas, gardens. 4. Spiraea prunifolia Sieb. & Zucc. E Fig. 873 bridalwreath meadowsweet. Spiraea prunifolia Sieb. & Zucc. var. plena Schneid. • CT, MA. Fields, roadsides, waste areas, gardens. 5. Spiraea thunbergii Sieb. ex Blume E Thunberg’s meadowsweet. CT; also reported from MA by Magee and Ahles (1999), but specimens are unknown. Fields, roadsides, waste areas, abandoned homesteads. Fig. 873  Inflorescence of Spiraea prunifolia.

6. Spiraea tomentosa L. N rosy meadowsweet. CT, MA, ME, NH, RI, VT. Fields, ditches, swamps, marshes, wetland margins, shorelines. Spiraea ‌ cantoniensis Lour. × S. trilobata L. Spiraea ×vanhouttei (Briott) Carr. is frequently planted (though rarely escaped) meadowsweet hybrid known from CT, MA. It is similar to S. chamaedryfolia in that the flowers are borne in peduncled umbels that are produced on short shoots. The hybrid differs from that species in its stamens not exceeding the petals in length (vs. stamens longer than the petals), its erect to spreading ‌vanhouttei is further sepals (vs. recurved), and glabrous follicles (vs. pubescent). Spiraea × characterized by its leaf blades that are often obscurely 3- to 5-lobed.

Rubiaceae Magee and Ahles (1999) noted a report of Canthium odoratum (G. Forst.) Seem. in MA but provided no details (e.g., location, source, validity of report). Specimens of this species from New England are unknown. Stenaria nigricans (Lam.) Terrel var. nigricans was reported from NH by Kartesz (1999), but the report was erroneous.

Rub i ac e a e   8 2 1

1a. Plants woody; inflorescence a densely flowered, spherical cluster [Fig. 874]; corolla slender-funnelform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephalanthus 1b. Plants herbaceous; inflorescence not a spherical cluster; corolla salverform, rotate, cupuliform, or funnelform 2a. Leaves 2 at each node (i.e., opposite) [Figs. 882, 883]; fruit fleshy or dehiscent; corolla pubescent on the adaxial (i.e., inside) surface (glabrous in Houstonia caerulea) 3a. Flowers united by their hypanthia [Fig. 883]; fruit a berry; plants creeping, with evergreen leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mitchella 3b. Flowers separate, not united [Fig. 882]; fruit a capsule or schizocarp; plants prostrate to erect, sometimes tufted, with deciduous leaves 4a. Fruit a schizocarp, separating into 2 mericarps; ovules solitary in each locule; flowers borne in the axils of leaves; stipules terminated by a fringe of erect bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diodia 4b. Fruit a capsule; ovules 4 or more in each locule; flowers borne at the apex of the stem, either solitary or in cymes; stipules entire, without a fringe. . . . . . . . . . Houstonia 2b. Leaves 3–12 at a node (i.e., whorled; rarely only 2 leaves at some of the nodes in Galium) [Figs. 877, 879]; fruit dry, schizocarpic, consisting of 2 indehiscent, 1-seeded carpels [Figs. 876, 881]; corolla glabrous (pubescent on the abaxial surface in Galium circaezans) 5a. Sepals triangular; corolla blue or pink, funnelform; inflorescence subtended by an involucre composed of a whorl of basally connate leaves . . . . . . . . . . . . . . . . . . . Sherardia 5b. Sepals obsolete; corolla white, green-white, yellow, yellow-green, green-purple, or purple, or blue-purple in Asperula usually rotate; inflorescence not subtended by an involucre or subtended by one composed of distinct leaves 6a. Corolla 5-merous; fruit fleshy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rubia 6b. Corolla 3- or 4-merous [Figs. 875, 879]; fruit dry 7a. Corolla blue-purple, funnelform, with a prolonged, slender, basal, connate portion 4–5.5 mm long that is longer than the lobes; flowers with closely subtending, narrow bracteoles, borne on pedicels shorter than or as long as the ovary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asperula 7b. Corolla various shades of white, yellow, or purple, rotate or cupuliform (funnelform in G. odoratum), the basal, connate portion up to ca. 1.5 mm long, shorter than or as long as the lobes; flowers without closely subtending bracteoles, often borne on pedicels longer than the ovary [Figs. 880, 881] . . . . . . . . . . . . . . . . . . . Galium

Asperula 1. Asperula arvensis L. E blue woodruff. CT, MA. Fields, roadsides.

Cephalanthus 1. Cephalanthus occidentalis L. N Fig. 874 common buttonbush. Cephalanthus occidentalis L. var. pubescens Raf. • CT, MA, ME, NH, RI, VT. Pond shores, slow-moving rivers, swamps, ditches. Fig. 874 Inflorescences and leaves of Cephalanthus occidentalis.

8 22   tricolpate s

Diodia 1a. Calyx with 2 sepals; corolla salverform, 8–11 mm long; gynoecium with 2 linear stigmas; fruit 6–9 mm long, glabrous or villous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. virginiana 1b. Calyx with 4 sepals; corolla funnelform, 5–6 mm long; gynoecium with a single, capitate stigma; fruit 2.5–6 mm long, strigose to hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. teres 1. Diodia teres Walt. var. teres n rough buttonweed. Diodella teres (Walt.) Small; Diodia teres Walt. var. setifera Fern. & Grisc. • CT, MA, RI. Coastal beaches, roadsides, sandy fields, waste areas. Native to CT and RI, nonnative in MA. 2. Diodia virginiana L. var. virginiana E Virginia buttonweed. Diodia hirsuta Pursh; D. tetragona Walt. • CT. Gardens, waste areas.

Galium Corolla measurements provided in the key were based on dried specimens. References: Ehrendorfer (1976), Puff (1976). 1a. Primary axis and main branches of plants with (2–) 4 leaves at each node [Figs. 875, 879] 2a. Principal leaves with 3 or 5 nerves [Fig. 875]; schizocarps bristly (sometimes glabrous in G. boreale) [Fig. 876] 3a. Corolla white to cream-white; leaf blades broad-linear, 1–2.5 mm wide; bristles of the schizocarp, when present, not hooked at the apex; angles of the stem commonly retrorse-scabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. boreale 3b. Corolla green-purple, purple, or yellow-green; leaf blades ovate to obovate or elliptic to oval, 5–25 mm wide; bristles of the schizocarp hooked at the apex [Fig. 876]; angles of the stem glabrous or pubescent, but not retrorse-scabrous 4a. Flowers all pedicellate at the tips of the inflorescence branches; leaf blades 10–30 mm long 5a. Flowering stems with 2–4 (–5) nodes; distal leaf blades larger than proximal leaf blades; corolla yellow-white . . . . . . . . . . . . . . . . . . . . . . . . . . . G. kamtschaticum 5b. Flower stems with more than 6 nodes; leaf blades of ± uniform size along the stem; corolla green-white to purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. pilosum 4b. Some of the flowers sessile along the branches of the inflorescence [Fig. 875]; leaf blades 20–80 mm long 6a. Leaf blades oval to elliptic, widest usually near the middle, obtuse at the apex [Fig. 875]; petals commonly pubescent on the abaxial surface, acute at the apex [Fig. 875] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. circaezans 6b. Leaf blades elliptic to lanceolate, widest usually below the middle, acute to acuminate at the apex; petals glabrous, acuminate at the apex . . . . G. lanceolatum 2b. Principal leaves with 1 nerve [Figs. 878, 880]; schizocarps without bristles [Fig. 880] 7a. Cymes branched 3 or more times, bearing 5 or more flowers [Fig. 879]; some of the pedicels spreading, especially in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) G. palustre 7b. Cymes branched 1 or 2 times, bearing 2–4 flowers [Figs. 880, 881]; pedicels mostly erect to ascending (often divaricately spreading in G. tinctorium) 8a. Corollas mostly 3-merous, 0.8–2.5 mm in diameter, the obtuse lobes as wide as or wider than long; stems often retrorse-scabrous on the angles [Fig. 881], usually not densely short-bearded just below the nodes; plants developing prostrate, basal offshoots late in the season

Rub i ac e a e   8 23

9a. Fruiting pedicels ± straight, stiff, lacking scabrules, usually some divaricately spreading (sometimes somewhat arcuate and/or with a few scabrules in G. tinctorium ssp. floridanum) [Fig. 880]; corolla 1.5–2.5 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. tinctorium 9b. Fruiting pedicels arcuate, scabrous or, less commonly, smooth, mostly erect to ascending [Fig. 881]; corolla 0.8–1.5 (–1.6) mm in diameter 10a. Corolla 0.8–1 mm in diameter; leaf blades 5–8 (–12) × (0.8–) 1–2 (–3) mm; mature schizocarps 1.8–2 (–2.3) mm wide; fruiting peduncles 1.5–5 mm long, smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. brevipes 10b. Corolla 1.2–1.5 (–1.6) mm in diameter; leaf blades (7–) 8–20 (–25) × (1–) 1.2– 5 (–6) mm; mature schizocarps (2–) 2.2–3.5 (–4) mm wide; fruiting peduncles (2–) 3–15 (–20) mm long, scabrous or smooth . . . . . . . . . . . . . . . . . . . . . G. trifidum 8b. Corollas 4-merous, 2–3.5 mm in diameter, the acute lobes longer than wide; stems not scabrous on the angles [Fig. 878], densely short-bearded just below the nodes [Fig. 878]; plants not developing basal offshoots in late season 11a. Leaf blades 10–30 × 1–6 mm, ascending to spreading; mericarps 4–5 mm in diameter, tuberculate, usually only 1 developing; stems 2–8 dm tall . . . . G. obtusum 11b. Leaf blades 8–15 × 1–3 mm, spreading to deflexed; mericarps 2–3 mm in diameter, smooth, usually both developing; stems 1–4 dm tall . . . . . G. labradoricum 1b. Primary axis and main branches with 5–12 leaves at each node [Fig. 877] 12a. Leaf blades blunt at the apex [Figs. 880, 881] 13a. Cymes branched 3 or more times, bearing 5 or more flowers; some of the pedicels spreading, especially in fruit; corollas 4-merous, 2.5–3.3 mm wide, the lobes longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) G. palustre 13b. Cymes branched 1 or 2 times, bearing 2–4 flowers; pedicels mostly erect or ascending; corollas mostly 3-merous, 0.8–2.5 mm wide, the lobes as wide as or wider than long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . go to couplet 9 12b. Leaf blades sharply pointed to cuspidate at the apex 14a. Corolla 0.5–1 mm in diameter, yellow-red; schizocarps 1–1.4 mm wide, finely papillose on the surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. divaricatum 14b. Corolla 1–7 mm in diameter, white, green-white, or yellow; schizocarps 2–6 mm wide, smooth or uncinate-bristly (papillose with pointed projections in G. tricornutum, but then the schizocarps 3–5 mm wide) 15a. Pedicels strongly recurved to deflexed post anthesis; schizocarps papillose with pointed projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. tricornutum 15b. Pedicels erect to spreading; schizocarps smooth or uncinate-bristly 16a. Stems scabrous on the angles of the internodes (at least on the proximal internodes), reclining to upright; inflorescences relatively few-flowered, each with 2–5 flowers, usually once- or twice-branched (note: a given plant can bear multiple such inflorescences) 17a. Leaves (6–) 8 per node; plants annual, with a short taproot; mericarps 2–4 (–5) mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. aparine 17b. Leaves (4–) 6 per node; plants perennial, with creeping rhizomes; mericarps 1.5–2 mm in diameter 18a. Schizocarps glabrous; leaf blades 8–20 × 2–6 mm, retrorsely scabrous on the margin; corolla white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. asprellum 18b. Schizocarps uncinate-bristly [Fig. 876]; leaf blades 15–85 × 4–13 mm, antrorsely scabrous on the margin; corolla green-white . . . . . . . . G. triflorum

8 24   tricolpate s

16b. Stems glabrous or pubescent on the internodes, but without scabrules, usually upright; inflorescences relatively many-flowered, each with 5–20 or more flowers, usually 2 or more times branched, collectively forming a large, open, cymose panicle [Fig. 877] 19a. Corolla funnelform, 4–7 mm in diameter; schizocarps covered with uncinate bristles; stems retrorsely hispid at the nodes . . . . . . . . . . . G. odoratum 19b. Corolla rotate, 2–3 (–5) mm in diameter; schizocarps smooth; stems glabrous or pubescent, but without a distinct ring of hairs at the nodes 20a. Leaf blades narrow-linear, 0.5–2 (–3) mm wide, the margins prominently revolute (sometimes the margins even recurved as far as the midrib), (6–) 8–12 per node; lobes of corolla merley acute at apex (rarely with an obscure apiculus in G. verum) 21a. Corollas yellow, rotate, 2–3.5 mm in diameter; schizocarps 1–1.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. verum 21b. Corollas white, cupuliform, 4–6 mm in diameter; schizocarps 1.5–2 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. glaucum 20b. Leaf blades linear to elliptic, (1–) 2–10 mm wide, not or only slightly revolute, 6–8 per node; lobes of corolla apiculate at apex or usually merely acute in G. sylvaticum 22a. Leaf blades 20–40 mm long, glaucous abaxially, distinctly paler than the adaxial surface; pedicels capillary, mostly 0.05–0.1 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. sylvaticum 22b. Leaf blades 10–25 (–40) mm long, not glaucous abaxially, only somewhat paler than the adaxial surface; pedicels relatively thicker, mostly 0.08–0.2 mm thick 23a. Corolla 2–3 (–3.3) mm in diameter; pedicels usually longer than the diameter of the flower; inflorescence lax and open, the branches divaricately spreading post anthesis . . . . . . . . . . . . . . . . . . . . G. mollugo 23b. Corolla (2.5–) 3–5 mm in diameter; pedicels usually shorter than the diameter of the flower; inflorescence relatively dense, the branches ascending to spreading post anthesis . . . . . . . . . . . . . . . . . . . . . . G. album 1. Galium album L. E white bedstraw. Galium erectum Huds., sensu Huds. 1778; G. mollugo L. ssp. erectum (Huds.) Briq.; G. mollugo L. var. erectum (Huds.) Domin. • CT, MA, ME, NH, VT. Fields, roadsides, waste places. 2. Galium aparine L. N scratch bedstraw. Galium aparine L. var. echinospermum (Wallr.) Farw.; G. aparine ssp. spurium (L.) Simonkai; G. aparine L. var. vaillantii (DC.) Koch; G. spurium L.; G. spurium L. var. echinospermum (Wallr.) Hayek; G. spurium L. var. vaillantii (DC.) G. Beck; G. vaillantii DC. • CT, MA, ME, NH, RI, VT. Forests, forest fragments, fields and field edges, swamps, roadsides, waste areas. Two subspecies are sometimes recognized—ssp. aparine with green-white corollas 1.5–1.7 mm in diameter and uncinate-bristly schizocarps (2.8–) 3–4 (–5) mm wide exclusive of the hairs and ssp. spurium with green-yellow corollas 0.8–1.3 mm in diameter and glabrous or uncinate-bristly schizocarps 2–3 mm wide exclusive of the hairs. Preliminary herbarium surveys suggest that these two taxa are not discrete and supposed diagnostic characters appear mixed on some specimens. Until further study contributes to our understanding of local plants, this species is recognized broadly without subspecific entities.

Rub i ac e a e   8 25

3. Galium asprellum Michx. N rough bedstraw. CT, MA, ME, NH, RI, VT. Swamps, riparian forests, stream banks, wetland margins. 4. Galium boreale L. ssp. septentrionale (Roemer & J.A. Schultes) Hara N northern bedstraw. Galium boreale L. var. hyssopifolium (Hoffmann) DC.; G. boreale L. var. intermedium DC.; G. boreale L. var. linearifolium Rydb.; G. boreale L. var. scabrum DC.; G. boreale L. ssp. septentrionale (Roemer & J.A. Schultes) Iltis; G. hyssopifolium Hoffman; G. septentrionale Roemer & J.A. Schultes • CT, MA, ME, NH, RI, VT. Fens, low meadows, fields, and woodlands in high-pH bedrock regions. Our subspecies of this plant differs from the typical European one in that it shows larger corollas, anthers, and fruits, more densely pubescent nodes, and differences in the shapes of bracts and inflorescences. This combined with the differences in chromosome number support treating our plants as distinct (Iltis 1957). 5. Galium brevipes Fern. & Wieg.

NC

limestone swamp bedstraw. Galium trifidum L. ssp. brevipes (Fern. & Wieg.) A. & D. Löve • MA, ME, NH, VT. Fens, swamps dominated by Thuja occidentalis, pond shores, and ditches in regions of high-pH bedrock and/or till. 6. Galium circaezans Michx. N Fig. 875 forest licorice bedstraw.  6a. Galium circaezans Michx. var. glabrum Britt.; G. circaezans Michx. var. typicum Fern.; G. rotundifolium L. var. circaezans (Michx.) Kuntze • CT, MA, ME, NH, RI, VT. Mesic to dry-mesic deciduous forests and woodlands, balds, ridges. 1a. Leaf blades abaxially glabrous or sparsely short-hispid on the veins, 15–25 (–40) × 7–14 (–18) mm; stems glabrous or sparsely pilose on the angles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6a. G. circaezans var. circaezans 1b. Leaf blades abaxially appressed-pilose on the surface and long-hispid on the veins, the larger blades 20–50 × 10–25 mm; stems pubescent on the angles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6b. G. circaezans var. hypomalacum Fern.

Fig. 875  Flowers and leaves of Galium circaezans.

Variety circaezans is known from CT, MA, ME, NH, RI, VT. Variety hypomalacum is known from CT, MA, ME, NH, RI, VT. 7. Galium divaricatum Pourret ex Lam. E Lamarck’s bedstraw. Galium anglicum Huds.; G. parisiense L. ssp. anglicum (Huds.) Gaud.; G. parisiense L. var. anglicum (Huds.) G. Beck; G. parisiense L. var. divaricatum (Pourret ex Lam.) Vis. • VT. Fields. This record rests on the following voucher: Pease 38486 (NEBC!). It is unusual for the species because its corolla is too large (wider than 1 mm in diameter). The identification should be considered tentative. 8. Galium glaucum L. E waxy bedstraw. Asperula glauca (L.) Bess. • CT. Fields. 9. Galium kamtschaticum Steller ex J.A. & J.H. Schultes

N C Fig. 876

boreal bedstraw. ME, NH, VT. Boreal and subalpine forests, seeps and stream banks in northern and/or mountainous areas, often associated with abundant bryophyte cover. 10. Galium labradoricum (Wieg.) Wieg.

NC

northern bog bedstraw. Galium tinctorium (L.) Scop. var. labradoricum Wieg. • CT, MA, ME, VT; restricted to western counties of southern New England states. Fens, swamps, often dominated by Thuja occidentalis, marshes, frequently in regions of high-pH bedrock and/ or till. Reports of this species in NH (e.g., Kartesz 1999) are based on specimens of Galium palustre, G. tinctorium, and G. trifidum (specimens at NEBC! and NHA!). 11. Galium lanceolatum Torr. N lance-leaved licorice bedstraw. Galium circaezans Michx. var. lanceolatum (Torr.) Torr. & Gray; G. rotundifolium L. var. lanceolatum (Torr.) Kuntze • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic, often rich, deciduous forests and woodlands.

Fig. 876  Schizocarp of Galium kamtschaticum.

8 26   tricolpate s

12. Galium mollugo L. E Fig. 877 whorled bedstraw. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, open shorelines. 12 × 22. This very rare bedstraw hybrid is known from MA. It most closely resembles Galium verum but can be recognized by its quandrangular stems (at least below; rather than terete with 4 raised lines), linear to narrow-lanceolate leaf blades with some usually wider than 1.5 mm that usually do not darken in drying (rather than linear, usually narrower than 1.5 mm, and usually darkening in drying), and a bright yellow to yellow-white corolla with somewhat apiculate lobes (rather than yellow and ‌pomeranicum Retz. merely acute at the apex of the lobes). It is very similar to G. × (G. album × G. verum; a nothospecies not yet documented from New England) except that the corollas are usually narrower than 3 mm (rather than usually wider than ‌pomeranicum in New England 3 mm). This hybrid is responsible for reports of G. × (e.g., Seymour 1982, Sorrie and Somers 1999). It may also be responsible for reports of that nothospecies from CT, but specimens were not available for study. 13. Galium obtusum Bigelow ssp. obtusum N Fig. 878 Fig. 877  Inflorescence and leaves of Galium mollugo.

blunt-leaved bedstraw. Galium obtusum Bigelow var. ramosum Gleason; G. trifidum L. var. latifolium Torr. • CT, MA, NH, RI, VT; also reported from ME by Campbell et al. (1995), but specimens are unknown. Swamps, lacustrine floodplain forests, wet-mesic forests. 14. Galium odoratum (L.) Scop. E sweet-scented bedstraw. Asperula odorata L. • MA, VT. Fields, lawns, gardens, waste areas. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI. 15. Galium palustre L. N Fig. 879 marsh bedstraw. CT, MA, ME, NH, RI, VT. Swamps, marshes, hydric fields and meadows. 16. Galium pilosum Ait.

NC

hairy bedstraw.  16a. Galium puncticulosum Michx. var. pilosum (Ait.) DC.; 16b. Galium punctatum Pers; G. puncticulosum Michx. • CT, MA, NH, RI, VT. Woodlands, ridges, fields, clearings. 1a. Pubescence of stem and leaves ± straight, ± spreading . . . . 16a. G. pilosum var. pilosum Fig. 878  Retrorsely pubescent node of Galium obtusum.

1b. Pubescence of stem and leaves of short, upwardly curved hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16b. G. pilosum var. puncticulosum (Michx.) Torr. & Gray Variety pilosum is known from CT, MA, NH, RI, VT. Variety puncticulosum is known from CT and is of regional conservation concern. 17. Galium sylvaticum L. E wood bedstraw. CT, MA, ME, VT. Fields, roadsides, waste areas, forest borders. 18. Galium tinctorium (L.) Scop. N Fig. 880 stiff three-petaled bedstraw. 18a. Galium tinctorium (L.) Scop. ssp. floridanum (Wieg.) Puff; 18b. Asperula tinctoria L.; Galium trifidum L. ssp. tinctorium (L.) Hara; G. trifidum L. var. tinctorium (L.) Torr. & Gray • CT, MA, ME, NH, RI, VT. Swamps, fens, marshes, hydric fields and meadows. 1a. Fruiting pedicels (4–) 5–8 (–12) mm long; schizocarps (2.5–) 3–4 (–5) mm wide; ultimate portion of inflorescence with 3 or more flowers; leaf blades (10–) 12–20 × 2–3 (–4) mm; plants (15–) 20–75 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18a. G. tinctorium var. floridanum Wieg. 1b. Fruiting pedicels (2–) 2.5–5 (–6) mm long; schizocarps (2–) 2.5–3 mm wide; ultimate portion of inflorescence with 3 flowers; leaf blades (6–) 10–15 (–18) × (1.5–) 2–2.5 (–2.8) mm; plants 15–50 (–60) cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18b. G. tinctorium var. tinctorium

Fig. 879  Inflorescence and leaves of Galium palustre.

Variety floridanum is known from CT, MA, RI. Variety tinctorium is known from CT, MA, ME, NH, RI, VT.

Rub i ac e a e   8 27

19. Galium tricornutum Dandy E rough-fruited bedstraw. Galium tricorne Stokes, pro parte • RI. Fields, waste areas. 20. Galium trifidum L. N Fig. 881 three-petaled bedstraw. 20a. Galium trifidum L. var. halophilum Fern. & Wieg. • CT, MA, ME, NH, RI, VT; nearly throughout. Swamps, fens, marshes, hydric fields and meadows, shorelines, saline and brackish marshes, Atlantic coast shorelines. 1a. Plants smooth or with a few scattered scabrules, slightly succulent, occurring in saline influenced habitats; schizocarps (2–) 2.5–3.5 (–5) mm wide; pedicels relatively thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20a. G. trifidum ssp. halophilum (Fern. & Wieg.) Puff 1b. Plants usually retrorsely scabrous on the stems and leaf blades [Fig. 881], never succulent, occurring in non-saline habitats; schizocarps 2–2.5 (–2.8) mm wide; pedicels filiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20b. G. trifidum ssp. trifidum

Fig. 880  Smooth, straight pedicels of Galium tinctorium.

Subspecies halophilum is known from MA, ME. It is restricted to saline habitats of the coastal region. Subspecies trifidum is known from CT, MA, ME, NH, RI, VT. It is widespread in New England in hydric and seasonally hydric communities. 21. Galium triflorum Michx. N fragrant bedstraw. Galium triflorum Michx. var. asprelliforme Fern. • CT, MA, ME, NH, RI, VT. Dry-mesic to wet-mesic forests, swamps, rocky slopes. 22. Galium verum L. E yellow bedstraw. 22b. Galium wirtgenii F.W. Schultz • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 1a. Leaf blades 15–30 (–35) × 0.5–1 (–2) mm, with prominently revolute margins such that the margins are recurved to the abaxial midrib, shorter than to longer than the associated internodes; inflorescence with dense, contiguous flowers, the branches longer than the corresponding stem internodes; flowers fragrant . . . . . . . . . . . . . . . . 22a. G. verum ssp. verum 1b. Leaf blades 25–40 × 1–3 mm, with revolute margins, but the margins not recurved to the abaxial midrib, shorter than the associated internodes; inflorescence often interrupted, the branches usually shorter then the corresponding stem internodes; flowers odorless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22b. G. verum ssp. wirtgenii (F.W. Schultz) Oborny

Fig. 881  Scabrous, arching pedicels of Galium trifidum ssp. trifidum.

Subspecies verum is known from CT, MA, ME, NH, RI, VT. Subspecies wirtgenii is known from CT, MA, ME, VT.

Houstonia Houstonia canadensis Willd. ex Roemer & J.A. Schultes was reported from ME by Gleason (1963), but specimens are unknown. Reference: Terrell (1996). 1a. Lower leaves borne on an evident petiole; corolla lobes horizontally spreading, glabrous; flowers borne singly on an elongate peduncle 20–70 mm long; capsule compressed; seeds globular, with a deep, round cavity on the inner face . . . . . . . . . . . . . . . . . . . . . . . . . . H. caerulea 1b. Lower leaves sessile or with an obscure petiole; corolla lobes ascending to recurved, pubescent within the basal, connate tube; flowers borne multiple together in a cyme on pedicels 2–5 mm long [Fig. 882]; fruit not compressed; seeds meniscoid, with a ridge across the depression on the inner face 2a. Stem leaf blades broad-lanceolate to narrow-oblong, (4–) 5–12 mm wide, rounded at the base; sepals 3–6 mm long at anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. purpurea 2b. Stem leaf blades broad-linear to narrow-oblong, 2–5 mm wide, narrowed to the base; sepals 1.5–2.5 (–2.7) mm long at anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. longifolia

Fig. 882  Flowers of Houstonia longifolia.

8 28   tricolpate s

1. Houstonia caerulea L. N little bluet. Hedyotis caerulea (L.) Hook.; Houstonia caerulea L. var. faxonorum Pease • CT, MA, ME, NH, RI, VT. Fields, lawns, roadsides, forests, clearings, ledges, open gravel, stream banks, sometimes ascending high into the mountains. Plants from high-elevation settings tend to have large capsules and seeds and have been segregated as var. faxonorum. 2. Houstonia longifolia Gaertn. N long-leaved bluet. Hedyotis longifolia (Gaertn.) Hook.; H. longifolia (Gaertn.) Hook. var. tenuifolia (Nutt.) Torr. & Gray; H. purpurea (L.) Torr. & Gray var. longifolia (Gaertn.) Fosberg; Houstonia longifolia Gaertn. var. glabra Terrell; H. longifolia Gaertn. var. tenuifolia (Nutt.) Wood; H. tenuifolia Nutt.; Oldenlandia purpurea (L.) Gray var. tenuifolia (Nutt.) Gray ex Chapman • ct, ma, me, nh, ri, vt. Cobble and ledge river shores, lake shore ledges, woodlands, fields, clearings. 3. Houstonia purpurea L. var. calycosa Gray E large bluet. Hedyotis lanceolata Poir.; H. purpurea (L.) Torr. & Gray var. calycosa (Gray) Fosberg; Houstonia calycosa (Gray) C. Mohr; H. lanceolata (Poir.) Britt. • CT, MA, ME, VT. Dry fields, meadows, and clearings.

Mitchella 1. Mitchella repens L. N Fig. 883 partridge-berry. CT, MA, ME, NH, RI, VT. Dry-mesic to wet-mesic forests and forest borders, usually with some component of evergreen trees.

Rubia Fig. 883  Flowers and leaves of Mitchella repens.

1. Rubia tinctoria L. E madder. MA. Gardens, waste areas.

Sherardia 1. Sherardia arvensis L. E blue field-madder. CT, MA, ME, RI, VT. Fields, lawns.

Rutaceae 1a. Flowers weakly zygomorphic; petals 20–25 mm long; stamens declined and upwardly curved near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dictamnus 1b. Flowers actinomorphic; petals up to 10 mm long; stamens ± straight 2a. Herbaceous plants with bipinnatifid leaves; fruit a 4- or 5-lobed capsule; flowers all bisexual, with yellow petals 8–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ruta 2b. Shrubs or trees with pinnately compound leaves; fruit a samara, drupe, or follicle; some or all of the flowers on a given plant unisexual (the unisexual flowers in Ptelea by imperfect or abortive stamens), with green-white, yellow-white, or yellow-green petals up to 7 mm long 3a. Leaves with 3 leaflets; fruit a ± orbicular samara; plants polygamous (i.e., some flowers bisexual, others unisexual, in the latter case the carpellate flowers functionally unisexual due to imperfect or abortive stamens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ptelea

Ru tac e a e   8 2 9

3b. Leaves with 5–13 leaflets; fruit a drupe or fleshy follicle; plants dioecious (i.e., all the flowers unisexual) 4a. Leaves opposite; branches unarmed; fruit a drupe . . . . . . . . . . . . . Phellodendron 4b. Leaves alternate; branches armed with prickles; fruit a fleshy follicle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zanthoxylum

Dictamnus 1. Dictamnus albus L. E gasplant. VT. Fields, roadsides, forest borders, gardens.

Phellodendron Reference: Ma et al. (2006). 1. Phellodendron amurense Rupr. E Amur corktree. Phellodendron amurense Rupr. var. sachalinense F. Schmidt; P. japonicum Maxim.; P. sachalinense (F. Schmidt) Sarg. • CT, MA, RI. Forests, forest fragments, roadsides. Phellodendron amurense is interpreted in the broad sense based on work by Ma et al. (2006) who showed that several morphological characters deemed to be of importance to past researchers were continuous and did not discriminate taxa.

Ptelea 1. Ptelea trifoliata L. ssp. trifoliata var. trifoliata E common hoptree. Ptelea baldwinii Torr. & Gray • CT, MA, ME, NH, VT. Roadsides, forest fragments and borders, waste areas. Reports of this species in ME (e.g., Seymour 1982, Kartesz 1999) are based on specimens taken from planted individual.

Ruta 1. Ruta graveolens L. E common rue. CT, RI, VT. Fields, roadsides, waste areas, gardens. This species was reported from most counties in New England by Magee and Ahles (1999), but specimens are unknown (and this is certainly erroneous).

Zanthoxylum 1. Zanthoxylum americanum P. Mill. n common prickly-ash. CT, MA, ME, NH, RI, VT. Fields, roadsides, forests, forest fragments and borders, woodlands, sometimes associated with rocky and/or ledgy soils. This species is introduced in ME and native to the other states.

Salicaceae 1a. Winter buds covered by more than one scale; aments drooping; flowers subtended by a cup-like disk, without basal glands, each staminate flower with 5–80 stamens . . . . . . Populus

830  tricolpate s

1b. Winter buds covered by 1 scale; aments erect to spreading [Figs. 887, 893]; flowers not subtended by a disk, with 1–4 basal glands, each staminate flower with 1–8 stamens . . . . Salix

Populus Species of Populus display leaf blade heterophylly along the branchlet over the growing season. Those leaves that overwinter in the winter in bud and expand in the spring are called first leaves (or preformed) vs. those leaves that are produced later in the season as the growing shoot continues to expand, which are called new leaves (or neoformed). As these leaves can vary in size, shape, and number of marginal teeth, it is important to collect both types when possible. Greatly enlarged leaves produced on vigorous shoots are not accounted for in the identification key. Reference: Eckenwalder (2010).

Key for reproductiv e material 1a. Floral scales with margins dentate or with 3–7 narrow-triangular, long-ciliate lobes; staminate flowers with 5–12 (–15) stamens; carpellate flowers with 2 slender, bifid stigmas 2a. Floral scales merely dentate; capsules 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . P. alba 2b. Floral scales with 3–7 narrow-triangular lobes; capsules 5–9 mm long 3a. Branchlets (especially about the nodes) and terminal winter bud pubescent; stigmas lacking basal lobes; floral scales of staminate aments with 5–7 lobes; ovary pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grandidentata 3b. Branchlets and terminal winter bud glabrous or nearly so; stigmas with recurved basal lobes; floral scales of staminate aments with 3–5 lobes (usually with more lobes in P. tremula); ovary glabrous (or sometimes scabridulous in P. tremula) 4a. Mature capsules 5–9 mm long; scales of staminate aments with 3–5 lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. tremuloides 4b. Mature capsules usually shorter than 5 mm; scales of staminate aments usually with more than 5 apical lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. tremula 1b. Floral scales with margins conspicuously fimbriate; staminate flowers with 12–80 stamens; carpellate flowers with 2–4 broadly dilated stigmas 5a. Stigmas borne on elongate styles; staminate flowers with 12–20 stamens; anthers apiculate at the apex; capsules on pedicels 10–15 mm long; terminal winter bud not or scarcely viscid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. heterophylla 5b. Stigmas sessile or nearly so; staminate flowers with (15–) 20–80 stamens; capsules on pedicels 2–10 mm long; anthers blunt or emarginate at the apex; terminal winter bud viscid 6a. Carpellate flowers with 3 or 4 stigmas; staminate flowers with (30–) 40–80 stamens; capsules 3- or 4-valved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. deltoides 6b. Carpellate flowers with 2 stigmas; staminate flowers with 20–30 stamens; capsules 2-valved 7a. Ovules 15–22 per placenta; floral scales of carpellate flowers fimbriate with many, very narrow segments; stigmas spreading, with coarsely toothed, ± orbicular lobes; trees with spreading to ascending branches . . . . . . . . . . . . . . . . . . . . . . . P. balsamifera 7b. Ovules 4–8 per placenta; floral scales of carpellate flowers fimbriate with 9–11 slender segments; stigmas reflexed and retrorsely appressed agaist the ovary, with entire, ± ovate lobes; trees often with strictly ascending branches producing a narrow, columnar form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. nigra

Key for mature vegetati ve material 1a. Petioles broad-elliptic to terete in cross-section (sometimes slightly flattened near the apex), usually with a shallow channel on the upper surface

S a l i c ac e a e   83 1

2a. Mature leaf blades of long shoots abaxially tomentose, the margins varying from 3- to 5-lobed or irregularly serrate with coarse teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. alba 2b. Mature leaf blades abaxially glabrous or with pubescence confined to the main veins, the margins regularly serrate or crenate-serrate 3a. Leaf blades obtuse to rounded at the apex, cordate at the base [Fig. 885], sometimes narrowed to a short, broad base prior to junction with petiole, without orange resin on the abaxial surface, permanently pubescent on the abaxial veins; terminal winter bud not or scarcely viscid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. heterophylla 3b. Leaf blades acute to short-acuminate at the apex, broad-cuneate to subcordate at the base, often with streaks of orange resin on the abaxial surface, glabrous; terminal winter bud very viscid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. balsamifera 1b. Petioles conspicuously compressed from the lateral surfaces, without a channel on the upper surface 4a. Leaf blades with a definite, though sometimes very narrow, translucent margin [Fig. 884] 5a. Leaf blades on long shoots 5–10 cm long, eciliate, crenate-serrate with teeth ca. 1 mm tall; apex of petiole without glands; branchlets terete; lateral winter buds appressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. nigra 5b. Leaf blades on long shoots 10–18 cm long, ciliate, serrate with teeth 2–5 mm tall; apex of petiole with 2–5 glands; branchlets often somewhat 4-angled; lateral winter buds ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. deltoides 4b. Leaf blades lacking a translucent border [Fig. 886] 6a. Leaf blades narrow-ovate to broad-ovate, (5–) 7–12 cm long; terminal winter buds pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grandidentata 6b. Leaf blades rhombic to suborbicular or reniform, 2–8 (–10) cm long; terminal winter buds glabrous to sparsely pubescent 7a. Leaf blade regularly denticulate with 18–30 (–40) teeth per margin up to 1 mm tall [Fig. 886] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. tremuloides 7b. Leaf blade coarsely sinuate-dentate with mostly 5–10 teeth per margin taller than 1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. tremula 1. Populus alba L. E white poplar. Populus alba L. var. bolleana Lauche • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, waste areas, railroads, near dwellings. 1‌ × 4. Populus ×rouleauiana Boivin is a rare poplar hybrid in New England known from CT, MA. It has leaves borne on variable petioles, with most only somewhat compressed. The leaf blades, especially those toward the apex of the shoot, are abaxially moderately to densely tomentose, the tomentum sometimes in patches leaving much of the proximal leaf blade glabrous (rarely with some leaves with the entire abaxial surface glabrous). The leaf blade margins most resemble P. grandidentata, though the teeth tend tend to be fewer and some blades show obscure lobes (the hybrid has 4–12 teeth per margin and P. grandidentata has 5–14 teeth per margin). 1‌ × 7. Populus ×canescens (Ait.) J.E. Smith is an uncommon hybrid poplar that has escaped from cultivation in CT, MA, ME, NH, RI, VT. It resembles P. alba in some aspects but has leaf blades that are merely dentate (i.e., the blades are not lobed or only scarcely so), the proximal ones sparsely to moderately tomentose abaxially with gray, brownwhite, or sordid-white hairs (rather than bright white), and petioles that vary from compressed to subterete. The floral scales are prominently lacerate with 7–9 lobes, and the staminate flowers have 8–15 stamens (rather than floral scales merely dentate with many apical teeth and staminate flowers with 5–10 stamens in P. alba).

832  tricolpate s

1‌ × 8. Populus ×heimburgeri Boivin is a rare poplar hybrid that is known from MA. It has first leaves that are tomentose on the abaxial surface of the blades with white-gray hairs and blade margins with (4–) 6–8 large, blunt teeth (only rare blades show any sign of the lobing of P. alba). The new leaves are similar in regard to pubescence but have mostly 11–17 small teeth (i.e., approaching the morphology of P. tremuloides). It is further characterized by petioles that vary from compressed to subterete. 2. Populus balsamifera L. ssp. balsamifera N balsam poplar. Populus balsamifera L. var. candicans (Ait.) Gray; P. balsamifera L. var. lanceolata Marsh.; P. balsamifera L. var. subcordata Hyl.; P. candicans Ait.; P. tacamahaca P. Mill. P. tacamahaca P. Mill. var. candicans (Ait.) Stout; P. tacamahaca P. Mill. var. lanceolata (Marsh.) Farw. • CT, MA, ME, NH, RI, VT; becoming infrequent in southern New England. River shores and banks, forests and forest borders, lake shores, successional fields, roadsides.

Fig. 884  Leaf of Populus deltoides showing translucent border and glands at apex of petiole.

‌2 × 3. Populus ×jackii Sarg. is an uncommon hybrid poplar that has escaped from cultivation in CT, MA, ME, NH, RI, VT. It has long been known by the name Populus ×‌gileadensis Rouleau. It can be recognized by its broad-ovate or triangular-ovate leaf blades that are acuminate at the apex and borne on weakly compressed petioles, visicid terminal winter buds, terete branchlets (rather than somewhat 4-angled in P. deltoides), staminate flowers with 25–40 stamens (rather than (30–) 40–80 stamens in P. deltoides), and 2- or 3-valved capsules with 7–14 (–20) seeds per placenta (rather than 2-valved capsules with 15–22 seeds per placenta in P. balsamifera). 3. Populus deltoides Bartr. ex Marsh. var. deltoides n Fig. 884 necklace poplar. Populus angulata Ait.; P. angulata Ait. var. missouriensis A. Henry; P. balsamifera L. var. missouriensis (A. Henry) Rehd.; P. deltoides Bartr. ex Marsh. var. angulata (Ait.) Sarg.; P. deltoides Bartr. ex Marsh. var. missouriensis (A. Henry) A. Henry; P. deltoides Bartr. ex Marsh. var. pilosa (Sarg.) Sudworth • CT, MA, ME, NH, RI, VT. River banks, riparian forests, lacustrine forests, roadsides, waste areas, about dwellings. This species is non-native in ME and native, at least in part, to other states in New England. As a native, it appears usually on shorelines and riverbanks and in lacustrine and riparian floodplains. 4. Populus grandidentata Michx. N big-toothed poplar. CT, MA, ME, NH, RI, VT. Forests, woodlands, successional fields, forest borders. ‌4 × 8. Populus ×smithii Boivin is a rare poplar hybrid known from MA (but is likely more common than currently realized). It is recognized by its intermediate leaf blades— the teeth are mostly 0.8–1.7 mm tall and number 16–20 per margin of the blade and the blades measure mostly 46–74 × 37–69 mm.

Fig. 885  Leaf of Populus heterophylla.

5. Populus heterophylla L.

N C Fig. 885

swamp poplar. CT, MA, RI. Swamps, pond shores. 6. Populus nigra L. E black poplar. Populus italica (Du Roi) Moench; P. nigra L. var. italica Du Roi • CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, near dwellings. The most frequent form of this plant seen in New England is the columnar form with erect to strongly ascending branches. 7. Populus tremula L. E European poplar. MA. Waste areas. 8. Populus tremuloides Michx. N Fig. 886 quaking poplar. Populus tremula L. ssp. tremuloides (Michx.) A. & D. Löve; P. tremuloides Michx. var. magnifica Victorin • CT, MA, ME, NH, RI, VT. Forests, woodlands, successional fields, forest borders.

Fig. 886  Leaf of Populus tremuloides.

S a l i c ac e a e   83 3

Salix Salix is a difficult genus that displays tremendous phenotypic plasticity. Though hybridization does occur (and can be occasional between some species pairs), most aberrant forms are the products of envinromental and seasonal variation rather than interspecific crossing. Therefore, not all hybrid combinations that have been reported are included here until further research confirms their identities. Early season leaves usually possess denser indument than later season leaves. Some species often produce red-brown hairs, which, when present, are very useful identifying characters. However, all species that produce such hairs can lack them altogether. The leaves that emerge from the buds are called first leaves (or preformed). As the branchlet continues to grow it produces additional leaves called new leaves (or neoformed). It is important to distinguish between these two types as they can differ morphologically, especially in the prominence of stipules. To assist with identifying features of the branch wood surface (decorticated branches), be sure to remove the bark when specimens are fresh, otherwise the bark adheres to the wood and becomes difficult to remove. Some species of Salix have brittle-based branchlets that break ± cleanly at the junction of annual growth. Other species have flexible-based branchlets that partially break and then must be torn free from the branch. This character (branch brittleness) is very useful for identification and is important to note on herbarium labels. Salix triandra L. was reported from ME by Argus (2010). However, the determination is qualified as “probably,” and the tree was originally determined as S. amygdalina L., a species planted in the former Fay Hyland Arboretum ‌smithiana Willd. from MA (Sorrie and Somers 1999) (University of Maine). Reports of Salix × and ME (Campbell et al. 1995) were based on collections of S. × ‌sericans (see discussion under that nothospecies). References: Argus (1986), Meikle (1984).

Key for carpellate reproductive material 1a. Ovaries and capsules pubescent [Figs. 891, 897] 2a. At least some of the carpellate aments opposite or subopposite . . . . . . . . . S. purpurea 2b. All carpellate aments alternate 3a. Floral bracts of aments pale yellow-brown, pale red-brown, or pale green-yellow, sometimes suffused with red near the apex [Fig. 887] 4a. Ovaries densely white-tomentose, the background color of the ovary nearly or fully concealed by hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. candida 4b. Ovaries moderately sericeous to villous, the background color of the ovary (usually green to light brown) visible through the hairs 5a. Aments precocious [Fig. 894], 15–37 mm long, borne on flowering branchlets 2.5–7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. aurita 5b. Aments coetaneous (sometimes also serotinous in S. exigua) [Fig. 888], 16.5–85 mm long, borne on flowering branchlets 1–65 mm long 6a. Stipes 0.5–0.8 (–1.5) mm long; floral bracts deciduous before maturation of fruit; branchlets of previous year usually glabrous; styles 0–0.2 mm long; aments sometimes branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. exigua 6b. Stipes (2–) 3–6 mm long; floral bracts persistent in fruit; branchlets of previous year usually pubescent; styles 0.1–0.4 mm long; aments never branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. bebbiana 3b. Floral bracts of aments brown or black, at least at the apex [Figs. 892, 897] 7a. Aments precocious (i.e., expanding before the leaves) 8a. Stipe 0.1–0.5 mm long (up to 0.8 mm long in S. planifolia, an alpine and subalpine species); styles 0.5–2 mm long 9a. Plants of alpine and subalpine areas, elevation near or exceeding 1000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. planifolia

Fig. 887  Carpellate flower of Salix ×fragilis (viewed from abaxial surface) showing pale floral bract.

834 tricolpate s

9b. Plants not of alpine areas, elevation less than 1000 m 10a. Branchlets glaucous; adaxial nectaries 0.25–1 mm long, shorter than to longer than the stipe; ovules 10–18 per ovary; native plants of northern New England shorelines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pellita 10b. Branchlets not glaucous; adaxial nectaries 0.9–1.4 mm long, longer than the stipe; ovules 18–19 per ovary; non-native plants escaping from cultivation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. viminalis 8b. Stipe 0.6–2.7 mm long; styles 0–0.5 (–0.6) mm long (up to 1 mm long in S. discolor) 11a. Ovaries and capsules short-beaked or blunt at the apex [Fig. 897]; capsules 2.5–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. sericea 11b. Ovaries and capsules distally tapering to a distinct beak [Fig. 891]; capsules 5–12 mm long 12a. Decorticated wood (i.e., with the bark removed) of 1- to 4-year-old branches with many prominent ridges, some longer than 20 mm [Fig. 890]; capsules 5–7 (–8) mm long 13a. Styles 0.2–0.5 mm long; floral bracts brown to dark brown (at least near the apex) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. cinerea 13b. Styles 0–0.25 mm long; floral bracts usually light brown to medium red-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. aurita 12b. Decorticated wood smooth, with small points, or with a few, short ridges generally shorter than 5 mm; capsules 5–12 mm long 14a. Stigma lobes 0.2–0.56 mm long; aments 9–47 mm long; styles 0.2–0.4 mm long; floral bracts 0.8–2 mm long 15a. Carpellate aments (8–) 15–32 mm long, with floral bracts 1.2–2 mm long; generally upright shrubs 0.3–3 m tall . . . . . . . S. humilis 15b. Carpellate aments 10–18 mm long, with floral bracts 0.8–1.4 mm long; decumbent shrubs 0.3–0.7 m tall . . . . . . . . . . . . . . . S. occidentalis 14b. Stigma lobes 0.4–0.72 mm long; aments (25–) 40–108 mm long; styles 0.3–1 mm long; floral bracts 1.4–4 mm long 16a. Aments (25–) 40–108 mm long; floral scales 1.4–2.5 mm long; styles 0.3–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. discolor 16b. Aments 27–64 mm long; floral scales 2–4 mm long; styles 0.3–0.6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. caprea 7b. Aments coetaneous or serotinous (i.e., expanding with or after the leaves, respectively) 17a. Plants 0.03–0.6 (–1) m tall, of alpine areas, elevation exceeding 1000 m 18a. Aments 11–20.5 mm long; capsules 2–4 mm long; stipes 1–2.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. argyrocarpa 18b. Aments 30–79 mm long; capsules 5–9 mm long; stipes 0.8–1.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. arctophila 17b. Plants 0.3–7 m tall, not of high elevation communities 19a. Ovary pubescent with bent and crimped, woolly hairs . (in part) S. candida 19b. Ovary pubescent with ± straight, silky hairs 20a. Stipes 0.1–0.5 mm long; styles 0.6–1.8 mm long . . . . (in part) S. viminalis 20b. Stipes 0.6–4 mm long; styles 0–0.5 mm long

S a l i c ac e a e   83 5

21a. Stipes 1.5–4 mm long; capsules 5–9 mm long; branches flexible, not breaking cleanly at the junction of yearly growth . . . . . . . . . . S. petiolaris 21b. Stipes 0.6–1.5 mm long; capsules 2.5–4 mm long; branches brittle at the base and breaking cleanly at the junction of yearly growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. sericea 1b. Ovaries and capsules glabrous [Fig. 887] 22a. Floral bracts brown to black (at least at the apex), persisting until after dehiscence of the capsule [Figs. 892, 897] 23a. Dwarf shrubs forming colonies by rhizomes or prostrate stems, rarely exceeding 0.15 m in height, of alpine habitats, elevation exceeding 1000 m; stipes 0.3–1.6 mm long 24a. Styles 0.35–1 mm long; floral bracts 1.1–1.8 mm long; aments 11–47 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. uva‑ursi 24b. Styles 0.2–0.4 mm long; floral bracts 0.5–1 mm long; aments 3.2–13 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. herbacea 23b. Upright to spreading shrubs, colonial or not, 0.2–6 m tall, not of alpine communities; stipes (1–) 1.2–3.4 mm long 25a. Floral bracts 0.8–1.6 mm long; styles 0.3–0.6 mm long; stigmas 0.16–0.28 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. eriocephala 25b. Floral bracts 1.2–3 mm long (mean=1.8 mm); styles (0.3–) 0.5–1.3 mm long; stigmas 0.24–0.56 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. myricoides 22b. Floral bracts pale yellow-brown to green-yellow or white-brown, sometimes suffused with red near the apex [Fig. 887], either persisting until after dehiscence of the capsule or deciduous prior to dehiscence the capsule 26a. Dwarf shrubs 0.01–0.15 m tall, of alpine habitats, elevation exceeding 1000 m; aments 3.2–13 mm long, with 2–11 flowers . . . . . . . . . . . . . . . . . . . . . . . (in part) S. herbacea 26b. Short shrubs to trees, 0.3–25 m tall, not of alpine communities; aments 17–80 mm long, with more than 11 flowers 27a. Aments loosely flowered; flowers ± in whorls about the rachis of the ament 28a. Stipes 0.5–1.5 mm long; branchlets brittle at base, breaking ± cleanly at the junction of yearly growth; expanding leaf blades green on both surfaces . . . . S. nigra 28b. Stipes (1–) 1.6–3.2 mm long; branchlets flexible at base, tearing when bent to failure; expanding leaf blades glaucous on the abaxial surface . . . . S. amygdaloides 27b. Aments densely flowered; flowers spirally arranged about the rachis of the ament 29a. Emerging leaf blades glaucous abaxially or the surfaces ± concealed by thick tomentum of white to gray hairs 30a. Floral scales 0.8–1.6 mm long, abaxially very sparsely pubescent with hairs concentrated near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pedicellaris 30b. Floral bracts (1–) 1.5–4 mm long, abaxially sparsely to densely pubescent throughout 31a. Emerging leaf blades membranaceous, translucent; stipes 1.8–3.5 mm long; capsules 7–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pyrifolia 31b. Emerging leaf blades thick-membranaceous to herbaceous, opaque or nearly so; stipes 0.2–2.4 mm long; capsules 3–5 mm long (7–12 mm long in S. serissima) 32a. Emerging leaf blades densely white- to gray-tomentose on the abaxial surface; floral bracts persisting after dehiscence of the capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. elaeagnos

836  tricolpate s

32b. Emerging leaf blades glabrous or pubescent with straight, silky hairs on the abaxial surface, but then the surface not concealed by the hairs; floral bracts deciduous prior to or during maturation of the capsule 33a. Aments 11–22 mm thick, 1–2.3 times as long as thick; capsules 7–12 mm long; stipes 1.2–2.4 mm long; native shrub to 5 m tall of fens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. serissima 33b. Aments 4–15 mm thick, 3.9–10.5 times as long as thick; capsules 3.5–5 mm long; stipes 0.2–1.5 mm long; non-native trees to 25 m tall of various habitats, though frequently naturalized in riparian and lakeside forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. alba 29b. Emerging leaf blades without bloom abaxially (though paler than the adaxial surface), generally pubescent, but the pubescence not dense enough to conceal the surface 34a. Styles 0–0.2 mm long; aments sometimes branched; expanding leaf blades remotely denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. exigua 34b. Styles 0.3–0.8 mm long; aments unbranched; expanding leaves closely serrulate 35a. Floral bracts moderately to densely pubescent throughout the abaxial surface; stipes 1.2–2.4 mm long . . . . . . . . . . . . . . . . . . . . . . (in part) S. serissima 35b. Floral bracts sparsely pubescent, the hairs mainly near the apex; stipes 0.5–1.6 (–2) mm long 36a. Adaxial nectaries 0.2–0.45 mm long, ⅕ to ⅓ as long as the stipe; abaxial nectaries absent; emerging leaves sometimes with red-brown hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lucida 36b. Adaxial nectaries 0.4–0.8 mm long, ½ to fully as long as the stipe; abaxial nectaries present or absent; emerging leaves never with redbrown hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pentandra

Key for s taminate repro ductive material 1a. Flowers with an abaxial nectary (or nectaries) in addition to the adaxial nectary (or nectaries) 2a. Flowers each with 3–10 stamens 3a. Rachis of anther sac curved; floral bracts entire at apex (sometimes toothed in S. amygdaloides; aments loosely flowered; flowers ± in whorls about the rachis of the ament 4a. Branchlets brittle at base, breaking ± cleanly at the junction of yearly growth; expanding leaf blades green on both surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nigra 4b. Branchlets flexible at base, tearing when bent to failure; expanding leaf blades glaucous on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. amygdaloides 3b. Rachis of anther sac straight; floral bracts entire, toothed, or erose at apex; aments densely flowered; flowers spirally arranged about the rachis of the ament 5a. Emerging leaves with stipules and often with red-brown hairs . . . . . . . . . S. lucida 5b. Emerging leaves exstipulate (sometimes with minute stipule rudiments) and without red-brown hairs 6a. Floral bracts sparsely pubescent, the hairs mainly near the apex; aments 27–81 mm long; emerging leaves with abaxially green blades; non-native trees 5–15 mm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pentandra 6b. Floral bracts moderately to densely pubescent throughout abaxial surface; aments 11–31 (–53) mm long; emerging leaves with abaxially glaucous blades (this sometimes difficult to detect on early emerging blades); native shrubs 1–5 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. serissima 2b. Flowers each with 1 or 2 stamens [Fig. 892]

S a l i c ac e a e   837

7a. Flowers each with 1 stamen, or appearing as 1 due to fusion of the filaments 8a. At least some of the aments opposite or subopposite; flowers with 2 stamens but with filaments connate their entire length, bearing 2 anthers at the apex; upright shrubs 1.5–5 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. purpurea 8b. Aments all alternate; flowers with 1 stamen (i.e., only 1 anther at the summit of the filament); dwarf shrubs rarely exceeding 0.15 m . . . . . . . . . . . . . . . . . . . . . S. uva‑ursi 7b. Flowers each with 2 stamens [Fig. 892] 9a. Aments 3–7.5 × 1.5–5 mm, with 5–12 flowers; flowering branchlets 0.2–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. herbacea 9b. Aments 10–61 × 4–16 mm, with more than 12 flowers; flowering branchlets (1–) 2–20 mm long 10a. Floral bracts brown to black, at least near the apex [Figs. 892, 897]; filaments glabrous throughout; aments 10–21.5 mm long . . . . . . . . . . . . . . . . . . S. argyrocarpa 10b. Floral bracts yellow-brown to green-yellow or white-brown, sometimes suffused with red near the apex [Fig. 887]; filaments pilose, at least in the basal portion; aments 20–61 mm long 11a. Petioles of emerging leaves eglandular; filaments densely pilose in the basal half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. exigua 11b. Petioles of emerging leaves glandular at the summit; filaments sparsely pilose only at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. alba 1b. Flowers with only an adaxial nectary 12a. Filaments pubescent (though sometimes sparsely), at least near the base 13a. Floral bracts yellow-brown or pale green-yellow, often suffused with red at the apex [Fig. 887] 14a. Filaments basally connate for up to slightly more than ½ their length 15a. Aments borne on flowering branchlets 1–1.5 mm long; expanding leaves mostly linear to narrow-oblong or narrow-oblanceolate . . . . . . . . . . . S. elaeagnos 15b. Aments borne on flowering branchlets 0.5–11 mm long; expanding leaves mostly elliptic or oblong to oblanceolate or obovate . . . . . . . . (in part) S. bebbiana 14b. Filaments distinct 16a. Decorticated wood smooth, with small points or with a few short ridges generally shorter than 5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. bebbiana 16b. Decorticated wood (i.e., with the bark removed) of 1- to 4-year-old branches with many prominent ridges, some longer than 20 mm . . . . . . . . . (in part) S. aurita 13b. Floral bracts brown to black, at least near the apex [Figs. 892, 897] 17a. Decorticated wood (i.e., with the bark removed) of 1- to 4-year-old branches with many prominent ridges, some longer than 20 mm [Fig. 890] 18a. Aments 15.5–21.5 × 10–15 mm; floral bracts 1–2.2 mm long; anthers 0.5–0.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. aurita 18b. Aments 26–39 × 12–26 mm; floral bracts 2–3 mm long; anthers 0.7–1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. cinerea 17b. Decorticated wood smooth, with small points, or with a few short ridges generally shorter than 5 mm 19a. Aments coetaneous (i.e., expanding with the leaves) [Fig. 888] 20a. Branchlets brittle at base, breaking ± cleanly at the junction of yearly growth; expanding leaves without red-brown hairs; aments 13.5–40 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sericea

838  tricolpate s

20b. Branchlets flexible at base, tearing when bent to failure; expanding leaves often with red-brown hairs; aments 12–29 mm long . . . . . . . . . . . . . S. petiolaris 19b. Aments precocious (i.e., expanding before the leaves) [Fig. 894] 21a. Floral bracts 2–4 mm long; anthers yellow; small trees 8–15 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. caprea 21b. Floral bracts 0.8–2.5 mm long; anthers yellow or purple initially, later turning yellow; dwarf shrubs to tall shrubs 0.3–4 (–8) m tall 22a. Anthers 0.5–1 mm long; aments 23–52 mm long; adaxial nectaries 0.6–1.1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. discolor 22b. Anthers 0.4–0.6 mm long; aments (5–) 6–20 (–34) mm long; adaxial nectaries 0.2–0.7 mm long 23a. Staminate aments (10–) 14–20 (–34) mm long, with floral bracts 1.4–1.6 mm long; generally upright shrubs 0.3–3 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. humilis 23b. Staminate aments (5–) 6–11 (–13) mm long, with floral bracts 0.8–1.6 mm long; decumbent shrubs 0.3–0.7 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. occidentalis 12b. Filaments glabrous throughout 24a. Aments precocious (i.e., expanding before the leaves) [Fig. 892] 25a. Plants of subalpine and alpine communities, elevation usually exceeding 900 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. planifolia 25b. Plants not of alpine areas, elevation less than 900 m 26a. Branchlets prominently glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pellita 26b. Branchlets not glaucous 27a. Anthers 0.6–0.8 mm long; adaxial nectaries 0.6–1.5 mm long; second year branches usually yellow to yellow-brown or gray-brown . . . . (in part) S. viminalis 27b. Anthers 0.4–0.6 mm long; adaxial nectaries 0.2–0.7 mm long; second year branches usually red-brown to dark red-brown 28a. Staminate aments (10–) 14–20 (–34) mm long, with floral bracts 1.4– 1.6 mm long; generally upright shrubs 0.3–3 m tall . . . . . . . . . (in part) S. humilis 28b. Staminate aments (5–) 6–11 (–13) mm long, with floral bracts 0.8–1.6 mm long; decumbent shrubs 0.3–0.7 m tall . . . . . . . . . . . . (in part) S. occidentalis 24b. Aments coaetaneous or serotinous (i.e., expanding with or after the leaves, respectively) [Fig. 888] 29a. Floral bracts yellow-brown or pale green-yellow, often suffused with red at the apex [Fig. 887] 30a. Second-year branches usually pubescent, dull to sublustrous; aments subglobose to short-obloid, 10–42 mm long . . . . . . . . . . . . . . . . (in part) S. bebbiana 30b. Second-year branches glabrous and lustrous; aments short-obloid to cylindric, 18.5–63 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pyrifolia 29b. Floral bracts brown to black, at least near the apex [Figs. 892, 897] 31a. Dwarf, colonial plants 0.03–0.15 m tall, of alpine communities, elevation exceeding 1000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. arctophila 31b. Short shrubs to small trees 0.3–7 m tall, not of alpine communities, of lower elevations

S a l i c ac e a e   83 9

32a. Branchlets and often the second-year branches conspicuously pubescent with white tomentum (i.e., the hairs crimped and tangled; glabrescent in rare forms) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. candida 32b. Branchlets and branches glabrous or pubescent with pilose, villous, or veluntinous hairs (i.e., the hairs not crimped and tangled) 33a. Expanding leaf blades from vegetative buds with serrulate margins (rarely entire in S. eriocephala); first leaves with foliaceous stipules (sometimes minute in S. myricoides) 34a. Floral bracts 0.8–1.5 (–1.6) mm long, pubescent with wavy hairs; anthers 0.4–0.64 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. eriocephala 34b. Floral bracts (1.2–) 1.5–3 mm long, pubescent with straight hairs; anthers 0.52–0.76 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. myricoides 33b. Expanding leaf blades with entire margins; first leaves exstipulate or with minute stipules 35a. Floral bracts 0.8–1.6 mm long, pubescent abaxially mainly near the apex; aments 11–21 mm long; anthers 0.4–0.6 mm long . . . S. pedicellaris 35b. Floral bracts 1.6–2.2 mm long, pubescent abaxially throughout the surface; aments 24–48 mm long; anthers 0.6–0.8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. viminalis

Key for mature vegetative material 1a. Leaf blades at least somewhat grayed or whitened on the abaxial surface by hairs or bloom (note: in some species the bloom is very thin) 2a. Leaf blade margins entire, undulate, or crenate, the teeth (when present) rounded at the apex and often somewhat irregular from leaf to leaf [Figs. 889, 896] 3a. Dwarf, trailing, colonial shrubs not exceeding 0.2 m in height, of alpine habitats exceeding 1000 m elevation 4a. Leaf blades 0.5–2 (–2.5) cm long, 1.7–3.6 times as long as wide; petioles 2–4 (–6) mm long; winter buds 1–3 (–4) mm long . . . . . . . . . . . . . . . (in part) S. uva‑ursi 4b. Leaf blades 2–4 cm long, mostly 1.2–2 times as long as wide; petioles mostly 5–12 mm long; winter buds 3–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. arctophila 3b. Upright or low-spreading shrubs or small trees (0.3–) 1–15 m tall, of various habitats 5a. At least some leaves on a given plant opposite or subopposite . . . . . S. purpurea 5b. Leaves alternate throughout 6a. Fresh leaf blades with revolute margins 7a. Leaf blades very soon glabrous, with strictly entire margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. pedicellaris 7b. Leaf blades pubescent on one or both surfaces, though sometimes becoming glabrate in late season, with entire to crenate margins 8a. Leaf blades 1.5–3 times as long as wide 9a. Decorticated wood (i.e., with the bark removed) of 1- to 4-year-old branches with many prominent ridges, some longer than 20 mm 10a. Shrubs 1–2.5 m tall; stipules foliaceous, persistent . . . . . . S. aurita 10b. Shrubs or small trees 2–12 m tall; stipules small and caducous to foliaceous and persistent . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. cinerea

8 40  tricolpate s

9b. Decorticated wood smooth, with small points, or with a few short ridges generally shorter than 10 mm 11a. Leaf blades 2.5–8 cm wide, 2–3 times as long as wide, pubescent with only white hairs; stems 8–15 m tall . . . . . . . . . . . . . . . . . . . S. caprea 11b. Leaf blades 0.3–2.3 (–3.4) cm wide, 2.3–9 times as long as wide, often with some red-brown hairs intermixed with the white hairs; stems 0.3–3 m tall 12a. Leaf blades 3–10 × 1–2.5 cm, stipulate (at least on later leaves); petioles 3–7 mm long; branchlets 2–5 mm thick; generally upright shrubs 0.3–3 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. humilis 12b. Leaf blades 2–5 × 0.3–1.2 cm, exstipulate; petioles 0.5–3 mm long; branchlets usually 2 mm thick or thinner; decumbent shrubs 0.3–0.7 m tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. occidentalis 8b. Leaf blades 3–40 times as long as wide 13a. Leaf blades elongate-linear, 1–8 mm wide, mostly 17–40 times as long as wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. elaeagnos 13b. Leaf blades narrow-lanceolate or narrow-oblong to oblanceolate or obovate, 7–30 mm wide, mostly 3–17 times as long as wide 14a. Leaf blades 2–5 cm long; dwarf shrubs up to 0.7 m tall; stipules absent 15a. Unfolding leaves densely sericeous with lustrous white hairs; red-brown hairs absent; alpine shrubs of elevations exceeding 1000 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. argyrocarpa 15b. Unfolding leaves pubescent with dull hairs that are bent, curled, and tangled; red-brown hairs frequently present on leaves, winter buds, and/or branchlets; shrubs of dry, well-drained soils of lower elevations . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. humilis 14b. Leaf blades 4–17 cm long; shrubs or small trees to 5 m tall; stipules commonly present 16a. Unfolding leaves pubescent with straight, neatly aligned, and lustrous hairs; stipules usually lacking (sometimes present in S. viminalis and then linear to narrow-lanceolate, mostly more than 7 times as long as wide) 17a. Branchlets brittle at base, breaking ± cleanly at the junction of yearly growth, becoming glaucous later in season; native plants of north-temperate and boreal shores . . . . . . . . . . S. pellita 17b. Branchlets flexible at base, tearing when bent to failure, not glaucous; introduced plants more common near civilization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. viminalis 16b. Unfolding leaves pubescent with bent, curled, and tangled hairs that are gray to white, but not lustrous; stipules usually present, lanceolate to narrow-ovate, mostly less than 7 times as long as wide 18a. Decorticated wood (i.e., with the bark removed) of 1–4 year old branches with many prominent ridges, some longer than 20 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. cinerea 18b. Decorticated wood smooth, with small points, or with a few, short ridges generally less than 10 mm in length

S a l i c ac e a e   8 41

19a. Branchlets pubescent largely with gray or brown villosetomentose hairs; red-brown hairs sometimes present on leaves, winter buds, and/or branchlets; plants mostly of dry, well-drained soils . . . . . . . . . . . . . . . . . . . . . . . (in part) S. humilis 19b. Branchlets usually pubescent with conspicuous white, yellow-white, or gray-white tomentum; red-brown hairs absent; plants of high-pH fens and swamps . . . . . . S. candida 6b. Fresh leaf blades plane, the margins not rolled under 20a. Branchlets frequently becoming glaucous later in season; plants of alpine habitats exceeding 1000 m elevation . . . . . . . . . . . . . . . . . . . . . . . . . . . S. planifolia 20b. Branchlets glabrous to pubescent, but not glaucous; plants of lower elevation 21a. Winter branchlets red, glabrous, and lustrous; leaf blades glabrous, thin and somewhat translucent, especially in early season, rounded to cordate at the base, persistently fragrant in drying for many years . . . (in part) S. pyrifolia 21b. Winter branchlets either pubescent or of color other than red; leaf blades glabrous to pubescent, membranaceous to coriaceous, cuneate at the base (rounded to rarely cordate in S. myricoides), not fragrant in drying 22a. Leaf blades strictly entire [Fig. 896], very soon glabrous; branchlets glabrous; petioles 2–7 mm long; low shrubs up to 1.5 m, of organic soil wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. pedicellaris 22b. Leaf blades usually with some form of dentition, pubescence, or both; branchlets often pubescent, minimally retaining patches of hairs near the nodes in most species; petioles 3–17 mm long; shrubs or small trees to 5 m, of various habitats 23a. Leaf blades coriaceous, prominently glaucous on the abaxial surface, tending to be more regularly toothed, the teeth numbering 5–9 per cm; leaves broad-cuneate to rounded or less frequently subcordate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. myricoides 23b. Leaf blades membranaceous to herbaceous, obscurely to prominently glaucous on the abaxial surface, tending to have fewer coarser, less regular teeth numbering 1–3 (–5) per cm; leaves narrowto broad-cuneate at the base 24a. Leaf blades usually impressed-veiny above and rugoseveiny below [Fig. 889]; branchlets persistently pubescent, even on second-year branches; herbage lacking red-brown hairs; winter branchlets and buds red to red-brown . . . . . . . . . . . . . . . . S. bebbiana 24b. Leaf blades not notably veiny; branchlets commonly glabrate with small patches of hairs near the nodes, the second-year branches usually glabrous; herbage frequently with red-brown hairs, these often persisting along the midrib on the adaxial surface; winter branchlets brown to yellow-brown; winter buds yellow to yellow-brown or red-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . S. discolor 2b. Leaf blade margins denticulate to serrulate, the teeth sharply pointed at the apex and, in most species, ± uniform through the margin [Figs. 893, 895] 25a. Trees to 25 m tall; apex of petiole with one or more dark glands, these often ± paired on opposite sides of the petiole (glands often absent in S. amygdaloides) [Fig. 895] 26a. Winter buds with a sharply pointed apex, the margins distinct and overlapping; leaf blades somewhat abruptly tapering to a long, thin apex; branchlets flexible at base, tearing when bent to failure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. amygdaloides

8 42   tricolpate s

26b. Winter buds blunt at apex, the margins connate; leaf blades gradually tapering to the apex; branchlets brittle at base, breaking ± cleanly at the junction of yearly growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. alba 25b. Shrubs to 4 m tall; apex of petiole lacking a pair of dark glands (present in S. serrisima) [Fig. 893] 27a. Winter branchlets red, glabrous, and lustrous; leaves thin and somewhat translucent, especially in early season, persistently balsam-fragrant in drying for many years . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. pyrifolia 27b. Winter branchlets usually yellow-brown, red-brown, green-brown, or brown, varying from glabrous to pubescent, dull to sublustrous (lustrous in S. serissima); leaves thick-herbaceous to coriaceous, not fragrant in drying 28a. Leaves very soon glabrous, lustrous on the adaxial surface; petioles with one or more dark glands, these often ± paired on opposite sides of the petiole; plants restricted to high-pH wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. serissima 28b. Leaves and branchlets pubescent when young and often remaining so until at least mid-season (or later); petioles lacking apical glands; plants of various habitats 29a. Leaf blades cuneate at base, sometimes rounded in forms of S. sericea, with neatly aligned and appressed hairs (i.e., sericeous) at least when young, frequently darkening in drying; stipules absent or up to 4 mm long 30a. Leaf blades 2.5–7 × 0.3–2 cm, sometimes nearly entire at the base, frequently with red-brown hairs intermixed with the white hairs; branches flexible, not breaking cleanly at the junction of yearly growth, commonly glabrate, sometimes glaucous; stipules usually wanting . . . . . . . S. petiolaris 30b. Leaf blades 4–15 × 1–4 cm, serrulate to the base, lacking red-brown hairs; branches brittle at the base and breaking cleanly at the junction of yearly growth, usually puberulent, at least remaining so near the nodes, not glaucous; stipules often present, especially on vigorous shoots . . . . S. sericea 29b. Leaf blades rounded to cordate at the base (rarely broad-cuneate) [Fig. 893], with short, ascending and curving hairs (i.e., villous) at least when young, not prominently darkening in drying; stipules usually present and longer than 4 mm 31a. Leaf blades lanceolate to more commonly narrow-ovate, ovate-oval, or obovate, with serrulate to crenate-serrate margins, prominently glaucous on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. myricoides 31b. Leaf blades lanceolate to oblong-lanceolate, with serrulate margins [Fig. 893], green to thinly glaucous on the abaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. eriocephala 1b. Leaf blades green beneath, sometimes paler than the adaxial surface but without bloom or dense hairs at maturity 32a. Tree to 20 m tall; winter bud, sharply pointed at apex, with distinct and overlapping margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nigra 32b. Dwarf shrubs to small trees to 4 m tall; winter buds blunt, with connate margins 33a. Leaf blades 0.5–3 cm long; dwarf shrubs to 15 cm tall of alpine habitats exceeding 1000 m elevation 34a. Leaf blades 1–2.5 cm wide, oval to suborbicular, rounded to retuse at the apex, cordate to truncate at the base; plants commonly found in hydric, mossy areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. herbacea 34b. Leaf blades 0.2–1 (–1.7) cm wide, narrow-elliptic to obovate, acute to broadobtuse at the apex, cuneate at the base; plants commonly found in gravels, cracks of bedrock, and folists . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. uva‑ursi 33b. Leaf blades (2–) 3.5–17 cm long; shrubs or small trees much taller than 5 cm, primarily of low elevation habitats

S a l i c ac e a e   8 43

35a. Leaves elliptic to oblanceolate or obovate, usually widest above the middle [Fig. 896], sometimes revolute; low shrubs up to 15 dm tall, of organic soil wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. pedicellaris 35b. Leaves linear to narrow-ovate, widest below the middle, not revolute; shrubs of various sizes and habitats 36a. Petioles 1–5 mm long; leaves amphistomatous (i.e., stomates present on both surfaces), somewhat remotely toothed, those teeth near the middle of blade margin often further than 3 mm apart . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. exigua 36b. Petioles (3–) 5–16 mm long; leaves hypostomatous (i.e., stomates lacking on adaxial surface), finely and closely toothed, those teeth near the middle of the blade margin closer than 3 mm 37a. Apex of petiole lacking dark glands [Fig. 893]; stipules mostly 4–10 mm long; branchlets usually flexible at base and tearing when bent to failure, infrequently brittle at base . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. eriocephala 37b. Apex of petiole with one or more dark glands, these often ± paired on opposite sides of the petiole [Fig. 895]; stipules absent or up to 6 mm long; branchlets brittle at base, breaking ± cleanly at the junction of yearly growth 38a. Leaf blades 5–17 cm long, frequently with red-brown hairs when unfolding, tapering to a long-acuminate to caudate apex on mature blades [Fig. 895]; stipules usually present, 1–6 mm long . . . . . . . . . . . . . . . . . . S. lucida 38b. Leaf blades 3.5–10 cm long, lacking red-brown hairs, tapering to an acuminate apex; stipules wanting . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pentandra 1. Salix alba L. E white willow. Salix alba L. ssp. caerulea (Sm.) Rechinger f.; S. alba L. var. caerulea (Sm.) Sm.; S. alba L. var. calva G.F.W. Mey.; S. alba L. ssp. vitellina (L.) Arcang.; S. alba var. vitellina (L.) Stokes; S. vitellina L.; • CT, MA, ME, NH, RI, VT. Riparian forests, river banks, roadsides, field edges, wetland margins, near dwellings. There are four forms of this willow that occur in New England. The “vitellina” form has bright yellow to bright orange-yellow winter branchlets and floral bracts commonly 3–3.5 mm long. The “caerulea” form has brown to green-brown winter branchlets, floral bracts 2.5–3 mm long, mature leaf blades often longer than 10 cm, 1.5–2 cm wide, and usually becomes glabrate on the abaxial surface. The “alba” form has branchlets and floral bracts similar to the preceeding, but has leaf blades 5–10 (–12) cm long, 0.5–1.5 cm wide, and often remain long-sericeous on the abaxial surface. A fourth form, which may be referable to “micans” of Europe, is similar to the “alba” form but has the leaf blades densely and persistently long-sericeous with white hairs. These forms are regarded as valid infraspecific taxa in Europe (where they occur as natives; e.g., Meikle 1984) but are treated as cultivars by some North American authors (e.g., Argus 1986). Until the appropriate rank to treat these taxa can be determined, they are here recognized informally. ‌ehrhartiana by various New ‌1 × 16. Salix ×jesupii Fern. has been incorrectly called S. × England authors (but that hybrid has S. alba and S. pentandra as parents and is not known to occur in New England). Salix × ‌jesupii is known from CT, MA, NH, VT. This hybrid is a tall shrub or small tree 4–8 m tall with abaxially glaucous leaf blades (rather than green and not glaucous in S. lucida) that show a mixture of white and red-brown hairs. The blades are mostly 2.8–4.8 times as long as wide (vs. mostly 4.2–7.3 times in S. alba). The staminate flowers have 3 or 4 stamens (rather than 2 in S. alba). 1‌ × Salix babylonica L. Salix ×sepulcralis Semonkai is primarily responsible for reports of S. babylonica in New England, which is an Asian species that is not hardy in the northeastern United States. Salix × ‌sepulcralis is known from CT, MA, ME, NH, VT. It is recognized by its conspicuously pendulous branches and branchlets and gold-yellow (or less frequently green-yellow) branchlets. It is very similar to another S. babylonica ‌pendulina; hybrid in New England called S. × however, that nothotaxon has green-brown (or less frequently yellow-brown, gray-brown, or red-brown) branchlets.

8 44   tricolpate s

1‌ × Salix euxina I.V. Belyaeva. Salix ×fragilis L. [Fig. 887] is a frequent willow hybrid known from CT, MA, ME, NH, RI, VT. It resembles S. alba but has sparsely pubescent to glabrous leaf blades, coarser, sometimes slightly irregular, teeth, and larger, asymmetrical stipules. From S. euxina it differs in its amphistomatous, usually abaxially sparsely pubescent leaves and relatively loosely flowered aments (leaves both glabrous and hypostomatous and aments relatively densely flowered in S. euxina). This hybrid, which is fully naturalized in riparian and lacustrine floodplains, is responsible for reports of Salix euxina in New England (note: S. euxina was formerly referred to as S. fragilis, and this hybrid was formerly referred to as S. × ‌rubens Schrank; Belyaeva 2009). The nothovarietal epithet “basfordiana” applies to our material; however, the appropriate combination has not yet been made. 2. Salix amygdaloides Anderss.

NC

peach-leaved willow. VT; northern portion of state. Lacustrine floodplain forests, swamp edges. 3. Salix arctophila Cockerell ex Heller

NC

northern willow. Salix arctica Liebm.; S. arctophila Cockerell ex Heller var. lejocarpa (Anderss.) Schneid.; S. groenlandica Lundstr.; S. groenlandica Lundstr. var. lejocarpa (Anderss.) Lange • ME. Alpine ravines and slides. Two staminate individuals are known to occur in a remote basin on Katahdin, Piscataquis County, ME. 4. Salix argyrocarpa Anderss.

NC

Labrador willow. Salix argyrocarpa Anderss. var. denudata Anderss. • ME, NH; northern portion of states. Alpine ravines, plateaus, and tarn shores. 4 ‌ × 24. Salix ×grayii Schneid. is a very rare willow hybrid known only from Mount Washington, Coos County, NH, in New England. It differs from S. argyrocarpa in being more robust, frequently with glaucous branchlets (later in season), and broader leaf blades that have sparser pubescence. From S. planifolia the hybrid differs in its revolute leaf blades and red to purple styles (those of S. planifolia are ± yellow). 5. Salix aurita L. E

Fig. 888  Carpellate ament of Salix bebbiana, each flower subtended by a pale floral bract.

eared willow. MA. Fields, roadsides, forest fragments and borders, waste areas. Specimens of Salix aurita from New England are questionable and appear to be S. cinerea. More work is needed to confirm if this species truly occurs in New England. 6. Salix bebbiana Sarg. N Fig. 888, 889 long-beaked willow. Salix bebbiana Sarg. var. capreifolia (Fern.) Fern.; S. bebbiana Sarg. var. projecta (Fern.) Schneid.; S. rostrata Richards.; S. rostrata Richards. var. capreifolia Fern.; S. rostrata Richards. var. projecta Fern.; S. starkeana Willd. ssp. bebbiana (Sarg.) Youngberg; S. vagans Hook. f. ex Anderss. var. occidentalis Anderss.; S. vagans Hook. f. ex Anderss. var. rostrata (Richards.) Anderss. • CT, MA, ME, NH, RI, VT; nearly throughout. Fields, roadsides, waste areas, shorelines, swamps, stream banks. ‌6 × 7. Salix ×cryptodonta Fern. is a very rare willow hybrid known from MA. It suggests S. candida in its tomentose branchlets and leaf blades and its leaves with revolute blades and prominent stipules, but it has blades 30–50 mm long (vs. 47–103 mm) with relatively more prominent teeth and more prominent rugose texture, floral bracts ca. 3 mm long (vs. 1.2–1.8 mm), and capsules 8–10 mm long (vs. 4–6 mm). 6 ‌ × 10. Salix ×beschelii Boivin is a rare willow hybrid known from MA. The plants generally resemble S. discolor, but the leaf blades have thicker texture with weak rugose veins abaxially, the floral bracts are red-brown (rather than nearly black at the apex), and the ovary stipes are 2.5–4.5 mm long (rather than 2–2.7 mm long).

Fig. 889  Crenate leaf blade of Salix bebbiana.

6 × 23. This rare willow hybrid is known from ME. It has the general appearance of Salix petiolaris, but the leaf blades are slightly wider relative to length, have somewhat irregular toothing (as to spacing of the teeth) with slightly coarser teeth, a somewhat rugose nature, and red-brown hairs (absent in S. bebbiana).

S a l i c ac e a e   8 45

7. Salix candida Fluegge ex Willd. N sage willow. Salix candida Fluegge ex Willd. var. denudata Anderss.; S. candida Fluegge ex Willd. var. tomentosa Anderss. • CT, MA, ME, VT. Fens and swamps in regions of high-pH bedrock and/or till. 7 ‌ × 12. Salix ×rubella Bebb is a very rare willow hybrid in New England that is known from MA. It has narrow-elliptic leaf blades with small, sharp, forward-pointing teeth and weakly revolute margins, sparsely pubescent ovaries, and two-toned floral bracts (light brown at the base and darker brown near the apex). 7 × 23. This very rare willow hybrid is known from MA. It has narrow-elliptic leaf blades that are acute at apex, sparsely pubescent after expansion, with small, forward-pointing teeth, and weakly revolute margins. The ovaries are mostly moderately villoustomentose at maturity, the background color showing through pubescence (background color obscured in Salix candida). The capsules are blunt, on stipes 0.9-2 mm, and subtended by light red-brown floral bracts. 8. Salix caprea L. E goat willow. CT. Roadsides, forest fragments, near dwellings.

Fig. 890  Elongate ridges on decorticated branches of Salix cinerea.

‌8 × 30. Salix ×sericans Tausch ex Kern. is a rare willow hybrid in New England known from ‌smithiana MA, ME. It is sometimes referred to by the name S. × Willd. (9 × 30), but that name properly refers to the hybrid between S. cinerea and S. viminalis (a hybrid that has not been documented in New England). It has lanceolate to broad-lanceolate leaf blades that are mostly 13–30 mm wide (vs. oblong to suborbicular and 25–80 mm wide in S. caprea and narrow-lanceolate to narrow-elliptic and 5–15 (–25) mm wide in S. viminalis). This hybrid is further characterized by dense gray tomentum on the abaxial blade surface, precocious flowering, and dark floral bracts that are densely pubescent. 9. Salix cinerea L. E Fig. 890 gray willow. 9b. Salix atrocinerea Brot.; S. cinerea L. ssp. atrocinerea (Brot.) Guinier • CT, MA, ME, RI; most frequent along the coastal plain. Fields, roadsides, waste areas, pond shores. Two subspecies are recognized (rather than two species) because most of the characters used in various European floras do not hold on the North American continent. 1a. Mature leaf blades pubescent with gray hairs on the abaxial surface (i.e., red-brown hairs absent), usually gray-green and dull on the adaxial surface; branchlets ± gray at maturity and usually persistently pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9a. S. cinerea ssp. cinerea

Fig. 891  Carpellate flower of Salix discolor (viewed from adaxial surface).

1b. Mature leaf blades pubescent with both gray and red-brown hairs on the abaxial surface, usually dark green and sometimes sublustrous on the adaxial surface; branchlets red to redbrown at maturity and usually becoming glabrate later in season . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9b. S. cinerea ssp. oleifolia (Sm.) Macreight Subspecies cinerea is known from MA, RI. Subspecies oleifolia is known from CT, MA, ME, RI. 10. Salix discolor Muhl. N Fig. 891, 892 pussy willow. Salix ancorifera Fern. • CT, MA, ME, NH, RI, VT; nearly throughout. Fields, roadsides, waste areas, swamps, wetland margins, shorelines. 10 × 12. This uncommon willow hybrid is known from MA, ME. It resembles S. discolor except that the leaf blades are ± elliptic (vs. elliptic to oblanceolate or obovate), and the marginal teeth are more numerous and relatively sharper. The ovaries show sparse pubescence during flower (vs. densely pubescent in S. discolor and glabrous in S. eriocephala). ‌10 × 15. Salix ×conifera Wangenh. is an uncommon willow hybrid known from MA, ME, NH, RI, VT. It mostly closely resembles S. humilis in the thicker, reticulate-veiny, and abaxially tomentose leaf blades and pubescent branchlets but has more elongate carpellate aments, longer styles, and longer petioles that are intermediate between the parental species (27–55 mm long, 0.3–0.5 mm long, and 5–16 mm long, respectively, in the hybrid vs. 8–32 mm long, 0.2–0.4 mm long, and mostly 3–7 mm, respectively, in S. humilis).

Fig. 892  Staminate flower of Salix discolor (viewed from abaxial surface) showing darktipped floral bract.

846  tricolpates

11. Salix elaeagnos Scop. E hoary willow. Salix incana Schrank • CT, MA, ME. Roadsides, river banks, areas of cultivation. 12. Salix eriocephala Michx. ssp. eriocephala var. eriocephala N Fig. 893 heart-leaved willow. Salix cordata Muhl.; S. cordata Michx. var. abrasa Fern.; S. cordata Michx. var. rigida (Muhl.) Boivin; S. cordata Muhl. var. ridiga (Muhl.) Carey; S. cordata Muhl. var. missouriensis (Bebb) Mackenzie & Bush; S. discolor Muhl. var. eriocephala (Michx.) Anderss.; S. missouriensis Bebb; S. rigida Muhl. • CT, MA, ME, NH, RI, VT; throughout. Fields, roadsides, shorelines, ditches, swamps. Reports of Salix cordata Michx. from New England (Fernald 1950b, Seymour 1982) were all based on specimens of Salix eriocephala. Many of the specimens in question were vigorous sprouts on northern, ice-scoured river shores where the leaf blade morphology was atypical. 12 × 23. This uncommon willow hybrid is known from MA, ME, and VT (and very likely found in other states as well). It is intermediate between the two species in terms of leaf blade outline and size and frequently shows small stipules at the nodes. The ovaries are usually sparsely pubescent but are sometimes nearly glabrous except for patches of hair near the base and apex. Fig. 893  Serrulate leaf blade of Salix eriocephala.

‌12 × 27. Salix ×bebbii Gandog. is an uncommon hybrid known from CT, MA, ME, NH, VT. Though it is generally intermediate between the two parental species in various morphological characters (e.g., leaf blade outline, leaf blade pubescence, prominence of stipules, pubescence on branchlets), it is morphologically variable, and extremes approach either parent. One of the most diagnostic characters is the pubescence on the ovary, which is sparse in this hybrid (though often absent near the base; absent altogether in S. eriocephala and dense throughout during flower in S. sericea). Most reports of S. myricoides from New England other than Aroostook County, ME, are based on this hybrid (i.e., the name Salix myricoides was misapplied to this hybrid by many authors, including Fernald 1950b and Magee and Ahles 1999). 13. Salix exigua Nutt. ssp. interior (Rowlee) Cronq. NC sandbar willow. Salix exigua Nutt. var. exterior (Fern.) C.F. Reed; S. exigua Nutt. var. pedicellata (Anderss.) Cronq.; S. exigua Nutt. var. sericans (Nees) Nesom; S. fluviatilis Nutt. var. sericans (Nees) Boivin; S. interior Rowlee; S. interior Rowlee var. exterior Fern.; S. interior Rowlee var. pedicellata (Anderss.) C. R. Ball; S. interior Rowlee var. wheeleri Rowlee; S. longifolia Muhl.; S. longifolia Muhl. var. interior (Rowlee) M.E. Jones; S. longifolia Muhl. var. sericans Nees; S. longifolia Muhl. var. wheeleri (Rowlee) C. K. Schneider; S. wheeleri (Rowlee) Rydb. • CT, MA, ME, NH, VT. Primarily along sand, gravel, and cobble shorelines of major rivers, less frequently on lake shores, rarely in borrow pits. Some recent occurrences along impounded lake shores and in borrow pits of ME and VT appear to be introductions. Most forms of this species in New England have elongate, linear to narrow-lanceolate leaf blades 60–160 × 4–11 mm. A rare form, found only on the Aroostook River in ME, has oblong to oblong-elliptic blades mostly 27–63 × 9–15 mm (named var. exterior). It is a distinctive morphology, but apparent intermediates occur in Quebec (specimens at CAN; images seen!). 14. Salix herbacea L. NC snow-bed willow. ME, NH. Alpine ravines and gullies, often in wet moss at the heads of gullies or at the base of cliffs (i.e., in areas where snow accumulates). ‌ × 29. Salix ×peasei Fern. is a very rare willow hybrid known from NH. It shows 14 abaxially glaucous leaf blades (similar to S. uva-ursi) but shows some rounded to slightly retuse tips (closer to S. herbacea, S. uva-ursi is usually acute at the apex). The floral bracts are sparsely to moderately pubescent on the abaxial surface (glabrous or rarely very sparsely pubescent in S. herbacea, densely pubescent in S. uva-ursi).

Fig. 894  Precocious aments of Salix humilis.

15. Salix humilis Marsh. N Fig. 894 prairie willow. Salix conifera Muhl.; S. humilis Marsh. var. angustifolia (Barratt) Anderss.; S. muehlenbergiana Willd. var. angustifolia Barratt; S. tristis Ait. var. glabrata (Anderss.) Anderss. • CT, MA, ME, NH, RI, VT; nearly throughout. Fields, roadsides, forest borders, sand plains,

S a l i c ac e a e   8 47

woodlands, often in well-drained soils. Salix humilis can be confused with S. cinerea ssp. oleifolia, especially when the ridges on decorticated branches are longer than usual (as does occasionally occur on New England material). The two taxa can be separated on the basis of plant height, leaf blade outline, stipules, anther color, and ovary pubescence. Salix humilis is 0.3–3 m tall, with leaf blades 2.3–7.5 times as long as wide, lacking or with rudimentary stipules on the first leaves (i.e., those that expand from the winter bud), has purple anthers that turn yellow in age, and has wavy or crimped hairs on the ovary. Salix cinerea ssp. oleifolia is 2–12 m tall, with leaf blades 1.8–4.3 times as long as wide, with foliaceous stipules on the first leaves, has yellow anthers, and ± straight hairs on the ovary. 16. Salix lucida Muhl. ssp. lucida N Fig. 895 shining willow. Salix lucida Muhl. var. angustifolia (Anderss.) Anderss.; S. lucida Muhl. var. intonsa Fern.; S. lucida Muhl. var. latifolia (Anderss.) Anderss.; S. pentandra L. var. lucida (Muhl.) Kuntze • CT, MA, ME, NH, RI, VT. Shorelines, wet ditches, swamps, wetland margins. 17. Salix myricoides Muhl. var. myricoides N bayberry willow. Salix cordata Michx. var. glaucophylla Bebb; S. cordata Michx. var. myricoides (Muhl.) Carey; S. glaucophylla (Bebb) Bebb; S. glaucophylloides Fern.; S. glaucophylloides Fern. var. glaucophylla (Bebb) Schneid. • ME; northern portion of state. Ice-scoured river shores in high-pH bedrock and/or till regions. 18. Salix nigra Marsh. N

Fig. 895  Serrulate leaf blade of Salix lucida showing glands at summit of petiole.

black willow. Salix denudata Raf.; S. falcata Pursh; S. nigra Marsh. var. brevifolia Anderss.; S. nigra Marsh. var. falcata (Pursh) Torr. • CT, MA, ME, NH, RI, VT; generally limited in ME to the southern half of the state. Shorelines, swamps, riparian forests, river banks. 19. Salix occidentalis Walt. N dwarf prairie willow. Salix humilis Marsh. var. microphylla (Anderss.) Fern.; S. humilis Marsh. var. tristis (Ait.) Griggs; S. muehlenbergiana Willd.; S. tristis Ait.; S. tristis Ait. var. microphylla Anderss. • CT, MA, ME, NH, RI. Sand plains, sandy roadsides and fields, woodlands. This species is frequently included in Salix humilis as a variety (var. tristis). However, it is distinct in vegetative and floral morphology and has a different range than S. humilis (contrary to the statements of Argus (1986); S. humilis ranges further north than does S. occidentalis). In New England, Salix occidentlis is predominantly found on the coastal plain (S. humilis is not at all restricted to the coastal plain). 20. Salix pedicellaris Pursh N Fig. 896 bog willow. Salix myrtilloides L. var. hypoglauca (Fern.) Ball; S. myrtilloides L. var. pedicellaris (Pursh) Anderss.; S. pedicellaris Pursh var. hypoglauca Fern.; S. pedicellaris Pursh var. tenuescens Fern. • CT, MA, ME, NH, RI, VT. Fens, swamps, peaty lake shores. 21. Salix pellita (Anderss.) Anderss. ex Schneid. N satiny willow. Salix chlorophylla Anderss. var. pellita Anderss.; S. sitchensis Sanson ex Bong. var. pellita (Anderss.) Jepson • ME, NH, VT; mainly in the northern portions of states. Lake shores, river shores, stream banks. 22. Salix pentandra L. E laurel willow. CT, MA, ME, NH, RI, VT. Fields, roadsides, shorelines, areas of cultivation. 23. Salix petiolaris Sm. N meadow willow. Salix gracilis Anderss.; S. gracilis Anderss. var. textoris Fern.; S. petiolaris Sm. var. angustifolia Anderss.; S. petiolaris Sm. var. gracilis (Anderss.) Anderss.; S. petiolaris Sm. var. subsericea Anderss.; S. sericea Tausch ex Kern. var. subsericea (Anderss.) Rydb.; S. subsericea (Anderss.) Schneid. • CT, MA, ME, NH, RI, VT; restricted in CT to the northern and/or western portions of state. Hydric fields, fens, shorelines, low ditches, wetland margins. 24. Salix planifolia Pursh ssp. planifolia

NC

tea-leaved willow. Salix chlorophylla Anderss.; S. chlorophylla Anderss. var. nelsonii (Ball) Flod.; S. nelsonii Ball; S. phylicifolia L. ssp. planifolia (Pursh) Hiitonen; S. planifolia Pursh var. nelsonii (Ball) Ball ex E.C. Sm. • ME, NH, VT. Tarns, brooksides, ravines, seeps, cliff bases, and talus in alpine areas.

Fig. 896  Entire leaf blade of Salix pedicellaris.

8 48   tricolpate s

25. Salix purpurea L. E basket willow. CT, MA, ME, NH, RI, VT. Fields, roadsides, river banks, wetland margins, areas of cultivation. 26. Salix pyrifolia Anderss. N balsam willow. Salix balsamifera (Hook.) Barratt ex Anderss.; S. balsamifera (Hook.) Barratt ex Anderss. var. lanceolata Bebb; S. cordata Muhl. var. balsamifera Hook.; S. pyrifolia Anderss. var. lanceolata (Bebb) Fern. • ME, NH, VT. Bogs, fens, swamps, peaty shores and ditches. 27. Salix sericea Marsh. N Fig. 897 silky willow. Salix coactilis Fern.; S. grisea Willd.; S. petiolaris Sm. var. grisea (Willd.) Torr.; S. petiolaris Sm. var. sericea (Marsh.) Anderss. • CT, MA, ME, NH, RI, VT; nearly throughout. Sandy, gravelly, or cobbly river and lake shores, less frequently on ledgy shorelines and in swamps and roadside ditches, rarely in fields. Rare forms of this willow from river shores in central and northern ME show larger leaf blades with dentate margins and non-sericeous pubescence abaxially (the “coactilis” form). These apparently intergrade with the typical form of Salix sericea. Fig. 897  Carpellate flower of Salix sericea (viewed from abaxial surface) showing blunt ovary and dark-tipped floral bract.

28. Salix serissima (Bailey) Fern. N autumn willow. Salix arguta Anderss. var. pallescens Anderss.; S. lucida Muhl. var. serissima Bailey • CT, MA. Fens, peaty meadows, and swamps in regions of high-pH bedrock. 29. Salix uva-ursi Pursh

NC

bearberry willow. Salix cutleri Tuckerman; S. cutleri Tuckerman var. labradorica (Anderss.) Anderss. • ME, NH, VT; northern portion of states. Alpine plateaus, ridges, and summits. 30. Salix viminalis L. E osier willow. CT, MA, ME, NH, RI, VT. River banks, near dwellings. Salix babylonica L. × Salix euxina I.V. Belyaeva. Salix ×pendulina Wenderoth is an ‌ uncommon willow hybrid in New England known from CT, MA, ME. It is very similar to 1a × S. babylonica (S. × sepulcralis), especially in its pendulous branches and branchlets, but has green-brown (or less frequently yellow-brown, gray-brown, or red-brown) branchlets rather than gold-yellow (or less frequently green-yellow) branchlets.

Sapindaceae 1a. Leaves alternate, pinnately compound; fruit a 3-locular or partially 3-locular, bladderyinflated capsule 2a. Vines climbing by axillary tendrils; leaves ternately or biternately compound; petals white to red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cardiospermum 2b. Woody plants without tendrils; leaves pinnately compound with 7–15 leaflets; petals yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Koelreuteria 1b. Leaves opposite, simple [Figs. 900, 901], palmately compound, or pinnately compound in 1 species; fruit a samara-like schizocarp or a 1- to 3-locular, non-inflated capsule 3a. Leaves simple or pinnately compound; perianth actinomorphic; fruit a samara-like schizocarp, without prickles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acer 3b. Leaves palmately compound; perianth zygomorphic; fruit a capsule, commonly with small prickles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aesculus

S a p i ndac e a e   8 49

Acer 1a. Leaves pinnately compound with 3–9 leaflets; nectary disk wanting; branchlets often glaucous (not glaucous in var. negundo), with a line from each leaf scar meeting in a raised point between the leaf scars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. negundo 1b. Leaves simple, palmately lobed [Figs. 900, 901]; nectary disk present (absent or vestigial in A. rubrum and A. saccharinum); branchlets not glaucous, with the line from each leaf scar not meeting in a raised point 2a. Leaf blades ± evenly toothed or double-toothed through the margin, the sinuses angled (i.e., V-shaped) [Fig. 901] 3a. Sinuses of leaf blades extending ⅔ or more the distance to the base of the blade; sepals red to purple, ca. 3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. palmatum 3b. Sinuses of leaf blades extending less than ½ the distance to the base of the blade [Fig. 901]; sepals not red or purple, except in A. rubrum and then 1–3 mm long 4a. Winter buds stalked, covered by 2 valvate scales 5a. Inflorescence an upright panicle, pubescent; petals 2–3 mm long; branchlets and bud scales pubescent; bark brown to gray-brown; leaf blades 6–12.5 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. spicatum 5b. Inflorescence a drooping raceme, ± glabrous [Fig. 898]; petals 5–10 mm long; branchlets and bud scales glabrous; bark green to green-brown, with slender, longitudinal, pale stripes; leaf blades 12–18 (–20) cm long . . . . . A. pensylvanicum 4b. Winter buds sessile, covered by 3–10 imbricate scales 6a. Inflorescence an upright panicle; mericarps nearly parallel, diverging at an angle of less than 30 degrees; leaf blades green on the abaxial surface 7a. Leaf blades dark green and lustrous on the adaxial suface, 3-lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ginnala 7b. Leaf blades light green and dull on the adaxial surface, unlobed or sometimes with 3 obscure lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. tataricum 6b. Inflorescence either a dense, umbel-like fascicle or a drooping panicle; mericarps diverging at an angle of 30–90 degrees; leaf blades thinly glaucous on the abaxial surface 8a. Flowers coetaneous or serotinous, in drooping panicles; sepals green, 4–5 mm long; leaf blades crenate-serrate, the secondary veins prominent on the adaxial surface; collateral winter buds absent . . . . . . . . A. pseudoplatanus 8b. Flowers precocious, in dense, umbel-like fascicles [Fig. 899]; sepals red, 1–3 mm long; leaf blades serrate, the secondary veins visible, but not prominent, on the adaxial surface; collateral winter buds often present on some branchlets on the plant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. rubrum 2b. Leaf blades entire through much of the margin, rarely with more than a few pairs of points on each major lobe (A. saccharinum often with more teeth, but these concentrated near the tips of the lobes), the sinues rounded (i.e., U-shaped; sometimes angled in A. campestre and A. saccharinum) [Fig. 900] 9a. Flowers with petals; mericarps usually diverging at an angle of 170–180 degrees; broken petioles yielding a milky latex; leaf blades green on the abaxial surface 10a. Leaf blades mostly 4–7 cm long, ciliate, abaxially pubescent, with short, blunt lobes; corymb pubescent; mericarps usually pubescent . . . . . . . . . . . . . . A. campestre 10b. Leaf blades (5–) 10–15 cm long, eciliate, abaxially glabrous or with hairs in the junctions of veins, with prominent, pointed lobes; corymb ± glabrous; mericarps glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. platanoides

8 50   tricolpate s

9b. Flowers without petals (i.e., with only sepals); mericarps usually diverging at an angle of 60–120 degrees; broken petioles yielding a watery latex; leaf blades glaucous on the abaxial surface (green on the abaxial surface in A. nigrum) 11a. Flowers precocious, borne in dense clusters on very short pedicels that barely exceed the associated bud scales (pedicels later elongating as fruits mature); ovary and fruit pubescent; nectary disk absent or vestigial; winter buds red, blunt at the apex; branchlets often with collateral buds; central lobe of the leaf more than 50% as long as the blade and narrowed near the base; bark on mature trees with long, curling plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. saccharinum 11b. Flowers ± coetaneous, evidently pedicellate; ovary and fruit glabrous; extrastaminal nectary disk present; winter buds brown, pointed at the apex; branchlets without collateral buds; central lobe of the leaf up to 50% as long as the blade and not narrowed near the base [Fig. 900]; bark on mature trees with furrows 12a. Leaf blades abaxially glaucous and glabrous (rarely green) at maturity (sometimes pubescent on the veins or rarely across the surface with appressed to ascending hairs), relatively plane (i.e., the sides not drooping); bark gray to graybrown, with loose-edged plates on mature trees; middle lobe of leaf blades often with ± parallel basal margins [Fig. 900]; petioles without foliaceous outgrowths, not pubescent through their length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. saccharum 12b. Leaf blades green and abaxially pubescent across the surface with spreading to erect hairs, the sides often drooping; bark dark brown and furrowed; middle lobe of leaf blades usually ± triangular and tapering from base to apex; petioles of vigorous shoots sometimes with foliaceous outgrowths that ± conceal the axillary buds, sometimes pubescent through their length . . . . . . . . . . . . . . . . . . . . . A. nigrum 1. Acer campestre L. E hedge maple. MA, VT; also reported from RI by George (1997), but specimens are unknown. Roadsides, waste areas, near dwellings. 2. Acer ginnala Maxim. E Amur maple. CT, MA, ME, NH, VT. Fields, roadsides, forest borders. 3. Acer negundo L. n ash-leaved maple.  3a. Negundo aceroides (L.) Moench; N. negundo (L.) Karst.; 3b. Acer violaceum (Kirchn.) Simonkai; Negundo aceroides (L.) Moench ssp. violaceum (Kirchn.) W.A. Weber; N. aceroides (L.) Moench var. violaceum Kirchn. • CT, MA, ME, NH, RI, VT. Riparian forests, river banks, roadsides, waste areas. This species is non-native in ME and RI, and is native, at least in part, to the other states in New England. Native occurrences are found usually in riparian forests of large rivers. 1a. Branchlets green, not glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . 3a. A. negundo var. negundo 1b. Branchlets green to purple, glaucous . . . 3b. A. negundo var. violaceum (Kirchn.) Jaeger Variety negundo is known from CT, MA, ME, NH, RI, VT. Variety violaceum is known from CT, MA, ME, NH, RI, VT. 4. Acer nigrum Michx. f. N black maple. Acer saccharum Marsh. ssp. nigrum (Michx. f.) Desmarais; A. saccharum Marsh. var. nigrum (Michx. f.) Britt.; Saccharodendron nigrum (Michx. f.) Small • CT, MA, NH, VT. Rich, mesic forests, riparian forests, often in regions of high-pH bedrock. Some occurrences of this species are believed to be planted (e.g., Salisbury, CT, see Graves et al. 1910). The distinctiveness of Acer nigrum and its treatment as a taxon separate from A. saccharum has been called into question by some researchers (Skepner and Krane 1997a, 1997b). However, the authors failed to supply an adequate discussion of the morphological differences between these two species (i.e., their difficulty identifying them may be based, in part, on using too few characters). Further, they made some false assumptions that led them to a potentially incorrect conclusion. Though their research does indicate a need for additional study, it is

S a p i ndac e a e   8 5 1

far from conclusive. Fall foliage of Acer nigrum is usually yellow-orange, in contrast to the usual orange-colored (often brightly so) foliage of A. saccharum. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it was not naturalized in RI. 5. Acer palmatum Thunb. E Japanese maple. CT, MA; also reported from RI by George (1997), but specimens are unknown. Forest borders, fields, roadsides. 6. Acer pensylvanicum L. N Fig. 898 striped maple. CT, MA, ME, NH, RI, VT. Deciduous and mixed evergreen-deciduous forests. 7. Acer platanoides L. E Norway maple. Acer platanoides L. var. schwedleri Nichols. • CT, MA, ME, NH, RI, VT. Roadsides, waste areas, forest fragments, areas of habitation. 8. Acer pseudoplatanus L. E

Fig. 898  Inflorescence of Acer pensylvanicum.

sycamore maple. CT, MA, RI. Roadsides, waste areas, forest borders, river banks. 9. Acer rubrum L. N Fig. 899 red maple. Acer carolinianum Walt.; A. rubrum L. var. tridens Wood; A. stenocarpum Britt.; Rufacer rubrum (L.) Small; R. carolinianum (Walt.) Small • CT, MA, ME, NH, RI, VT; nearly throughout. Deciduous and mixed evergreen-deciduous forests, swamps, riparian forests, shorelines, successional fields. ‌9 × 10. Acer ×freemanii Murray is a rare maple hybrid known from MA, ME, NH, VT. The leaf blades are similar to A. saccharinum but have V-shaped sinuses, a shorter central lobe, and teeth distributed over the apical 50–75% of the lobes. 10. Acer saccharinum L. N

Fig. 899  Carpellate flowers of Acer rubrum showing precocious flowering.

silver maple. Acer dasycarpum Ehrh.; A. saccharinum L. var. laciniatum Pax; Argentacer saccharinum (L.) Small • CT, MA, ME, NH, RI, VT. Riparian forests, lacustrine flood plains, swamps. 11. Acer saccharum Marsh. var. saccharum N Fig. 900 sugar maple. Acer nigrum Michx. f. var. glaucum (F. Schmidt) Fosberg; A. nigrum Michx. f. var. saccharophorum (K. Koch) Clausen; A. saccharum Marsh. var. glaucum (F. Schmidt) Sarg.; Saccharodendron saccharum (Marsh.) Moldenke • CT, MA, ME, NH, RI, VT. Mesic forests, including uplands, rocky slopes, and high-terrace floodplains. 12. Acer spicatum Lam. N Fig. 901 mountain maple. CT, MA, ME, NH, RI, VT; rare or absent in southeastern New England. Forests, woodlands, talus slopes, ledges, stream banks. 13. Acer tataricum L. E Tatarian maple. MA; also reported from RI by Magee and Ahles (1999), but specimens are unknown. Roadsides, forest fragments, near dwellings.

Fig. 900  Leaf of Acer saccharum.

Aesculus 1a. Petals 5, white marked with red or yellow near the base of the blade, all but the lowermost petal with a cordate-based claw; leaves with (5–) 7 (–9) leaflets; winter buds resinous; fruit 5–6 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. hippocastanum 1b. Petals 4, yellow to green-yellow, all with blades tapering to the claw; leaves with 5 (–7) leaflets; winter buds not resinous; fruit 3–5 cm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. glabra 1. Aesculus glabra Willd. E Ohio buckeye. NH, also reported from ME by Campbell et al. (1995), but specimens are unknown. Forest fragments and borders.

Fig. 901  Leaf of Acer spicatum.

8 52   tricolpate s

2. Aesculus hippocastanum L. E horse-chestnut. CT, MA, ME, VT; also reported from RI by George (1992), but specimens from naturalized populations are unknown. Roadsides, forest borders and fragments, near dwellings.

Cardiospermum 1. Cardiospermum halicacabum L. E balloon-vine. CT, MA. Waste areas, gardens.

Koelreuteria 1. Koelreuteria paniculata Laxm. E golden rain-tree. MA. Roadsides, forest borders and fragments, waste areas.

Sarraceniaceae Sarracenia 1. Sarracenia purpurea L. ssp. purpurea N Fig. 902 purple pitcherplant. Sarracenia heterophylla Eat.; S. pupurea L. var. heterophylla (Eat.) Torr. • CT, MA, ME, NH, RI, VT. Peatlands, peaty shores. Two color forms of this species occur in our

area. Most plants show red to purple coloration (to a lesser or greater extent) on the leaves and petals and stigma apices. Rare plants show entirely pale green leaves with pale yellow petals and stigma apices. Fig. 902  Pitcher-like leaves of Sarracenia purpurea.

Saxifragaceae 1a. Perianth 4-merous; petals absent; flowers with 4 or 8 stamens that are set in the notches of an 8-lobed disk; at least the lower leaves opposite . . . . . . . . . . . . . . . . . . . . Chrysosplenium 1b. Perianth 5-merous; petals present; flowers with 5 or 10 stamens; leaves alternate or all basal (opposite in Saxifraga oppositifolia) 2a. Petals conspicuously pectinately fringed [Fig. 904]; inflorescence a slender, simple raceme with flowers borne on short pedicels 1–5 mm long . . . . . . . . . . . . . . . . . . . . . . . Mitella 2b. Petals entire [Fig. 906]; inflorescence a raceme, panicle, cyme, or thyrse, or of solitary flowers at the tips of stems, the flowers variously arranged on short to long pedicels 3a. Ovary 2-locular, the placentation axile; stamens shorter than to scarcely longer than the petals 4a. Stems with leaves (note: these sometimes small but are not confined to branching points of the stem) [Fig. 906]; leaf blades lacking foliar crystals, with one or more of the following features: palmately lobed outline, with 1 or more lime encrusted pores, prominently ciliate along the margin, and/or entire through most or all of margin; seeds smooth, tuberculate, or papillate . . . . . . . . . . . . . . . . . . . . Saxifraga

S a x i fr agac e a e   8 5 3

4b. Stems without leaves (note: bracts found on the main axis of the plant are only present at branching points) [Fig. 903]; leaf blades with foliar crystals, the principal ones crenate or dentate through much of the margin (except M. foliolosa), lacking all of the features present in 4a (except M. foliolosa, which is entire through most of the margin); seeds ribbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micranthes 3b. Ovary 1-locular, the placentation parietal; stamens long-exserted beyond the petals 5a. Flowers with 10 stamens; inflorescence a raceme; valves of the capsule very unequal, one larger than the other . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tiarella 5b. Flowers with 5 stamens; inflorescence a panicle-like thyrse; valves of the capsule ± equal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heuchera

Chrysosplenium 1. Chrysosplenium americanum Schwein. ex Hook. N golden-saxifrage. CT, MA, ME, NH, RI, VT. Swamps, brooks, seeps.

Heuchera 1. Heuchera americana L. var. americana N common alum-root. CT. Forests, forest borders, banks.

Micranthes Reference: Brouillet and Elvander (2009a). 1a. Inflorescence with usually a single, terminal flower, the lower flowers modified into vegetative bulbils; ovary superior, the hypanthium not adnate to the carpels; leaf blades toothed only near apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. foliolosa 1b. Inflorescence with normal, petal-bearing flowers throughout [Fig. 903]; ovary partly inferior, the hypanthium adnate to the base of the carpels; leaf blades often toothed throughout much of the margin 2a. Leaf blades (5–) 10–20 (–30) cm long; stems 30–100 cm tall; sepals soon reflexed; petals green-white to purple; plants of wet, low ground . . . . . . . . . . . . . . . . . . . . . . M. pensylvanica 2b. Leaf blades (0.8–) 1–5 (–8) cm long; stems 5–40 cm tall; sepals ascending to spreading; petals white; plants of rocky woods, ledges, and cliffs . . . . . . . . . . . . . . . . . . . . M. virginiensis 1. Micranthes foliolosa (R. Br.) Gornall

NC

naked-bulbil small-flowered-saxifrage. Hydatica foliolosa (R. Br.) Small; Saxifraga foliolosa R. Br.; S. stellaris L. var. comosa Poir.; Spatularia foliolosa (R. Br.) Small • ME; northern portion of state. Moist to wet alpine cliffs and gullies. 2. Micranthes pensylvanica (L.) Haw. N swamp small-flowered-saxifrage. Saxifraga forbesii Vasey; S. pensylvanica L.; S. pensylvanica L. var. forbesii (Vasey) Engl. & Irmsch.; S. pensylvanica L. var. purpuripetala (A.M. Johnson) Bush • CT, MA, ME, NH, RI, VT. Swamps, peaty meadows, fens. Two forms of this species are found in New England—one with white-yellow to green-yellow petals and one with red-purple petals (this latter form is much less common). 3. Micranthes virginiensis (Michx.) Small N Fig. 903 early small-flowered-saxifrage. Saxifraga virginiensis Michx. • CT, MA, ME, NH, RI, VT. Cliffs, ledges, balds, rocky slopes.

Fig. 903  Habit of Micranthes virginiensis with small leaves confined to the branching points of the inflorescence.

8 54   tricolpate s

Mitella 1a. Stem leaves 1 or none; basal leaf blades rounded to obtuse at the apex; petals light greenyellow, 3–5 mm long; pedicels 1–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. nuda 1b. Stem leaves 2–4; basal leaf blades obtuse to acute at the apex; petals white, 1.8–2.2 mm long; pedicels 1–2 mm long 2a. Stem leaves 2, opposite, positioned below the lowest flower . . . . . . . . . . . . . M. diphylla 2b. Stems leaves 4, alternate, the uppermost one subtending a flower . . . . . . M. prostrata 1. Mitella diphylla L. N two-leaved bishop’s-cap. CT, MA, NH, VT; most frequent in western New England. Mesic forests, often rich, rocky, and deciduous. 2. Mitella nuda L. N Fig. 904 naked bishop’s-cap. CT, MA, ME, NH, VT. Swamps, wooded fens, stream borders, usually associated with abundant bryophyte cover. The inclusion of Mitella prostrata in the concept of M. nuda by Soltis and Freeman (2009) is not supported by the morphology of these species. 3. Mitella prostrata Michx. Fig. 904  Flower of Mitella nuda.

NC

prostrate bishop’s-cap. CT, VT. Swamps. Mitella prostrata is a species of unknown origin. It has the flowers and basal leaves of M. diphylla (however, the leaf blades are frequently not as sharply pointed at the apex) but with a different number and arrangement of stem leaves. The VT report is approximate because the original collection by Michaux did not provide enough information to know whether the population occurred in NY or VT.

Saxifraga Soltis et al. (2001) showed that the traditionally defined Saxifraga is not monophyletic and that several of our species must be segregated. These taxa not related to the core species of Saxifraga are placed in the genus Micranthes. Reference: Brouillet and Elvander (2009b). 1a. Leaves opposite and borne on a stem, closely imbricate; petals purple . . . S. oppositifolia 1b. Leaves alternate or ± all basal [Fig. 905], those borne on a stem not closely imbricate; petals white or yellow 2a. Primary leaves with orbicular to reniform blades that have 3–7 palmately arranged, tooth like lobes and are borne on evident petioles [Fig. 906] 3a. Upper leaves with vegetative bulbils in their axils; inflorescence a solitary, terminal flower; petals 6–10 mm long; stems solitary or a few together, 8–20 cm tall . . S. cernua 3b. Upper leaves without vegetative bulbils; inflorescence of 1–4 flowers; petals 3–5 mm long; stems cespitose, 2–10 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rivularis 2b. Primary leaves with narrow-oblong or lanceolate to to obovate or obovate-oblong blades, entire or toothed, sessile or tapering to the base [Fig. 905] 4a. Leaf blades entire, without a conspicuous, lime-encrusted pores; petals yellow, often orange-dotted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. aizoides 4b. At least the larger leaf blades toothed, each tooth with a conspicuous, white, limeencrusted pore [Fig. 905]; petals white, without orange dots . . . . . . . . . . . . S. paniculata 1. Saxifraga aizoides L.

NC

yellow mountain saxifrage. Leptasea aizoides (L.) Haw. • VT; northern portion of state. Boreal and subalpine cliffs in regions of high-pH bedrock. 2. Saxifraga cernua L. nodding saxifrage. NH; northern portion of state. Wet, alpine cliffs and gullies.

NC

S a xi f r agac e a e   8 5 5

3. Saxifraga oppositifolia L. ssp. oppositifolia

NC

purple mountain saxifrage. Antiphylla oppositifolia (L.) Fourr. • VT; northern portion of state. Boreal and subalpine cliffs in regions of high-pH bedrock. 4. Saxifraga paniculata P. Mill. ssp. neogaea (Butters) D. Löve

N C Fig. 905

white mountain saxifrage. Chondrosea paniculata (P. Mill.) A. Löve ssp. neogaea (Butters) A. Löve; Saxifraga aizoon Jacq. ssp. neogaea (Butters) A. & D. Löve; S. aizoon Jacq. var. neogaea Butters • ME, NH, VT; primarily in the northern portion of states. Boreal to alpine cliffs and gullies in regions of high-pH bedrock. 5. Saxifraga rivularis L. ssp. rivularis

N C Fig. 906

alpine-brook saxifrage. S. rivularis L. ssp. flexuosa (Sternb.) Gjaerev.; S. rivularis L. var. debilis (Engelm. ex Gray) Dorn; S. rivularis L. var. flexuosa (Sternb.) Engl. & Irmsch. • NH; northern half of state. Mossy, alpine summits and gullies.

Tiarella 1. Tiarella cordifolia L. var. cordifolia N foam-flower. CT, MA, ME, NH, VT; also reported from RI by Kartesz (1999), but specimens are unknown. Mesic forests, swamps.

Fig. 905  Habit of Saxifraga paniculata showing limeencrusted pores along margin of leaf blades.

Scrophulariaceae 1a. Amphibious plants of tidal shores with horizontal stems emitting from the nodes clusters of linear leaves and sometimes also peduncles bearing a solitary flower with a ± actinomorphic, 5-merous corolla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limosella 1b. Terrestrial plants with upright stems and variable leaves wider than linear, with inflorescences borne on a stem bearing flowers with weakly to conspicuously zygomorphic corollas (actinomorphic in Buddleja, but then 4-merous) 2a. Plants woody; flowers actinomorphic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Buddleja 2b. Plants herbaceous (though often with firm, persistent stems); flowers weakly to conspicuously zygomorphic [Fig. 907, 908] 3a. Foliage leaves all basal or alternate; corolla weakly zygomorphic, rotate [Fig. 908]; androecium composed of 5 stamens, all of which bear pollen; plants biennial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Verbascum 3b. Foliage leaves opposite; corolla zygomorphic, bilabiate [Fig. 907]; androecium composed of 5 stamens—4 pollen-bearing, 1 sterile; plants perennial . . . . Scrophularia

Buddleja 1. Buddleja davidii Franch. E orange-eye butterfly-bush. CT, MA; also reported from RI by George (1997), but specimens are unknown. Roadsides, forest borders, waste areas.

Limosella 1. Limosella australis R. Br. N Atlantic mudwort. Limosella aquatica L. var. tenuifolia Hoffmann; L. subulata Ives • CT, MA, ME, NH, RI. Fresh to brackish-tidal river shores.

Fig. 906  Palmately lobed leaf blades of Saxifraga rivularis.

8 56   tricolpate s

Scrophularia 1a. Sterile filament yellow-green near the apex; capsules 6–10 mm long; pedicels 0.4–0.7 mm thick; leaf blades broad-cuneate to broad-rounded or truncate at the base . . . . S. lanceolata 1b. Sterile filament dark brown to purple-brown near the apex; capsules 4–7 mm long; pedicels 0.3–0.5 mm thick; leaf blades cordate to broad-rounded at the base 2a. Leaf blades acute at the apex; petiole wing-margined, less than ⅓ as long as the blade; corolla 7–10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nodosa 2b. Leaf blades acuminate at the apex; petiole not or scarcely wing-margined, ⅓ to ½ as long as the blade; corolla 5–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. marilandica 1. Scrophularia lanceolata Pursh N Fig. 907 lance-leaved figwort. Scrophularia leporella Bickn.; S. pectinata Raf. • CT, MA, ME, NH, RI, VT. Fields, meadows, roadsides, forest borders, frequently in proximity to rivers and ponds. Fig. 907  Flower of Scrophularia lanceolata showing anthers opening by transverse slits.

2. Scrophularia marilandica L. N eastern figwort. CT, MA, ME, NH, RI, VT. Fields, roadsides, forest fragments, woodlands, river banks, railroads. 3. Scrophularia nodosa L. E woodland figwort. MA, ME, RI; also reported from CT by Hultén and Fries (1986), but specimens are unknown. Roadsides, dumps, forest fragments, waste areas, gardens.

Verbascum Collections of putative Verbascum lychnitis × V. thapsus from Nantucket County, MA, require more study. The specimens—MacKeever N672 (GH!) and MacKeever N822 (NEBC!) appear referable to V. phlomoides. Reference: Ferguson (1972). 1a. Plants pubescent with simple, glandular hairs; filaments pubescent with purple, knobtipped hairs; capsule subglobose, pubescent with minute hairs or glabrous . . . . . . V. blattaria 1b. Plants pubescent with compound, eglandular hairs; some of the filaments pubescent, usually the 3 upper (i.e., shorter) ones pubescent with white to yellow hairs and the 2 lower (i.e., longer) ones glabrous or nearly so (pubescent with purple hairs in V. nigrum); capsules ellipsoid to ovoid, densely pubescent 2a. Filaments pubescent with purple hairs; bracts 2–5 mm long; basal leaf blades cordate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. nigrum 2b. At least the upper 3 filaments pubescent with white to yellow hairs; bracts 8–40 mm long; basal leaf blades cuneate to rounded at the base 3a. Axis of inflorescence branched; calyx (1.5–) 2.5–4 mm long; anther reniform, not decurrent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. lychnitis 3b. Axis of inflorescence unbranched (though a given plant may have more than 1 inflorescences); calyx 5–12 mm long; anther decurrent on the filament 4a. Stigma capitate; inflorescence crowded at maturity, the flowers concealing the axis of the spike; corolla 10–25 (–35) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . V. thapsus 4b. Stigma spatulate, decurrent on the sides of the style; inflorescence less dense at maturity, the flowers spaced enough to expose the axis of the spike [Fig. 908]; corolla (20–) 25–45 (–55) mm wide 5a. Leaves decurrent on the stem nearly or completely to the next leaf; floral bracts 15–40 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. densiflorum 5b. Leaves not, or only very shortly, decurrent on the stem; floral bracts 9–15 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. phlomoides

S c ro p hu l a r i ac e a e   8 57

1. Verbascum blattaria L. E moth mullein. CT, MA, ME, NH, RI, VT. Dry fields, roadsides, waste areas, and clearings. 2. Verbascum densiflorum Bertol. E dense-flowered mullein. Verbascum thapsiforme Schrad. • MA. Wool waste. 3. Verbascum lychnitis L. E white mullein. CT, MA, NH, RI, VT. Fields, roadsides, waste areas. 4. Verbascum nigrum L. E black mullein. MA, NH. Fields, roadsides, waste areas, wool waste. 5. Verbascum phlomoides L. E Fig. 908 orange mullein. CT, MA, ME, RI, VT. Fields, roadsides, waste areas, wool waste. 6. Verbascum thapsus L. E common mullein. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, railroads, ledges, clearings.

Simaroubaceae Ailanthus 1. Ailanthus altissima (P. Mill.) Swingle E tree-of-heaven. Ailanthus glandulosa Desf. • CT, MA, RI, vt; also reported from ME by Campbell et al. (1995), but specimens are unknown. Forests, forest edges and fragments, roadsides, waste areas.

Solanaceae Callibrachoa parviflora (Juss.) D’Arcy was reported from CT, MA, ME, NH, and RI by Magee and Ahles (1999). However, the report was erroneous given that this species has not been collected in New England, and the authors included as a synonym Petunia integrifolia (which is a distinct taxon belonging to another genus). Magee and Ahles (1999) also reported Solandra grandiflora Sw. for CT; however, the voucher specimens (at CONN) were taken from cultivated plants (i.e., Solandra grandiflora is not naturalized in New England). 1a. Inflorescences (2–) 3- to many-flowered, with both peduncles and pedicels (note: the inflorescence is leafy-bracteate in Hyoscyamus) [Figs. 910, 911, 915] 2a. Corolla zygomorphic, the upper lip 2-lobed; stamens 4, fewer than the lobes of the corolla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Browallia 2b. Corolla ± actinomorphic; stamens 5, numbering the same as the corolla lobes 3a. Corolla ± rotate [Figs. 915, 916]; anthers dehiscing by terminal pores or small clefts (dehiscing by longitudinal slits in S. lycopersicon); fruit a berry . . . . . . . . . . . . . . . Solanum 3b. Corolla funnelform or salverform [Figs. 910, 911]; anthers dehiscing by longitudinal slits; fruit a capsule or pyxis 4a. Androecium with 4, didynamous stamens; corolla lobes conspicuously emarginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Salpiglossis

Fig. 908  Portion of inflorescence of Verbascum phlomoides showing visible raceme axis.

8 58   tricolpate s

4b. Androecium with 5, equal stamens that are not paired; corolla lobes truncate to acute 5a. Inflorescence a leafy-bracteate, ± secund raceme; corolla dull yellow to greenyellow with a purple basal, interior region, conspicuously reticulate-veined with purple veins; fruit a pyxis, circumscissile above the middle . . . . . . . . . . Hyoscyamus 5b. Inflorescence a panicle or secund raceme (but then with very small bracts) [Figs. 910, 911]; corolla variously colored, but without purple reticulations; fruit a capsule that dehisces by 2–4 apical valves . . . . . . . . . . . . . . . . . . . . . . . . . . . Nicotiana 1b. Inflorescence consisting of solitary flowers in the axils of leaves (rarely in pairs or trios), with only pedicels, the peduncles absent or obsolete [Fig. 909] 6a. One anther conspicuously smaller than the other 4; calyx connate only at the very base; leaves opposite; fruit a 2-valved capsule; . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Petunia 6b. All 5 anthers of similar size; calyx connate at least half its length (connate only near the base in Nicandra, but then each sepal with 2 basal auricles); leaves alternate or falsely appearing opposite in some genera; fruit a berry or 4-valved capsule 7a. Flowers 7–24 cm long; fruit berry-like capsule armed with long, soft prickles, with 4 locules (at least in the basal portion), solitary on the stem . . . . . . . . . . . . . . . . . . . Datura 7b. Flowers 0.6–2.5 cm long (15–20 cm long in Solandra, but then each of the 5 corolla lobes crenate); fruit a 2-locular berry (with 3–5 locules in Nicandra), usually more than 1 per stem (solitary in Nicandra) 8a. Calyx connate only at the very base, each sepal with 2 basal auricles; corolla 20–25 mm long; gynoecium with 3–5 carpels . . . . . . . . . . . . . . . . . . . . . . . . . . . Nicandra 8b. Calyx connate at least half its length, the sepals lacking basal auricles; corolla 6–20 mm long; gynoecium with 2 carpels 9a. Plants woody, usually armed with thorns; corolla blue to purple; leaves borne on petioles 3–10 mm long, the blades with entire margins . . . . . . . . . . . . . . . . Lycium 9b. Plants herbaceous, unarmed; corolla yellow to white; principal leaves either borne on petioles longer than 10 mm or the blades toothed or both 10a. Fruiting calyx usually inflated and larger than the berry it contains [Fig. 912], essentially closed at the apical end, membranaceous to scarious, often with 10 prominent ribs; corolla campanulate to open-campanulate . . . Physalis 10b. Fruiting calyx not inflated, filled by the berry it contains, open at the apical end, herbaceous, without prominent ribs; corolla ± rotate . . . . . . Leucophysalis

Browallia 1. Browallia americana L. E Jamaican-forget-me-not. MA. Dumps.

Datura 1a. Corolla 7–10 cm long, 5-toothed at the apex [Fig. 909]; calyx 3–5 cm long, evidently angled, the angles sometimes with thin wing-margins; capsule erect, dehiscent by 4 valves; stems and principal branches glabrous or sparsely pubescent . . . . . . . . . . . . . . . D. stramonium 1b. Corolla 16–24 cm long, 5-toothed or 10-toothed at the apex; calyx 5–15 cm long, the connate portion without angles or with obscure angles; capsule ± nodding, irregularly dehiscent; stems and principal branches evidently pubescent

S o l a n ac e a e   8 5 9

2a. Corolla 16–20 cm long, entirely white, with 5 primary apical teeth alternating with 5 secondary apical teeth for a total of 10 teeth; mature leaf blades subsucculent and densely velvety-pubescent with crisp hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. inoxia 2b. Corolla 20–24 cm long, entirely white or white with a pale purple limb, with only 5 apical teeth; mature leaf blades relatively thinner and often subglabrous . . . . . . D. wrightii 1. Datura inoxia P. Mill. E downy thorn-apple. Datura meteloides, auct. non Dunal pro parte • CT, MA, ME, NH, RI. Dumps, waste areas, shorelines, roadsides. 2. Datura stramonium L. E Fig. 909 thorn-apple. Datura inermis Juss. ex Jacq.; D. stramonium L. var. inermis (Juss. ex Jacq.) Fern.; D. stramonium L. var. tatula (L.) Torr.; D. tatula L. • CT, Ma, Me, NH, RI, VT. Fields, pastures, roadsides, waste areas, railroads, dumps, coastal beaches. This species has a white or light blue-purple corolla. 3. Datura wrightii Regel E sacred thorn-apple. Datura inoxia P. Mill. ssp. quinquecuspida (Torr.) Barcl.; D. metel L. var. quinquecuspida Torr. • MA. Dumps. Report of this species in CT, ME, NH, and RI by Kartesz (1999) are based on specimens of Datura inoxia.

Hyoscyamus 1. Hyoscyamus niger L. E black henbane. CT, MA, ME, VT; also reported from NH by Magee and Ahles (1999), but specimens are unknown. Fields, waste areas, Atlantic coast shorelines.

Leucophysalis 1. Leucophysalis grandiflora (Hook.) Rydb.

NC

large false ground-cherry. Chamaesaracha grandiflora (Hook.) Fern.; Physalis grandiflora Hook. • VT; northern portion of state. Lake shores, clearings, quarries.

Lycium 1a. Calyx usually 2-lobed with each lobe 2- or 3-toothed near the apex, 3–4 mm long; corolla lobes subglabrous, the lobes obviously shorter than the basal, connate portion; leaf blades 20–30 × 3–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. barbarum 1b. Calyx 3- to 5-parted to near half way to base, 4–5 mm long; corolla lobes pubescent near the margin, the lobes as long as or longer than the basal, connate portion; leaf blades 15–50 (–100) × 5–25 (–40) mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. chinense 1. Lycium barbarum L. E matrimony-vine. Lycium halimifolium P. Mill. • CT, MA, ME, NH, RI, VT. Forest fragments and borders, roadsides, waste areas, near dwellings. 2. Lycium chinense P. Mill. E Chinese matrimony-vine. CT, MA, ME, NH, VT. Roadsides, waste areas.

Nicandra 1. Nicandra physalodes (L.) Gaertn. E apple-of-Peru. Atropa physalodes L.; Physalodes physalodes (L.) Britt. • CT, MA, ME, NH, RI, VT. Waste areas, fields, dumps.

Fig. 909  Flower and leaves of Datura stramonium.

8 60   tricolpate s

Nicotiana Reference: Goodspeed (1954). 1a. Inflorescence a panicle-like thyrse [Fig. 911] 2a. Petiole slender, not decurrent on the stem; pedicels 3–4 mm long in flower, elongating to 7 mm in fruit; basal, tubular, connate portion of the corolla (i.e., excluding the lobes) 12–20 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. rustica 2b. Petiole broadly winged and shortly decurrent on the stem; pedicels 5–10 (–15) mm long in flower, elongating to 20 (–25) mm in fruit; basal, tubular, connate portion of the corolla (i.e., excluding the lobes) (30–) 35–55 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. tabacum 1b. Inflorescence appearing to be a raceme [Fig. 910] 3a. Corolla mostly more than 5 times as long as the calyx, the basal, connate, tubular portion 40–120 mm long and 1.5–2.5 mm wide near its midpoint . . . . . . . . . . . . N. longiflora 3b. Corolla 5 times as long as the calyx or shorter, the basal, connate tubular portion 13–100 mm long and 2–8 mm wide near its midpoint (note: corolla tube length 4a. Corolla 50–100 mm long, the lobes ± acute, extending more than 50% of the way to the corolla opening [Fig. 910] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. alata 4b. Corolla 18–50 mm long; the lobes broad-acute to ± truncate, extending less than 50% of the way to the corolla opening 5a. Corolla (33–) 35–50 mm long, white to violet-tinged, with evident, broad-acute to obtuse lobes; flowering calyx 15–20 mm long; stem leaves neither claspling nor decurrent at the base; pollen yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. quadrivalvis 5b. Corolla 18–28 mm long, green-yellow to green, with obscure, ± truncate lobes; flowering calyx 8–10 mm long; stem leaves clasping and decurrent; pollen blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. langsdorffii 1. Nicotiana alata Link & Otto E Fig. 910 Jasmine tobacco. Nicotiana affinis T. Moore; N. alata Link & Otto var. grandiflora Comes • CT, MA, ME. Fields, waste areas, dumps. This species is commonly cultivated and shows many different corolla colors but white is common. The stem leaves are both clasping and shortly decurrent.

Fig. 910 Raceme-like inflorescence of Nicotiana alata.

‌1 × Nicotiana forgetiana hort. ex Hemsley. Nicotiana ×sanderae hort. ex W. Wats. is the red-flowered garden tobacco. It is a very rare tobacco hybrid known from MA. It is similar in many regards to N. alata but has a calyx 8–13 mm long, red corolla lobes that change to dark blue in drying, and the basal connate portion of the corolla is ca. 3 times as long as the calyx (vs. calyx 15–25 mm long, commonly ± white corolla lobes, and the basal connate portion of the corolla 4–5 times as long as the calyx). 2. Nicotiana langsdorffii Schrank E flowering tobacco. MA. Dumps. 3. Nicotiana longiflora Cav. E long-flowered tobacco. CT, NH. Fields, waste areas, dumps. The report of this species in VT by Magee and Ahles (1999) was erroneous. They stated the species occurred in Strafford County, VT, but there is no such county in that state; however, there is a Strafford County, NH, and a specimen exists at NHA! for that county. Most collections labeled as Nicotiana longiflora in regional herbaria (including some from CT and all those from MA) are N. alata. In addition to characters used in the identification key, the two species can be separated by differences in leaf morphology and corolla tube width. Nicotiana longiflora has clasping but non-decurrent stem leaves and a corolla tube 1.5–2.5 mm wide near the midpoint of the corolla (vs. clasping and short-decurrent stem leaves and a corolla 3–8 mm wide near the midpoint in N. alata).

S o l a nac e a e   8 6 1

4. Nicotiana quadrivalvis Pursh var. bigelovii (Torr.) DeWolf E Indian tobacco. Nicotiana bigelovii (Torr.) S. Wats. • MA. Wool waste. 5. Nicotiana rustica L. E Fig. 911 Aztec tobacco. CT, MA. Fields, waste areas, dumps. The report of this species in NH by Magee and Ahles (1999) is based on specimens taken from cultivated plants—22 Sep 1864, Blake s.n. (NHA!) and 2 Sep 1865, Blake s.n. (NHA!). 6. Nicotiana tabacum L. E cultivated tobacco. CT. Waste areas. Reports of this species from MA (specimens at NEBC!) are based on collections from cultivated material (i.e., this species is not naturalized in MA).

Petunia ‌hybrida Report ‌atkinsiana of Petunia × D. Don ex Loud. (as P. × Vilm.) by Sorrie and Somers (1999) was based on a synonymy that incorrectly included both P. axillaris and P. integrifolia ‌atkinsiana are unknown from New England. (as P. violacea). Specimens of P. × 1a. Corolla white, with a nearly cylindric basal tube 40–60 mm long that is only slightly expanded upward; stamens inserted near the middle of the basal tube of the corolla and extending nearly to the throat; stems usually upright . . . . . . . . . . . . . . . . . . . . . . . . . . . P. axillaris 1b. Corolla purple to light purple or pink-red, with a funnelform basal tube 29–36 mm long that is conspicuously expanded upward; stamens inserted near the base of the basal tube of the corolla and barely extending past the midpoint of the tube; stems sometimes prostrate in basal portion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. integrifolia 1. Petunia axillaris (Lam.) B.S.P. E garden petunia. Petunia nyctaginiflora Juss. • CT, MA. Fields, roadsides, waste areas. 2. Petunia integrifolia (Hook.) Schinz & Thellung E violet-flowered petunia. Petunia violacea Lindl. • CT, MA, ME, NH, RI, VT. Waste areas, dumps, railroads, gardens.

Physalis Physalis hederifolia Gray var. hederifolia was reported from RI and VT by Kartesz (1999), but specimens are unknown. Physalis lanceolata Michx. was reported from NH by Magee and Ahles (1999), but specimens are unknown. This species is endemic to the southeastern United States and was likely reported from New England based on misapplication of names by the authors. Physalis peruviana L. was reported from MA by Sorrie and Somers (1999). The specimen on which this record rests was cultivated—Kidder s.n. (NEBC!). References: Waterfall (1958), Sullivan (2004). 1a. Plants annual from a taproot; corolla 4–10 mm long or up to 15 mm long in P. philadelphica; flowering calyx 3–7 (–10) mm long 2a. Apical half of stem villous or viscid-villous; leaf blades villous 3a. Stems, young growth, and major veins of the leaf blades provided with villous pubescence intermixed with sessile glands [Fig. 912]; leaf blades gray-green, coarsely dentate to the base, with conspicuous reticulate venation (note: most visible on the abaxial surface), frequently drying orange or with orange spots; anthers yellow, sometimes with blue tinged; fruiting calyx ± as long as broad; mature berry orange to brown-orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. grisea 3b. Stems, young growth, and major veins of leaf blades with fine, non-viscid pubescence; leaf blades green, entire to obscurely dentate and then the teeth often confined to the distal half of the blade, without conspicuous reticulate venation, drying green or brown; anthers blue to purple; fruiting calyx longer than broad; mature berry green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pubescens

Fig. 911 Panicle-like inflorescence of Nicotiana rustica.

862  tricolpate s

2b. Apical half of stem glabrous or sparsely strigillose; leaf blades glabrous or essentially so 4a. Corolla yellow, 4–10 mm long; anthers 1–2.3 mm long, not or scarcely twisted after dehiscence; berry yellow, 8–11 mm wide; flowering calyx 3–5 mm long . . . . . P. angulata 4b. Corolla yellow with a 5 dark spots near the center, 7–15 mm long; anthers mostly 2.5–4 mm long, strongly twisted after dehiscence; berry usually purple or purplestreaked, 12–40 mm wide; flowering calyx 5–7 (–10) mm long . . . . . . . . . P. philadelphica 1b. Plants perennial from rhizomes; corolla 10–20 mm long; flowering calyx (4–) 6–12 (–15) mm long 5a. Corolla white, with 5 evident lobes; fruiting calyx bright red . . . . . . . . . . . . . P. alkekengi 5b. Corolla yellow, often with 5 purple spots within, scarcely lobed [Fig. 913]; fruiting calyx green or yellow, or becoming brown in drying 6a. Apical half of the stem pubescent and often other parts of the plant pubescent with stipitate-glands intermixed with short, eglandular hairs and longer, eglandular, multicellular hairs; leaf blades ovate to rhombic, broad-rounded to cordate at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. heterophylla 6b. Apical half of the stem pubescent entirely (or nearly entirely) with eglandular hairs of various lengths, these often short, stiff, appressed or decurved hairs; leaf blades narrow-lanceolate to ovate, narrow-cuneate to broad-cuneate at the base 7a. Pubescence of the stem mostly appressed; connate portion of the calyx pubescent in 10 vertical strips along the nerves with minute, appressed hairs up to 0.5 mm long; anthers 3–4 mm long; berry yellow . . . . . . . . . . . . . . . . . . . . . . P. longifolia 7b. Pubescence of the stem mostly decurved; connate portion of the calyx pubescent throughout with long, spreading hairs usually 0.5–1.5 mm long; anthers 2–3 mm long; berry orange-red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. virginiana 1. Physalis alkekengi L. E strawberry ground-cherry. CT, MA, ME, VT. Waste areas, gardens. This species is the “Chinese lantern” of horticulture. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it was not naturalized in RI. 2. Physalis angulata L. E cut-leaved ground-cherry. Physalis angulata L. var. lanceifolia (Nees) Waterfall; P. angulata L. var. pendula (Rydb.) Waterfall; P. lanceifolia Nees; P. pendula Rydb. • CT, MA. Fields, dumps. Fig. 912  Densely viscid-villous fruiting calyx and stem of Physalis grisea.

3. Physalis grisea (Waterfall) M. Martinez N Fig. 912 downy ground-cherry. Physalis pruinosa, auct. non L.; P. pubescens L. var. grisea Waterfall • CT, MA, ME, RI, VT. Fields, waste areas. Physalis grisea has a confusing taxonomic history

and, in part due to the missaplication of the name P. pruinosa L., has been confused with P. pubescens in various treatments (see Martinez 1993 for correct application of the name P. pruinosa). 4. Physalis heterophylla Nees var. heterophylla N Fig. 913 clammy ground-cherry. Physalis ambigua (Gray) Britt.; P. heterophylla Nees var. ambigua (Gray) Rydb.; P. heterophylla Nees var. clavipes Fern.; P. heterophylla Nees var. nyctaginea (Dunal) Rydb.; P. heterophylla Nees var. villosa Waterfall; P. nyctaginea Dunal; P. sinuata Rydb. • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 5. Physalis longifolia Nutt. var. subglabrata (Mackenzie & Bush) Cronq. N Fig. 913  Flower of Physalis heterophylla.

long-leaved ground-cherry. Physalis subglabrata Mackenzie & Bush; P. virginiana P. Mill. var. subglabrata (Mackenzie & Bush) Waterfall • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas. 6. Physalis philadelphica Lam. var. immaculata Waterfall E Mexican ground-cherry. Physalis ixocarpa, auct. non Brot. ex Hornem. • MA, VT. Waste areas, dumps. This is the “tomatillo” of agriculture.

S o l a nac e a e   8 63

7. Physalis pubescens L. var. integrifolia (Dunal) Waterfall N hairy ground-cherry. Physalis hirsuta Dunal var. integrifolia (Dunal) Waterfall • MA; also reported for RI and VT by Kartesz (1999), but specimens are unknown. Waste areas. This species was also reported from CT by Kartesz (1999), based on Dowhan (1979). However, Dowhan reported Physalis pubescens var. grisea, a nomenclatural synonym of P. grisea, and not P. pubescens var. integrifolia. 8. Physalis virginiana P. Mill. var. virginiana n Virginia ground-cherry. Physalis intermedia Rydb.; P. monticola C. Mohr • CT, MA, ME, NH. Fields, waste areas, woodlands, clearings, wool waste. This species is native in CT and non-native elsewhere. Reports of this species in VT (e.g., Dole 1937, Magee and Ahles 1999) are based on collections of Physalis longifolia var. subglabrata (a plant that has been referred to by the nomenclatural synonym P. virginiana var. subglabrata).

Salpiglossis 1. Salpiglossis sinuata Ruiz & Pavón E paintedtongue. CT. Waste areas.

Solanum Reference: Schilling (1981). 1a. Leaf blades pubescent with compound hairs; stems and/or leaf blades armed with slender prickles; anthers tapering to the apex (usually not tapering in S. melongena) 2a. Leaf blades subentire to sinuate-lobed or toothed with large teeth, but not obviously pinnately lobed; calyx not enclosing fruit at maturity; calyx armature variable, often lacking or sparse; plants perennial (sometimes annual in S. melongena) 3a. Inflorescence with mostly 3–6 flowers; corolla 20–30 mm wide, light purple to white; berry 1–1.5 cm long, yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. carolinense 3b. Inflorescence with 1 (–3) flowers; corolla 30–50 mm wide, purple to light purple; berry 5–30 cm long, black, purple, pink, brown, yellow, or white . . . . . . . . . S. melongena 2b. Leaf blades conspicuously 1- or 2-times pinnately lobed; calyx loosely to tightly enclosing fruit at maturity; calyx armature always present, of numerous slender prickles [Fig. 916]; plants annual 4a. Petals yellow; inflorescence pubescent with eglandular hairs . . . . . . . . . S. rostratum 4b. Petals blue to light purple; inflorescence pubescent with both glandular and eglandular hairs 5a. All the athers yellow, ± equal; berry loosely enclosed by the accrescent calyx; fruiting pedicels usually spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sisymbriifolium 5b. One of the anthers tinged with light purple, much longer, and curved compared with the other 4 yellow anthers; berry tightly enclosed by the accrescent, often adherent, calyx; fruit pedicels usually erect to ascending . . . . . . . . . . . S. citrullifolium 1b. Leaf blades glabrous or pubescent with simple hairs (sometimes with a few stellate hairs in S. pseudocapsicum); stems and leaf blades unarmed; anthers not tapering to the apex 6a. Leaf blades pinnately compound; plants perennial (annual in S. lycopersicon) 7a. Plants climbing or twining; larger leaf blades frequently with 1 or 2 pairs of small basal lobes or leaflets; berry ellipsoid to ovoid . . . . . . . . . . . . . . . . . (in part) S. dulcamara 7b. Plants upright, though often reclining under the weight of fruits; larger leaf blades with 5–9 primary leaflets (often also with smaller leaflets between the larger ones); berry ± globose

8 64   tricolpate s

8a. Plants perennial, from tubers; plants without aromatic, glandular hairs; anthers dehiscing by terminal pores; berry 2–4 cm in diameter, green to purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tuberosum 8b. Plants annual; plants pubescent with aromatic, glandular hairs; anthers dehiscing by longitudinal slits; berry 2–10 cm in diameter, red to yellow . . . . . . . S. lycopersicon 6b. Leaf blades simple, subentire to lobed; plants annual 9a. Leaf blades evidently pinnately lobed with mostly 5–9 linear to narrow-oblong lobes; inflorescence with (1–) 2 or 3 flowers; berry green . . . . . . . . . . . . . . . . . . S. triflorum 9b. Leaf blades subentire to toothed or the larger frequently with a pair of small basal lobes or leaflets in S. dulcamara; inflorescence with 1–25 flowers; berry black, green to green-brown, or yellow to red 10a. Plants climbing or twining; larger leaf blades frequently with 1 or 2 pairs of small basal lobes or leaflets [Fig. 914]; inflorescence with (8–) 10–25 flowers; berry ellipsoid to ovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) S. dulcamara 10b. Plants erect to spreading; leaf blades simple; inflorescence with 1–10 flowers; berry ± globose 11a. Plants woody; stems often with some stellate hairs; berry 12–15 (–20) mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pseudocapsicum 11b. Plants herbaceous; stems lacking stellate hairs; berry 5–13 mm in diameter 12a. Stems glabrous or sparsely pubescent; berry black 13a. Berries 8–13 mm wide, with 15–60 seeds 1.8–2.2 mm long, without sclerotic granules, dull on the outside surface; anthers 1.8–2.5 mm long; pollen 25–35 μm in diameter; style 3.5–4.5 mm long; inflorescence with usually 5–7 (–10) flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nigrum 13b. Berries 5–9 mm wide, with 50–110 seeds 1.4–1.8 mm long, with 4–15 sclerotic granules, dull or lustrous; anthers 1.3–2 mm long; pollen 17–23 μm in diameter; style 2.5–3.2 mm long; inflorescence with usually 1–4 flowers [Fig. 915] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. ptycanthum 12b. Stems moderately to densely pubescent; berry green, yellow, or red 14a. Stems pubescent with spreading hairs; calyx accrescent, concealing up to 50% of the fruit at maturity; berry green to green-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. physalifolium 14b. Stems densely pubescent with appressed hairs or ascending hairs; calyx not accrescent; berry yellow to red . . . . . . . . . . . . . . . . . . . . . . . . . . . S. villosum 1. Solanum carolinense L. var. carolinense E Carolina nightshade. CT, MA, ME, NH, RI, VT. Fields, waste areas. 2. Solanum citrullifolium A. Braun var. citrullifolium E watermelon nightshade. MA. Waste areas. 3. Solanum dulcamara L. E Fig. 914 climbing nightshade. CT, MA, ME, NH, RI, VT. Swamps, stream banks, shorelines, forests fragments, roadsides, waste areas. 1a. Branchlets and leaf blades glabrous or nearly so . . . . . . 3a. S. dulcamara var. dulcamara 1b. Branchlets and leaf blades conspicuously pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3b. S. dulcamara var. villosissimum Desv. Fig. 914  Infructescence and leaves of Solanum dulcamara.

Variety dulcamara is known from CT, MA, ME, NH, RI, VT. Variety villosissimum is known from CT, MA, ME, VT; also reported from RI by George (1992), but specimens are unknown.

S o l a n ac e a e   8 6 5

4. Solanum lycopersicon L. var. lycopersicon E garden tomato. Lycopersicon esculentum P. Mill.; L. lycopersicum (L.) Karst. ex Farw. • CT, MA, NH, RI, VT. Fields, dumps, river banks and shorelines, waste areas. 5. Solanum melongena L. E eggplant. MA. Dumps, waste areas. 6. Solanum nigrum L. ssp. nigrum E European black nightshade. MA, ME. Atlantic coast shorelines and near shoreline areas. 7. Solanum physalifolium Rusby var. nitidibaccatum (Bitter) Edmonds E ground-cherry nightshade. Solanum sarrachoides, auct. non Sendtner • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, dumps, gardens. Solanum physalifolium has been confused with S. sarrachoides in many treatments that include these plants. The former species (specifically var. nitidibaccatum) has (3–) 4- to 8 (–10)-flowered inflorescences, white to purple corollas with a distinct purple (rarely brown) and yellow central region, corolla lobes ± as long as wide, and translucent green to green-brown berries that contain brown seeds and usually 2 sclerotic granules. The latter species has 3- or 4 (–5)-flowered inflorescences, white corollas with a yellow central region, corolla lobes wider than long, and opaque, pale green berries that contain yellow seeds and usually 4–6 sclerotic granules. See Edmonds (1986) for discussion. 8. Solanum pseudocapsicum L. E

Fig. 915  Inflorescence of Solanum ptycanthum.

Jerusalem-cherry nightshade. MA; also reported from CT by Magee and Ahles (1999), but specimens are unknown. Dumps, waste areas. 9. Solanum ptycanthum Dunal N Fig. 915 eastern black nightshade. Solanum americanum, auct. non P. Mill. • CT, MA, ME, NH, RI, VT. Forests, swamps, shorelines, coastal beaches. This species has long gone by the name Solanum americanum in regional literature. However, that species is restricted (in this country) to the southern United States (Schilling 1981). 10. Solanum rostratum Dunal E Fig. 916 horned nightshade. Androcera rostrata (Dunal) Rydb.; Solanum cornutum, auct. non Lam. • CT, MA, ME, NH, RI, VT. Fields, gardens, dumps, waste areas. 11. Solanum sisymbriifolium Lam. E sticky nightshade. MA. Waste areas, dumps.

Fig. 916  Flowers of Solanum rostratum.

12. Solanum triflorum Nutt. E cut-leaved nightshade. CT, MA. Waste areas. 13. Solanum tuberosum L. E Irish potato. CT, MA, ME, NH, RI, VT. Fields, dumps, waste areas. 14. Solanum villosum L. E hairy nightshade. Solanum nigrum L. var. villosum L. • MA. Dumps.

Staphyleaceae Staphylea 1. Staphylea trifolia L. N Fig. 917 American bladdernut. CT, MA, NH, VT. Forest borders and fragments, woodlands, rocky slopes, roadsides.

Fig. 917  Capsules and leaves of Staphylea trifolia.

8 66   tricolpate s

Styracaceae Halesia 1. Halesia carolina L. E Carolina silverbell. MA, RI. Forest fragments and roadsides in areas of habitation.

Symplocaceae Symplocos 1. Symplocos paniculata (Thunb.) Miq. E Asiatic sweetleaf. Myrtus chinensis Lour.; Prunus paniculata Thunb.; Symplocos chinensis (Lour.) Druce • MA. Forest fragments.

Tamaricaceae Tamarix Tamarix gallica L. was reported from CT by Magee and Ahles (1999), but specimens are unknown. 1. Tamarix parviflora DC. E small-flowered tamarisk. Tamarix tetrandra, auct. non Pallas • CT, MA. Roadsides, waste areas.

Theophrastaceae Samolus 1. Samolus valerandi L. ssp. parviflorus (Raf.) Hultén N seaside brookweed. Samolus floribundus Kunth; S. parviflorus Raf. • CT, MA, ME, NH, RI, VT; coastal plain, but disjunct in VT. Mainly on fresh to brackish-tidal river shores, rarely found away from the coastal region along rivers and in swamps.

Thymelaeaceae 1a. Sepals well-developed, spreading, petaloid; hypanthium pubescent; stamens and style included within the perianth; flowers white to pink to purple; drupes 6–10 mm long; leaf blades narrow-oblanceolate to oblong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Daphne

Thy m e l a e ac e a e   8 67

1b. Sepals minute, represented by tiny lobes at the summit of the tubular hypanthium [Fig. 918]; hypanthium glabrous; stamens and style exserted beyond the perianth [Fig. 918]; flowers yellow; drupes 12–15 mm long; leaf blades ovate to broad-oblong-obovate or obovate . . . . . . . . . . . Dirca

Daphne 1a. Leaves evergreen, with blades 10–18 (–25) × (2–) 3–5 (–6) mm; flowers coetaneous to serotinuous, in groups of 6–10 (–20) at the apex of branchlets; drupe brown-yellow; low plants with procumbent stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cneorum 1b. Leaves deciduous, with blades 30–80 × 8–25 mm; flowers precocious, appearing in groups of 2–4 in the axils of the previous year’s leaves; drupe red; upright plants with erect to ascending stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. mezereum 1. Daphne cneorum L. E rose daphne. MA. Roadsides, abandoned homesteads. 2. Daphne mezereum L. E February daphne. CT, MA, ME, NH, RI, VT. Forests, forest fragments, roadsides.

Dirca 1. Dirca palustris L. N Fig. 918 eastern leatherwood. CT, MA, ME, NH, VT. Mesic, mostly deciduous, forests.

Tropaeolaceae Tropaeolum ........................................................................................ 1. Tropaeolum majus L. E garden-nasturtium. CT, MA, NH. Waste areas, roadsides.

Ulmaceae Ulmus Reference: Sherman-Broyles (1997). 1a. Flowers serotinous, appearing in late summer; samaras not winged; mature leaf blades 1.5–2.5 cm wide, with (4–) 5 or more lateral veins forking well before reaching margin per half of leaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. parvifolia 1b. Flowers precocious, appearing in early spring; samaras winged; mature leaf blades 2–8 (–10) cm wide, when less than 3 cm wide then no more than 3 lateral veins forking well before reaching margin per half of leaf 2a. Mature leaf blades 2–3.5 cm wide, ± single serrate on the margin; samaras glabrous and eciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. pumila

Fig. 918  Precocious flowers of Dirca palustris.

8 68   tricolpate s

2b. Mature leaf blades 2.5–8 (–10) cm wide, double serrate (at least in the distal half) [Fig. 920]; samaras pubescent and/or ciliate (though the pubescence sometimes limited to a small region of the fruit) 3a. Lateral veins of leaf blade not forking until near the margin (rarely some leaf blades with 1 vein forking well before reaching margin) [Fig. 919]; samara marginally ciliate through most or all of its margin 4a. Inflorescence a dense fascicle usually shorter than 2.5 cm; samaras glabrous over the surface of body and wing; branches lacking irregular corky wings . . . . U. americana 4b. Inflorescence a raceme-like cyme, up to 5 cm long; samaras pubescent over the surface of body and wing; branches often irregularly winged with 2 or more corky plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. thomasii 3b. Usually 2 or more lateral veins of leaf blade forking well before reaching the margin [Figs. 920, 921]; samaras eciliate or ciliate only near the apex (though pubescent on surface of body or wing) 5a. Leaf blades with marginal cilia; winter bud scales red, pubescent with red tomentum near the margins; samara pubescent on only the body . . . . . . . . . U. rubra 5b. Leaf blades without marginal cilia; winter bud scales red-brown to brown or dark brown, with white or pale cilia near the margins; samara pubescent on only the central vein of wing or near apical margin 6a. Leaf blade with a strongly inequilateral base, with one side partially concealing the petiole, the apex sometimes with 3 acuminate lobes [Fig. 920]; branches lacking corky wings; calyx with prominent lobes, the sinuses between them extending ca. half the distance to the base; samara elliptic to obovate, pubescent on central vein of wing, with the seed positioned near the middle . . . . . U. glabra

Fig. 919  Leaf blade of Ulmus americana, usually with no veins forking well before reaching margin.

6b. Leaf blade with a weakly to moderately inequilateral base, neither side concealing the petiole, the apex without additional lobes; branches often irregularly winged with 2 or more corky plates; calyx with short lobes, the sinuses between them extending much less than half the distance to the base; samara ± orbicular at maturity (narrower during development), pubescent near apical margin, with the seed positioned distal to middle . . . . . . . . . . . . . . . . . . . . U. procera 1. Ulmus americana L. N Fig. 919 American elm. CT, MA, ME, NH, RI, VT. Upland and riparian, deciduous forests, swamps, field edges, roadsides, river banks. See Ulmus rubra for an additional character to separate vegetative individuals of these two species. 2. Ulmus glabra Huds. E Fig. 920 Scotch elm. CT, MA, ME, VT. Roadsides, areas of habitation. 3. Ulmus parvifolia Jacq. E Chinese elm. MA. Roadsides, forest borders. Most collections determined to be this species in New England (including all those from ME and most of those from MA) were based on specimens of Ulmus pumila. 4. Ulmus procera Salisb. E

Fig. 920  Leaf blade of Ulmus glabra.

English elm. Ulmus campestris L., pro parte; U. minor, auct. non P. Mill. • CT, MA, RI, VT. Roadsides, forest borders, areas of habitation. Ulmus minor P. Mill. and U. procera are sometimes combined as one species (under the name U. minor). However, the two species have different morphologies and are maintained as separate taxa where they are native (Tutin 1993b). 5. Ulmus pumila L. E Siberian elm. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, forest borders, lake shores.

U l m ac e a e   8 6 9

6. Ulmus rubra Muhl. N Fig. 921 slippery elm. Ulmus fulva Michx. • CT, MA, ME, NH, RI, VT; absent from much of ME. Rich, deciduous and riparian forests, rocky slopes. Ulmus rubra is sometimes confused with U. americana when collected later in the growing season (i.e., when vegetative). The outer bark is a useful character to separate these two species. The outer bark of U. rubra has contrasting light and dark layers when viewed in cross-section, that of U. americana does not. 7. Ulmus thomasii Sarg.

NC

rock elm. NH, VT; also reported for CT by Magee and Ahles, but specimens are unknown. Woodlands and ledges in regions of high-pH bedrock, riparian forests, swamps.

Urticaceae Reference: Boufford (1997). 1a. Leaves alternate

Fig. 921  Leaf blade of Ulmus rubra, usually with some veins forking well before reaching the margin.

2a. Plants with conspicuous, stinging hairs; leaf blades serrate; carpellate sepals distinct or nearly so, only 2 of the 4 sepals persistent in fruit, not enclosing the achene in fruit; staminate flowers each with 5 sepals and stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . Laportea 2b. Plants without stinging hairs (or sometimes with a few minute stinging hairs); leaf blades entire; carpellate sepals connate at the base, 4-lobed, accrescent, enclosing the achene in fruit; staminate flowers each with 4 sepals and stamens . . . . . . . . . . . . Parietaria 1b. Leaves opposite 3a. Plants with conspicuous stinging hairs; carpellate calyx of 4 distinct sepals . . . . Urtica 3b. Plants without stinging hairs (or sometimes with a few minute stinging hairs); carpellate calyx of 3 sepals or 4 connate sepals 4a. Inflorescence a spike [Fig. 922]; carpellate sepals connate at the base, 4-lobed, accrescent, enclosing the achene in fruit; leaf blades with rounded cystoliths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Boehmeria 4b. Inflorescence a panicle; carpellate sepals distinct or nearly so, 3-lobed, not enclosing the achene in fruit; leaf blades with linear cystoliths . . . . . . . . . . . . . . . . . . Pilea

Boehmeria 1. Boehmeria cylindrica (L.) Sw. N Fig. 922 small-spiked false nettle. Urtica cylindrica L. • CT, MA, ME, NH, RI, VT. Rich, mesic forests, riparian and lacustrine forests, swamps.

Laportea 1. Laportea canadensis (L.) Weddell N Canada wood-nettle. Urtica canadensis L.; Urticastrum divaricatum (L.) Kuntze • CT, MA, ME, NH, RI, VT. Rich, mesic forests, riparian forests.

Parietaria Parietaria floridana Nutt was reported from NH by several sources (e.g., Seymour 1982, Magee and Ahles 1999). The voucher specimen was determined to be P. hespera Hinton var. hespera, a species of the southwestern United States (David Boufford, personal communication). The occurrence of this species in New England is doubtful and the label information is likely erroneous (i.e., a label mix-up occurred during preparation of the specimen).

Fig. 922  Inflorescence of Boehmeria cylindrica.

870 tricolpate s

1a. Plants annual; achenes light brown, obtuse at the apex, with an apical mucro; tepals 1.5–2 mm long, shorter than the bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pensylvanica 1b. Plants perennial; achenes dark brown, acute at the apex, without an apical mucro; tepals mostly 2–3.5 mm long, longer than the bracts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. judaica 1. Parietaria judaica L. E spreading pellitory. MA. Roadsides, waste areas. 2. Parietaria pensylvanica Muhl. ex Willd. n Pennsylvania pellitory. Parietaria obtusa Rydb. ex Small; P. occidentalis Rydb.; P. pensylvanica Muhl. ex Willd. var. obtusa (Rydb. ex Small) Shinners • CT, MA, ME, NH, VT. Rich, often rocky or ledgy, forests and woodlands in regions of high-pH bedrock, waste areas, sea beaches. This species is introduced to ME on rocks of a seawall and native (at least in part) to other states in New England that it occurs in. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it could be in RI and was unaware of any collections.

Pilea 1a. Achenes 1–1.5 mm wide, dark purple to purple-black except for a narrow, pale, marginal band, with raised, rounded tubercles on the surface [Fig. 923]; leaf blades relatively less lustrous and less translucent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. fontana 1b. Achenes 0.6–1.1 mm wide, largely green to yellow-green, often also with slightly raised purple marks [Fig. 924]; leaf blades relatively more lustrous and more translucent . . . . P. pumila 1. Pilea fontana (Lunell) Rydb. N Fig. 923 Fig. 923  Achene of Pilea fontana.

lesser clearweed. Adicea fontana Lunell; Pilea opaca (Lunell) Rydb. • CT, MA, RI, VT. Rich, mesic forests, riparian forests. 2. Pilea pumila (L.) Gray var. pumila N Fig. 924 Canada clearweed. Adicea pumila (L.) Raf.; Urtica pumila L. • CT, MA, ME, NH, RI, VT. Mesic, deciduous forests, riparian forests, shaded seeps, stream banks, wet logging roads, shaded lawns.

Urtica 1a. Stipules 5–15 mm long, erect; staminate and carpellate flowers usually in separate inflorescences; inflorescence exceeding the length of the petioles of the subtending leaves; plants perennial from rhizomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. dioica Fig. 924  Achene of Pilea pumila.

1b. Stipules 1–3 mm long, spreading to deflexed; staminate and carpellate flowers usually in the same inflorescence; inflorescence often shorter than the length of the petioles of the subtending leaves; plants annual from a taproot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . U. urens 1. Urtica dioica L. n stinging nettle.  1a. Urtica gracilis Ait. var. latifolia Farw.; 1b. Urtica dioica L. var. angustifolia Schlecht.; U. dioica L. var. gracilis (Ait.) C.L. Hitchc.; U. dioica L. var. procera (Muhl. ex Willd.) Weddell; U. gracilis Ait.; U. procera Muhl. ex Willd.; U. viridis Rydb. • CT, MA, ME, NH, RI, VT. Riparian forests, stream banks, forest borders, roadsides, waste areas. 1a. Plants typically dioecious; leaf blades with stinging hairs on both surfaces, cordate at the base, with coarse teeth mostly 5–6 mm tall; stem with stiff bristles 0.75–2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. U. dioica ssp. dioica 1b. Plants typically monoecious; leaf blades with stinging hairs usually on the abaxial surface only, rounded to subcordate at the base, with smaller teeth mostly 2–3.5 mm tall; stem glabrous or pubescent with shorter, softer hairs; bristles lacking or very sparse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. U. dioica ssp. gracilis (Ait.) Seland.

U rt i c ac e a e   871

Subspecies dioica is known from CT, MA, ME, NH and is non-native. The report of this species from VT by Seymour (1982) was based on a specimens of U. dioica ssp. gracilis. Subspecies gracilis is known from CT, MA, ME, NH, RI, VT and is native. 2. Urtica urens L. E burning nettle. CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, yards, about dwellings.

Verbenaceae 1a. Calyx usually more than twice as long as the mature fruit, the portion extending beyond the fruit often contorted; style more than 3 times as long as the ovary; sterile style lobe protruding beyond stigmatic surface; connective of upper pair of anthers usually with a glandular appendage; schizocarps black; limb of corolla 11–24 mm wide . . . . . . . . Glandularia 1b. Calyx usually less than twice as long as the mature fruit, the portion extending beyond the fruit usually not contorted; style less than 3 times as long as the ovary; sterile style lobe not protruding beyond stigmatic surface; connective of anthers without appendages; schizocarps red-brown; limb of corolla 2–7 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Verbena

Glandularia Reports of Glandularia ×‌hybrida (Grönland & Rümpler) Nesom from New England (e.g., Magee and Ahles 1999, Sorrie and Somers 1999) were based on a cultivated specimen—7 Aug 1929, Kidder s.n. (ECON!). Reference: Umber (1979). 1. Glandularia canadensis (L.) Nutt. E rose mock-vervain. Glandularia drummondii (Lindl.) Small; G. lambertii (Sims) Small; Verbena canadensis (L.) Britt.; V. canadensis (L.) Britt. var. lambertii (Sims) Thellung; V. lambertii Sims • CT. Roadsides, gardens, areas of habitation. Reports of this species in MA were based on collections taken from a cultivated setting—Botanical Garden of Harvard University, 1873, unknown s.n. (GH!).

Verbena Reference: Perry (1933). 1a. Leaf blades 1- or 2-times pinnatifid, cleft into 3–5 segments, or with deep sinuses producing lobe-like teeth [Fig. 925] 2a. Floral bracts 8–15 mm long; calyx 3–4 mm long; spikes 2–10 (–15) × (0.6–) 1–1.5 (–2) cm in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. bracteata 2b. Floral bracts ca. 1–1.3 mm long; calyx 2–2.5 mm long; spikes (7–) 10–25 × 0.2–0.5 cm in fruit. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. officinalis 1b. Leaf blades entire to serrate or coarsely dentate [Figs. 926, 927] 3a. Spikes solitary or less frequently in trios at the tips of the stem and branches [Fig. 926] 4a. Leaf blades slightly tapered to the base, sometimes with an broad, ill-defined, petiole-like base; spikes obloid to cylindric, mostly 3–7.5 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. hispida 4b. Leaf blades conspicuously tapered and/or very narrow at the base, often with a short, but evident, petiole; spikes cylindric to narrow-cylindric, (6.5–) 7.5 or more times as long as wide

872  tricolpate s

5a. Stems sparsely strigillose; leaf blades linear or narrow-oblong to lanceolate or narrow-spatulate, the primary ones 3–15 mm wide; limb of corolla 4–6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. simplex 5b. Stems hirsute, often densely so; leaf blades elliptic to ovate or suborbicular [Fig. 926], 15–50 (–60) mm wide; limb of corolla 7–9 mm wide . . . . . . . . . . . . . V. stricta 3b. Spikes few to numerous, in panicles near the apex of the plant [Fig. 927] 6a. Leaves sessile and clasping the stem; spikes (0.5–) 2–5 cm long in fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. bonariensis 6b. Leaves petiolate, not clasping; spikes 3–30 cm long in fruit 7a. Corolla white, the limb usually 2–2.5 mm wide; flowering spikes slender, 2–4 (–5) mm wide, relatively less crowded, the flowers not contiguous [Fig. 927]; styles mostly 0.4–0.6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. urticifolia 7b. Corolla blue, the limb 2.5–4.5 mm wide; flowering spikes thicker, mostly (5–) 6–8 mm wide, with crowded and contiguous flowers; styles mostly 1–1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. hastata 1. Verbena bonariensis L. var. conglomerata Briq. E purple-topped vervain. MA. Fields, waste areas. 2. Verbena bracteata Lag. & Rodr. E Fig. 925 prostrate vervain. Verbena bracteosa Michx. • CT, MA, ME, RI, VT. Waste areas, about mills. 3. Verbena hastata L. var. hastata N blue vervain. CT, MA, ME, NH, RI, VT. Fields, meadows, roadsides, marshes, shorelines, stream banks. Fig. 925  Leaves of Verbena bracteata.

‌ × 6. Verbena ×blanchardii Moldenke is a very rare vervain hybrid known from 3 VT. It has 1–5 spikes at the summit of the stems or branches, moderately strigose stems (sparsely strigose in V. simplex), a corolla averaging larger than V. hastata (limb 3.8–5.3 mm wide), that is light blue or light purple (not deep blue as in V. hastata). 3 ‌ × 7. Verbena ×rydbergii Moldenke is a very rare vervain hybrid known from CT. It is similar to V. hastata in that the flowers are borne is spikes that are paniculately arranged (i.e., numerous). However, the fruiting calyx is larger, mostly 3–4 mm long (vs. 2.5–3 mm in V. hastata and 5.5–6.5 mm in V. stricta) and the calyx hairs are longer, mostly 0.2–0.4 mm long (vs. 0.1–0.2 mm in V. hastata and 0.5–1 mm in V. stricta). This nothospecies also shows slightly more developed petioles than in V. stricta. ‌ × 8. Verbena ×engelmanii Moldenke is a rare vervain hybrid known from CT, MA, NH, 3 VT. It can be recognized by its flowering spikes 4–6.5 mm wide with overlapping, but not crowded, flowers, and corolla tinged with blue. 4. Verbena hispida (Ruiz & Pavón) Ruiz & Pavón E hairy mock-vervain. Glandularia hispida Ruiz & Pavón • MA. Dumps, gardens. 5. Verbena officinalis L. E European vervain. CT, MA. Fields, roadsides, waste areas. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it had questionable naturalization in RI and was unaware of any collections. 6. Verbena simplex Leym.

NC

narrow-leaved vervain. Verbena angustifolia Michx. • CT, MA, VT. Dry fields, sandplains, ledges, quarries, road cuts. This species was reported from Grafton County, NH, by Magee and Ahles (1999). However, the herbarium specimens that voucher these occurrences (at least those formerly at HNH) are without locality information save for the pre-printed mention of Hanover, NH, on the Jesup label. It is doubtful that these collections originated in NH. Fig. 926  Inflorescence and upper leaves of Verbena stricta.

7. Verbena stricta Vent. E Fig. 926 hoary vervain. CT, MA, VT. Fields, roadsides, waste areas, dumps, city streets.

V e r b e n ac e a e   87 3

8. Verbena urticifolia L.

N C Fig. 927

white vervain. CT, MA, ME, NH, RI, VT. Fields, roadsides, forest borders, open rights-of-way, riparian forests. 1a. Leaf blades densely veluntinous abaxially with short hairs up to 0.3 mm long; calyx puberulent, 1.7–2 mm long in fruit; bracts 0.5–1 mm long; mericarps relatively smooth on the outer surface, ca. 1.5 mm long . . . . . . . . . . . . . . . . . 8a. V. urticifolia var. leiocarpa Perry & Fern. 1b. Leaf blades hirtellous with hairs 1–1.3 mm long abaxially or ± glabrous; calyx strigose, 2–2.3 mm long in fruit; bracts 1–1.5 mm long; mericarps corrugated or ribbed on the outer surface, ca. 2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8b. V. urticifolia var. urticifolia Variety leiocarpa is known from CT as is of regional conservation concern. Variety urticifolia is known from CT, MA, ME, NH, RI, VT.

Violaceae 1a. Sepals lacking basal auricles; corolla green-white; stamens connate, forming a sheath that surrounds the carpels, none of them appendaged; leafy-stemmed plants 30–90 (–100) cm tall [Fig. 928] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hybanthus

Fig. 927  Inflorescence and upper leaves of Verbena urticifolia.

1b. Sepals with basal auricles; corolla white, yellow, blue to purple, or multicolored (but not green-white); stamens connivent but not fused, the two lower ones with appendages that are prolonged backwards into the nectary spur; scapose or leafy-stemmed plants 2–45 cm tall [Figs. 933, 935, 936] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Viola

Hybanthus 1. Hybanthus concolor (T.F. Forst.) Spreng.

N C Fig. 928

green-violet. Cubelium concolor (T.F. Forst.) Raf.; Viola concolor T.F. Forst. • CT, VT. Rich, deciduous forests and woodlands, rocky slopes.

Viola The genus Viola is very complex, in large part due to the frequency of interspecific hybridization. This is especially common within the Boreali-americanae (i.e., the acaulescent, blue-flowered species). Early-generation hybrids are usually intermediate in the characters that discriminate the progenitor species. Further, they very often grow is close proximity to the parents and are largely sterile (capsules usually contain only a few well-developed seeds). This is in contrast to occasional individuals that may display an odd character state (e.g., a plant of V. cucullata that has ciliate sepals but is otherwise typical of V. cucullata). Such plants may be later-generation products of past hybridization events. They sometimes occur within a single-species population and are probably best referred to as the species they most resemble (this approach is considered appropriate given that the morphology is an indication of the underlying genetic composition). Though many features of violet plants used for identification are straightforward, some are not intuitive and/or are rarely noted. Most species of Viola have two types of flowers and fruits: spring-appearing chasmogamous flowers and early to mid-summer-appearing cleistogamous flowers. Style morphology allows these two types of fruits to be differentiated. The chasmogamous flowers produce a capsule with an elongate style [Fig. 934], whereas the cleistogamous ones produce capsules with a tightly coiled style [Fig. 929]. Acaulescent species of Viola can be described as homophyllous or heterophyllous. Homophyllous species produce similar types of leaves throughout the season (in regard to lobed or unlobed), whereas heterophyllous species produce dissimilar leaves—

Fig. 928  Flowers and leaves of Hybanthus concolor.

874  tricolpates

the first ones are unlobed and later ones are lobed. Note that the first set of leaves to be produced by many acaulescent species with unlobed blades are not typical (i.e., they may differ in outline from the later leaves) and are not always provided for in the identification key. Haines (2001b) provided a relatively thorough review of the morphological characters used to identify members of the genus Viola. Viola hirsutula × V. sororia was reported from CT by Kartesz (1999) based on Brainerd (1924). This hybrid was not explicitly attributed to New England by Brainerd, but V. hirsutula × V. papilionacea was. Kartesz synonymized V. papilionacea with V. sororia in part (and therefore the origin of the report of V. hirsutula × V. sororia in CT). However, the correct application of the epithet “papilionacea” has not been satisfactorily determined. Therefore, this report remains ambiguous until further work is done. Viola × ‌modesta House (V. lanceolata × V. primulifolia) was reported for MA and ME by various authors; however, the specimens require further study to confirm their identities. References: Brainerd (1921, 1924), Russell (1965), Haines (2001b).

Key for chasmo gamous f lowering material 1a. Plants acaulescent, the leaves and peduncles arising directly from rhizomes or stolons [Figs. 933, 936] 2a. Petals entirely yellow; stipules entire; stigma essentially lacking a beak .V. rotundifolia 2b. Petals not entirely yellow, mostly purple, violet, or white; stipules ciliate, toothed, or lacerate on the margin; stigma with a conical, scoop-shaped, or hook-shaped beak 3a. Style terminating in a slender, downward-oriented, hook-shaped beak; ovary (and later the capsule) pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. odorata 3b. Style dilated at apex, terminating in a scoop-shaped or conical beak; ovary and capsule glabrous (capsule rarely pubescent in V. affinis) 4a. Spur elongate, 4–10 mm long, 2 or more times as long as wide; flowers with purple, glabrous petals; stipules connate to the petioles 5a. Leaf blades cordate-ovate to cordate-suborbicular, 0.8–1.3 times as long as wide, with a deep basal sinus, not decurrent on the petiole; spur 4–6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. selkirkii 5b. Leaf blades oblong to broad-lanceolate, becoming narrow-triangular to narrowovate post anthesis, 1.5–2.7 times as long as wide, broad-cuneate to subcordate at the base, decurrent on the petiole; spur 5–10 mm long . . . . . . . . . . . . . . . V. japonica 4b. Spur shorter, up to 3.2 mm long and less than 2 times as long as wide; flowers with purple or white petals, when purple then at least the lateral petals pubescent; stipules distinct 6a. Flowers relatively smaller, mostly 7–12 mm long, with white petals (pale blue in one alpine species); rhizomes slender, mostly less than 3 mm thick, usually horizontal; stolons produced, at least by summer (not produced in V. renifolia) 7a. Leaf blades 1.5–7 times as long as wide, narrow-cuneate to subcordate at the base 8a. Leaf blades 3–7 times as long as wide, narrowly tapering at the base, the margins denticulate, each tooth with a distinct red-brown to black gland at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. lanceolata 8b. Leaf blades 1.5–2.5 times as long as wide, broader at the base, widely tapering to truncate or subcordate at the base, the margins crenate, the teeth with inconspicuous, pale glands . . . . . . . . . . . . . . . . . . . . . . V. primulifolia 7b. Leaf blades up to 1.5 times as long as wide, cordate at the base 9a. Corolla violet or at least tinged with violet; lateral petals sparsely pubescent; plants restricted to alpine areas over 900 m elevation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. palustris

V i o l ac e a e   87 5

9b. Corolla white; lateral petals glabrous or pubescent; plants found mostly at lower elevation, some species casually subalpine 10a. Leaf blades low-crenate or subentire along the margins, thin, strictly glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. pallens 10b. Leaf blades low-serrate along the margins, thicker, commonly with some pubescence (sometimes glabrous in V. renifolia with characteristically wide leaves—see description) 11a. Leaf blades cordate-orbicular to reniform, the midrib usually equaling 50–77 percent of the total blade width; stolons not produced; pubescence variable, but when restricted to a single side of the blade, then found only on the abaxial (i.e., lower) surface . . . . . . . . V. renifolia 11b. Leaf blades broad cordate-ovate (to cordate-orbicular), the midrib usually equaling 75–90 percent of the total blade width; stolons present, at least later in season; pubescence variable, but when restricted to a single side of the blade, then found only on the adaxial (i.e., upper) surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. blanda 6b. Flowers relatively larger, mostly 13–22 mm long, with purple petals (rare albino forms said to exist); rhizomes stouter, mostly 4–6 mm thick, usually ascending; stolons not produced 12a. Leaf blades with conspicuous lobes (though the first leaves to appear in V. palmata are unlobed) [Figs. 933, 938] 13a. Leaf blades oblong-lanceolate to ovate-triangular, 1.5–3 times as long as wide, coarsely serrate to divided into short lobes at the base [Fig. 936]; lobes, if present, less than 50% of the total length of the leaf blade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (in part) V. sagittata 13b. Leaf blades ± ovate in outline, less than 1.5 times as long as wide, with evident lobes (except for the first leaves to emerge which may be unlobed to weakly lobed); lobes often greater than 50% of the total length of the leaf blade 14a. Lateral petals glabrous; style lacking a beak, with an asymmetrical concave summit; corolla rotate (i.e., wide open), the stamens clearly visible; corolla either bicolorous with the upper two petals dark purple or concolorous and all the petals light violet . . . . . . . . . . . . . . . . . . . . V. pedata 14b. Lateral petals pubescent on inner surface; style with a conical beak on the lower side of the expanded apex; corolla only distally spreading, the stamens somewhat concealed; corolla concolorous with violet to purple petals 15a. Sepals lanceolate, narrow-acute to acuminate at the apex, eciliate [Fig. 930]; leaf blades and petioles essentially glabrous . . . . V. brittoniana 15b. Sepals ovate, rounded to obtuse at the apex, usually ciliate [Fig. 937]; leaf blades and petioles pubescent 16a. Plants heterophyllous, the early- and late-season leaves unlobed, the middle season leaves lobed [Fig. 933]; leaf blades divided into 3–5 segments, the middle lobe usually without smaller lateral lobes [Fig. 933] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. palmata 16b. Plants homophyllous, all of the leaves with evident lobes [Fig. 938]; leaf blades with 5–16 lobes that become progressively smaller and separated by shallower sinuses toward the basal margins, the middle lobe usually with smaller lateral lobes [Fig. 938] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. subsinuata 12b. Leaf blades all unlobed

876   tricolpates

17a. Leaf blades 1.5–3 times as long as wide, cuneate to subcordate at the base, often more coarsely serrate or even somewhat lobed at the base of the blade (especially in later season leaves) [Fig. 936] . . . . . . . (in part) V. sagittata 17b. Leaf blades commonly less than 1.75 times as long as wide, definitely cordate at the base, the teeth at the base of the blade no more obvious than those along the middle of the blade margin (except in V. pectinata) 18a. Lateral petals pubescent with hairs shorter than 1 mm and are knobshaped at the apex [Fig. 931]; spurred petal glabrous [Fig. 931] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. cucullata 18b. Hairs of three lower petals longer than 1 mm and tapering to broadly rounded the apex [Fig. 932]; spurred petal sparsely pubescent (rarely glabrous in V. sororia) [Fig. 932] 19a. Sepals ciliate (sometimes eciliate in V. hirsutula) [Fig. 937]; leaf blades usually as wide as or wider than long and pubescent 20a. Pubescence of leaves consisting of silver hairs concentrated on the adaxial surface, the abaxial surface and petioles glabrous or nearly so; leaves usually rounded at apex, often suffused with purple abaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. hirsutula 20b. Pubescence of leaves variable, but not concentrated on the adaxial surface and often found on the petioles; leaves rounded, to more commonly, obtusely to acutely pointed at apex, without purple coloration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. sororia 19b. Sepals eciliate [Figs. 930, 934]; leaf blades either conspicuously longer than wide or essentially glabrous or both 21a. Sepals lanceolate, narrow-acute to acuminate at apex [Figs. 930, 934] 22a. Leaf blades triangular at chasmogamous anthesis, becoming broad-triangular, later with prominent, coarse teeth near the basal margins that gradually become shorter and narrower toward the apex . . . . . . . . . . . . . . . . . . . . . . . V. pectinata 22b. Leaf blades narrow-ovate, with rather uniform teeth along the margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. affinis 21b. Sepals ovate to narrow-ovate or ovate-oblong, obtuse to rounded at apex [Fig. 937] 23a. Leaf blades longer than wide, often narrow cordate-ovate to triangular; lateral petals spreading, the flowers wide open in life; leaves commonly pubescent . . . . . . . . . . . . . . . . V. novae‑angliae 23b. Leaf blades as wide as or wider than long, often broadcordate-ovate; lateral petal directed forward, the flowers narrow in life; leaves glabrous . . . . . . . . . . . . . . . . . . . . . . . . V. nephrophylla 1b. Plants with leafy aerial stems, the peduncles arising from the axils of leaves [Fig. 935] 24a. Plants annual, with taproots; stipules foliaceous, prominently lobed in pinnatifid fashion; style dilated upward into a globose, hollow stigma with a wide orifice on the lower side 25a. Flowers longer than 15 mm; petals exceeding the sepals by 2 mm or more; corolla variously colored, often yellow-white or yellow-orange with a yellow center and the upper petals dark blue on the apical half or sometimes dark blue throughout with a yellow center . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. tricolor 25b. Flowers up to 10 mm long; petals shorter than the sepals or up to 2 mm longer than the sepals; corolla variously colored, often yellow-white with a yellow center . . . . . . . V. arvensis

Vi o l ac e a e   87 7

24b. Plants perennial from rhizomes; stipules scarious to herbaceous, entire to toothed or lacerate [Fig. 935]; style either slender at the apex or capitate, but not globose 26a. Petals white to cream or violet to purple, the lateral ones pubescent with hairs that taper to the apex (glabrous in V. rostrata); spur 3–18 mm long, usually 2 or more times as long as wide; style slender throughout or only slightly widened apically, not capitate, bent at the apex (straight in V. rostrata) 27a. Petals white to cream; sepals ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. striata 27b. Petals violet to purple; sepals eciliate 28a. Spur 8–15 mm long; lateral petals glabrous; style ± straight . . . . . . V. rostrata 28b. Spur 2.8–7 mm long; lateral petals pubescent on inner surface; style bent near tip 29a. Leaf blades commonly with sparse to dense pubescence, the hairs never restricted to the adaxial surface, the upper blades ovate to triangularlanceolate and mostly truncate at the base; petioles distinctly winged; style slightly expanded apically, terminating in a stout, bent tip as long as the diameter of the style . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. adunca 29b. Leaf blades commonly glabrous, when pubescent the hairs restricted to the adaxial surface, the upper blades usually ovate to orbicular, with a cordate base; petioles not distinctly winged; style of nearly uniform diameter throughout, terminating in a bent tip nearly twice as long as the diameter of the style . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. labradorica 26b. Petals yellow or white with yellow at the base, the lateral ones pubescent with hairs that are dilated and rounded or knob-shaped at the apex; spur less than 2 mm long and less than 2 times as long as wide; style capitate at the apex 30a. Petals entirely yellow; stipules herbaceous, green . . . . . . . . . . . . . . V. pubescens 30b. Petals white with a yellow base, tinged with purple on the outside; stipules scarious or subscarious, pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. canadensis

Key for fruiting and cleis to gamous flowering material 1a. Plants acaulescent, the leaves and peduncles borne directly from rhizomes or stolons [Figs. 933, 936] 2a. Plants producing stolons, these naked or with tiny leaves, often also with cleistogamous flowers from the nodes 3a. Stolons leafy; capsules densely pubescent; seeds 3.4–4 mm long; rhizomes mostly 2–4 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. odorata 3b. Stolons naked or with few leaves; capsules glabrous; seeds 1–2.1 mm long; rhizomes 1–2 (–3) mm wide 4a. Leaf blades 1.5–7 times as long as wide, narrow-cuneate to subcordate at the base 5a. Leaf blades 3–7 times as long as wide, narrowly tapering at the base, the margins denticulate, each tooth with a distinct red-black to black gland at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. lanceolata 5b. Leaf blades 1.5–2.5 times as long as wide, broader at the base, widely tapering to truncate or subcordate at the base, the margins crenate, the teeth with inconspicuous, pale glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. primulifolia 4b. Leaf blades up to 1.5 times as long as wide, cordate at the base 6a. Leaf blades pubescent on one or both surfaces, sometimes sparsely so; capsules spotted or suffused with purple to purple-brown [Fig. 929], the cleistogamous ones borne on prostrate to low-arching peduncles; seeds 1.6–2.1 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. blanda

878  tricolpate s

6b. Leaf blades glabrous on both surfaces; capsules green (often with minute orange dots in V. pallens) [Fig. 934], the cleistogamous ones borne on ascending to arching peduncles; seeds 1–1.7 mm long 7a. Seeds 1–1.4 mm long, brown to nearly black . . . . . . . . . . . . . . . . . . . . V. pallens 7b. Seeds 1.5–1.7 mm long, light gray to light brown . . . . . . . . . . . . . . . V. palustris 2b. Plants not producing stolons 8a. Rhizomes horizontal to ascending, elongate, slender, mostly 1–3 mm thick 9a. Leaf blades cordate-ovate to cordate-suborbicular, acute to obtuse at the apex; basal sinus of leaf blade either narrow or with converging to overlapping lobes; stipules connate to the petiole for a short distance; seeds 1.5–1.9 mm long . . . . . . . . . . V. selkirkii 9b. Leaf blades cordate-orbicular to reniform, rounded to obtuse at the apex; basal sinus of leaf blade wide open; stipules distinct from the petiole; seeds 1.9–2.4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. renifolia 8b. Rhizomes usually ascending, short, stout, mostly 4–6 mm thick 10a. Leaf blades with conspicuous lobes (though the first leaves to appear in V. palmata are unlobed) 11a. Leaf blades oblong-lanceolate to ovate-triangular, 1.5–3 times as long as wide, coarsely serrate to divided into short lobes at the base [Fig. 936]; lobes, if present, less than 50% of the total length of the leaf blade . . . . . . . . . . (in part) V. sagittata 11b. Leaf blades ± ovate in outline, less than 1.5 times as long as wide, with evident lobes (except for the first leaves to emerge which may be unlobed to weakly lobed); lobes often greater than 50% of the total length of the leaf blade 12a. Capsules green [Fig. 934]; sepals lanceolate, narrow-acute to acuminate at the apex, eciliate or ciliate 13a. Foliage pubescent; rhizome fleshy, barrel-like; cleistogamous flowers not produced; sepals usually ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. pedata 13b. Foliage glabrous; rhizome ± firm, not barrel-like; cleistogamous flowers produced—these recognized by the coiled style; sepals eciliate [Fig. 930] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. brittoniana 12b. Capsules spotted or suffused with purple to purple-brown on a green background [Fig. 929]; sepals ovate, rounded to obtuse at the apex, usually at least sparsely ciliate [Fig. 937] 14a. Plants heterophyllous, the early- and late-season leaves unlobed, the middle-season leaves lobed [Fig. 933]; leaf blades divided into 3–5 segments, the middle lobe usually without smaller lateral lobes [Fig. 933] . . . . V. palmata 14b. Plants homophyllous, all of the leaves with evident lobes [Fig. 938]; leaf blades with 5–16 lobes that become progressively smaller and separated by shallower sinuses toward the basal margins, the middle lobe usually with smaller lateral lobes [Fig. 938] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. subsinuata 10b. Leaf blades all unlobed 15a. Leaf blades 1.5–3 times as long as wide, truncate to subcordate at the base, usually more coarsely serrate or even somewhat lobed at the base of the blade (especially in later season leaves; not so in V. japonica) 16a. Stipules distinct; roots white-brown to yellow-brown; leaf blade not, or scarcely, decurrent on the petiole . . . . . . . . . . . . . . . . . . . . . . (in part) V. sagittata 16b. Stipules connate to the petiole; roots dark brown; leaf blade decurrent on petiole . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. japonica

Vi o l ac e a e   87 9

15b. Leaf blades commonly less than 1.75 times as long as wide, definitely cordate at the base, the teeth at the base of the blade no more obvious than those along the middle of the blade margin (except in V. pectinata) 17a. Leaves on spreading petioles, nearly flat to the ground; stipules entire; cleistogamous flowers borne on a horizontal, late-appearing stem, usually 2 or more, the stem appearing raceme-like . . . . . . . . . . . . . . . . . . . V. rotundifolia 17b. Leaves mostly on ascending petioles, held above the ground; stipules ciliate-toothed on the margin; cleistogamous flowers borne singly at the summit of a peduncle 18a. Leaf blades with prominent, coarse (i.e., almost lobe-like) teeth near the basal margins that gradually become shorter and narrower toward the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. pectinata 18b. Leaf blades with rather uniform teeth along the margin 19a. Leaf blades longer than wide, acute to short-acuminate at the apex; sepals eciliate [Figs. 930, 934] 20a. Sepal auricles 2–5.3 mm long on cleistogamous fruits; cleistogamous flowers borne on ascending peduncles; capsules green, 10–15 mm long; seeds dark brown to black-red or olive-black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. cucullata 20b. Sepal auricles 0.5–1.6 mm long; cleistogamous flowers borne on peduncles that are at first prostrate, later ascending from a declined base; capsules spotted or suffused with purple, 6–10 mm long; seeds light yellow-brown to brown (infrequently to dark brown in V. sororia) 21a. Sepals narrow-ovate, rounded to obtuse at apex; peduncles often pubescent; capsules glabrous; seeds brown, sometimes with darker brown markings (rarely yellow-brown) . . . . V. novae‑angliae 21b. Sepals lanceolate, narrow-acute to acuminate at apex; peduncles glabrous [Figs. 930, 934]; capsules sometimes pubescent; seeds light yellow-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. affinis 19b. Leaf blades as wide as or wider than long, mostly rounded to obtuse at the apex; sepals usually ciliate (eciliate in V. nephrophylla) [Fig. 937] 22a. Leaf blades glabrous; peduncles of cleistogamous capsules ascending, not declined at base; capsules green. . . . . . V. nephrophylla 22b. Leaf blades usually with some hairs on one or both surfaces; peduncles of cleistogamous capsules at first prostrate (or even subterranean), later ascending from a declined base; capsules spotted or suffused with purple 23a. Pubescence of leaves consisting of silver hairs concentrated on the adaxial surface, the abaxial surface and petioles glabrous or nearly so; leaves usually rounded at apex, often suffused with purple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. hirsutula 23b. Pubescence of leaves variable, but not concentrated on adaxial surface and often found on the petioles; leaves rounded, to more commonly, obtusely to acutely pointed at apex, without purple coloration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. sororia 1b. Plants with leafy aerial stems, the peduncles arising from the axils of leaves [Fig. 935] 24a. Plants annual from an erect taproot; stipules foliaceous and prominently lobed in a pinnatifid fashion

8 80   tricolpate s

25a. Lower leaf blades usually truncate to subcordate at the base; capsules (6–) 6.5– 9.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. tricolor 25b. Lower leaf blades usually broad-cuneate to truncate at the base; capsules 4–6.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. arvensis 24b. Plants perennial from rhizomes; stipules scarious to herbaceous or subfoliaceous, entire to lacerate-toothed along the margin [Fig. 935] 26a. Stipules scarious to herbaceous, entire to dentate or undulate; capsules 5–12 mm long; stems often produced singly at the summit of the rhizome 27a. Stipules scarious or subscarious, pale, entire; capsules 5–10 mm long; seeds 1.5–2.2 × 1.2–1.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. canadensis 27b. Stipules herbaceous, green, mostly dentate or undulate; capsules 10–12 mm long; seeds 2.5–3.2 × 1.5–1.8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. pubescens 26b. Stipules herbaceous or subfoliaceous, fimbriate to lacerate-toothed (entire in high-elevation plants of V. labradorica); capsules 4–6 mm long; stems often clustered together at the summit of the rhizome 28a. Sepals ciliate; leaf blade margins with closely set, crenulate teeth . . . . V. striata 28b. Sepals eciliate [Figs. 930, 934]; leaf blades variously subentire to remotely or regularly crenulate 29a. Upper stem leaf blades narrow-ovate, tending to be acuminate at the apex; seeds mostly 1.1–1.4 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. rostrata 29b. Upper stem leaf blades triangular-lanceolate to ovate or orbicular, rounded to acute at the apex; seeds mostly 0.8–1.1 mm thick 30a. Leaves commonly sparsely to densely pubescent, the hairs never restricted to the adaxial (i.e., upper) surface only, the blades dark blue-green to dark gray-green in life, subcoriaceous, often with a revolute margin, the upper blades ovate to triangular-lanceolate and mostly truncate at the base; petioles distinctly winged; seeds dark brown to olive-black, mostly 0.8–1.0 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. adunca 30b. Leaves commonly glabrous, when pubescent the hairs restricted to the adaxial surface, the blades light green in life, thin, not or barely revolute, the upper blades usually ovate to orbicular, with a cordate base; petioles not distinctly winged; seeds light brown to brown (dark brown), mostly 1–1.1 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. labradorica 1. Viola adunca Sm. var. adunca N hook-spurred violet. Viola arenaria DC., sensu early American authors; V. aduncoides A. & D. Löve; V. subvestita Greene • MA, ME, NH, VT. Woodlands, sandy fields, roadsides, open rights-of-way, sandplains, clearings. 1 × 10. This very rare violet hybrid is known from MA, NH, VT. It is difficult to detect due to the similarity of the parental species. This hybrid is sometimes best distinguished by the lack of distinguishing characters found in the parents. It frequently shows foliage pubescence closer to Viola adunca but with shorter, thicker spurs (closer to V. labradorica), and is more robust than either parent. Leaf blade outline is typically intermediate. 2. Viola affinis Le Conte N Le Conte’s violet. Viola latiuscula Greene; V. sororia Willd. ssp. affinis (Le Conte) R.J. Little; V. sororia Willd. var. affinis (Le Conte) McKinney • CT, MA, RI, VT. Swamps, stream banks, wetmesic to hydric meadows, shorelines, riparian forests. The distribution presented here is tentative due to tremendous confusion with Viola papillionacea (note: many specimens labeled as this were in fact hybrids between V. cucullata and V. sororia, among other taxa).

Vi o l ac e a e   8 8 1

Viola affinis is the only acaulescent, blue-flowered violet to sometimes produce capsules that are minutely pubescent (all other species in New England have glabrous capsules). ‌2 × 7. Viola ×consocia House is a rare violet hybrid known from CT, VT. It has relatively glabrous herbage and differs from V. cucullata with more cylindrical or weakly clavate hairs of the lateral petals, the sparsely pubescent spurred petal, and capsules spotted with purple. It differs from V. affinis in the more ascending cleistogamous peduncles, the corolla with a dark purple ring around the basal white area (similar to V. cucullata), and the elongate sepal auricles on cleistogamous fruits. ‌2 × 12. Viola ×subaffinis House is a very rare violet hybrid known from VT. It has a broader leaf blade than V. affinis (broad-ovate in the hybrid vs. narrow-ovate), capsules spotted with purple as in V. affinis (green in V. nephrophylla), and seeds that are olivebrown (more like V. nephrophylla, with yellow-brown to olive-brown or dark brown seeds vs. light yellow-brown in V. affinis). 2 ‌ × 25a. Viola ×hollickii House is a very rare violet hybrid known from CT. It has relatively glabrous herbage, narrow-triangular leaf blades with coarse teeth at the base (intermediate between the parental taxa), and relatively upright cleistogamous peduncles (prostrate in V. affinis, ascending in V. sagittata var. ovata). The capsules are sometimes pubescent (as in V. affinis). ‌2 × 27. Viola ×filicetorum Greene is a very rare violet hybrid known from VT. It has leaf blades with less pubescence than V. sororia, sepals usually with some cilia (but sparser than V. sororia), and capsules with a mixture of seed colors, including light yellow-brown (as in V. affinis) to dark brown (as sometimes in V. sororia). 3. Viola arvensis Murr. E European field violet. Viola tricolor L. var. arvensis (Murr.) Boiss. • CT, MA, ME, NH, VT; also reported from RI by George (1992), but specimens are unknown. Fields, waste areas, lawns. 4. Viola blanda Willd. N Fig. 929 sweet white violet.  4b. Viola incognita Brainerd; V. incognita Brainerd var. forbesii Brainerd • CT, MA, ME, NH, RI, VT. Upland and riparian forests, swamps, shaded roadsides, ditches, lawns.

1a. Leaf blades acute at the apex, nearly glabrous, the hairs often restricted to the basal lobes on the adaxial surface, often with a narrow basal sinus, commonly 0.8–1 times as long as wide; peduncles usually tinged or entirely suffused with red and ± glabrous; lateral petals commonly glabrous; upper petals often strongly reflexed . . . . . . . . 4a. V. blanda var. blanda

Fig. 929  Cleistogamous capsule of Viola blanda with a coiled style

1b. Leaf blades obtuse to broad-rounded at the apex, with sparse to dense pubescence on one or both surfaces, often with a wide basal sinus, commonly 0.75–0.85 times as long as wide; peduncles usually green and pubescent; lateral petals commonly pubescent; upper petals not strongly reflexed . . . . . . . . . . . . . . . . . . . . . . . . 4b. V. blanda var. palustriformis Gray Variety blanda is known from CT, MA, ME, NH, RI, VT. Variety palustriformis is known from CT, MA, ME, NH, RI, VT. In general, var. palustriformis is the more frequently encountered variety, especially in the northern half of New England. 5. Viola brittoniana Pollard

N C Fig. 930

coast violet. Viola atlantica Britt.; V. pedatifida G. Don ssp. brittoniana (Pollard) McKinney • CT, MA. Fields, meadows, trail edges, and forest clearings adjacent to rivers and coastal marshes, also peaty river shores. Previous reports of this species in ME were based on collections of a hybrid between Viola brittoniana and V. sororia (specimens at GH!, NEBC!). ‌5 × 7. Viola ×notabilis Bickn. is a very rare violet hybrid known from CT, MA, ME (the ME population is an escape from cultivation, the others are naturally occurring hybrids in wild populations). This hybrid is recognized by its ± glabrous herbage, robust size (characteristic of many V. cucullata hybrids), its variably lobed leaves with lobes broader and shorter than in V. brittoniana, and its longer, more cylindrical hairs on the lateral petals (not conspicuously knob-shaped as in V. cucullata).

Fig. 930  Flower of Viola brittoniana showing lanceolate, pointed, eciliate sepals.

8 82   tricolpate s

5 × 11. This very rare violet hybrid is known from MA. It is relatively easy to distinguish due to the very different morphologies of the parental species. It is a relatively glabrous plant with cyanic corollas, short stolons later in the season, broad-lanceolate to narrowovate leaf blades that are lobed with an elongate central lobe and short lateral lobes (rather than triangular-ovate to suborbicular and with elongate lobes nearly as long as the entire blade in V. brittoniana). 5 × 18. This very rare violet hybrid is known from CT, MA. It has lobed leaf blades somewhat similar to Viola brittoniana except that the lobes are broader relative to length and the sinuses are not as deep. Further, the early leaves are usually less lobed relative to the later leaves. Given the floral and fruiting similarity of V. brittoniana and V. pectinata, the flowers appear as in both taxa (i.e., there is no altering of the reproductive morphology as there would be in other hybrids; e.g., shorter and broader tipped petal hairs in hybrids with V. cucullata, relatively broader sepals with some cilia in hybrids with V. sororia). ‌5 × 25a. Viola ×mulfordae Pollard is a very rare violet hybrid known from CT, MA. It resembles V. brittoniana in its conspicuous lobed leaf blades, but the lobes are shorter and broader than in V. brittoniana and often with an elongate central lobe (contributing to the longer-than-wide aspect of the leaf blade). Further, the petioles and leaf blades are pubescent (glabrous in V. brittoniana). 5 ‌ × 27. Viola ×insolita House is a very rare violet hybrid known from CT, ME. It has variably lobed leaves (with broader and shorter segments than in V. brittoniana) with a prominent central lobe that are at least sparsely pubescent, sepals with cilia (at least on the auricles), prostrate to ascending cleistogamous peduncles, green or purple-spotted capsules, and brown seeds (pale brown to orange-yellow in V. brittoniana, light yellowbrown to brown or dark brown in V. sororia). 6. Viola canadensis L. var. canadensis N Canada white violet. CT, MA, ME, NH, RI, VT; most frequent in western New England, confined to extreme western ME. Rich, mesic, often rocky, deciduous forests. 7. Viola cucullata Ait. N Fig. 931 blue marsh violet. Viola cucullata Ait. var. microtitis Brainerd; V. obliqua Hill • CT, MA, ME, NH, RI, VT; nearly throughout. Swamps, hydric meadows, stream banks, seeps, shorelines, mesic to hydric lawns. Fig. 931  Flower of Viola cucullata showing distribution of short, clavate hairs.

‌7 × 12. Viola ×insessa House is a very rare violet hybrid known from VT. This ± glabrous plant is recognized by its intermediacy and, sometimes, combination of characters from the parental species, including cylindrical petal hairs that may be slightly expanded at the apex, a variably pubescent spurred petal (glabrous in V. cucullata, pubescent in V. nephrophylla), narrow-ovoid to ovoid capsules (mostly 2.2–2.5 times as long as wide in V. cucullata, mostly 1.2–1.5 times as long as wide in V. nephrophylla), and broadlanceolate to narrow-ovate sepals (lanceolate in V. cucullata, ovate to ovate-oblong in V. nephrophylla). ‌7 × 16. Viola ×greenmanii House is a very rare violet hybrid known from CT, MA. It is a sparsely to moderately pubescent plant with variable leaves—some unlobed and some with small lateral lobes and a large middle lobe. The cleistogamous peduncles are variably prostrate to ascending, and the cleistogamous flowers have few (if any) cilia on the sepals and rather elongate sepal auricles. ‌7 × 25a. Viola ×porteriana Pollard is a rare violet hybrid known from CT, MA, ME, NH, VT. It is a sparsely to moderately pubescent plant with narrow-ovate leaf blades, which are transitional in shape between V. cucullata and V. sagittata var. ovata. The sepals usually have some cilia (at least on the auricles), and the petal hairs are often slightly expanded at the apex (i.e., intermediate between the two parents). ‌7 × 27. Viola ×bissellii House is an uncommon violet hybrid known from CT, MA, ME, NH, VT. It is locally common in human-disturbed areas. It is a sparsely pubescent, robust plant with sparsely ciliate sepals (at least on the auricles), slightly expanded petal

V i o l ac e a e   8 83

hairs (i.e., intermediate between the two parents), variably prostrate to ascending cleistogamous peduncles, and a spurred petal that sometimes shows a few hairs. The corolla of this hybrid can be largely blue or is sometimes largely white with a blue ring around the white center. Descriptions of V. papilionacea (as well as many specimens determined as such) fit very well for this hybrid. ‌7 × 29. Viola ×ryoniae House is a very rare violet hybrid known from CT. It is a sparsely pubescent plant with lobed leaf blades (the lobes separated by shallower sinuses than in V. subsinuata). The sepals are usually ciliate (at least sparsely) on the auricles, and the cleistogamous flowers are variably borne on prostrate to ascending peduncles. This plant might be confused with V. palmata or V. subsinuata, but the more elongate sepal auricles, fewer cilia on the sepals, presence of some ascending cleistogamous peduncles, and sparse pubescence during flowering would all point to hybridization with V. cucullata. 8. Viola hirsutula Brainerd NC southern woodland violet. CT; western portion of state. Dry-mesic forests, woodlands, and ridges, dry to seasonally dry fields. Most collections of this plant in regional herbaria are not textbook specimens of Viola hirsutula and appear to be forms of V. sororia. 9. Viola japonica Langsd. ex Ging. E Japanese violet. MA. Gardens, waste areas. This species is responsible for reports of Viola chinensis G. Don in New England (i.e., the voucher specimens were misidentified). 10. Viola labradorica Schrank N American dog violet. Viola adunca Sm. var. minor (Hook.) Fern.; V. conspersa Reichenb. • CT, MA, ME, NH, RI, VT. Forests, forest clearings, fields, roadsides, swamps, ascending to high into the mountains on cliffs and other open areas. See Haines (2001b) for a brief taxonomic history. ‌10 × 23. Viola ×malteana House is a very rare violet hybrid known from CT, MA, VT. It differs from V. labradorica in its longer spur (mostly 8–10 mm long vs. mostly 2.8–4.7 mm) and from V. rostrata in its pubescent lateral petals and spur that is often bent at the tip (rather than glabrous lateral petals and straight spur). ‌10 × 28. Viola ×eclipes H.E. Ballard is a very rare violet hybrid known from MA, RI. It is identified by its very pale blue corolla with a large white center and sparsely ciliate sepals (both character states are intermediate between the parents). It is further distinguished by its heavily fringed stipules, long sepal auricles, and abruptly acuminate leaf blades (in these traits the hybrid is more like V. striata). 11. Viola lanceolata L. ssp. lanceolata N lance-leaved violet. CT, MA, ME, NH, RI, VT. Sandy and/or peaty shorelines, borrow pits, open rights-of-way, ditches, and marshes. 12. Viola nephrophylla Greene N Fig. 932 northern bog violet. CT, MA, ME, NH, VT. Fens, meadows, and river shores in regions of high-pH bedrock and/or till. All reports of this species from RI and many of those in MA were based on misidentified specimens (mainly collections of Viola sororia with relatively sparse pubescence on the foliage and peduncles). ‌12 × 27. Viola ×napae House is a very rare violet hybrid known from VT. It is a sparsely pubescent plant with broad-ovate leaf blades and cleistogamous peduncles variably prostrate to ascending. It differs from V. nephrophylla in its pubescent foliage and sparsely ciliate sepals (at least on the auricles). It differs from V. sororia in its broader leaf blades, sparse sepal cilia, and presence (usually) of some ascending cleistogamous peduncles. 13. Viola novae-angliae House NC New England violet. Viola sororia Willd. var. novae-angliae (House) McKinney • ME; also reported from NH by Pease (1964) and from VT by Atwood et al. (1973), but specimens are unknown. River shores, mainly on ledges and outcrops, less frequently on sand and gravel.

Fig. 932  Flower of Viola nephrophylla showing distribution of elongate, pointed hairs.

8 84   tricolpate s

14. Viola odorata L. E English violet. CT, MA, ME, RI. Fields, roadsides, waste areas, lawns, areas of cultivation. Reports of this species in VT are based on misidentified collections. 15. Viola pallens (Banks ex DC.) Brainerd N smooth white violet. Viola macloskeyi Lloyd ssp. pallens (Banks ex DC.) M.S. Baker; V. rotundifolia Michx. var. pallens Banks ex DC. • CT, MA, ME, NH, RI, VT; nearly throughout. Swamps, seeps, shorelines, ditches, lawns, marshes, ascending high into the mountains along stream banks, pond shores, and mossy areas.

Fig. 933  Leaves of Viola palmata showing heterophyllous habit.

16. Viola palmata L. N Fig. 933 wood violet. Viola palmata L. var. dilatata Ell.; V. triloba Schwein. • CT, MA, ME, NH, RI, VT; restricted in northern New England to southern ME and NH and western VT. Dry-mesic forests, woodlands, clearings, rocky slopes, and fields. Rare forms of this plant have deeply cleft blades (the dilatata form) and could be confused with Viola subsinuata. These can be separated by their 3–7 principal segments, unlobed middle lobes, and occasional presence of petiolules on the outer (i.e., lower) pair(s) of segments (vs. 5–16 principal segments, lobed middle lobes, and absence of petiolules on the outer pair(s) of segments in V. subsinuata). The dilitata form is known from CT and VT. ‌ × 25a. Viola ×robinsoniana House is a very rare violet hybrid known from CT, MA, 16 NH. It has pubescent foliage and narrow-ovate to ovate-oblong leaf blades, some with 1–3 lateral lobes near the base and a prolonged middle lobe (the first leaves and late season leaves unlobed). It differs from V. palmata in the relatively narrower leaf blades and reduced lateral lobes. It differs from V. sagittata in its relatively broader leaf blades and more prominent lateral lobes. ‌16 × 27. Viola ×populifolia Greene is a very rare violet hybrid known from CT, MA, VT. It has pubescent foliage, and usually some of the leaf blades show obscure lateral lobes separated by short sinuses (the first leaves and late season leaves are unlobed). Some forms of this plant (including the type of Viola populifolia) have broad-ovate leaf blades with a shallow basal sinus, a form that could easily be confused with V. sororia, but the cleistogamous capsules will show very few well-formed seeds. Such plants usually also show a broad, but shallow, indentation on each lateral margin of the leaf blade, suggesting a very weak lobing of the blade. 16 ‌ × 29. Viola ×variabilis Greene is a very rare violet hybrid known from CT. It has pubescent foliage, often showing the first few leaves without lobed blades (as in V. palmata), and subsequent leaves with 5- to 7-lobed blades that are more similar to V. subsinuata, and the lateral lobes are broadly dilated and without the characteristic, progressively smaller lobes toward the basal margins of the leaf blade. Late season leaf blades of this hybrid may be scarcely or not at all lobed. 17. Viola palustris L. var. palustris NC northern marsh violet. ME, NH; northern portion of states. Brooks, mossy seeps, meadows, tarn shores, and ravines in alpine areas. The report of this species in VT by Magee and Ahles (1999) is based on a collection of Viola nephrophylla—Ahles 78926 (MASS!). Collections from ME all originate from Katahdin (specimens at CONN!, GH!, MAINE!, NEBC!) and are problematic in that they all white-flowered (unlike fresh specimens from elsewhere in the northeast). Field searches of the collection sites on Katahdin revealed Viola pallens to be very common (i.e., the herbarium collections from ME are likely not V. palustris). Unfortunately, the specimens are all in flower and fruiting characteristics, which would shed light on this problem, are unavailable.

Fig. 934 Chasmogamous capsule of Viola pectinata with a straight style.

18. Viola pectinata Bickn. N C Fig. 934 pectinate-leaved violet. Viola brittoniana Pollard var. pectinata (Bickn.) Alexander • CT, MA. Riverside meadows, upper edge of coastal marshes, pond shores, clearings and trail edges in riparian areas. Viola pectinata has been considered to be merely a leaf-variant form of V. brittoniana (Brainerd 1921, Russell 1965) This is because it is similar in flower and fruit morphology to V. brittoniana and often (but not always) occurs with that species. However, these two species differ dramatically in leaf blade morphology, both in outline and margin.

Vi o l ac e a e   8 8 5

Further, these two species form recognizable hybrids with intermediate leaf characters, indicating additive inheritance, which further indicates the differences seen in leaf blade morphology are not under the control of one or a few genes (i.e., the differences are more profound). Viola pectinata is rarer than V. brittoniana. 19. Viola pedata L. N bird-foot violet. Viola pedata L. var. concolor Holm ex Brainerd; V. pedata L. var. lineariloba DC. • CT, MA, NH, RI; also reported from ME by Gleason and Cronquist (1991), but specimens are unknown. Fields, roadsides, clearings, barrens. 20. Viola primulifolia L. N primrose-leaved violet. Viola primulifolia L. var. acuta (Bigelow) Torr. & Gray • CT, MA, ME, NH, RI; also reported from VT by Russell (1965), but specimens are unknown. Low fields, meadows, shorelines, lawns. 21. Viola pubescens Ait. N Fig. 935 yellow forest violet. 21a. Viola eriocarpon (Nutt.) Schwein.; V. pensylvanica Michx.; V. pubescens Ait. var. eriocarpon Nutt.; 21b. Viola eriocarpon (Nutt.) Schwein. var. leiocarpon Fern. & Wieg.; V. pensylvanica Michx. var. leiocarpon (Fern. & Wieg.) Fern. V. pubescens Ait. var. leiocarpon (Fern. & Wieg.) Seymour • CT, MA, ME, NH, RI, VT. Dry-mesic to mesic, usually deciduous, forests, including upland and riparian types. 1a. Stems solitary, with 0 or 1 basal leaves; leaf blades densely pubescent; stipules broadovate, with an obtuse apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21a. V. pubescens var. pubescens

Fig. 935  Flowers and leaves of Viola pubescens var. scabriuscula.

1b. Stems 2 or more from the apex of the rhizome, with 1–3 basal leaves; leaf blades glabrous or sparsely pubescent; stipules lanceolate to narrow-ovate, with an acute apex [Fig. 935] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21b. V. pubescens var. scabriuscula Torr. & Gray Variety pubescens is known from CT, MA, ME, NH, RI, VT. Variety scabriuscula is known from CT, MA, ME, NH, RI, VT. The latter variety is generally more common in New England, particularly in the northern states, and has an affinity for more mesic (rather than dry-mesic) sites. 22. Viola renifolia Gray N kidney-leaved violet. Viola renifolia Gray var. brainerdii (Greene) Fern. • CT, MA, ME, NH, VT; also reported from RI by Kartesz (1999), but specimens are unknown; becoming rare in southern New England and limited in CT to the northwestern portion of the state. Evergreen to deciduous forests, swamps, trail edges. 23. Viola rostrata Pursh N long-spurred violet. CT, MA, NH, VT; western New England. Rich, mesic, deciduous forests. 24. Viola rotundifolia Michx. N round-leaved violet. Viola clandestina Pursh • CT, MA, ME, NH, RI, VT. Mesic, deciduous, or infrequently mixed evergreen-deciduous, forests and forest edges. 25. Viola sagittata Ait. N C Fig. 936 arrowhead violet.  25a. Viola fimbriatula Sm.; 25b. Viola emarginata (Nutt.) Le Conte; V. emarginata (Nutt.) Le Conte var. acutiloba Brainerd; V. sagittata Ait. var. subsagittata (Greene) Pollard • CT, MA, ME, NH, RI, VT. Fields, roadsides, waste areas, woodlands, clearings, meadows, shorelines, lawns, swamps. 1a. Leaf blades densely short-pubescent, spreading to somewhat ascending, usually ovate to ovate-oblong, serrate near the base of the blade, usually coarsely so; petiole shorter than the blade (sometimes equal to in later season); sepals ciliate; cleistogamous peduncles prostrate to ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25a. V. sagittata var. ovata (Nutt.) Torr. & Gray 1b. Leaf blades glabrous to somewhat short-pubescent, erect or strongly ascending, usually triangular-lanceolate, with small lobes near the base [Fig. 936]; petiole equal to or longer than the blade; sepals eciliate; cleistogamous peduncles erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25b. V. sagittata var. sagittata Variety ovata is known from CT, MA, ME, NH, RI, VT. Variety sagittata is known from CT, MA, ME and is of conservation concern. Reports of this variety from RI and VT (e.g., Seymour 1982) are

Fig. 936  Flower and leaves of Viola sagittata var. sagittata.

8 86   tricolpate s

based on specimens of Viola sagittata var. ovata (though it is expected to occur in RI). Variety sagittata, though often from dry-mesic habitats, is more often found on mesic to wet-mesic substrate than is var. ovata. ‌25a × 25b. Viola ×conjugens Greene is a very rare violet hybrid known from CT, MA. The plants usually have pubescent foliage and vary from displaying intermediacy in characters to combining different character states from the parental taxa. Given that the two varieties forming this hybrid are not always sharply distinct (i.e., intermediate plants are infrequently collected), sterility of capsules is an important character to use to determine hybrid origin (in addition to intermediate or confounding character states). ‌25a × 27. Viola ×fernaldii House is a rare violet hybrid known from MA, ME, NH, VT. It has pubescent foliage with leaf blades that are narrow-ovate (intermediate between the parental taxa). The sepals are ciliate (as in the parents) and are generally more elongate than in V. sororia. The fruits vary from green and obloid to marked with purple and subglobose. 25a × 29. Viola ×convicta House is a very rare violet hybrid known from CT. It has ‌ pubescent foliage with narrow-ovate to ovate leaf blades that show a broad, sometimes elongate, terminal lobe and 3–6 short, sometimes tooth-like, lobes along the basal margin. The bases of the leaf blades vary from cordate to nearly truncate. Sepal characteristics tend to be intermediate between the parents (lanceolate and acute to acuminate at the apex in V. sagittata, ± broad-lanceolate and obtuse at the apex in V. subsinuata). 26. Viola selkirkii Pursh ex Goldie N great-spurred violet. CT, MA, ME, NH, VT. Mesic, deciduous forests and borders, often rocky and/ or rich. 27. Viola sororia Willd. N Fig. 937

Fig. 937  Flower of Viola sororia showing ovate, blunt, ciliate sepals.

woolly blue violet. Viola floridana Brainerd; V. latiuscula Greene; V. palmata L. var. sororia (Willd.) Pollard; V. septentrionalis Greene • CT, MA, ME, NH, RI, VT. Forests, woodlands, riparian terraces, logging roads, forest borders, fields, clearings, roadsides, lawns. Viola sororia is a variable species. In the north, it often shows oblong-ovate to broad-ovate sepals with an obtuse to rounded apex and abundant cilia, hairs on the spurred petal, subglobose to shortovoid capsules, and leaf blades with hairs more dense on the lower surface. These forms have been called V. septentrionalis and can appear distinct from southern forms (V. sororia s.s. with oblong to ovate sepals with an obtuse to acute apex, no hairs on the spurred petal, ovoid capsules, and leaf blades with hairs ± equally dense on both surfaces). However, rangewide variation makes separation of these two morphologies impractical. Relatively grabrous forms are also known and are usually labeled V. papilionacea (these forms have been responsible for recent reports of V. nephrophylla in MA). 27 × 29. This very rare violet hybrid is known from MA. It would most likely be confused with Viola subsinuata but has reduced lobes on the later leaves (leaf blades with median lobes 4–12 mm long rather than 6–37 mm in V. subsinuata). Further, the early leaves tend to be obscurely lobed (rather than more evidently lobed in V. subsinuata). 28. Viola striata Ait. E striped cream violet. CT, MA, NH, RI. Deciduous upland and riparian forests, river banks. Reports of this species from VT are based on cultivated specimens. 29. Viola subsinuata Greene N Fig. 938

Fig. 938  Leaves of Viola subsinuata showing homophyllous habit.

early blue violet. Viola palmata, auct. non L. • CT, MA, RI, VT. Woodlands, rocky ridges, trail edges, especially frequent on trap rock (southern locations) and limestone (northern locations). This species has been confused in the literature and treated as Viola palmata by many authors. Haines (2002) discusses the nomenclature, identification, and distribution of this species. 30. Viola tricolor L. E garden violet. CT, MA, ME, NH, RI, VT. Fields, lawns, gardens, waste areas, near dwellings.

Vi s c ac e a e   8 87

Viscaceae Arceuthobium 1. Arceuthobium pusillum Peck N Fig. 939 dwarf mistletoe. Razoumofskya pusilla (Peck) Kuntze • CT, MA, ME, NH, RI, VT. Parasitic on branches on Picea, Larix, and Pinus.

Vitaceae

Fig. 939  Habit of Arceuthobium pusillum growing on a branch of Picea.

1a. Petals separate at the base but cohering at the apex, deciduous together; bark exfoliating in longitudinal strands, without lenticels [Fig. 943]; pith brown; seeds usually pyriform . . . Vitis 1b. Petals distinct, falling separately; bark close, not exfoliating, with lenticels [Fig. 940]; pith white; seeds ± triangular-obovoid 2a. Nectary disk entirely adnate to the ovary; tendrils terminating in expanded, adhesive disks (without disks in P. vitacea); leaf blades palmately compound (often merely palmately lobed in P. tricuspidata) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parthenocissus 2b. Nectary disk free from ovary and appearing as a cup-shaped ring; tendrils not terminating in adhesive disks; leaf blades simple, toothed to lobed . . . . . . . . . . . Ampelopsis

Ampelopsis Ampelopsis aconitifolia Bunge was reported from CT by Kartesz (1999). The report was erroneous. 1a. Leaf blades toothed to obscurely lobed; branchlets glabrous . . . . . . . . . . . . . . . . . A. cordata 1b. Leaf blades toothed to evidently lobed, at least some on a given plant with 3 or 5 lobes; branchlets pubescent (at least when young) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. glandulosa 1. Ampelopsis cordata Michx. E heart-leaved peppervine. Cissus ampelopsis Pers. • CT. Roadsides, forest fragments. 2. Ampelopsis glandulosa (Wallich) Momiy. var. brevipedunculata (Maxim.) Momiy. E Amur peppervine. Ampelopsis brevipedunculata (Maxim.) Trautv.; A. heterophylla (Thunb.) Sieb. & Zucc. var. amurensis Planch.; Ampelopsis heterophylla (Thunb.) Sieb. & Zucc. var. brevipedunculata C.L. Li; Cissus brevipedunculata Maxim.; Vitis brevipedunculata (Maxim.) Dippel. • CT, MA, NH, RI. Roadsides, waste areas, forest fragments, pond shores, upper edge of tidal marshes.

Parthenocissus 1a. Leaves simple or sometimes compound with 3 leaflets; plants introduced, normally found growing on buildings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. tricuspidata 1b. Leaves compound, usually with 5 leaflets; plants native, normally growing on the ground or on other plants 2a. Tendrils contacting substrate only slightly expanded near apex, the expanded region narrow-oblong in outline and not more than 3 times the width of the ultimate tendril

Fig. 940  Close bark of Ampelopsis glandulosa.

8 88   tricolpate s

segments [Fig. 941, L]; inflorescence mostly 10- to 60-flowered, without a central axis, normally with 2 subequal branches; berry 8–10 mm long; leaflets of mature leaves (i.e., those from the proximal portion of season’s growth) borne on petiolules 5–20 (–30) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. inserta 2b. Tendrils contacting substrate conspicuously expanded near apex, the expanded region oblong to circular in outline and more than 3 times the width of the ultimate tendril segments (though tendrils near shoot apex may lack expanded regions) [Fig. 941, R]; inflorescence mostly 25- to 200-flowered, with a central axis; berry 5–7 mm long; leaflets of mature leaves sessile or borne on petiolules to 10 mm long . . . . . . . . . . . P. quinquefolia 1. Parthenocissus inserta (Kerner) Fritsch N Fig. 941L thicket-creeper. Ampelopsis quinquefolia (L.) Michx. var. vitacea Knerr; Parthenocissus vitacea (Knerr) A.S. Hitchc.; Psedera vitacea (Knerr) Greene • CT, MA, ME, NH, RI, VT. Forests, roadsides, river shores, talus slopes. The specific epithet “vitacea” has been applied to this species by several authors; however, the epithet “inserta” applies to this taxon and has priority (James Pringle, personal communication). Fig. 941  Tendril apices of Parthenocissus inserta (left) and P. quinquefolia (right).

2. Parthenocissus quinquefolia (L.) Planch. N Fig. 941R Virginiana-creeper. Hedera quinquefolia L.; Ampelopsis quinquefolia (L.) Michx.; Parthenocissus quinquefolia (L.) Planch. var. hirsuta (Pursh) Planch.; Psedera quinquefolia (L.) Greene; Vitis quinquefolia (L.) Lam. • CT, MA, ME, NH, RI, VT. Forests, roadsides, river shores, talus slopes. 3. Parthenocissus tricuspidata (Sieb. & Zucc.) Planch. E Boston-ivy. Ampelopsis tricuspidata Sieb. & Zucc. • CT, MA, ME. Buildings and other humanmade structures, rocks, cliffs.

Vitis The leaf blades of Vitis have two very different types of hairs, and distinguishing these pubescence types is important for species recognition. One type of hair lies parallel with the surface of the leaf blade. It is a fine, tangled hair and can be categorized as tomentose. The other type of hair is ± spreading (i.e., it does not lie parallel with the leaf blade surface). This hair is relatively thick and straight compared with the tomentum. Reference: Moore (1991). 1a. Tendrils and/or panicles produced from 3–7 successive nodes; berries 10–20 (–25) mm thick; abaxial surface of leaf blades persistently pubescent with red-brown (later turning gray), tomentose hairs, the surface of the leaf blade largely or completely hidden by the hairs; axis of the inflorescence tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. labrusca 1b. Tendrils and/or panicles produced from no more than 2 successive nodes; berries 5–12 mm thick (to 22 mm thick in V. vinifera); abaxial surface of leaf blades glabrate to sparsely pubescent at maturity, much of the surface of the leaf blade clearly visible (one variety of V. aestivalis more heavily pubescent); axis of the inflorescence glabrous to sparsely pubescent 2a. Abaxial surface of leaf blade not green, either glaucous or somewhat concealed by a thin layer of red-brown tomentum; nodes of branchlets often glaucous . . . . . . V. aestivalis 2b. Abaxial surface of leaf blades green, the pubescence, if present, primarily restricted to the veins or occurring in patches; nodes of branchlets not glaucous 3a. Leaf blades orbicular to suborbicular, often prominently lobed with deep sinuses, commonly with some patches of thin tomentum on the abaxial surface (especially in early season); exocarp of fresh fruit separated from mesocarp only with difficulty . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. vinifera 3b. Leaf blades ovate to orbicular, unlobed or with obscure to evident lobes, lacking tomentose hairs (though usually with straight hairs); exocarp of fresh fruit readily separated from mesocarp

V i tac e a e   8 8 9

4a. Nodal diaphragms 0.5–1 (–2) mm thick; leaf blades with large teeth that taper to thin tips, the teeth mostly 5–20 mm long and with one or both tooth margins concave [Fig. 944]; growing branchlet tips enveloped by enlarged, unfolding leaves; berry prominently glaucous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. riparia 4b. Nodal diaphragms 2–5 mm thick; leaf blades with shorter teeth that do not taper to thin tips, the teeth usually less than 5 mm long and with straight or convex margins [Fig. 942]; growing branchlet tips not enveloped by unfolding leaves; berry not or only thinly glaucous 5a. Branchlets usually green, maturing gray to brown (if with any purple or red coloration, this restricted to one side of the branchlet); leaf blades acute to shortacuminate at the apex; seeds 3–5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . V. vulpina 5b. Branchlets red to purple throughout, maturing red-brown to dark red-brown; leaf blades acuminate to long-acuminate at the apex; seeds 4–7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. palmata 1. Vitis aestivalis Michx. N summer grape.  1a. Vitis lincecumii Buckl. var. glauca Munson; 1b. Vitis aestivalis Michx. var. argentifolia (Munson) Fern.; V. argentifolia Munson; V. bicolor Le Conte; V. lecontiana House • CT, MA, ME, NH, RI, VT. Forests, woodlands, forest fragments, roadsides, rocky and ledgy openings, talus.

Fig. 942  Leaf blade of Vitis labrusca.

1a. Mature leaf blades slightly to strongly pubescent abaxially with a thin layer of tomentum; nodes usually not glaucous; nodal diaphragms usually more than 2 mm thick; mature 3- or 4seeded berries 9–14 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1a. V. aestivalis var. aestivalis 1b. Mature leaf blades glabrous to glabrate abaxially; nodes usually glaucous; nodal diaphragms usually less than 2 mm thick; mature 3- or 4-seeded berries 8–9 mm thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1b. V. aestivalis var. bicolor Deam Variety aestivalis is known from CT, MA, RI. Variety bicolor is known from CT, MA, ME, NH, RI, VT. 2. Vitis labrusca L. N Fig. 942 fox grape. Vitis labrusca L. var. subedentata Fern. • CT, MA, ME, NH, RI, VT. Forests and forest borders, roadsides, clearings, river banks. 2 ‌ × 4. Vitis ×novae-angliae Fern. is an uncommon grape hybrid known from CT, MA, ME, NH, RI, VT. It is distinguished by its abaxially tomentose young leaf blades that become glabrate in later season (sometimes with patches of tomentum remaining), tendrils and/ or panicles produced from 3–5 successive nodes, and berries 12–17 mm thick.

Fig. 943  Exfoliating bark of Vitis riparia

3. Vitis palmata Valh E catbird grape. Vitis rubra Michx. • CT. Railroads, yards. 4. Vitis riparia Michx. N Fig. 943, 944 river grape. Vitis riparia Michx. var. praecox Engelm. ex Bailey; V. riparia Michx. var. syrticola (Fern. & Wieg.) Fern.; V. vulpina L. var. praecox (Engelm. ex Bailey) Bailey; V. vulpina L. ssp. riparia (Michx.) Clausen; V. vulpina L. var. syrticola Fern. & Wieg. • CT, MA, ME, NH, VT. Forests and forest borders, roadsides, clearings, river banks. This species was reported from RI by Kartesz (1999), based on George (1992); however, George (1999) stated it could be in RI based on a misidentified collection made by Richard Champlin (the specimen is Vitis aestivalis var. aestivalis)—24 Jun 2001, Champlin s.n. (Champlin Herb.). 5. Vitis vinifera L. E European grape. Vitis cordifolia Michx. • MA, NH. Dumps, waste areas. 6. Vitis vulpina L. E frost grape. MA. Roadsides, forest fragments and edges.

Fig. 944  Leaf blade of Vitis riparia.

8 90   tricolpate s

Zygophyllaceae Tribulus 1. Tribulus terrestris L. E puncture-vine. MA. Wool waste.

891

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9 17

Index Note: Accepted names are indicated in bold. Synonyms and common names are non-bold.

A

Abelmoschus, 663 esculentus, 663 Abies, 77 balsamea, 77 phanerolepis, 77 concolor, 77 homolepis, 77 menziesii, 79 Abutilon, 663 pictum, 663 striatum, 663 theophrasti, 663 Acadian quillwort, 41 Acalypha, 564 gracilens, 564 delzii, 564 fraseri, 564 rhomboidea, 564 virginica, 565 rhomboidea, 564 Acanthaceae, 307 Acanthopanax ricinifolius, 343 sieboldianus, 341 Acanthospermum, 371 australe, 371 Acer, 849 campestre, 850 carolinianum, 851 dasycarpum, 851 ×freemanii, 851 ginnala, 850 negundo, 850 nigrum, 850 palmatum, 851 pensylvanicum, 851 platanoides, 851 schwedleri, 851 pseudoplatanus, 851 rubrum, 851 tridens, 851 saccharinum, 851 laciniatum, 851 saccharum, 851 nigrum, 850 spicatum, 851 stenocarpum, 851 tataricum, 851 violaceum, 850 Acerates viridiflora, 350 Acetosa oblongifolia, 743 Acetosella corniculata, 697 Achillea, 371 borealis, 371 filipendulina, 371 lanulosa, 371

ligustica, 371 millefolium borealis, 371 lanulosa, 371 nigrescens, 371 occidentalis, 371 ptarmica, 372 tomentosa, 372 Achyranthes repens, 314 Acinos arvensis, 636 Acnida altissima, 317 cannabina, 316 Aconitum, 749 napellus, 750 uncinatum, 750 Aconogonum polystachyum, 736 Acoraceae, 87 Acorus, 87 americanus, 87 calamus, 87 americanus, 87 Acosta diffusa, 387 Acrocentron scabiosa, 387 Acrocystis, 112–114 Acrolophus diffusus, 387 Acrostichum alpinum ilvense, 74 ilvense, 74 Actaea, 750 alba, 750 arguta, 750 ludovici, 750 pachypoda, 750 racemosa, 750 rubra, 750 spicata arguta, 750 rubra, 750 Actinella odorata, 413 Actinidia, 307 arguta, 307 Actinidiaceae, 307 Actinomeris alternifolia, 445 Adam’s-needle, 87 adder’s-mouth, 205–206 Adiantum, 68 aleuticum, 68 hispidulum, 67

pedatum, 68 aleuticum, 68 caldera, 68 viridimontanum, 68 Adicea fontana, 870 pumila, 870 Adlumia, 699 fungosa, 699 Adoxaceae, 307–311 Aegilops, 226 cylindrica, 226 Aegopodium, 337 podagraria, 337 variegata, 337 Aesculus, 851 glabra, 851 hippocastanum, 852 Aethionema saxatile, 475 Aethulia uniflora, 426 Aethusa, 337 cynapium, 337 African-daisy, 376 agalinis, 689–690 Agalinis, 689 acuta, 689 besseyana, 690 flava, 691 maritima, 690 maritima, 690 neoscotica, 690 obtusifolia, 689 paupercula, 690 borealis, 690 neoscotica, 690 paupercula, 690 pedicularia, 691 caesariensis, 691 carolinensis, 691 typica, 691 purpurea, 690 neoscotica, 690 parviflora, 690 tenuifolia, 690 macrophylla, 690 parviflora, 690 tenuifolia, 690 virginica, 691 Agaloma marginata, 568 Agastache, 634 anethiodora, 635 foeniculum, 635 nepetoides, 635 scrophulariifolia, 635 mollis, 635 Agavaceae, 87

Ageratina, 372 altissima, 372 aromatica, 372 Ageratum, 372 altissimum, 372 conyzoides, 372 houstonianum, 372 Agrimonia, 767 bicknellii, 768 eupatoria, 768 gryposepala, 768 microcarpa, 768 mollis, 768 parviflora, 768 platycarpa, 768 pubescens, 768 microcarpa, 768 pumila, 768 rostellata, 768 striata, 768 agrimony, 768 Agropyron, 226 caninum pubescens, 255 cristatum, 226 desertorum, 226 desertorum, 226 pectiniforme, 226 pycnanthum, 285 repens, 255 smithii, 271 trachycaulum, 255 glaucum, 255 majus, 255 Agrostemma, 508 coeli-rosa, 511 coronaria, 513 githago githago, 508 Agrostis, 226 alba, 227 palustris, 228 stolonifera, 228 altissima, 228 borealis, 228 canina, 227 capillaris, 227 elliottiana, 227 exarata, 227 geminata, 228 gigantea, 227 hyemalis, 227 scabra, 228 tenuis, 228 interrupta, 231 latifolia, 242 mertensii, 228 mexicana, 268 minima, 265

918  index

Agrostis, cont. nigra, 227 perennans, 228 racemosa, 268 scabra, 228 septentrionalis, 228 semiverticillata, 277 serotina, 268 sobolifera, 268 spica-venti, 231 stolonifera, 228 gigantea, 227 sylvatica, 268 tenuiflora, 268 tenuis, 227 verticillata, 277 viridis, 277 Ailanthus, 857 altissima, 857 glandulosa, 857 Aira, 228 atropurpurea, 287 canescens, 242 caryophyllea caryophyllea, 228 cespitosa, 244 elongata, 244 flexuosa, 245 melicoides, 262 praecox, 228 spicata, 286 Aizoaceae, 311 Aizopsis, 540 aizoon, 540 ellacombeana, 540 kamtschatica, 540 Ajuga, 635 genevensis, 635 reptans, 635 Akebia, 653 ×pentaphylla, 653 quinata, 653 trifoliata, 653 Albae, 114 Albizia, 575 julibrissin, 575 Alcea, 664 rosea, 664 Alchemilla, 769 monticola, 769 pratensis, 769 vulgaris, 769 xanthochlora, 769 alder, 451 alder-leaved buckthorn, 763 Aletris, 198 farinosa, 198 alfalfa dodder, 534 alga-like pondweed, 295 Alexanders, 347 Alisma, 88 brevipes, 88 gramineum, 88 plantago-aquatica parviflorum, 88 subcordatum, 88 subulatum, 90 triviale, 88 angustissimum, 88 Alismataceae, 88–90 alkali grass, 278

Allegheny crowfoot, 757 Allegheny monkey-flower, 705 Allegheny plum, 800 Allegheny-vine, 699 Alliaceae, 91–92 Alliaria, 474 alliaria, 474 officinalis, 474 petiolata, 474 Allionia albida, 675 hirsuta, 675 linearis, 676 nyctaginea, 676 Allium, 91 burdickii, 92 canadense canadense, 91 robustum, 91 cepa, 91 viviparum, 91 oleraceum, 91 sativum sativum, 91 schoenoprasum, 92 laurentianum, 92 sibiricum, 92 tricoccum, 92 burdickii, 92 tricoccum, 92 vineale, 92 Allocarya californica, 466 reticulata, 466 all-seed flax, 658 Alnus, 450 alnus, 451 crispa mollis, 451 fallacina, 451 glutinosa, 451 incana, 451 americana, 451 rugosa, 451 noveboracensis, 451 rugosa, 451 americana, 451 serrulata, 451 serrulata, 451 subelliptica, 451 viridis, 451 viridis, 451 Alopecurus, 228 aequalis, 229 agrestis, 229 carolinianus, 229 geniculatus, 229 monspeliensis, 277 myosuroides, 229 pratensis, 229 alpine arctic-cudweed, 421 alpine-azalea, 556 alpine-bearberry, 552 alpine bistort, 728 alpine bitter-cress, 481 alpine blueberry, 563 alpine-brook saxifrage, 855 alpine clubsedge, 171 alpine dock, 742 alpine fescue, 259 alpine milk-vetch, 576

alpine northern-rockcress, 479 alpine sweet grass, 231 alpine sweet-vetch, 582 alsike clover, 597 Alsine aquatica, 514 borealis, 522 graminea, 522 holostea, 523 humifusa, 523 longifolia, 523 media, 523 tennesseensis, 522 uliginosa, 522 Alternanthera, 314 pungens, 314 repens, 314 alternate-flowered watermilfoil, 620 alternate-leaved dogwood, 538 alternate milkwort, 726 Althaea, 664 officinalis, 664 rosea, 664 trimestris, 665 Altingiaceae, 311 alum-root, 853 alyssum, 474, 476 Alyssum, 474 alyssoides, 474 calycinum, 474 incanum, 476 maritimum, 488 mutabile, 476 saxatile, 475 Amaranthaceae, 312–329 amaranth, 316–317 interior, 661 subalpina, 373 Amaranthus, 314 albus, 316 pubescens, 316 altissimus, 317 ambigens, 317 ascendens polygonoides, 316 blitoides, 316 blitum, 316 emarginatus, 316 polygonoides, 316 cannabinus, 316 caudatus, 316 chlorostachys erythrostachys, 317 pseudo-retroflexus, 317 cruentus, 316 deflexus, 316 dubius, 316 edulis, 316 flavus, 317 gracilis, 317 graecizans, 316 hybridus, 316 cruentus, 316 hybridus, 316 hypochondriacus, 317 hypochondriacus, 317 leucocarpus, 317 macrocarpus, 317 palmeri, 317 paniculatus, 316

powellii, 315 powellii, 317 pseudo-retroflexus powellii, 317 pumilus, 317 retroflexus, 317 salicifolius, 317 rudis, 317 sanguineus, 316 spinosus, 317 tuberculatus, 317 prostratus, 317 subnudus, 317 viridis, 317 Amaryllidaceae, 92–93 Amberboa, 372 moschata, 372 ambiguous spikesedge, 159 Ambrina ambrosioides, 326 Ambrosia, 373 artemisiifolia, 373 elatior, 373 paniculata, 373 bidentata, 373 coronopifolia, 373 elatior, 373 ×helenae, 373 integrifolia, 373 trifida, 373 monophylla, 373 paniculata, 373 psilostachya, 373 coronopifolia, 373 trifida integrifolia, 373 Amelanchier, 769 arborea, 770 laevis, 771 bartramiana, 770 canadensis, 770 botryapium, 771 oblongifolia, 771 oligocarpa, 770 pauciflora, 770 rotundifolia, 772 gaspensis, 771 humilis, 771 compacta, 771 exserrata, 771 intermedia, 771 laevis, 771 lucida, 771 micropetala, 771 nantucketensis, 771 ×neglecta, 770 oblongifolia, 771 micropetala, 771 oligocarpa, 770 racemosa, 788 sanguinea, 772 gaspensis, 771 spicata, 772 stolonifera, 772 micropetala, 771 ×wiegandii, 771 American alpine speedwell, 720 American-aster, 438–442 American barnyard grass, 252 American beach grass, 229

in d e x  9 19

American beech, 602 American bird’s-foot-treefoil, 588 American bittersweet, 524 American bladdernut, 865 American burnweed, 395 American bur-reed, 303 American chestnut, 601 American climbing fern, 61 American cow-parsnip, 342 American cow-wheat, 694 American cup-scale, 279 American dog violet, 883 American dragonhead, 637 American elm, 868 American false hellebore, 198 American false pennyroyal, 638 American featherfoil, 746 American field chickweed, 510 American germander, 652 American ginseng, 344 American globe-flower, 761 American golden dock, 744 American hazelnut, 454 American hog-peanut, 575 American holly, 352 American honeysuckle, 501 American hornbeam, 454 American hydrangea, 623 American larch, 77 American-laurel, 556 American licorice, 582 American linden, 667 American lop-seed, 705 American lotus, 675 American lyme grass, 264 American manna grass, 262 American marsh-pennywort, 342 American mountain-ash, 818 American plum, 800 American pokeweed, 705 American reed, 273 American sea-blite, 329 American sea-rocket, 479 American shinleaf, 558 American shoreweed, 712 American slough grass, 234 American speedwell, 718 American spikenard, 339 American spongeplant, 176 American spurred-gentian, 611 American squaw-root, 692 American sycamore, 720 American trout-lily, 193 American twinflower, 500 American water-awlwort, 492 American water-horehound, 640 American water-willow, 307 American waterwort, 549 American wild mint, 643 American wisteria, 600 American witch-hazel, 622 American yellow-rocket, 476 American yew, 80 Amerorchis, 200 rotundifolia, 200 amethyst eryngo, 341

Amica mollis petiolata, 376 Ammannia ramosior interior, 661 Ammoglochin, 106 siccata, 115 Ammophila, 229 breviligulata, 229 breviligulata, 230 champlainensis, 229 champlainensis, 229 Amorpha, 575 fruticosa, 575 angustifolia, 575 oblongifolia, 575 tennesseensis, 575 Ampelopsis, 887 cordata, 887 glandulosa brevipedunculata, 887 quinquefolia vitacea, 888 Ampelygonum perfoliatum, 735 Amphicarpaea, 575 bracteata, 575 comosa, 575 comosa, 575 monoica, 575 pitcheri, 575 Amphicarpum, 230 amphicarpon, 230 purshii, 230 Amsinckia, 458 barbata, 458 eastwoodiae, 458 echinata, 458 intermedia, 458 eastwoodiae, 458 echinata, 458 lycopsoides, 458 menziesii, 458 intermedia, 459 menziesii, 459 Amsonia, 347 glaberrima, 347 tabernaemontana tabernaemontana, 347 Amur corktree, 829 Amur honeysuckle, 502 Amur maple, 850 Amur peppervine, 887 Amur silvergrass, 266 Anacardiaceae, 329–331 Anacharis canadensis, 176 densa, 176 nuttallii, 176 Anagallis arvensis, 673 caerulea, 673 caerulea, 673 minima, 674 Anaphalis, 373 margaritacea, 373 angustior, 373 occidentalis, 373 subalpina, 373

Anchusa, 459 arvensis, 459 azurea, 459 officinalis, 459 procera, 459 Andrew’s bottle gentian, 610 Andromeda, 552 baccata, 555 floribunda, 557 polifolia glaucophylla, 552 Andropogon, 230 gerardii, 230 glomeratus, 230 littoralis, 279 nutans, 281 scoparius, 280 ducis, 280 frequens, 280 littoralis, 279 neomexicanus, 280 vimineum, 265 virginicus, 230 androsace, 746 Androsace, 746 arizonica, 746 occidentalis, 746 arizonica, 746 Anemone, 750 acutiloba, 751 americana, 751 blanda, 751 canadensis, 751 cylindrica, 752 alba, 752 globosa, 753 groenlandica, 754 hudsoniana, 752 multifida, 751 globosa, 752 hudsoniana, 752 multifida, 752 nemorosa, 752 bifolia, 752 quinquefolia, 751 bifolia, 752 interior, 752 quinquefolia, 752 riparia, 752 thalictroides, 761 virginiana, 752 alba, 752 riparia, 752 virginiana, 752 Anemonella thalictroides, 761 anemone meadow-rue, 761 Anemonidium canadense, 751 Anethum, 337 graveolens, 337 Angelica, 338 Angelica, 337 atropurpurea, 338 lucida, 338 venenosa, 338 villosa, 338 angelica-tree, 339 Anisantha rigida, 239 rubens, 239

sterilis, 239 tectorum, 239 anise, 343, 345 anise burnet-saxifrage, 345 Anisophyllum dentatum, 567 annual bastard-cabbage, 489 annual blue grass, 275 annual fleabane, 396 annual hair grass, 244 annual hedge-nettle, 651 annual honesty, 488 annual Jacob’s-ladder, 724 annual knawel, 517 annual mercury, 569 annual phlox, 724 annual rabbit’s-foot grass, 277 annual rose-gentian, 612 annual saltmarsh Americanaster, 442 annual sea-blite, 329 annual sea-purslane, 311 annual wall-rocket, 483 annual woolly bean, 594 annual wormwood, 378 anoda, 664 Anoda, 664 cristata, 664 Anomalae, 115 Antennaria, 374 ambigens, 375 brainerdii, 375 campestris, 375 canadensis, 374 fallax, 375 calophylla, 375 howellii, 374 canadensis, 374 neodioica, 374 petaloidea, 374 munda, 375 neglecta, 375 campestris, 375 canadensis, 374 neodioica attenuata, 374 canadensis, 374 petaloidea, 374 rupicola, 374 parlinii, 375 fallax, 375 farwellii, 375 parlinii, 375 petiolata, 375 plantaginifolia, 375 ambigens, 375 arnoglossa, 375 parlinii, 375 petiolata, 375 Antenoron virginianum, 736 Anthemis, 375 arvensis, 375 agrestis, 375 cotula, 375 tinctoria, 392 Anthopogon crinitum, 610

92 0   i n d e x

Anthoxanthum, 230 hirtum, 231 monticola monticola, 231 nitens, 231 odoratum, 231 ovatum aristatum, 231 Anthriscus, 338 cerefolium, 338 sylvestris, 338 Anthyllis, 575 vulneraria, 575 Anticlea, 196 elegans glaucus, 196 Anticosti American-aster, 438 Antiphylla oppositifolia, 855 Antirrhinum, 707 dalmaticum, 712 elatine, 711 majus, 707 minus, 708 orontium, 712 pinifolium, 712 repens, 712 reticulata, 712 reticulatum, 712 sparteum, 712 spurium, 711 Anychia canadensis, 515 dichotoma, 515 nuttallii, 515 polygonoides, 515 Apargia autumnalis, 426 Apera, 231 interrupta, 231 spica-venti, 231 Aphyllon uniflorum, 694 Apiaceae, 331–347 Apios, 576 americana, 576 turrigera, 576 tuberosa, 576 Apium, 338 graveolens, 338 petroselinum, 344 Aplectrum, 200 hyemale, 200 Appalachian barren-strawberry, 791 Appalachian bristle fern, 60 Appalachian gooseberry, 618 Appalachian polypody, 67 Appalachian sandplant, 513 Appalachian sand plum, 801 Appalachian sedge, 142 Appalachian white-aster, 394 apple, 793 apple-of-Peru, 859 appressed bog-clubmoss, 46 Apocynaceae, 347–351 Apocynum, 348 ambigens, 348 androsaemifolium, 348 glabrum, 348

cannabinum, 348 hypericifolium, 348 nemorale, 348 pubescens, 348 ×floribundum, 348 pumilum, 348 sibiricum, 348 Aquifoliaceae, 351–352 Aquilegia, 752 australis, 752 canadensis, 752 australis, 752 coccinea, 752 eminens, 752 latiuscula, 752 vulgaris, 752 Arabidopsis, 475 lyrata lyrata, 475 novae-angliae, 479 thaliana, 475 Arabis, 475 brachycarpa, 477 canadensis, 477 drummondii, 478 brachycarpa, 477 falcata, 477 falcata, 492 hirsuta pycnocarpa, 475 holboellii brachycarpa, 477 laevigata, 477 lyrata, 475 glabra, 475 missouriensis, 477 procurrens, 475 pycnocarpa pycnocarpa, 475 thaliana, 475 Araceae, 93–97 Arachis, 576 hypogaea, 576 Aralia, 338 elata, 339 hispida, 339 nudicaulis, 339 pentaphylla, 341 racemosa racemose, 339 spinosa, 339 Arbutus, 554 alpina, 552 Arceuthobium, 887 pusillum, 887 arching blackberry, 816 Arctanthemum, 375 arcticum polare, 375 arctic bur-reed, 304 arctic-cudweed, 421 arctic-daisy, 375 arctic hair grass, 287 Arctium, 375 lappa, 376 minus, 375–376 nemorosum, 376 tomentosum, 376 Arctostaphylos, 552 adenotricha, 552 alpina, 552

uva-ursi adenotricha, 552 coactilis, 552 Arctotis, 376 grandis, 376 stoechadifolia, 376 grandis, 376 Arctous, 552 alpina, 552 Arenaria, 508 canadensis, 521 caroliniana, 513 glabra, 513 groenlandica, 514 glabra, 513 laricifolia marcescens, 514 lateriflora, 514 leptoclados, 508 macrophylla, 514 marcescens, 514 media, 521 michauxii, 514 peploides robusta, 512 rubella, 514 rubra, 521 serpyllifolia, 508 leptoclados, 508 serpyllifolia, 508 stricta, 514 verna rubella, 514 Arethusa, 200 bulbosa, 200 medeoloides, 205 trianthophora, 212 verticillata, 205 Argemone, 699 alba, 699 albiflora albiflora, 699 intermedia, 699 leiocarpa, 699 mexicana, 699 albiflora, 699 spinosa, 699 vulgaris, 699 Argentina, 772 anserina, 772 concolor, 772 egedii groenlandica, 772 Arisaema, 94 acuminatum, 94 atrorubens, 94 stewardsonii, 94 dracontium, 94 pusillum, 94 stewardsonii, 94 triphyllum, 94 pusillum, 95 triphyllum, 94 Aristida, 232 basiramea, 232 dichotoma dichotoma, 232 geniculata, 232 glauca, 233 gracilis, 232 intermedia, 232

longespica, 232 geniculata, 232 longespica, 232 nealleyi, 233 oligantha, 232 purpurascens purpurascens, 232 purpurea, 233 tuberculosa, 233 Aristolochia, 81 clematitis, 81 convolvulacea, 81 durior, 82 macrophylla, 82 serpentaria, 81 hastata, 81 tomentosa, 82 Aristolochiaceae, 81–82 Armoracia, 475 amphibia, 490 aquatica, 490 armoracia, 475 lacustris, 490 lapathifolia, 475 rusticana, 475 arnica, 376 Arnica, 376 lanceolata lanceolata, 376 Arnoglossum incanum, 716 Arnoseris, 376 minima, 376 Aronia, 773 arbutifolia, 773 atropurpurea, 773 glabra, 773 nigra, 773 atropurpurea, 773 floribunda, 773 melanocarpa, 773 nigra, 773 prunifolia, 773 pyrifolia, 773 Arrhenatherum, 233 elatius, 233 arrow-arum, 96 arrow-grass, 192 arrow-feather threeawn, 232 arrowhead, 90 arrow-head rattlebox, 578 arrowhead violet, 885 arrow-leaved American-aster, 442 arrow-leaved tearthumb, 735 arrowwood, 309–311 Arsenococcus ligustrinus, 557 Artemisia, 376 abrotanum, 378 absinthium, 378 annua, 378 biennis, 378 campestris, 378 canadensis, 378 campestris, 378 caudata, 378 canadensis, 378 carruthii, 378 wrightii, 378

i n d e x  9 2 1

Artemisia, cont. caudata calvens, 378 dracunculus, 377 glauca, 378 frigida, 378 glauca dracunculina, 378 gnaphalodes, 378 ludoviciana, 378 americana, 378 gnaphalodes, 378 ludoviciana, 378 minima, 388 pontica, 379 procera, 378 stelleriana, 379 tridentata, 379 vulgaris, 378 vulgaris, 379 Arthraxon, 233 hispidus, 233 Arum virginicum, 96 Aruncus, 774 allegheniensis, 774 dioicus dioicus, 774 Arundinaria japonica, 277 Arundo canadensis, 240 cinnoides, 240 langsdorfii, 240 Asarum, 81 acuminatum, 81 canadense, 81 acuminatum, 81 reflexum, 81 Asclepias, 348 amplexicaulis, 349 bicknellii, 349 exaltata, 349 incarnata, 349 incarnata, 349 neoscotica, 349 pulchra, 349 intermedia, 350 phytolaccoides, 349 purpurascens, 349 quadrifolia, 349 syriaca, 350 kansana, 350 tuberosa, 350 interior, 350 tuberosa, 350 variegata, 350 verticillata, 350 viridiflora, 350 lanceolata, 350 linearis, 350 Ascyrum hypericoides, 626 ash, 677 Ashe’s variable rosettepanicgrass, 248 ash-leaved maple, 850 ashy cinquefoil, 796 ashy hydrangea, 623 ashy sunflower, 407 Asian bittersweet, 524

Asian virgin’s-bower, 753 Asiatic dayflower, 99 Asiatic sweetleaf, 866 Asiatic tearthumb, 735 Asparagaceae, 98 asparagus, 98 Asparagus, 98 officinalis, 98 Asperugo, 459 procumbens, 459 Asperula, 821 arvensis, 821 odorata, 826 asphodel, 302 Aspidium braunii, 58 cristatum clintonianum, 56 goldianum, 57 intermedium, 57 Aspleniaceae, 51–52 Asplenium, 51 acrostichoides, 73 ×clermontae, 52 cryptolepis, 52 ×ebenoides, 51 montanum, 51 nipponicum, 71 platyneuron, 52 pycnocarpon, 73 rhizophyllum, 52 ruta-muraria, 52 cryptolepis, 52 trichomanes, 52 ×lusaticum, 52 quadrivalens, 52 ramosum, 52 trichomanes, 52 trichomanes-ramosum, 52 tutwilerae, 51 viride, 52 Aspris caryophyllea, 228 praecox, 228 aster Tartarian, 379 Aunt Lucy, 461 Aster, 379 acuminatus, 420 anticostensis, 438 asteroides, 427 borealis, 438 brachyactis, 438 castaneus, 400 chinensis, 384 ciliolatus, 438 commixtus, 400 concolor, 439 cordifolius furbishiae, 439 incisus, 439 laevigatus, 440 polycephalus, 439 sagittifolius, 439 coridifolius, 439 curvescens, 400 divaricatus, 400 dumosus coridifolius, 439 gracilentus, 439 strictior, 439

subulifolius, 439 ericoides, 439 pringlei, 441 prostratus, 439 excavatus, 400 firmus, 442 fragilis, 442 frondosus, 439 glomerata, 400 glomeratus, 400 hirsuticaulis, 440 ianthinus, 400 infirmus, 394 interior, 440 laevis, 440 lanceolatus interior, 440 latifolius, 440 simplex, 440 lateriflorus, 440 angustifolius, 440 hirsuticaulis, 440 linariifolius, 413 victorinii, 413 lindleyanus, 438 loriformis, 442 lowrieanus, 440 lucidulus, 442 macrophyllus, 400 apricensis, 400 excelsior, 400 ianthinus, 400 pinguifolius, 400 velutinus, 400 multiflorus, 439 prostratus, 439 nemoralis, 420 novae-angliae, 440 novi-belgii, 439 ontarionis, 441 glabratus, 441 paniculatus simples, 440 patens, 441 paternus, 427 pilosus demotus, 441 praealtus angustior, 441 prenanthoides, 442 ptarmicoides, 421 puniceus firmus, 442 lucidulus, 442 racemosus, 442 radula, 400 strictus, 400 sagittifolius, 442 urophyllus, 442 saxatilis, 442 schreberi, 400 simplex, 440 interior, 440 ramosissimus, 440 solidagineous, 427 spectabilis, 400 suffultus, 400 strictus, 400 subulatus, 442 obtusifolius, 442 tataricus, 379

tenebrosus, 400 tenuifolious, 442 tradescantii, 442 saxatilis, 442 umbellatus, 394 undulatus, 442 loriformis, 442 vimineus saxatilis, 442 subdumosus, 442 Asteraceae, 352–447 Asterogeum laciniatum, 715 Astragalus, 576 alpinus brunetianus, 576 canadensis canadensis, 576 carolinianus, 576 longilobus, 576 contortuplicatus, 576 eucosmus, 576 glycyphyllos, 577 jesupii, 577 parviflorus, 576 robbinsii, 577 blakei, 576 jesupii, 577 minor, 577 robbinsii, 577 Astrantia, 339 major, 339 Atelophragma elegans, 576 Athyrium, 71 angustum, 71 asplenioides, 71 filix-femina, 71 asplenioides, 71 nipponicum, 71 pycnocarpon, 73 thelypterioides, 73 Atlantic manna grass, 262 Atlantic mock bishop-weed, 345 Atlantic mudwort, 855 Atlantic ninebark, 793 Atlantic white cedar, 75 Atocion, 508 armeria, 508 Atriplex, 317 acadiensis, 319 arenaria, 319 cristata, 319 glabriuscula, 319 acadiensis, 319 hortensis, 319 littoralis, 319 patula, 319 hastata, 319 littoralis, 319 triangularis, 319 subspicata, 320 pentandra, 319 arenaria, 319 prostrata, 319 rosea, 320 sibirica, 320 subspicata, 320 tatarica, 320 triangularis, 319

92 2   i n de x

Atropa physalodes, 859 Aunt Lucy, 461 Aureolaria, 691 flava flava, 691 typica, 691 pedicularia, 691 caesariensis, 691 carolinensis, 691 intercedens, 691 pedicularia, 691 typica, 691 virginica, 691 auricled twayblade, 206 Aurinia, 475 saxatilis, 475 Australian stork’s-bill, 614 Australian wedgeleaf, 384 Austrian pine, 79 Austrian yellow-cress, 490 autumn bentgrass, 228 autumn-crocus, 98 autumn-olive, 548 autumn water-starwort, 708 autumn willow, 848 Avena, 233 dubia, 287 fatua, 233 sativa, 233 flavescens, 286 pubescens, 233 sativa, 233 strigosa, 233 Avenella flexuosa montana, 245 avens, 790–792 Avenula, 233 pubescens, 233 awl American-aster, 441 awl-fruited sedge, 151 awl-leaved arrowhead, 90 awned flatsedge, 155 awnless barnyard grass, 252 awnless wild-rye, 254 axillary goldenrod, 431 Axyris, 320 amaranthoides, 320 azalea, 560 Azalea calendulacea, 560 canadensis, 560 lapponica, 560 prinophylla, 560 procumbens, 556 viscosa, 560 Azolla, 52 caroliniana, 52 Azollaceae, 52 Aztec marigold, 443 Aztec tobacco, 861 azure-blue sage, 648

B baby’s breath, 511–512 Baccharis, 379 halimifolia, 379 angustior, 379 Back’s sedge, 143 Baeothryon alpinum, 171 cespitosum, 171 clintonii, 171 verecundum, 171 Baeria chrysostoma, 416 gracilis, 416 minor, 416 Baeumerta nasturtium-aquaticum, 488 Bailey’s sedge, 149 baked-apple-berry, 813 bald brome, 239 Balkan campion, 519 Balkan clover, 597 ballast toadflax, 712 ball-mustard, 488 balloon-vine, 852 Ballota, 635 borealis, 635 nigra, 635 alba, 635 foetida, 635 nigra, 635 ruderalis, 635 balsam fir, 77 balsam groundsel, 422 Balsaminaceae, 447–448 Balsamita, 379 major, 379 tanacetoides, 379 balsam poplar, 832 balsam willow, 848 Baltic rush, 186 bamboo, 273 bane, 394 baneberry, 750 Baptisia, 577 australis australis, 577 bracteata glabrescens, 577 leucophaea, 577 tinctoria, 577 crebra, 577 projecta, 577 Barbarea, 476 americana, 476 arcuata, 476 orthoceras, 476 dolichocarpa, 476 stricta, 476 verna, 476 vulgaris, 476 arcuata, 476 barber-pole bulrush, 170 barberry, 449 barley, 263 barnyard grass, 252 Barratt’s sedge, 134 barren-strawberry, 791

Bartonia, 608 iodandra, 608 lanceolata, 609 paniculata, 609 intermedia, 608 iodandra, 608 sabulonensis, 608 virginica, 609 paniculata, 609 sabulonensis, 608 bartsia, 694 Bartsia, 694 coccina, 692 bashful clubsedge, 171 basil, 645 basil bee-balm, 644 basil-thyme, 636 basin cat’s-eye, 459 basket willow, 848 Bassia, 320 hirsuta, 320 hyssopifolia, 320 bastard-cabbage, 490 bastard speedwell, 720 bastard-toadflax, 530 Batidea strigosa, 815 Bayard’s adder’s-mouth, 205 bayberry, 671, 847 bayberry willow, 847 bayonet rush, 188 beach-head iris, 180 beach pinweed, 528 beach plum, 801 beach rose, 807 beach vetchling, 584 beach wormwood, 379 beadgrass, 271 beaked corn-salad, 505 beaked ditch-grass, 299 beaked hawk’s-beard, 393 beaked hawkweed, 411 beaked hazelnut, 454 beaked sedge, 149 beaked spikesedge, 161 beaksedge, 164 bean, 592, 594 bearberry, 552 bearberry willow, 848 beard, 445, 774 bearded sedge, 149 bearded sprangletop, 264 beardless rabbit’s-foot grass, 277 beard-lip beardtongue, 713 beardtongue, 713–714 beautybush, 500 Bebb’s sedge, 123 Beckmannia, 235 erucaeformis, 234 syzigachne, 234 Beck’s water-marigold, 382 bedstraw, 822–827 bee-balm, 643–644 bee-blossom, 686–687 beech, 70, 601–602 beech-drops, 692 bee-plant, 529 beet, 320 beetleweed, 546 beggar-tick, 382–383

Behen vulgaris, 520 Belamcanda chinensis, 179 Bellard’s knotweed, 739 bellflower, 495 Bellis, 379 perennis, 379 bellwort, 99 Benthamia lycopsoides, 458 menziesii, 458 kousa, 537 Benthamidia, 537 florida, 537 japonica, 537 bentgrass, 231 bent sedge, 133 Benzoin aestivale, 83 Berberidaceae, 448–449 Berberis, 448 aquifolium, 449 koreana, 449 ×ottawensis, 449 thunbergii, 449 vulgaris, 449 Berchtold’s pondweed, 294 Bermuda grass, 243 Berteroa, 476 incana, 476 mutabilis, 476 Beta, 320 hortensis, 320 maritima, 320 vulgaris, 320 Bethlehem lungwort, 466 Betula, 451 ainus glutinosa, 451 alba cordifolia, 453 alleghaniensis, 453 fallax, 453 macrolepis, 453 alnus rugosa, 451 ×caerulea, 453 cordifolia, 453 crispa, 451 glandulifera, 454 glandulosa, 453 glandulosa glandulifera, 454 lenta, 453 lutea macrolepis, 453 minor, 453 nigra, 453 papyracea minor, 453 papyrifera, 453 cordifolia, 453 pendula, 454 populifolia, 454 pubescens minor, 453 pubescens, 454 pumila, 454

in d e x  9 23

Betula, cont. pumila glandulifera, 454 renifolia, 454 ×sandbergii, 454 saxophila, 453 serrulata, 451 verrucosa, 454 Betulaceae, 450–455 Bicknell’s hawthorn, 782 Bicknell’s sedge, 123 Bicolores, 115 Bicuculla canadensis, 701 cucullaria, 701 eximia, 701 Bidens, 379 alba, 382 radiata, 382 aristosa, 382 mutica, 382 retrorsa, 382 beckii, 382 bipinnata, 382 biternatoides, 382 cernua, 382 anomala, 382 dentata, 382 elliptica, 382 integra, 382 minima, 382 oligodonta, 382 comosa, 383 connata, 382 anomala, 382 fallax, 382 coronata, 383 brachyodonta, 383 tenuiloba, 383 trichosperma, 383 discoidea, 382 eatonii, 382 fallax, 382 interstes, 382 kennebecensis, 382 major, 382 mutabilis, 382 simulans, 382 elegans, 383 frondosa, 383 anomala, 383 caudata, 383 puberula, 383 stenodonta, 383 heterodoxa agnostica, 383 hyperborea, 383 cathancensis, 383 colpophila, 383 laurentiana, 383 svensonii, 383 involucrata, 383 laevis, 381 ×multiceps, 382 odorata, 383 pilosa, 383 minor, 383 polylepis, 383 tenuisecta, 380 trichosperma, 383

tripartita, 383 comosa, 383 vulgata, 383 puberula, 383 biennial bee-blossom, 687 biennial wormwood, 378 Bifolium auriculatum, 206 australe, 206 convallarioides, 206 Bifora, 339 radians, 339 big bluestem, 230 big-bracted-dogwood, 537 big cordgrass, 282 Bigelow’s sedge, 139 big-leaved holly, 352 big-leaved pondweed, 294 Bignonia radicans, 455 tomentosa, 702 Bignoniaceae, 455 big sagebrush, 379 big-toothed poplar, 832 Bilderdykia aubertii, 730 baldschuanica, 730 cilinodis, 730 convolvulus, 730 cristata, 730 dumetorum, 730 scandens, 730 Billings’ sedge, 126 billion-dollar barnyard grass, 252 Biltia vaseyi, 560 Biltmore hawthorn, 782 bindweed, 531–532 birch, 451–455 bird-foot violet, 885 bird’s-eye gilia, 722 bird’s-eye pearlwort, 517 bird’s-eye primrose, 747 bird’s-eye speedwell, 719 bird’s foot, 591 bird’s-foot-treefoil, 588 bird vetch, 599 birthwort, 81 bishop, 339 bishops cap, 854 bishop’s goutweed, 337 bishop-weed, 345 Bismalva alcea, 666 erecta, 666 moschata, 666 pusilla, 666 rotundifolia, 666 sylvestris mauritiana, 666 bistort, 728 Bistorta, 728 officinalis, 728 vivipara, 728 bit, 196 bitter-cress, 481–482 bitter dock, 743 bitter fleabane, 396 bitter-melon, 545 bitternut hickory, 628

bitter panicgrass, 269 bittersweet, 524 Biventraria variegata, 350 black ash, 677 blackberry, 807–817 blackberry-lily, 179 black bindweed, 730 black bugbane, 750 black cherry, 801 black chokeberry, 773 black crowberry, 554 black-cumin, 755 black-edged sedge, 114 black elderberry, 307 black-eyed coneflower, 425 black-fruited spikesedge, 160 black-girdled woolsedge, 169 blackhaw, 311 black henbane, 859 black highbush blueberry, 563 black-horehound, 635 black huckleberry, 555 black jetbead, 802 black knapweed, 387 black locust, 593 black maple, 850 black medick, 590 black mullein, 857 black mustard, 478 black oak, 606 black poplar, 832 black raspberry, 816 black-seeded mountain-rice grass, 273 black-seeded spear grass, 274 black spruce, 78 black swallowwort, 350 blackthorn plum, 801 black tupelo, 538 black walnut, 629 black willow, 847 bladder campion, 520 bladdernut, 865 bladder-pod lobelia, 496 bladder-senna, 578 bladderwort, 654–657 bland sweet-cicely, 344 blanket-flower, 402 blazing star, 417 Blechnaceae, 53 Blechnum virginicum, 53 bleeding-heart, 701 Blephariglottis blephariglottis, 208 ciliaris, 208 cristata, 209 grandiflora, 209 lacera, 209 leucophaea, 210 psycodes, 210 Blephilia, 635 ciliata, 636 hirsuta, 636 glabrata, 636 hirsuta, 636 blessed knapweed, 386 blessed milk-thistle, 428

Blitum capitatum, 323 virgatum, 323 blood amaranth, 316 blood milkwort, 726 bloodroot, 173 blood-root, 702 blood-twig dogwood, 539 bloody crane’s-bill, 616 bluebead-lily, 193 bluebells, 463 blueberry, 561–563 bluebuttons, 500 blue cohosh, 449 bluecurls, 653 blue-eyed grass, 180–181 blue-eyed Mary, 709 blue field-madder, 828 blue flax, 658 blue forget-me-not, 464 blue gilia, 722 blue grama, 234 bluegrass, 275–276 blue ground-cedar, 45 blue huckleberry, 555 blue iris, 180 blue Jacob’s-ladder, 724 blue lupine, 589 blue marsh violet, 882 blue plantain-lily, 173 Blue Ridge false lupine, 595 Blue Ridge sedge, 114 blue sedge, 129 bluestar, 347 bluestem, 230, 279–280 bluet, 827–828 blue vervain, 872 blue water speedwell, 718 blue wild indigo, 577 blue woodruff, 821 blunt broom sedge, 125 blunt-flower rush, 188 blunt-fruited sweet-cicely, 344 blunt-leaved bedstraw, 826 blunt-leaved bog-orchid, 210 blunt-leaved grove-sandwort, 514 blunt-leaved pondweed, 296 blunt-leaved rabbit-tobacco, 424 blunt-leaved waterleaf, 462 blunt-lobed cliff fern, 74 blunt-lobed grapefern, 65 blunt-lobed hepatica, 751 blunt spikesedge, 160 Bocconia cordata, 701 Boebera papposa, 394 Boechera, 477 brachycarpa, 477 canadensis, 477 divaricarpa, 477 grahamii, 477 laevigata, 477 missouriensis, 477 stricta, 478 Boehmeria, 869 cylindrica, 869 bog American-laurel, 556 bog birch, 454

92 4   in de x

bog-clubmoss, 46–47 bog goldenrod, 434 bog Jacob’s-ladder, 725 bog-mat, 97 bog muhly, 268 bog nodding-aster, 420 bog-orchid, 208–210 bog-rosemary, 552 bog stitchwort, 522 bog willow, 847 bog willow-herb, 683 bog yellow-eyed-grass, 305 Bolboschoenus, 102 fluviatilis, 103 glaucus, 103 maritimus, 103 maritimus, 103 paludosus, 103 novae-angliae, 103 robustus, 103 Boltonia, 384 asteroides, 384 latisquama, 384 recognita, 384 latisquama, 384 microcephala, 384 occidentalis, 384 recognita, 384 recognita, 384 boneset thoroughwort, 399 Boott’s rattlesnake-root, 419 borage, 459 Boraginaceae, 456–467 Borago, 459 officinalis, 459 border forsythia, 677 border privet, 678 boreal bedstraw, 825 boreal bog sedge, 134 boreal sandplant, 514 boreal stitchwort, 522 Bosc’s rosette-panicgrass, 248 Boston-ivy, 888 Bothrocaryum alternifolia, 538 Botrychium, 62 angustisegmentum, 63 ascendens, 64 campestre, 64 dissectum, 64 obliquum, 64 lanceolatum angustisegmentum, 64 lunaria, 64 matricariifolium, 65 onondagense, 64 matricariifolium, 65 minganense, 65 multifidum, 65 intermedium, 65 oneidense, 65 onondagense, 64 pallidum, 65 rugulosum, 65 simplex, 65 teneborsum, 65 tenebrosum, 65 virginianum, 65 bottle-brush grass, 254 bottle gourd, 545

Bouteloua, 234 curtipendula curtipendula, 234 filiformis, 234 gracilis, 234 hirsuta, 234 repens, 234 rigidiseta, 234 simplex, 234 Bowman’s-root, 792 Brachiaria texana, 287 Brachyactis angusta, 438 ciliata, 438 angusta, 438 Brachyelytrum, 235 aristosum, 235 erectum, 235 septentrionale, 235 septentrionale, 235 Brachypodium, 235 pinnatum, 235 bracken fern, 54 bracted orache, 319 bracted plantain, 715 bracted strawflower, 447 Bradford pear, 802 Brainerd’s hawthorn, 782 branched bur-reed, 303 branched centaury, 609 branched Indian clover, 597 Brasenia, 84 peltata, 84 schreberi, 84 brassbuttons, 392 Brassica, 478 alba, 491 arvensis, 491 campestris campestris, 478 erucastrum, 485 hirta, 491 juncea, 478 crispifolia, 478 kaber, 491 pinnatifida, 491 napus, 478 nigra, 478 oleracea, 478 acephala, 478 capitata, 478 rapa, 478 Brassicaceae, 467–492 Brauneria pallida, 395 purpurea, 395 Braun’s holly fern, 58 Braxilia minor, 558 Braya, 479 humilis humilis, 479 leiocarpa, 479 novae-angliae, 479 Brazilian water-meal, 97 Brazilian-waterweed, 176 bread wheat, 287 Breea arvensis, 389

bridalwreath meadowsweet, 820 bristle-fruited hedge-parsley, 346 bristle-leaved sedge, 114 bristle-mallow, 666 bristly blackberry, 814 bristly crowfoot, 759 bristly greenbrier, 301 bristly hawkbit, 416 bristly hawk’s-beard, 393 bristly locust, 593 bristly ox-tongue, 409 bristly rose, 805 bristly sarsaparilla, 339 bristly sheepburr, 462 bristly smartweed, 735 bristly swamp currant, 618 bristly wolfstail, 267 brittle-stemmed hemp-nettle, 637 brittle waternymph, 177 Briza, 235 media, 235 minor, 235 broad beech fern, 70 broad-leaved cat-tail, 304 broad-leaved enchanter’snightshade, 681 broad-leaved helleborine, 203 broad-leaved ironweed, 446 broad-leaved pepperweed, 487 broad-leaved rosettepanicgrass, 249 broad-leaved sedge, 116 broad-leaved Solomon’s-seal, 300 broad-leaved speedwell, 719 broad-leaved spurge, 568 broad-leaved twayblade, 206 broad loose-flowered sedge, 133 broad vetch, 599 broad-winged sedge, 123 brome, 237–239, 287 brome-like sedge, 125 brome six-weeks grass, 287 Bromus, 235 altissimus, 238 arvensis, 237 breviaristatus, 239 briziformis, 238 canadensis, 238 carinatus linearis, 239 marginatus, 239 ciliatus, 238 intonsus, 238 commutatus, 238 dertonensis, 287 diandrus rigidus, 239 erectus, 238 hordeaceus, 238 hordeaceus, 238 pseudothominei, 238 thominei, 238 inermis, 236 inermis, 238 japonicus, 238 kalmii, 236, 238

latiglumis, 238 lepidus, 239 marginatus, 239 mollis, 238 pubescens, 239 pumpellianus, 238 purgans, 239 racemosus, 239 rigidus, 236 rubens, 239 secalinus, 239 velutinus, 239 squarrosus, 239 sterilis, 239 tectorum, 239 thominei, 238 brook cinquefoil, 797 brook lobelia, 496 brookweed, 866 broom, 124–125, 579 broom-crowberry, 553 broom-leaved toadflax, 712 broom-like ragwort, 427 broom-rape, 694 broomsedge bluestem, 230 Broussonetia, 670 papyrifera, 670 Browallia, 858 americana, 858 brown beaksedge, 164 brown bog sedge, 145 brown cudweed, 403 brown flatsedge, 154 brown-fruited rush, 188 brownish beaksedge, 164 brownish sedge, 126 brown knapweed, 387 Brunet’s hawthorn, 785 Bruniera columbiana, 97 Bryocles ventricosa, 173 Bryodesma rupestre, 49 buck-bean, 669 buckeye, 851–852 buck’s-horn plantain, 715 buckthorn, 763 buckwheat, 727 budding pondweed, 295 Buddleja, 855 davidii, 855 buffalo-berry, 548 bugbane, 750 bugle, 635 bugloss, 459–461 bugloss fiddleneck, 458 Buglossoides, 459 arvensis arvensis, 459 bulbil fragile fern, 72 bulblet-bearing water-hemlock, 340 Bulbostylis, 104 capillaris, 104 bulbous bitter-cress, 481 bulbous blue grass, 275 bulbous crowfoot, 758 bulbous wood rush, 191 bullhead pond-lily, 86

in d e x  9 25

Bulliarda aquatica, 541 bulrush, 166–170 bunch-flower, 198 bunch-flowered soft rush, 187 Bunias, 479 edentula, 479 orientalis, 479 Buphthalmum helianthoides, 408 Bupleurum, 339 lancifolium, 340 odontites, 340 rotundifolium, 340 burdock, 376 burgrass, 286 burhead, 88 bur medick, 590 burnet, 817 burnet-saxifrage, 345 burning nettle, 871 burnweed, 395 burr-cucumber, 545 bur-reed sedge, 143 bur-reed, 303–304 burr oak, 605 burrowing clover, 598 Bursa bursa-pastoris, 479 bush-clover, 535, 586–588 bush-clover dodder, 535 bush-honeysuckle, 500 Bush’s sedge, 144 bushy American-aster, 439 bushy bluestem, 230 bushy frostweed, 526 bushy St. John’s-wort, 626 Burshia humilis, 621 Butomaceae, 98 Butomus, 98 umbellatus, 98 butter-and-eggs toadflax, 712 butterbur sweet-coltsfoot, 423 butterfly-bush, 855 butterfly milkweed, 350 butterwort, 654 buttonbush, 534, 821 buttonbush dodder, 534 button eryngo, 341 button grass, 243 button sedge, 149 buttonweed, 822 Buxaceae, 493

C cabbage, 478–479 Cabomba, 85 caroliniana pulcherrima, 85 pulcherrima, 85 Cacalia suaveolens, 427 Cactaceae, 493 Cactus humifusus, 493

Cakile, 479 americana, 479 edentula edentula, 479 lanceolata, 479 maritima americana, 479 Calamagrostis, 239 canadensis, 240 canadensis, 240 langsdorfii, 240 macouniana, 240 cinnoides, 239–240 coarcta, 240 epigejos, 239 georgica, 240 fernaldii, 240 inexpansa novae-angliae, 240 lacustris, 241 langsdorfii, 240 nubila, 240 pickeringii, 240 stricta, 240 inexpansa, 241 stricta, 241 Calamintha acinos, 636 vulgaris, 636 Calandrinia, 745 ciliata, 745 Calendula, 384 officinalis, 384 Calestania palustris, 344 calico American-aster, 440 California goldfields, 416 California oatgrass, 243 California-poppy, 701 California privet, 678 California suncup, 680 calla, 95 Calla, 95 palustris, 95 Calliopsis basalis, 391 Callistephus, 384 chinensis, 384 hortensis, 384 Callitriche, 707 anceps, 708 austinii, 708 autumnalis, 708 deflexa, 708 austinii, 708 hermaphroditica, 708 heterophylla, 708 heterophylla, 708 palustris, 708 verna, 708 stagnalis, 708 terrestris, 708 Calluna, 553 vulgaris, 553 Calopogon, 201 pulchellus, 201 tuberosa tuberosus, 201 Calotis, 384 cuneifolia, 384

Caltha, 752 asarifolia, 752 palustris, 752 asarifolia, 752 flabellifolia, 752 Calycanthaceae, 82 Calycanthus, 82 fertilis, 82 floridus, 82 floridus, 82 glaucus, 82 glaucus, 82 Calylophus serrulatus, 687 Calypso, 201 bulbosa americana, 201 Calystegia, 531 americanus, 531 fraterniflora, 532 japonica, 531 pellitus anestius, 531 pubescens, 531 sepium, 531 americana, 532 angulata, 532 appalachiana, 532 fraterniflorus, 532 repens, 531 sepium, 532 silvatica, 532 fraterniflora, 532 spithamaea, 532 spithamaea, 532 camas, 196 Camelina, 479 microcarpa, 479 sativa microcarpa, 479 sativa, 479 Camissonia, 680 bistorta, 680 campestris campestris, 680 Campanula, 494 aparinoides, 495 carpatica, 495 divaricata, 495 flexuosa, 495 glomerata, 495 lactifolia, 495 lanata, 495 latifolia, 495 patula, 495 persicifolia, 495 punctata, 495 rapunculoides, 495 rotundifolia, 495 trachelium, 495 uliginosa, 495 Campanulaceae, 493–497 Campe barbarea, 476 stricta, 476 verna, 476 camphor false golden-aster, 409 camphorweed, 423 campion, 519–520, 523

Campsis, 455 radicans, 455 Camptosorus rhizophyllus, 52 Canada buffalo-berry, 548 Canada burnet, 817 Canada clearweed, 870 Canada dwarf-dogwood, 537 Canada fleabane, 396 Canada frostweed, 526 Canada goldenrod, 431 Canada hawkweed, 412 Canada honewort, 341 Canada lily, 194 Canada-mayflower, 300 Canada milk-vetch, 576 Canada moonseed, 669 Canada mountain-rice grass, 273 Canada plum, 801 Canada reed grass, 240 Canada rockcress, 478 Canada rush, 187 Canada sand-spurry, 521 Canada sanicle, 345 Canada shore quillwort, 42 Canada white violet, 882 Canada wild ginger, 81 Canada windflower, 751 Canada wood-nettle, 869 Canadian single-spike sedge, 145 Canby’s bulrush, 166 cancerwort, 711 candytuft, 486 canescent whitlow-mustard, 484 Cannabaceae, 497–498 Cannabis, 497 sativa, 497 indica, 497 cantaloupe, 545 capitate sedge, 115 Capituligerae, 115 Capnoides, 699 aureum, 700 flavulum, 700 glauca, 699 sempervirens, 699 Capnorchis canadensis, 701 cucullaria, 701 eximia, 701 formosa, 701 Caprifoliaceae, 498–505 Capsella, 479 bursa-pastoris, 479 Caragana, 577 arborescens, 577 Carara coronopus, 488 didyma, 487 caraway, 340 Cardamine, 479 bellidifolia, 480 bellidifolia, 481 bulbosa, 481 concatenata, 481 dentata, 482 diphylla, 482 douglassii, 482

92 6   in de x

Cardamine, cont. flexuosa, 482 hirsuta, 482 impatiens, 482 incisa, 482 longii, 482 maxima, 482 parviflora, 481 arenicola, 482 pensylvanica, 482 pratensis, 482 Cardaria draba, 487 latifolia, 487 Cardiospermum, 852 halicacabum, 852 Carduus, 384 acanthoides, 384 altissimus, 389 crispus, 384 discolor, 390 flodmanii, 390 lanceolatus, 390 muticus, 390 nutans, 385 leiophyllus, 385 palustris, 390 pumilus, 390 spinosissimus, 390 carelessweed, 394 Carey’s smartweed, 734 Carex, 104, 115 abacta, 145 abdita, 114 abscondita, 116 glauca, 116 acuta nigra, 140 sparsiflora, 141 acutiformis, 136 adusta, 123 aenea, 119 aestivalis, 130 alata, 122 alata, 123 albicans, 113 albolutescens, 123 albursina, 132 alopecoidea, 150 alpina, 145 amphibola, 129 turgida, 129 annectens, 135 appalachica, 143 aquatilis, 138 altior, 139 substricta, 139 arcta, 126 arctata, 130 faxonii, 130 arctogena, 115 argyrantha, 123 artitecta, 113 asa-grayi, 134 atherodes, 115 atlantica, 146 capillacea, 146 atrata atratiformis, 145 atratiformis, 145 aurea, 115

backii, 143 subrostrata, 143 baileyi, 149 barrattii, 134 bebbii, 123 bicknellii, 123 bigelowii, 139 billingsii, 126 blanda, 133 brachyglossa, 135 brevior, 123 bromoides bromoides, 125 brunnea, 127 nakiri, 127 brunnescens, 126 brunnescens, 127 sphaerostachya, 126 bullata, 149 bushii, 144 buxbaumii, 145 camporum, 125 canescens, 127 alpicola, 126 canescens, 127 disjuncta, 127 polystachya, 126 capillaris, 118 elongata, 118 robustior, 118 capitata arctogena, 115 caroliniana cuspidata, 144 caryophyllea, 135 castanea, 131 cephaloidea, 142 cephalophora, 142 mesochorea, 142 chordorrhiza, 117 clivicola, 114 collinsii, 118 communis, 112 communis, 113 comosa, 149 complanata hirta, 144 conoidea, 129 contigua, 143 costata, 144 crawei, 128 crawfordii, 119, 123 crinita, 139 brevicrinis, 139 crinita, 140 gynandra, 140 mitchelliana, 138 paleacea, 140 simulans, 140 cristatella, 120, 123 cryptolepis, 117 cumulata, 123 davisii, 131 debilis, 131 allegheniensis, 131 debilis, 131 intercursa, 131 interjecta, 131 rudgei, 131 strictior, 131

deflexa, 114 demissa, 117 deweyana, 125 diandra, 129 ramosa, 129 digitalis, 116 dioica, 143 gynocrates, 143 disperma, 125 echinata, 146 echinata, 147 echinodes, 123 elachycarpa, 147 emmonsii, 113 muehlenbergii, 113 emoryi, 140 exilis, 147 festucacea, 123 flacca, 147 flaccosperma, 129 flava fertilis, 117 gaspensis, 117 flexilis, 131 flexuosa, 131 foenea, 115 foenea, 119 folliculata, 145 formosa, 131 fuscidula, 118 garberi, 115 bifaria, 115 glauca, 147 glaucodea, 129 gracilescens, 133 gracillima, 131 granularis, 128 haleana, 128 grayi, 134 hispidula, 134 grisea, 129 gynandra, 140 gynocrates, 143 haydenii, 140 hirsutella, 144 hirta, 116 hirtifolia, 129 hitchcockiana, 129 hormathodes, 124 houghtoniana, 136 howei, 146 hystericina, 149 incomperta, 146 inops, 114 interior, 147 intumescens, 134 fernaldii, 134 globularis, 134 katahdinensis, 129 kattegatensis, 141 ×knieskernii, 131 kobomugi, 135 lacustris, 136 laevivaginata, 150 lanuginosa, 136 americana, 136 lasiocarpa americana, 136 latifolia, 136 laxiculmis, 116 laxiflora, 132

angustifolia, 133 blanda, 133 gracillima, 133 latifolia, 132 leptonervia, 133 ormostachya, 133 serrulata, 132 lenticularis, 140 albimontana, 140 blakei, 140 leporina, 124 leptalea, 133 leptonervia, 133 limosa, 134 rariflora, 134 livida, 137 grayana, 137 radicaulis, 137 longii, 124 longirostris, 131 lucorum, 114 lupuliformis, 134 lupulina, 135 pedunculata, 135 lurida, 149 mackenziei, 127 macrocephala kobomugi, 135 magellanica, 133 irrigua, 134 marginata, 114 meadii, 137 media, 145 merritt-fernaldii, 124 mesochorea, 142 michauxiana, 145 miliaris, 149 millegrana, 140 mirabilis tincta, 125 mitchelliana, 138 molesta, 122 monile, 150 muehlenbergii, 142 enervis, 142 muehlenbergii, 142 muricata cephaloidea, 142 sterilis, 147 nigra, 141 strictiformis, 140 nigromarginata, 114 elliptica, 114 muehlenbergii, 113 normalis, 122 norvegica, 127 inferalpina, 145 novae-angliae, 114 oederi, 117 pumila, 117 oklahomensis, 151 oligocarpa, 129 hitchcockiana, 129 oligosperma, 149 ×olneyi, 149 ormostachya, 132 oronensis, 124 ovalis, 124 ovata, 145 paleacea, 140 transatlantica, 140

i n d e x  9 27

Carex, cont. pallescens, 144 neogaea, 144 panicea, 137 pauciflora, 133 paupercula, 134 irrigua, 134 peckii, 114 pedunculata, 118 pellita, 136 pensylvanica, 114 distans, 114 plana, 142 plantaginea, 116 platylepis, 124 platyphylla, 116 polygama, 145 polymorpha, 137 porteri, 139 praegracilis, 125 prairea, 129 prasina, 131 praticola, 124 projecta, 124 pseudocyperus, 149 ×pseudohelvola, 127 pubescens, 129 radiata, 143 rariflora, 134 recta, 141 retroflexa, 143 retrorsa, 149 reznicekii, 114 rhomalea, 149 richardsonii, 118 rigida, 139 bigelowii, 139 rosea, 143 radiata, 143 rostrata, 149 utriculata, 150 ×ruedtii, 117 rugosperma, 114 tonsa, 114 salina kattegatensis, 141 saltuensis, 137 saxatilis, 149 miliaris, 149 rhomalea, 149 scabrata, 115 schweinitzii, 150 scirpoidea scirpoidea, 145 scirpoides, 147 scoparia, 124 tesselata, 124 seorsa, 147 setacea, 135 ambigua, 135 silicea, 124 sparganioides, 143 cephaloidea, 142 spicata, 143 sprengelii, 131 squarrosa, 145 stellulata conferta, 146 radiata, 143

×stenolepis, 150 sterilis, 147 stipata laevivaginata, 150 oklahomensis, 151 stipata, 151 straminea, 120 brevior, 123 crawei, 123 cumulata, 123 invisa, 124 striata, 136 brevis, 136 striatula, 133 stricta, 141 decora, 140 elongata, 140 emoryi, 140 haydenii, 140 strictior, 141 styloflexa, 133 subloliacea, 127 substricta, 139 super-goodenoughii, 141 swanii, 144 sychnocephala, 125 sylvicola, 143 tenera, 125 echinodes, 123 tenuiflora, 125 tetanica, 137 meadii, 137 tincta, 125 tonsa, 114 rugosperma, 114 tonsa, 114 torta, 141 composita, 141 tribuloides, 125 ×trichina, 127 trichocarpa, 116 trisperma, 127 billingsii, 126 tuckermanii, 150 typhina, 146 umbellata, 114 tonsa, 114 utriculata, 150 vacillans, 141 vaginata, 137 vesicaria, 150 jejuna, 150 laurentiana, 150 monile, 150 raeana, 150 vestita, 136 virescens, 144 swanii, 144 viridula, 117 oedocarpa, 117 viridula, 117 vulpinoidea, 135 walteriana, 136 wiegandii, 147 willdenowii, 143 woodii, 137 xanthocarpa annectens, 135 Careyanae, 116 Carolina bloodroot, 173

Carolina bristle-mallow, 666 Carolina crane’s-bill, 616 Carolina fanwort, 85 Carolina meadow-foxtail, 229 Carolina mosquito fern, 52 Carolina nightshade, 864 Carolina rhododendron, 560 Carolina rose, 806 Carolina sea-lavender, 721 Carolina silverbell, 866 Carolina spring-beauty, 745 Carolina whitlow-mustard, 484 carp grass, 233 carpet bugle, 635 carpetweed, 669 Carpinus, 454 caroliniana virginiana, 454 Carpogymnia continentale, 73 dryopteris, 73 carrion-flower, 301 carrot, 341 Carruth’s wormwood, 378 Carthamus, 385 lanatus, 385 tinctorius, 385 Carum, 340 carvi, 340 carved quillwort, 42 Carya, 627 alba, 629 cordiformis, 627 latifolia, 628 demareei, 628 glabra, 628 hirsuta, 628 megacarpa, 628 laciniosa, 628 laneyi, 628 ovalis, 628 hirsuta, 628 ovata, 627, 629 pubescens, 629 tomentosa, 629 subcoriacea, 629 Caryophyllaceae, 505–523 Case’s ladies’-tresses, 211 Cassandra calyculata angustifolia, 553 latifolia, 553 Cassia chamaecrista, 578 fasciculata, 578 hebecarpa, 594 nictitans, 578 obtusifolia, 594 Cassiope hypnoides, 555 Castalia leibergii, 86 odorata, 86 tuberosa, 86 Castanea, 601 alnifolia, 601 floridana, 601 ashei, 601 dentata, 601 floridana, 601 mollissima, 601

pumila, 601 ashei, 601 sativa, 601 Castilleja, 691 coccinea, 692 exserta, 692 pallida septentrionalis, 692 septentrionalis, 692 castor-aralia, 343 castor-bean, 569 catalpa, 455 Catalpa, 455 bignonioides, 455 catalpa, 455 ovata, 455 speciosa, 455 Catapodium rigidum, 245 Catawba rosebay, 560 catbird grape, 889 catchfly, 508, 511 Cathartolinum catharticum, 658 curtissii, 658 intercursum, 658 medium, 658 striatum, 658 multijugum, 658 sulcatum, 658 virginianum, 658 catnip, 635, 645 catnip giant-hyssop, 635 cat’s ear, 413 cat’s eye, 459 cat-tail, 304 cattail sedge, 146 Caulinia flexilis, 177 guadalupensis, 177 minor, 177 Caulophyllum, 449 giganteum, 449 thalictroides, 449 giganteum, 449 Ceanothus, 762 americanus, 762 intermedius, 762 pitcheri, 762 herbaceus, 762 pubescens, 762 intermedium, 762 ovatus, 762 pubescens, 762 cedar, 75–76 cedar glade St. John’s-wort, 626 celandine, 700 Celastraceae, 523–525 Celastrus, 523 alatus, 524 orbiculatus, 524 scandens, 524 celery, 338 Celosia, 321 argentea cristata, 321 cristata, 321

92 8   in d e x

Celtis, 497 occidentalis, 497 canina, 497 pumila, 497 Cenchrus, 241 carolinianaus, 241 echinatus, 241 incertus, 241 longispinus, 241 pauciflorus, 241 racemosa, 286 spinifex, 241 Centaurea, 385 adulterina, 386 biebersteinii, 388 benedicta, 386 calcitrapa, 388 cyanus, 387 debeauxii thuillieri, 387 diffusa, 388 dubia, 387 nigrescens, 387 vochinensis, 387 jacea, 387 decipiens, 387 pratensis, 387 maculosa, 388 melitensis, 388 ×moncktonii, 387 montana, 387 moschata, 372 nigra, 387 nigrescens, 387 phrygia, 387 pratensis, 387 ×psammogena, 387 pulchra, 387 scabiosa, 387 segetum, 387 solstitialis, 388 stoebe, 388 micranthos, 388 thuillieri, 387 transalpina, 387 vochinensis, 387 Centaurella paniculata, 609 Centaurium, 609 erythraea, 609 pulchellum, 609 spicatum, 612 umbellatum, 609 centaury, 609, 612 Centipeda, 388 minima, 388 centipede grass, 258 Centunculus minimus, 674 Cephalanthus, 821 occidentalis, 821 pubescens, 821 Cephalaria, 499 gigantea, 499 Cerastium, 508 angustatum, 510 aquaticum, 514 arvense, 509 angustifolium, 510 latifolium, 510 strictum, 510

campestre, 510 diffusum, 509 fontanum, 509 glomeratum, 510 apetalum, 510 longepedunculatum, 510 nutans, 509 nutans, 510 pubescens, 510 pumilum, 509 pumilum, 510 semidecandrum, 510 strictum, 510 tetrandrum, 509 tomentosum, 510 vestitum, 510 viscosum, 510 vulgatum, 509 Cerasus avium, 800 vulgaris, 800 Ceratochaete aquatica, 288 Ceratocystis, 117 Ceratophyllaceae, 82–83 Ceratophyllum, 82 apiculatum, 82 demersum, 82 demersum echinatum, 83 echinatum, 83 Ceratoschoenus macrostachys inundata, 164 Ceratoxalis cymosa, 697 Cercidiphyllaceae, 525 Cercidiphyllum, 525 japonicum, 525 Cercis, 578 canadensis canadensis, 578 Cerefolium cerefolium, 338 Cespa aquatica, 172 Chaenactis, 388 glabriuscula glabriuscula, 388 Chaenomeles, 774 japonica, 774 lagenaria, 774 speciosa, 774 Chaenorhinum, 708 minus, 708 Chaerophyllum sylvestre, 338 chaff-seed, 695 chaffweed pimpernel, 674 chaffy sedge, 140 chain-tree, 583 chair-maker’s bulrush, 166 Chamaecistus procumbens, 556 Chamaecrista, 578 fasciculata, 578 nictitans nictitans, 578 procumbens, 578 Chamaecyparis, 75 thyoides, 75, 118

Chamaedaphne, 553 calyculata, 553 angustifolia, 553 Chamaelirium, 196 luteum, 196 Chamaepericlymenum, 537 canadense, 537 Chamaesaracha grandiflora, 859 Chamaesyce glyptosperma, 568 maculata, 568 nutans, 568 polygonifolia, 568 prostrata, 568 rafinesquei, 568 serpyllifolia, 568 Chamerion, 681 angustifolium circumvagum, 681 platyphyllum, 681 chamomile, 375, 392, 418 Chamomilla chamomilla, 418 inodora, 445 maritima, 445 recutita, 418 suaveolens, 418 Chapman’s blue grass, 275 charlock, 491 cheat brome, 239 checkered rattlesnake-plantain, 204 Cheilanthes, 68 lanosa, 68 vestita, 68 Cheiranthus capitatus, 485 Cheirinia inconspicua, 485 repanda, 485 Chelidonium, 700 majus, 700 laciniatum, 700 plenum, 700 Chelone, 709 glabra, 709 lyonii, 709 obliqua obliqua, 709 Chenopodium, 321 album, 323 lanceolatum, 323 ambrosioides, 326 anthelminticum, 326 anthelminticum, 326 berlandieri, 323 bushianum, 322 macrocalycium, 323 zschackei, 323 bonus-henricus, 323 boscianum, 324 botrys, 326 bushianum, 323 capitatum, 323 carinatum, 326 cristatum, 326 desiccatum, 324 leptophylloides, 324 foggii, 323 foliosum, 323

gigantospermum, 324 standleyanum, 324 glaucum, 323 glaucum, 323 graveolens, 326 hirsutum, 320 humile, 324 hybridum gigantospermum, 324 simplex, 324 lanceolatum, 323 leptophyllum, 323 macrocalycium, 323 missouriense, 323 multifidum, 326 murale, 322, 324 opulifolium, 323 polyspermum, 324 acutifolium, 324 obtusifolium, 324 polyspermum, 324 pratericola, 322, 324 leptophylloides, 324 pumilio, 326 rubrum, 324 humile, 324 rubrum, 324 scoparium, 326 simplex, 322, 324 standleyanum, 324 strictum, 322 glaucophyllum, 325 subspicatum, 320 urbicum, 322, 325 virgatum, 323 viride, 323 cherry, 453, 800–801 cherry birch, 453 cherry plum, 800 chervil, 338 chestnut, 601 chestnut sedge, 131 chick-pea, 578 chickweed, 509–510, 512, 514 chicory, 388 childing-pink, 515 Chile tarplant, 417 Chimaphila, 553 corymbosa, 553 maculata, 553 umbellata cisatlantica, 553 Chinese-aster, 384 Chinese bindweed, 730 Chinese buckthorn, 763 Chinese bush-clover, 586 Chinese catalpa, 455 Chinese chestnut, 601 Chinese elm, 868 Chinese foxtail, 280 Chinese hemlock-parsley, 340 Chinese hound’s-tongue, 460 Chinese matrimony-vine, 859 Chinese mustard, 478 Chinese peony, 698 Chinese privet, 679 Chinese silvergrass, 266 Chinese smartweed, 734 Chinese wisteria, 600 Chinese yam, 172

in d e x  9 2 9

Chiogenes hispidula, 555 Chionanthus, 676 virginicus, 676 maritimus, 676 Chionodoxa, 174 forbesii, 174 luciliae, 174 Chironia amoena, 612 angularis, 612 campanulata, 612 chloroides, 612 dodecandra, 612 stellata, 612 chives, 92 Chloris, 241 gayana, 241 virgata, 241 Chlorostachyae, 118 chocolate-vine, 653 chokeberry, 773 choke cherry, 801 Chondrosea paniculata neogaea, 855 Chondrosum gracile, 234 prostratum, 234 Chordorrhizae, 117 Chorispora, 483 tenella, 483 Chorizanthe, 728 pungens pungens, 727 Christmas fern, 58 Chrysanthemum, 388 arcticum polare, 375 balsamita, 379 tanacetoides, 379 bipinnatum huronense, 443 carinatum, 403 coronarium, 403 indicum, 388 leucanthemum, 416 morifolium, 388 nipponicum, 420 parthenium, 443 segetum, 403 serotinum, 416 uliginosum, 443 Chrysopsis, 388 falcata, 423 mariana, 388 Chrysosplenium, 853 americanum, 853 churchmouse threeawn, 232 Cicer, 578 arietinum, 578 Cichorium, 388 endivia, 388 intybus, 388 Cicuta, 340 bulbifera, 340 maculata maculata, 340 Cimicifuga racemosa, 750 serpentaria, 750

Cinna, 242 arundinacea, 242 latifolia, 242 cinnamon fern, 66 cinnamon rose, 806 cinquefoil, 788, 794–798, 818 Circaea, 681 alpina alpina, 681 canadensis canadensis, 681 virginiana, 681 latifolia, 681 lutetiana canadensis, 681 quadrisulcata, 681 ×sterilis, 681 Cirsium, 388 altissimum, 389 arvense horridum, 389 integrifolium, 389 mite, 389 bigelovii, 390 discolor, 390 flodmanii, 390 horridulum, 390 lanceolatum, 390 muticum, 390 monticola, 390 odoratum, 390 palustre, 390 pumilum, 390 spinosissimus, 390 vulgare, 390 Cistaceae, 525–529 Citrullus, 544 citrullus, 544 lanatus, 544 vulgaris, 544 city goosefoot, 325 Cladium, 151 mariscoides, 151 Cladrastris, 578 kentukea, 578 lutea, 578 clammy azalea, 560 clammy-campion, 523 clammy glandular-goosefoot, 326 clammy ground-cherry, 862 clammy hedge-hyssop, 710 clammy locust, 593 clammy waxweed, 660 clammyweed, 529 Clandestinae, 118 clarkia, 681 Clarkia, 681 pulchella, 681 rhomboidea, 681 clasping heliotrope, 462 clasping-leaved pondweed, 297 clasping-leaved twistedstalk, 195 clasping-leaved Venus’-lookingglass, 497 clasping milkweed, 349 clasping pepperweed, 487 clasping water-horehound, 640 Clausenellia ternata, 543

Claytonia, 745 caroliniana, 745 lewisii, 745 spathulifolia, 745 fontana, 745 linearis, 746 robusta, 745 sibirica, 745 bulbifera, 745 virginica, 745 hammondiae, 745 cleareye sage, 648 clearweed, 870 cleft-leaved hawthorn, 787 Clematis, 753 canadensis, 753 dioscoreifolia, 753 robusta, 753 florida, 753 japonica, 753 missouriensis, 753 occidentalis occidentalis, 753 paniculata, 753 recta, 753 terniflora, 753 verticillaris, 753 grandiflora, 753 virginiana, 753 missouriensis, 753 vitalba, 753 Cleomaceae, 529–530 Cleome hassleriana, 530 serrulata, 529 angusta, 529 spinosa, 530 Clethra, 530 alnifolia, 530 tomentosa, 530 Clethraceae, 530 cliff-brake, 68 climbing bindweed, 731 climbing dayflower, 99 climbing hempvine, 418 climbing nightshade, 864 climbing rose, 807 climbing spindle-tree, 525 Clingman’s hedge-nettle, 651 Clinopodium, 636 acinos, 636 vulgare, 636 Clintonia, 193 borealis, 193 Clinton’s clubsedge, 171 Clinton’s wood fern, 56 closed-headed sedge, 145 clover, 597–598 clover dodder, 534 club-awn grass, 242 clubmoss, 43–47 clubsedge, 171 clustered bellflower, 495 clustered clover, 597 clustered moonshine-daisy, 394 clustered mountain-mint, 647 clustered sanicle, 346 clustered sedge, 123 clustered wheat grass, 226 clustered yellowtops, 402

cluster-headed pink, 511 Clypeola alyssoides, 474 maritimum, 488 Cnicus benedictus, 386 coastal goldfields, 416 coastal jointed knotweed, 738 coastal plain blue-eyed-grass, 181 coastal plain grass-leavedgoldenrod, 401 coastal plain Joe-Pye weed, 401 coastal sandbur, 241 coastal sweet-pepperbush, 530 coastal tidytips, 416 coast barnyard grass, 252 coast cinquefoil, 797 coast Douglas-fir, 79 coast violet, 881 Cochlearia armoracia, 475 coronopus, 488 Cochranea anchusifolia, 462 cocklebur, 446 cock’s comb, 321 cockspur hawthorn, 783 Coeloglossum, 201 viride, 201 Coelopleurum actaeifolium, 338 gmelinii, 338 lucidum, 338 gmelinii, 338 coffee-tree, 582 cohosh, 449 Coix dactyloides, 286 Colchicaceae, 98–99 Colchicum, 98 autumnale, 98 Coleogeton filiformis alpinus, 298 occidentalis, 298 pectinatus, 298 colic-root, 198 collared dodder, 535 Collinsia, 709 parviflora, 709 Collinsiae, 118 Collinsonia, 636 canadensis, 636 Collins’ sedge, 118 Collomia, 722 linearis, 722 coltsfoot, 445 Columbian water-meal, 97 columbine, 752 Colutea, 578 arborescens, 578 Colymbada scabiosa, 387 Comandra, 530 richardsiana, 530 umbellata umbellata, 530 Comandraceae, 530

93 0   i n d e x

Comarum, 774 palustre, 774 comb-leaved mermaid-weed, 622 comfrey, 466–467 Commelina, 99 communis, 99 ludens, 99 diffusa diffusa, 99 dubia, 289 longicaulis, 99 Commelinaceae, 99–100 common agrimony, 768 common alum-root, 853 common arrowhead, 90 common barberry, 449 common barley, 263 common barnyard grass, 252 common bird’s-foot, 591 common blackberry, 811 common blanket-flower, 402 common bleeding-heart, 701 common brassbuttons, 392 common bugloss, 459 common burdock, 376 common buttonbush, 821 common canary grass, 272 common clarkia, 681 common clubmoss, 47 common comfrey, 467 common corncockle, 508 common cotton-rose, 402 common creeping-cudweed, 397 common crupina, 393 common daffodil, 93 common dandelion, 444 common dock, 742 common dodder, 534 common dog-mustard, 485 common dropwort, 789 common duck-meal, 97 common duckweed, 96 common eastern wild-rye, 256 common evening-primrose, 686 common eyebright, 693 common fig, 670 common flowering-quince, 774 common fox sedge, 135 common fumitory, 701 common glasswort, 327 common globe-amaranth, 326 common golden Alexanders, 347 common gorse, 598 common grape-hyacinth, 174 common grass-leavedgoldenrod, 401 common greenbrier, 301 common ground-nut, 576 common hackberry, 497 common hawkweed, 412 common hedge-mustard, 492 common henbit, 638 common hop, 498 common hoptree, 829 common hornwort, 82 common hound’s-tongue, 460

common interrupted-clubmoss, 48 common juniper, 75 common kidney-vetch, 575 common lilac, 679 common lowbush blueberry, 562 common mallow, 666 common mare’s-tail, 710 common marsh-mallow, 664 common milkweed, 350 common moonwort, 64 common morning-glory, 536 common mullein, 857 common nipplewort, 415 common pear, 802 common pearlbrush, 788 common peony, 698 common persimmon, 547 common plantain, 716 common prickly-ash, 829 common purslane, 746 common quickweed, 402 common ragweed, 373 common ragwort, 427 common reed, 273 common rose-gentian, 612 common rue, 829 common safflower-thistle, 385 common selfheal, 646 common shepherd’s-cress, 492 common silver-hairgrass, 228 common silverweed, 772 common snowberry, 503 common soapwort, 517 common soft rush, 187 common sow-thistle, 434 common speedwell, 719 common spikesedge, 160 common star-grass, 178 common stitchwort, 523 common St. John’s-wort, 626 common stork’s-bill, 613 common St. Paul’s-wort, 427 common strawberry, 789 common sunflower, 406 common tansy, 443 common thistle, 390 common three-seededMercury, 564 common Timothy, 272 common valerian, 504 common velvet grass, 262 common vetch, 600 common viper’s-bugloss, 461 common water-hyacinth, 289 common watermoss, 69 common water-primrose, 684 common waterweed, 176 common winterberry, 352 common wood rush, 191 common woolsedge, 169 common wormwood, 379 common wrinkle-leaved goldenrod, 433 common yarrow, 371 common yellow-cress, 490 common yellow flax, 658 common yellow wood sorrel, 697 compact dodder, 534

Comptonia, 671 peregrina, 671 coneflower, 395, 424–425 Conioselinum, 340 chinense, 340 Conium, 340 maculatum, 340 Conopholis, 692 americana, 692 Conringia, 483 orientalis, 483 Consolida, 753 ajacis, 754 ambigua, 754 orientalis, 754 regalis, 754 Convallaria, 299 biflora, 300 latifolia, 300 majalis, 299 pubescens, 300 racemosa, 300 stellata, 300 trifolia, 300 Convolvulaceae, 530–536 Convolvulus, 532 arvensis, 532 fraterniflorus, 532 panduratus, 536 pubescens, 531 purpurea, 536 sepium, 531 spithamaeus, 532 Conyza asteroides, 427 canadensis, 396 pusilla, 396 linifolia, 427 Coprosmanthus herbaceus, 301 Coptidium, 754 lapponicum, 754 Coptis, 754 groenlandica, 754 trifolia, 754 groenlandica, 754 coral necklace, 512 Corallorhiza, 201 maculata, 201 maculata, 201 occidentalis, 201 multiflora, 201 odontorhiza, 201 odontorhiza, 202 pringlei, 202 pringlei, 201 trifida, 202 coral-root, 201–202 Corchorus scandens, 802 cordgrass, 282–283 Corema, 553 conradii, 553 Coreopsis, 390 alternifolia, 445 aristosa, 382 basalis, 391 cardaminifolia, 391 drummondii, 391 grandiflora, 391 pilosa, 391

heterogyna, 391 lanceolata, 391 villosa, 391 major, 391 palmata, 391 pubescens, 391 robusta, 391 rosea, 391 tinctoria, 392 imminuta, 392 tripteris, 392 smithii, 392 subrhomboidea, 392 verticillata, 392 coriander, 340 Coriandrum, 340 sativum, 340 cork spindle-tree, 525 corktree, 829 corn, 239, 288, 375, 403, 445, 491, 520, 702, 718 Cornaceae, 536–539 corn brome, 239 corn chamomile, 375 corn charlock, 491 corncockle, 508 corn daisy, 403 corn-gromwell, 459 Cornish heath, 554 corn poppy, 702 corn-salad, 505 corn scentless-chamomile, 445 corn speedwell, 718 corn spurry, 520 Cornucopiae hyemalis, 227 Cornus alba stolonifera, 539 alternifolia, 538 amomum, 538 canadensis, 537 florida, 537 foemina racemosa, 539 kousa, 537 paniculata, 539 racemosa, 539 rugosa, 539 sanguinea, 539 sericea, 539 stolonifera, 539 stolonifera, 539 obliqua, 538 schuetzeana, 538 Coronaria flos-cuculi, 513 Coronilla, 578 scorpioides, 578 Coronopus didymus, 487 squamatus, 488 vulgaris, 715 Corrigiola, 510 litoralis litoralis, 510 corydalis, 700 Corydalis, 700 aurea, 700 bulbosa, 700 canadensis, 701

i n d e x  9 3 1

Corydalis, cont. flavula, 700 glauca, 699 sempervirens, 699 solida, 700 washingtoniana, 700 Corylus, 454 americana, 454 indehiscens, 454 cornuta cornuta, 454 rostrata, 454 Corynephorus, 242 canescens, 242 cosmos, 392 Cosmos, 392 bipinnatus, 392 parviflorus, 392 sulphureus, 392 costmary, 379 Cota, 392 tinctoria, 392 Cotinus, 329 coggygria, 329 cotoneaster, 774 Cotoneaster, 774 divaricatus, 774 cotton, 664 cotton-rose, 402, 417 cottonsedge, 161–162 cotton-thistle, 421 cotton-weed, 326 Cotula, 392 coronopifolia, 392 cowcockle, 523 cow-parsnip, 342 cowslip primrose, 747 cow-wheat, 694 Cracca latidens, 595 crab apple, 792–793 crabgrass, 251 crabweed, 670 cranberry, 563 highbush, 310 cranefly orchid, 212 crane’s-bill, 615–616 Crassula, 541 aquatica, 541 Crassulaceae, 539–543 Crataegus, 774 affinis, 787 alnifolia, 818 ambrosia, 787 ampla, 784 anomala, 784 apiomorpha, 784 ardula, 787 arnoldiana, 787 ascendens, 786 baxteri, 782 beckwithae, 786 bicknellii, 779 biltmoreana, 782 bissellii, 782 blandita, 786 boyntonii, 782 brainerdii, 782 scabrida, 787 brunetiana, 785 caerulescens, 786

caesariata, 785 chadsfordiana, 784 champlainensis, 787 chrysocarpa, 782 bicknellii, 782 chrysocarpa, 782 phoenicea, 783 praecox, 782 chrysocarpa, 785 caesariata, 785 coccinea, 783 coccinioides, 783 dilatata, 783 cognata, 786 compta, 786 conjuncta, 786 cordata, 786 crudelis, 783 crus-galli, 783 culta, 786 damei, 786 demissa, 786 dilatata, 783 dissimilis, 786 dissona, 783 dodgei, 783 flavida, 783 lumaria, 785 rotundata, 783 dumicola, 785 eamesii, 784 egglestonii, 783 exclusa, 786 exigua, 783 faxonii, 783 praecoqua, 782 praetermissa, 783 fernaldii, 785 ferta, 787 fertilis, 785 festiva, 786 flabellata, 783 flabellata, 783 grayana, 783 flavida, 783 florea, 786 florifera, 787 fluviatilis, 784 foetida, 782 forbesae, 784 fretalis, 784 fulgens, 785 fuscosa, 786 gemmosa, 787 genialis, 786 glabrata, 787 glaucophylla, 786 gravesii, 783 gravis, 786 grayana, 783 handyae, 784 hargeri, 782 holmesiana, 784 villipes, 784 ideae, 784 incisa, 784 insolens, 785 intricata, 784 intricata, 784 straminea, 784

iracunda populnea, 786 stolonifera, 786 irrasa, 776 blanchardii, 784 jesupii, 784 jonesiae, 785 keepii, 785 kennedyi, 785 laurentiana, 785 brunetiana, 785 fernaldii, 785 leiophylla, 786 lemingtonensis, 783 levis, 785 littoralis, 786 lobulata, 783 lucorum, 785 lumaria, 785 macracantha, 785 macracantha, 785 occidentalis, 785 macrosperma, 786 acutiloba, 786 demissa, 786 matura, 786 pentandra, 786 roanensis, 786 matura, 786 media, 786 modesta, 782 monogyna, 786 monstrata, 786 napaea, 786 oakesiana, 786 parvula, 786 peckii, 782 pedicellata, 783 pennsylvanica, 787 pequatorum, 786 phaenopyrum, 786 philadelphica, 786 picta, 787 pisifera, 786 populifolia, 786 populnea, 786 porteri, 786 caerulescens, 786 praecoqua, 782 praecox, 782 praetermissa, 783 pringlei, 786 exclusa, 786 lobulata, 783 pruinosa, 786 dissona, 783 latisepala, 786 leiophylla, 786 parvula, 786 philadelphica, 786 porteri, 787 pruinosa, 787 punctata punctata, 787 racemosa, 771 rhombifolia, 785 robbinsiana, 786 rotundata, 783 rotundifolia bicknellii, 782 faxonii, 783

scabrida, 787 schizophylla, 787 schuettei, 787 basilica, 787 searsii, 785 seelyana, 782 shirleyensis, 782 spatiosa, 784 spicata, 772 stolonifera, 786 stonei, 787 straminea, 784 submollis, 787 succulenta, 787 gemmosa, 787 pisifera, 786 succulenta, 787 tardipes, 784 umbratilis, 788 venulosa, 787 villipes, 784 virilis, 787 Crawe’s sedge, 128 Crawford’s sedge, 123 cream-colored avens, 792 cream pincushions, 503 cream vetchling, 584 creeping bellflower, 495 creeping bentgrass, 228 creeping birthwort, 81 creeping bladderwort, 656 creeping bush-clover, 588 creeping cinquefoil, 797 creeping crowfoot, 758 creeping-cudweed, 397 creeping juniper, 76 creeping mazus, 704 creeping phlox, 724 creeping spicy-wintergreen, 555 creeping spikesedge, 160 creeping St. John’s-wort, 625 creeping thistle, 389 creeping velvet grass, 262 creeping waxweed, 660 creeping wild-rye, 255 creeping yellow-cress, 491 creeping yellow-loosestrife, 674 creeping yellow wood sorrel, 697 Crepis, 392 biennis, 393 capillaris, 393 foetida, 393 nicaeensis, 393 setosa, 393 tectorum, 393 vesicaria haenseleri, 393 crested anoda, 664 crested bindweed, 730 crested cock’s-comb, 321 crested dogtail grass, 243 crested glandular-goosefoot, 326 crested iris, 179 crested late-summer-mint, 637 crested orange bog-orchid, 209 crested sedge, 123 crested wheat grass, 226

93 2   i n de x

crested wood fern, 56 Cretan viper’s-bugloss, 461 crimson clover, 598 crimson weigela, 505 Critesion brachyantherum, 263 jubatum, 263 murinum, 263 pusillum, 263 Crocanthemum, 526 bicknellii, 526 canadense, 527 dumosum, 526 majus, 526 propinquum, 527 crocus, 178 Crocus, 178 vernus, 178 crooked-stemmed Americanaster, 442 crossflower, 483 cross-leaved heath, 554 Crotalaria, 578 sagittalis, 578 oblonga, 578 croton, 565 Croton, 565 capitatus, 565 capitatus, 565 texensis, 565 willdenowii, 565 Crotonopsis, 565 elliptica, 565 crowberry, 553–554 crowfoot, 754–759 crow garlic, 92 crownbeard, 445 crown daisy, 403 crowned beggar-ticks, 383 crown-vetch, 578, 593 crupina, 393 Crupina, 393 vulgaris, 393 Crypsis, 242 schoenoides, 242 Cryptantha, 459 ambigua, 459 Cryptogramma, 68 stelleri, 68 Cryptotaenia, 341 canadensis, 341 Cubelium concolor, 873 Cucubalis pumilio, 517 stellatus, 520 Cucubertia citrullus, 544 cucumber, 194, 545 cucumber-leaved sunflower, 406 cucumber root, 194 cucumber-tree, 84 Cucumis, 544 anguria, 544 melo, 545 myriocarpus, 545 sativus, 545 Cucurbita, 545 maxima, 545 pepo, 545

Cucurbitaceae, 543–545 cudweed, 397, 403, 421 cultivated apple, 793 cultivated endive, 388 cultivated fenugreek, 598 cultivated flax, 658 cultivated garlic, 91 cultivated lettuce, 415 cultivated purslane, 746 cultivated radish, 489 cultivated rye, 280 cultivated tobacco, 861 Culver’s-root, 720 cumin, 341 Cuminum, 341 cyminum, 341 Cunigunda purpurea, 402 Cunila pulegioides, 638 cup-scale, 279 Cuphea, 660 petiolata, 660 procumbens, 660 viscosissima, 660 cup-plant rosinweed, 428 Cupressaceae, 75–76 Cupressus thyoides, 75 curly blue grass, 277 curly dock, 743 curly plumeless-thistle, 384 curly pondweed, 295 curly-top gum-weed, 404 currant, 618–619 cursed crowfoot, 759 Cuscuta, 532 approximata, 533 urceolata, 534 arvensis, 535 calycina, 534 campestris, 534 cephalanthi, 533 chlorocarpa, 535 compacta, 533 adpressa, 533 compacta, 533 coryli, 533 epithymum, 534 europaea, 533 glomerata adpressa, 533 gronovii, 534 gronovii, 535 latiflora, 535 indecora, 535 indecora, 535 neuropetala, 535 neuropetala, 535 pentagona, 535 calycina, 533 planiflora, 533 polygonorum, 535 pulcherrima, 535 saururi, 533 vulgivaga, 533 cushion-plant, 546 cut-leaved blackberry, 816 cut-leaved crane’s-bill, 616

cut-leaved evening-primrose, 687 cut-leaved germander, 652 cut-leaved glandulargoosefoot, 326 cut-leaved ground-cherry, 862 cut-leaved medick, 590 cut-leaved nightshade, 865 cut-leaved selfheal, 646 cut-leaved teasel, 500 cut-leaved toothwort, 481 cut-leaved water-milfoil, 621 cut-leaved windflower, 752 Cutler’s goldenrod, 432 Cyanococcus canadensis, 563 corymbosus, 563 fuscatus, 563 pallidus, 563 vacillans, 563 Cyclachaena, 394 xanthiifolia, 394 Cyclolepis platyphylla, 325 Cycloloma, 325 atriplicifolium, 325 platyphylla, 325 Cydonia, 788 japonica, 774 lagenaria, 774 lagenaria, 774 oblonga, 788 vulgaris, 788 Cylindropyrum cylindricum, 226 Cymbalaria, 709 muralis, 709 Cymbidium hyemale, 200 Cynanchum, 350 louiseae, 350 medium, 350 nigrum, 350 rossicum, 350 Cynodon, 243 aristiglumis, 243 dactylon dactylon, 243 Cynoglossum, 460 amabile, 460 boreale, 460 microglochin, 460 officinale, 460 virginianum, 460 boreale, 460 virginianum, 460 Cynosurus, 243 aegyptius, 243 cristatus, 243 indicus, 252 Cynoxylon floridum, 537 Cyperaceae, 100–171 Cyperoideae, 118–125 Cyperus, 151 acuminatus, 153 aristatus, 155 arundinaceus, 155 bipartitus, 153 brevifolioides, 163 cylindricus, 155

dentatus, 153 diandrus, 153 echinatus, 152 engelmannii, 154 erythrorhizos, 153 esculentus, 153 angustispicatus, 154 leptostachyus, 154 ferruginescens, 154 filicinus, 154 filiculmis macilentus, 154 oblitus, 154 flavescens, 154 fuscus, 154 grayi, 154 halei, 153 houghtonii, 153 inflexus, 155 iria, 154 microiria, 154 lupulinus, 154 lupulinus, 154 macilentus, 154 ×mesochorus, 154 microiria, 152 odoratus, 154 ovularis, 153 polystachyos, 155 filicinus, 154 texensis, 155 pseudovegetus, 155 retrorsus, 155 rivularis, 153 schweinitzii, 155 squarrosus, 155 strigosus, 155 robustior, 155 tuberosus, 154 virens arenicola, 155 cypress, 326 cypress-like sedge, 149 cypress spurge, 567 Cypress-vine morning-glory, 536 Cypripedium, 202 acaule, 202 arietinum, 202 bulbosum, 201 calceolus parviflorum, 202 pubescens, 202 parviflorum, 202 makasin, 203 parviflorum, 203 pubescens, 202 reginae, 203 spectabile, 203 Cyrtorhyncha cymbalaria, 758 Cystopteris, 72 bulbifera, 72 dickieana, 72 fragilis, 72 laurentiana, 72 mackayi, 73 protrusa, 73 laurentiana, 72 protrusa, 73 tenuis, 73

in d e x  9 3 3

Cytisus, 579 ×praecox, 579 scoparius scoparius, 579

D

Dactylis, 243 glomerata, 243 Dactyloctenium, 243 aegyptium, 243 radulans, 243 Dactylorhiza viridis, 201 daffodil, 93 Dahlburg-daisy, 444 Dahurian buckthorn, 763 daisy, 403, 416, 420, 444 daisy-leaved moonwort, 65 Dalea, 579 alopecuroides, 579 candida, 579 leporina, 579 Dalibarda fragarioides, 791 repens, 813 Dalmatian toadflax, 712 darnel, 265 dame’s-rocket, 486 Damson plum, 800 dandelion, 444 Danthonia, 243 alleni, 244 californica, 243 compressa, 244 spicata, 244 pinetorum, 244 daphne, 867 Daphne, 867 cneorum, 867 mezereum, 867 dark-green bulrush, 169 dark red helleborine, 203 Darwinia exaltata, 594 Dasiphora, 788 floribunda, 788 fruticosa floribunda, 788 Dasistoma flava, 691 pedicularia, 691 virginica, 691 Dasystephana andrewsii, 610 grayi, 610 Datisca hirta, 330 Datura, 858 inermis, 859 inoxia, 859 quinquecuspida, 859 metel quinquecuspida, 859 meteloides, 859 stramonium, 859 inermis, 859 tatula, 859 tatula, 859 wrightii, 859

Daucus, 341 carota, 341 Davis’ sedge, 131 dayflower, 99 day-lily, 173 Decachaena baccata, 555 frondosa, 555 Decodon, 660 verticillatus, 661 laevigatus, 660 deerberry, 563 deer-tongue rosettepanicgrass, 248 Deinandra, 394 fasciculata, 394 delicate quill sedge, 125 Delphinium, 754 ajacis, 754 ambiguum, 754 consolida, 754 exaltatum, 754 orientale, 754 Demidovia tetragonioides, 311 Dendranthema arcticum, 375 grandiflorum, 388 hultenii, 375 morifolium, 388 Dendrobenthamia japonica, 537 Dendrolycopodium, 43 dendroideum, 44 hickeyi, 44 obscurum, 44 Dennstaedtia, 53 punctilobula, 53 Dennstaedtiaceae, 53–54 dense-flowered mullein, 857 dense silky bentgrass, 231 Dentaria incise, 482 diphylla, 482 laciniata, 481 maxima, 482 Deparia, 73 acrostichoides, 73 Deptford pink, 511 Deschampsia, 244 anadyrensis, 244 atropurpurea, 287 cespitosa, 244 glauca, 244 parviflora, 244 danthonioides, 244 elongata, 244 flexuosa, 245 montana, 245 glauca, 244 Descurainia, 483 incana, 483 irio, 492 pinnata, 483 brachycarpa, 483 richardsonii, 483 sophia, 483 desert goosefoot, 324 Desmazeria, 245 rigida, 245

Desmodium, 579 acuminatum, 582 canadense, 580 canescens, 580 ciliare, 580 cuspidatum, 580 glabellum, 580 glutinosum, 582 grandiflorum, 580 humifusum, 581 longifolium, 580 marilandicum, 580 nudiflorum, 583 obtusum, 581 paniculatum, 581 perplexum, 581 rotundifolium, 581 sessilifolium, 581 Deutzia, 622 scabra, 622 plena, 622 devil-in-the-bush, 755 Devil’s beggar-ticks, 383 devil’s bit, 196 Devil’s-bit, 503 dewdrop, 813 dew-drop crane’s-bill, 616 Deweyanae, 125 Deyeuxia langsdorfii, 240 macouniana, 240 neglecta, 240 nuttalliana, 240 pickeringii, 240 preslii, 240 Dianthera americana, 307 Dianthus, 510 armeria, 511 atrorubens, 511 barbatus, 511 carthusianorum, 511 deltoides, 511 plumarius, 511 prolifer, 515 saxifragus, 515 Diapensia, 546 lapponica lapponica, 546 Diapensiaceae, 546 Dicentra, 700 canadensis, 701 cucullaria, 701 eximia, 701 formosa, 701 occidentalis, 701 spectabilis, 701 Dichanthelium, 245 acuminatum, 247 acuminatum, 247 columbianum, 247 fasciculatum, 247 implicatum, 247 lindheimeri, 247 spretum, 247 boreale, 248 boscii, 248 clandestinum, 248 columbianum, 247

commutatum ashei, 248 depauperatum, 248 dichotomum, 249 dichotomum, 249 mattamuskeetense, 249 microcarpon, 249 lanuginosum, 247 latifolium, 249 linearifolium, 249 oligosanthes, 246 scribnerianum, 249 ovale, 249 pseudopubescens, 249 villosissimum, 250 pseudo-esula, 567 scabriusculum, 250 ×scoparioides, 248 scoparium, 250 sphaerocarpon, 250 wrightianum, 250 xanthophysum, 250 Dictamnus, 829 albus, 829 Didier’s tulip, 195 Diervilla, 500 diervilla, 500 floribunda, 505 lonicera, 500 hypomalaca, 500 Digitalis, 709 ambigua, 710 grandiflora, 710 lanata, 710 lutea, 710 purpurea, 710 purpurea, 710 Digitaria, 250 cognata, 251 filiformis, 251 filiformis, 251 laeviglumis, 251 ischaemum, 251 laeviglumis, 251 sanguinalis, 251 violascens, 251 Dilatris caroliniana, 173 Dilepyrum aristosum, 235 dill, 337 Dinebra repens, 234 Diodia, 822 hirsuta, 822 teres teres, 822 virginiana virginiana, 822 Diodella teres, 822 dioecious sedge, 147 Dioscorea, 172 batatas, 172 hirticaulis, 172 polystachya, 172 villosa, 172 Dioscoreaceae, 172

93 4   in de x

Diospyros, 547 mosieri, 547 virginiana, 547 mosieri, 547 platycarpa, 547 pubescens, 547 Diphasiastrum, 44 complanatum, 45 digitatum, 45 ×habereri, 45 ×issleri, 45 ×sabinifolium, 45 sitchense, 45 tristachyum, 45 ×verecundum, 45 ×zeilleri, 45 Diphryllum bifolium, 206 Diplachne fascicularis, 264 Diplazium, 73 acrostichoides, 73 pycnocarpon, 73 Diplopappus chinensis, 384 Diplotaxis, 483 muralis, 483 tenuifolia, 484 Dipsacus, 500 fullonum, 500 sylvestris, 500 laciniatus, 500 sylvestris, 500 Dirca, 867 palustris, 867 Dispermae, 125 dissected grapefern, 64 distant scorpion-weed, 465 ditchgrass, 299 ditch-stonecrop, 703 Distichlis, 251 spicata, 251 Dittrichia, 394 graveolens, 394 Divisae, 125 Dobie-pod, 492 dock, 740–744 dock-leaved smartweed, 734 dodder, 532–535 Dodge’s hawthorn, 783 Doellingeria, 394 infirma, 394 ptarmicoides, 421 umbellata umbellata, 394 dogbane, 348 dog bentgrass, 227 dog-laurel, 556 dog-mustard, 485 dog rose, 806 dogtail grass, 243 dogwood, 538–539 Dolichos lobatus, 592 polystachyus, 592 doll’s-daisy, 384 Dondia americana, 329 calceoliformis, 329 depress, 329 linearis, 329

maritima, 329 squarrosa, 404 doormat grass, 268 dooryard knotweed, 738 Doronicum, 394 pardalianches, 394 dotted hawthorn, 787 dotted smartweed, 735 doubtful knight’s-spur, 754 Douglas-fir, 79 Douglas’ knotweed, 739 Douglas’ silverpuffs, 418 dove pincushions, 503 dove’s-foot crane’s-bill, 616 downy agrimony, 768 downy alpine oat grass, 233 downy arrowwood, 311 downy birch, 454 downy cinquefoil, 797 downy false foxglove, 691 downy goldenrod, 432 downy ground-cherry, 862 downy phlox, 724 downy rattlesnake-plantain, 204 downy shadbush, 770 downy thorn-apple, 859 downy wild-rye, 255 downy willow-herb, 684 downy wood-mint, 636 Draba, 484 arabisans, 484 cana, 484 canadensis, 484 breweri, 484 cana, 484 cana, 484 canadensis, 484 caroliniana, 484 daurica, 484 glabella, 484 megasperma, 484 orthocarpa, 484 incana, 484 arabisans, 484 lanceolata, 484 megasperma, 484 micrantha, 484 reptans, 484 micrantha, 484 verna, 484 aestivalis, 484 boerhaavii, 484 umbellata, 484 Dracocephalum, 636 moldavica, 636–637 parviflorum, 637 thymiflorum, 637 virginianum, 645 Dracontium foetidum, 97 dragonhead, 637, 645 dragon’s-mouth, 200 dragon wormwood, 378 dropseed, 284–285 drooping sedge, 131 drooping woodland sedge, 130 dropwort, 788 Drosera, 546 anglica, 547

filiformis, 547 intermedia, 547 linearis, 547 longifolia, 547 rotundifolia, 547 comosa, 547 Droseraceae, 546–547 drum-heads milkwort, 726 dry land sedge, 115 Drymocallis, 788 agrimonioides, 788 arguta, 788 Dryopteridaceae, 54–58 Dryopteris, 54 austriaca, 57 ×benedictii, 55 ×boottii, 56 ×burgessii, 56 campyloptera, 55 carthusiana, 57 clintoniana, 56 ×correllii, 56 cristata, 56 clintoniana, 56 ×dowellii, 56 filix-mas, 57 fragrans, 57 goldiana, 57 hexagonoptera, 70 intermedia, 57 marginalis, 58 ×mickelii, 56 ×montgomeryi, 57 ×neowherryi, 57 phegopteris, 70 ×pittsfordensis, 56 simulata, 69 ×slossoniae, 57 spinulosa, 57 americana, 55 intermedia, 57 ×triploidea, 56 ×uliginosa, 55 Duchesnea indica, 797 duck-meal, 95, 97 duckweed, 96 Dudley’s rush, 187 Dulichium, 155 arundinaceum, 155 dune bluestem, 279 Durban crowfoot grass, 243 Dutch crocus, 178 Dutchman’s-breeches, 701 Dutchman’s pipe, 82 Dysphania, 325 ambrosioides, 326 anthelmintica, 326 botrys, 326 cristata, 326 graveolens, 326 graveolenst, 325 multifida, 326 pumilio, 326 Dyssodia, 394 papposa, 394 tenuiloba, 444 dwarf amaranth, 317 dwarf birch, 453 dwarf blueberry, 562 dwarf-bulrush, 163

dwarf burhead, 88 dwarf chestnut, 601, 606 dwarf chestnut oak, 606 dwarf cinquefoil, 796 dwarf-dandelion, 414 dwarf-dogwood, 537 dwarf-gentian, 610 dwarf ginseng, 344 dwarf glasswort, 327 dwarf huckleberry, 555 dwarf iris, 180 dwarf mistletoe, 887 dwarf plantain, 716 dwarf prairie willow, 847 dwarf raspberry, 816 dwarf rattlesnake-plantain, 204 dwarf sand plum, 801 dwarf scouring-rush, 60 dwarf shadbush, 772 dwarf-snapdragon, 708 dwarf St. John’s-wort, 626 dwarf umbrella-sedge, 163 dwarf water-lily, 86 dyer’s greenweed, 581 dyer’s mignonette, 762

E Eames’ toothwort, 482 eared-rockcress, 475 eared willow, 844 early azalea, 560 early blue cohosh, 449 early blue violet, 886 early coral-root, 202 early crowfoot, 758 early goldenrod, 432 early meadow-rue, 760 early sand grass, 265 early silver-hairgrass, 228 early small-flowered-saxifrage, 853 early wild-rye, 255 early yellow-rocket, 476 eastern black currant, 618 eastern black nightshade, 865 eastern bluebells, 463 eastern blue-eyed-grass, 181 eastern bluestar, 347 eastern bottle-brush grass, 254 eastern false gromwell, 465 eastern false willow, 379 eastern featherbells, 196 eastern figwort, 856 eastern gamagrass, 286 eastern grasswort, 343 eastern hay-scented fern, 53 eastern hemlock, 80 eastern leatherwood, 867 eastern narrow-leaved sedge, 128 eastern prairie white fringed bog-orchid, 210 eastern prickly gooseberry, 618 eastern prickly-pear, 493 eastern purple bladderwort, 656 eastern purple coneflower, 395 eastern red cedar, 76 eastern riverbank wild-rye, 255

i n d e x  9 3 5

eastern rough sedge, 115 eastern shadbush, 771 eastern silver American-aster, 439 eastern smooth beardtongue, 714 eastern spicy-wintergreen, 555 eastern spindle-tree, 525 eastern star sedge, 143 eastern straw sedge, 124 eastern sweetshrub, 82 eastern tansy, 443 eastern waterleaf, 462 eastern white beardtongue, 714 eastern white oak, 604 eastern white pine, 79 eastern willow-herb, 683 eastern woodland sedge, 133 Eastwood’s fiddleneck, 458 Eatonia dudleyi, 283 intermedia, 283 nitida, 283 obtusata, 283 pensylvanica, 283 Eaton’s beggar-ticks, 382 Ebenaceae, 547 ebony spleenwort, 51 Echinacea, 394 pallida, 395 purpurea, 395 Echinochloa, 251 colona, 252 crus-galli, 252 frumentacea, 252 frumentacea, 252 microstachya, 252 muricata, 252 microstachya, 252 muricata, 252 pungens microstachya, 252 wiegandii, 252 walteri, 252 Echinocystis, 545 lobata, 545 Echinodorus parvulus, 88 tenellus, 88 Echinops, 395 sphaerocephalus, 395 Echinospermum lappula, 462 Echium, 460 australe, 461 creticum, 461 plantagineum, 461 pustulatum, 461 vulgare, 461 Eclipta, 395 prostrata, 395 Ectrus mexicanus, 699 edging candytuft, 486 eel-grass, 305 Egeria, 176 densa, 176 Eggleston’s hawthorn, 783 eggplant, 865

Eichhornia, 289 crassipes, 289 speciosa, 289 eight-flowered six-weeks grass, 288 Elaeagnaceae, 547–548 Elaeagnus, 547 angustifolia, 548 parvifolia, 548 pungens, 548 umbellata parvifolia, 548 Elatinaceae, 548–549 Elatine, 548 americana, 549 minima, 549 triandra, 549 americana, 549 elderberry, 307 elegant milk-vetch, 576 Eleocharis, 155 acicularis, 159 submersa, 159 aestuum, 159 ambigens, 159 calva, 159 capitata borealis, 159 pseudoptera, 161 compressa, 159 diandra, 159 depressa, 159 elliptica, 159 compressa, 159 elliptica, 159 engelmannii, 159 detonsa, 159 engelmannii, 159 equisetoides, 159 erythropoda, 159 fallax, 160 flavescens, 156 olivacea, 160 halophila, 161 intermedia, 160 melanocarpa, 160 microcarpa, 158 filiculmis, 160 monticola leviseta, 159 nitida, 160 obtusa, 160 detonsa, 159 engelmanii, 159 heuseri, 160 jejuna, 160 obtusa, 160 peasei, 160 olivacea, 160 ovata, 160 aphanactis, 159 obtusa, 160 pallidostachys, 159 palustris, 160 palustris, 160 vigens, 160 parvula, 160 anachaeta, 160 pauciflora, 161 quadrangulata, 161 crassior, 161

quinqueflora, 156 fernaldii, 161 robbinsii, 161 rostellata, 161 smallii, 160 tenuis, 161 pseudoptera, 161 tenuis, 161 tricostata, 161 tuberculosa, 161 uniglumis, 161 halophila, 161 Eleusine, 252 indica, 252 radulans, 243 Eleutherococcus, 341 pentaphyllus, 341 elk horn clarkia, 681 elk sedge, 115 elkweed, 609 Elliott’s bentgrass, 227 Elliott’s goldenrod, 432 elliptic-leaved rushfoil, 565 elliptic-leaved shinleaf, 558 elliptic spikesedge, 159 Ellisia, 461 nyctelea, 461 elm, 867–869 elm-leaved goldenrod, 434 Elodea, 176 canadensis, 176 fraseri, 627 nuttallii, 176 Elsholtzia, 637 ciliata, 637 Elymus, 252 arenarius, 264 arkansanus, 255 canadensis, 254 glaucifolius, 254 canadensis riparius, 255 villosus, 255 capitatus, 264 caput-medusae, 285 curvatus, 254 glabriflorus, 254 australis, 254 glabriflorus, 254 halophilus, 256 hystrix, 254 bigeloviana, 254 hystrix, 254 jejunus, 256 macgregorii, 255 mollis, 264 pungens pycnanthus, 285 pycnanthus, 285 repens, 255 riparius, 255 striatus, 256 submuticus, 254 trachycaulus, 255 glaucus, 255 trachycaulus, 255 villosus, 255 arkansanus, 255 villosus, 255 virginicus, 256 glabriflorus, 254

halophilus, 256 hirsutiglumis, 256 intermedius, 256 jejunus, 256 submuticus, 254 virginicus, 256 wiegandii, 256 Elytrigia pycnantha, 285 repens, 255 Emex, 728 spinosus, 728 Emory’s sedge, 140 Empetrum, 553 atropurpureum, 553 conradii, 553 eamesii atropurpureum, 553 hermaphroditum, 554 hermaphroditum, 554 nigrum, 554 atropurpureum, 553 hermaphroditum, 554 purpureum, 553 rubrum atropurpureum, 553 empress-tree, 702 enchanter’s-nightshade, 681 endive, 388 Endodeca, 81 serpentaria, 81 hastata, 81 Engelmann’s arrowhead, 90 Engelmann’s quillwort, 41 Engelmann’s spikesedge, 159 English cinquefoil, 796 English elm, 868 English holly, 352 English-ivy, 342 English lavender, 639 English oak, 606 English plantain, 716 English sundew, 547 English violet, 884 English yew, 80 Enslen’s blackberry, 814 entangled hawthorn, 784 entire-leaved skullcap, 649 Epidendrum domesticum, 179 Epifagus, 692 virginiana, 692 Epigaea, 554 repens, 554 glabrifolia, 554 Epilobium, 682 adenocaulon, 683 ecomosum, 683 occidentale, 683 alpinum, 683 lactiflorum, 683 nutans, 683 americanum, 683 anagallidifolium, 683 pseudoscaposum, 683 angustifolium abbreviatum, 681 circumvagum, 681 macrophyllum, 681 platyphyllum, 681

93 6   in de x

Epilobium, cont. boreale, 683 ciliatum, 683 ciliatum, 683 ecomosum, 683 glandulosum, 683 coloratum, 683 densum, 684 glandulosum adenocaulon, 683 ecomosum, 683 hirsutum, 683 hornemannii, 682 hornemannii, 683 lactiflorum, 683 lactiflorum, 683 leptophyllum, 683 lineare, 683 nesophilum, 683 sabulonense, 683 oliganthum, 683 palustre, 683 gracile, 683 grammadophyllum, 683 labradoricum, 683 sabulonense, 683 parviflorum, 683 strictum, 684 Epimedium, 449 ×youngianum, 449 Epipactis, 203 atrorubens, 203 convallarioides, 206 helleborine, 203 atrorubens, 203 latifolia, 203 Epithymum arvense, 535 cephalanthi, 534 coryli, 534 Equisetaceae, 58–60 Equisetum, 58 arvense, 59 ×ferrissii, 60 fluviatile, 59 hyemale, 60 affine, 60 laevigatum, ferrissii, 60 ×litorale, 59 ×mackaii, 60 palustre, 60 pratense, 60 scirpoides, 60 sylvaticum, 60 variegatum, 60 Eragrostis, 256 capillaris, 257 cilianensis, 257 curvula, 257 diffusa, 258 frankii, 257 hypnoides, 258 intermedia, 258 major, 257 megastachya, 257 mexicana, 258 minor, 258 multicaulis, 258

orcuttiana, 258 pectinacea, 257 pectinacea, 258 pilosa pilosa, 258 poaeoides, 258 spectabilis, 258 virescens, 258 Erechtites, 395 hieraciifolius hieraciifolius, 395 megalocarpus, 395 megalocarpus, 395 erect cinquefoil, 796 erect hedge-parsley, 346 Eremochloa, 258 ophiuroides, 258 Erica, 554 cinerea, 554 tetralix, 554 vagans, 554 Ericaceae, 549–563 Erigeron, 395 acris, 395 kamtschaticus, 396 angulosus kamtschaticus, 396 annuus, 396 discoideus, 396 canadensis, 396 pusillus, 396 graveolens, 394 hyssopifolius, 396 philadelphicus, 396 glaber, 396 philadelphicus, 397 provancheri, 397 provancheri, 396 scaturicola, 396 provancheri, 396 pulchellus, 396, 397 purpureus, 396 pusillus, 396 ramosus beyrichii, 397 strigosus, 397 septentrionalis, 397 strigosus, 397 Eriocaulaceae, 172 Eriocaulon, 172 aquaticum, 172 parkeri, 172 pellucidum, 172 Eriophorella alpina, 171 alpinum hudsonianus, 171 Eriophorum, 161 alpinum, 171 angustifolium angustifolium, 162 gracile, 162 latifolium viridicarinatum, 162 spissum, 162 tenellum, 162 vaginatum, 161 spissum, 162 virginicum, 162 viridicarinatum, 162

Erodium, 612 botrys, 613 ciconium, 613 cicutarium, 613 cicutarium, 613 cygnorum, 614 laciniatum bovei, 614 malacoides, 614 moschatum, 614 stephanianum, 614 Eruca, 484 vesicaria sativa, 484 Erucastrum, 485 gallicum, 485 Erxlebenia minor, 558 Eryngium, 341 amethystinum, 341 planum, 341 yuccifolium yuccifolium, 341 eryngo, 341 Erysimum, 485 alliaria, 474 arkansanum, 485 asperum purshii, 485 barbarea, 476 capitatum, 485 capitatum, 485 purshii, 485 cheiranthoides, 485 hieraciifolium, 485 inconspicuum inconspicuum, 485 irio, 492 loeselii, 492 odoratum, 485 officinale, 492 pannonicum, 485 parviflorum, 485 repandum, 485 strictum, 485 Erythronium, 193 albidum, 193 americanum americanum, 193 Eschscholzia, 701 californica californica, 701 procera, 701 estuary sedge, 141 Eubotrys, 554 elongata, 554 racemosa, 554 elongata, 554 Euchiton, 397 involucratus, 397 Euclidium, 485 syriacum, 485 Eudianthe, 511 coeli-rosa, 511 Eulalia viminea, 265 Eunanus brevipes, 705

Euonymus, 524 alatus, 524 atropurpureus atropurpureus, 525 europaeus, 525 fortunei, 525 radicans, 525 vegetus, 525 phellomana, 524 phellomanus, 525 radicans radicans, 525 Eupatoriadelphus dubius, 401 fistulosus, 401 maculatus, 402 purpureus, 402 Eupatorium, 397 album, 398 album, 399 glandulosum, 399 altissimum, 399 amoenum, 402 aromaticum, 372 dubium, 401 fistulosum, 401 glandulosum, 399 hyssopifolium, 399 calcaratum, 399 laciniatum, 399 leucolepis novae-angliae, 399 maculatum, 402 foliosum, 402 novae-angliae, 399 perfoliatum, 399 colpophilum, 399 petaloideum, 399 pilosum, 399 pubescens, 399 purpureum, 401 maculatum, 402 rotundifolium, 399 ovatum, 399 saundersii, 399 rugosum, 372 chlorolepis, 372 tomentellum, 372 villicaule, 372 saltuense, 399 scandens, 418 serotinum, 399 sessilifolium, 399 brittonianum, 399 torreyanum, 399 Euphorbia, 565 corollata, 567 cyparissias, 567 dentata, 567 linearis, 567 esula, 567 orientalis, 569 tommasiniana, 569 uralensis, 569 glyptosperma, 568 helioscopia, 568 intercedens, 567 ipecacuanhae, 568 lathyris, 568 maculata, 568 marginata, 568

i n d e x  9 37

Euphorbia, cont. nutans, 568 peplus, 568 platyphyllos, 568 polygonifolia, 568 prostrata, 568 ×pseudo-esula, 567 serpens, 568 serpyllifolia, 566 serpyllifolia, 568 uralensis, 569 vermiculata, 566 virgata, 569 orientalis, 569 Euphorbiaceae, 564–569 Euphrasia, 692 americana, 693 brevipila, 693 canadensis, 693 disjuncta, 693 micrantha, 693 mollis laurentiana, 693 nemorosa, 693 oakesii, 693 officinalis, 693 nemorosa, 693 purpurea, 693 randii, 693 farlowii, 693 reeksii, 693 rigidula, 693 stricta, 693 suborbicularis, 693 tetraquetra, 693 williamsii, 693 vestita, 693 Eurasian campion, 520 Eurasian water-milfoil, 621 Eurasian waterwort, 549 European alder, 451 European ash, 677 European beech, 602 European black nightshade, 865 European buckthorn, 763 European centaury, 609 European chickweed, 510 European cinquefoil, 798 European columbine, 752 European corn-salad, 505 European cow-parsnip, 342 European field violet, 881 European frog’s-bit, 176 European gooseberry, 619 European grape, 889 European gromwell, 463 European heliotrope, 462 European honeysuckle, 502 European knotweed, 738 European larch, 77 European lily-of-the-valley, 299 European meadow rush, 188 European mountain-ash, 818 European plum, 800 European poplar, 832 European privet, 679 European sea-heath, 607 European smoketree, 329 European speedwell, 719 European spindle-tree, 525

European stonecrop, 543 European turkey oak, 605 European vervain, 872 European wallflower, 485 European wand loosestrife, 661 European water-clover, 61 European water-horehound, 640 European weeping birch, 454 European windflower, 752 European yellow-rattle, 695 Eurybia, 399 divaricata, 400 glomerata, 400 ×herveyi, 400 macrophylla, 400 radula, 400 schreberi, 400 spectabilis, 400 Euthamia, 400 caroliniana, 401 galetorum, 401 graminifolia, 401 major, 401 nuttallii, 401 microcephala, 401 remota, 401 septentrionalis, 401 Eutrochium, 401 aculeatum, 402 dubium, 401 fistulosum, 401 maculatum, 402 purpureum, 401 purpureum, 402 Euxolus macrocarpus, 317 viridis, 317 evening-primrose, 686–688 evergreen rose, 807 evergreen winterberry, 352 evergreen wood fern, 57 everlasting, 373, 584 everlasting vetchling, 584 Exochorda, 788 racemosa, 788 eyebane sandmat, 568 eyebright, 692–693

F

Fabaceae, 569–600 Fagaceae, 601–606 Fagopyrum, 728 esculentum, 727 fagopyrum, 727 sagittatum, 727 tataricum, 727 vulgare, 727 Fagus, 601 grandifolia, 602 caroliniana, 602 sylvatica, 602 Fagus-castanea dentata, 601 fairy-slipper, 201 Falcaria, 341 vulgaris, 341 soides, 341 Falcata comosa, 575

fall coral-root, 201 fall crabgrass, 251 fall-dandelion, 426 Fallopia, 729 aubertii, 730 baldschuanica, 730 ×bohemica, 730 cilinodis, 730 convolvulus, 730 cristata, 730 dumetorum, 730 japonica, 729 japonica, 730 sachalinensis, 730 scandens, 731 fall panicgrass, 270 fall phlox, 724 fall sneezeweed, 404 false bartsia, 689 red, 694 false daisy, 395 false flax, 479 false ground-cherry, 859 false heather, 527 false hop sedge, 134 false Indian-plantain, 427 false indigo-bush, 575 false melic grass, 279 false mermaidweed, 657 false pimpernel, 659 false spiraea, 818 false stonecrop, 542 false toadflax, 530 false water-pepper smartweed, 734 fanpetals, 667 fanwort, 85 Farwell’s water-milfoil, 620 Fatoua, 670 villosa, 670 Faxon’s hawthorn, 783 featherbells, 196 featherfoil, 746 feathertop reed grass, 240 feather windmill-grass, 242 feathery false Solomon’s-seal, 300 feathery false threadleaf, 426 February daphne, 867 fen grass-of-Parnassus, 702 fennel, 341 fenugreek, 598 ferns, 52–53, 55–58, 60–61, 65–66, 68–74 fern flatsedge, 154 fern grass, 245 fern-leaved false foxglove, 691 Fern-leaved yarrow, 371 fescue, 259–260 fescue sedge, 123 Festuca, 258 brachyphylla brachyphylla, 259 bromoides, 287 capillata, 259 dertonensis, 287 duriuscula, 259 elatior pro parte, 279 filiformis, 259 heteromalla, 259

megalura, 288 multiflora, 259 myuros, 288 nigrescens, 259 obtusa, 260 octoflora glauca, 288 tenella, 288 ovina, 259 alpina, 259 brachyphylla, 259 brevifolia, 259 duriuscula, 260 rubra, 259 prolifera, 259 rubra, 259 commutata, 260 commutata, 259 densiuscula, 259 fallax, 259 juncea, 259 prolifera, 259 pruinosa, 259 rubra, 259 subverticillata, 260 tenella, 288 glauca, 288 trachyphylla, 260 fetid glandular-goosefoot, 326 fetid-marigold, 394 fetterbush, 557 feverfew, 422 feverfew tansy, 443 few-flowered nutsedge, 170 few-flowered rosettepanicgrass, 249 few-flowered sedge, 133 few-flowered spikesedge, 161 few-nerved cottonsedge, 162 few-seeded sedge, 149 fibrous-rooted sedge, 113 Ficaria, 754 verna bulbifera, 754 Ficus, 670 carica, 670 fiddle dock, 744 fiddlehead fern, 61 fiddleneck, 458–459 field beadgrass, 271 field bindweed, 532 field brome, 237 field chickweed, 509–510 field dodder, 534 field forget-me-not, 464 field garlic, 91 field hedge-nettle, 651 field horsetail, 59 field-madder, 828 field meadow-foxtail, 229 field penny-cress, 492 field pepperweed, 487 field pussytoes, 375 field sow-thistle, 434 field thistle, 390 field wood rush, 191 field wormwood, 378 fig, 670 fig-crowfoot, 754 figwort, 856

93 8   in d e x

Filago, 402 germanica, 402 minima, 417 vulgaris, 402 Filipendula, 788 denudata, 788 hexapetala, 789 rubra, 788 ulmaria, 788 denudata, 788 vulgaris, 789 fimbry, 162 Fimbristylis, 162 autumnalis, 162 mucronulata, 162 fine-leaved sheep fescue, 259 fine-leaved tansy-mustard, 483 fir, 77 fireberry hawthorn, 782 fireweed, 681 firmoss, 39–40 fistulous goat’s beard, 445 five-horned smotherweed, 320 five-leaved-aralia, 341 five-stamened chickweed, 510 five-stamened spurry, 520 flaccid manna grass, 262 flame azalea, 560 flanged brome, 238 flat-branched tree-clubmoss, 44 flat-leaved bladderwort, 656 flatsedge, 153–155 flat-spined sheepburr, 462 flat-stalked pondweed, 295 flat-stemmed blue grass, 275 flat-stemmed spikesedge, 159 flat-stem pondweed, 298 flattened oatgrass, 244 flat-topped-goldenrod, 421 Flaveria, 402 trinervia, 402 flax, 479, 658 flax-leaved stiff-aster, 413 fleabane, 396, 397 fleshy hawthorn, 787 floating bladderwort, 656 floating bur-reed, 304 floating-heart, 669 floating manna grass, 262 floating pondweed, 296 Flodman’s thistle, 390 Floerkea, 657 proserpinacoides, 657 floodplain avens, 791 florist’s daisy, 388 flowering big-bracteddogwood, 537 flowering-quince, 774 flowering raspberry, 816 flowering-rush, 98 flowering spurge, 567 flowering tobacco, 860 flowering yellow wood sorrel, 697 fly honeysuckle, 502 foam-flower, 855 Foeniculum, 341 foeniculum, 341 vulgaris, 341 Fogg’s goosefoot, 323

fool’s-parsley, 337 Forbes’ glory-of-the-snow, 174 Forbes’ hawthorn, 784 forest goldenrod, 431 forest licorice bedstraw, 825 forest lousewort, 695 forget-me-not, 464, 858 forked bluecurls, 653 forked campion, 519 forked rosette-panicgrass, 249 forked rush, 187 fork-tipped threeawn, 232 forsythia, 677 Forsythia, 676 intermedia, 677 suspensa, 677 viridissima, 677 Fragaria, 789 americana, 789 ×ananasa ananasa, 789 canadensis, 789 glauca, 789 grayana, 789 indica, 797 multicipita, 789 vesca, 789 alba, 789 americana, 789 vesca, 789 virginiana, 789 canadensis, 789 glauca, 789 grayana, 789 illinoensis, 789 terrae-novae, 789 virgniana, 789 four-flowered yellowloosestrife, 674 four-leaved manyseed, 516 four-leaved milkweed, 349 four-seeded vetch, 600 four-stamened chickweed, 509 fowl blue grass, 276 Fowler’s knotweed, 739 fowl manna grass, 262 foxglove, 691, 710 foxglove beardtongue, 713 fox grape, 889 fox-tail barley, 263 foxtail bog-clubmoss, 46 foxtail brome, 239 foxtail, 229, 280–281 fox-tail prairie-clover, 579 fox-tail sedge, 150 fragile fern, 72–73 fragrant bedstraw, 827 fragrant orchid, 204 fragrant sumac, 330 fragrant wallflower, 485 fragrant wood fern, 57 Frangula, 762 alnus, 762 Frankenia, 607 pulverulenta, 607 Frankeniaceae, 607 Frasera, 609 albicaulis nitida, 609 nitida, 609

Fraser’s marsh-St. John’s-wort, 627 Fraxinus, 677 americana, 677 biltmoreana, 677 microcarpa, 677 biltmoreana, 677 excelsior, 677 nigra, 677 pennsylvanica, 678 austinii, 678 integerrima, 678 lanceolata, 678 subintegerrima, 678 free-flowered waterweed, 176 freeway sedge, 125 French marigold, 443 French rose, 806 fringed bindweed, 730 fringed brome, 238 fringed-gentian, 610 fringed milkwort, 726 fringed redmaids, 745 fringed sedge, 139 fringed willow-herb, 683 fringed yellow-loosestrife, 674 fringe-tree, 676 Froelichia, 326 gracilis, 326 frogs bit, 176 frosted hawthorn, 786 frost grape, 889 frostweed, 526–527 Fuirena, 162 pumila, 163 squarrosa, 163 hispida, 163 pumila, 163 fuller’s teasel, 500 Fumaria, 701 cucullaria, 701 eximia, 701 formosa, 701 fungosa, 699 glauca, 699 officinalis officinalis, 701 sempervirens, 699 fumitory, 701 Furbish’s lousewort, 695 fuzzy pride-of-Rochester, 622

G

Gadellia, 495 lactiflora, 495 Gagea, 193 fistula, 193 fragifera, 193 liotardii, 193 Gaillardia, 402 aristata, 402 drummondii, 402 picta, 402 pulchella picta, 402 pulchella, 402 Galanthus, 92 nivalis nivalis, 92

Galarhoeus cyparissias, 567 esula, 567 helioscopius, 568 lathyris, 568 maculata, 568 nutans, 568 platyphyllos, 568 supina, 568 Galax, 546 aphylla, 546 urceolata, 546 Galearis, 203 rotundifolia, 200 spectabilis, 203 Galega, 581 officinalis, 581 Galeobdolon argentatum, 639 Galeopsis, 637 bifida, 637 ladanum, 637 latifolia, 637 tetrahit, 637 bifida, 637 Galinsoga, 402 aristulata, 402 bicolorata, 402 caracasana, 402 ciliata, 402 parviflora, 402 semicalva, 402 quadriradiata, 402 Galium, 822 album, 824 anglicum, 825 aparine, 824 echinospermum, 824 spurium, 824 vaillantii, 824 asprellum, 825 boreale, 822 hyssopifolium, 825 intermedium, 825 linearifolium, 825 scabrum, 825 septentrionale, 825 brevipes, 825 circaezans, 825 circaezans, 825 glabrum, 825 hypomalacum, 825 divaricatum, 825 erectum, 824 glaucum, 825 hyssopifolium, 825 kamtschaticum, 825 labradoricum, 825 lanceolatum, 825 mollugo, 826 erectum, 824 obtusum, 826 obtusum, 826 ramosum, 826 odoratum, 826 palustre, 826 parisiense anglicum, 825 divaricatum, 825

in d e x  9 3 9

Galium, cont. pilosum, 826 pilosum, 826 punticulosum, 826 punctatum, 826 septentrionale, 825 spurium, 824 echinospermum, 824 vaillantii, 824 sylvaticum, 826 tinctorium, 827 labradoricum, 825 tricornutum, 826 floridanum, 826 tricornutum, 826 trifidum, 827 brevipes, 825 latifolium, 826 halophilum, 827 trifidum, 827 triflorum, 827 asprelliforme, 827 vaillantii, 824 verum, 827 verum, 827 wirtgenii, 827 wirtgenii, 827 gamagrass, 286 Gamochaeta, 403 pensylvanica, 403 purpurea, 404 garden bird’s-foot-trefoil, 588 garden black currant, 618 garden buckwheat, 728 garden chervil, 338 garden cosmos, 392 garden evening-primrose, 687 garden knapweed, 387 garden lovage, 343 garden mignonette, 762 garden monkshood, 750 garden-nasturtium, 867 garden onion, 91 garden orache, 319 garden orpine, 541 garden pea, 592 garden pepperweed, 488 garden petunia, 861 garden pink, 511 garden red currant, 619 garden rhubarb, 740 garden-rocket, 484 garden smartweed, 736 garden snapdragon, 707 garden thyme, 653 garden tomato, 865 garden violet, 886 garden yellow-loosestrife, 675 garden yellow-rocket, 476 garlic, 91–92 garlic-mustard, 474 Gaspe Peninsula arrow-grass, 192 gasplant, 829 Gastridium, 260 phleoides, 260 ventricosum, 260 Gaultheria, 554 hispidula, 555 procumbens, 554

Gaura biennis, 687 mollis, 686 parviflora, 686 Gaylussacia, 555 baccata, 555 bigeloviana, 555 dumosa bigeloviana, 555 frondosa, 555 Gemmingia chinensis, 179 Genista, 581 tinctoria, 581 gentian, 610 Gentian pulchella, 609 Gentiana, 609 amarella, 610 andrewsii, 609 andrewsii, 610 clausa, 610 crinita, 610 linearis, 610 latifolia, 610 quinquefolia, 610 rotata, 611 rubricaulis, 610 saponaria linearis, 610 Gentianaceae, 607–612 Gentianaceae, 29 Gentianella, 610 acuta, 610 amarella acuta, 610 stricta, 610 amarelloides, 610 quinquefolia quinquefolia, 610 Gentianopsis, 610 crinita, 610 Geocaulon, 530 lividum, 530 Georgia bulrush, 169 Geraniaceae, 612–616 Geranium, 614 aequale, 615 bicknellii, 616 longipes, 616 botrys, 613 carolinianum, 616 confertiflorum, 616 sphaerospermum, 616 ciconium, 613 cicutarium, 613 columbinum, 616 dissectum, 616 laxum, 616 maculatum, 616 malacoides, 614 molle, 616 moschatum, 614 nepalense thunbergii, 616 pratense, 616 pusillum, 616 pyrenaicum, 616 robertianum, 616

sanguineum, 616 sibiricum, 616 thunbergii, 616 Gerardia acuta, 689 besseyana, 690 flava, 691 maritima, 690 neoscotica, 690 paupercula typica, 690 pedicularia intercedens, 691 purpurea, 690 borealis, 690 grandiflora, 690 parviflora, 690 parvula, 690 paupercula, 690 tenuifolia, 690 humilis, 690 macrophylla, 690 parviflora, 690 typica, 690 virginica, 691 germander, 652, 719, 820 germander meadowsweet, 820 germander speedwell, 719 German hedge-nettle, 651 German iris, 180 German madwort, 459 Geum, 790 aleppicum, 791 strictum, 791 canadense, 791 brevipes, 791 camporum, 791 canadense, 791 fragarioides, 791 laciniatum, 791 laciniatum, 791 trichocarpum, 791 macrocarpon, 791 macrophyllum, 791 peckii, 790 ×pulchrum, 791 rivale, 790 strictum, 791 urbanum, 790 vernum, 790 virginianum, 791 gherkin, 544 giant bellflower, 495 giant-chickweed, 514 giant cow-parsnip, 342 giant-daisy, 416 giant-hyssop, 635 giant knotweed, 730 giant needle-leaf, 327 giant ragweed, 373 giant rattlesnake-plantain, 204 giant rye grass, 279 giant-scabious, 499 giant-seeded goosefoot, 324 giant spider-flower, 530 gilia, 722 Gilia, 722 achilleifolia multicaulis, 722 diffusa tricolor, 722

interior, 722 multicaulis, 722 alba, 722 tenuiflora interior, 722 tricolor diffusa, 722 Gillenia, 792 trifoliata, 792 Gill-over-the-ground, 637 ginger wild, 81 ginger mint, 642 ginseng, 344 Gisopteris palmata, 61 glade-mallow, 666 glandular baby’s-breath, 512 glandular birch, 453 glandular-goosefoot, 326 Glandularia, 871 angustifolia, 872 canadensis, 871 drummondii, 871 lambertii, 871 Glareosae, 125 glasswort, 327–328 Glaucium, 701 flavum, 701 glaucous blue grass, 276 glaucous hair grass, 244 glaucous hawkweed, 412 glaucous-leaved greenbrier, 301 glaucous rattlesnake-root, 419 glaucous sedge, 147 glaucous tuber-bulrush, 103 Glaux maritima, 674 angustifolius, 674 obtusifolia, 674 Glebionis, 403 carinata, 403 coronaria, 403 segetum, 404 Glechoma, 637 hederacea, 637 micrantha, 637 parviflora, 637 Gleditsia, 582 triacanthos, 582 inermis, 582 globe-amaranth, 326 globe candytuft, 486 globe flatsedge, 153 globe-flower, 761 globe-mallow, 667 globe-thistle, 395 globe yellow-cress, 490 glory-of-the-snow, 174 Glossostigma, 704 cleistanthum, 704 diandrum, 704 glossy false buckthorn, 762 Glyceria, 260 acutiflora, 261 borealis, 261 canadensis, 261 fernaldii, 285 fluitans, 261

940 in dex

Glyceria, cont. grandis grandis, 262 laxa, 262 maxima, 262 americana, 262 grandis, 262 melicaria, 262 obtusa, 262 pallida, 285 paupercula, 278 septentrionalis septentrionalis, 262 spectabilis, 262 striata, 262 stricta, 262 Glycine, 582 apios, 576 floribunda, 600 max, 582 sinensis, 600 Glycyrrhiza, 582 glutinosa, 582 lepidota, 582 glutinosa, 582 Gnaphalium, 403 decurrens, 424 helleri micradenium, 424 involucratum, 397 macounii, 424 margaritaceum, 373 obtusifolium micradenium, 424 pensylvanicum, 403 plantaginifolium, 375 purpureum spathulatum, 403 supinum, 421 sylvaticum, 421 uliginosa, 404 goat-grass, 226 goat’s beard, 445, 774 goat’s rue, 595 goat willow, 845 golden-aster, 388, 409 golden chain-tree, 583 golden-chamomile, 392 golden-club, 96 golden corydalis, 700 golden crownbeard, 445 golden currant, 618 golden dock, 743–744 goldeneye, 408 golden-fruited sedge, 115 golden groundsel, 422 golden hedge-hyssop, 710 golden mane tickseed, 391 golden rain-tree, 852 goldenrod, 400–401, 421, 428–434 golden-saxifrage, 853 goldenseal, 754 golden tickseed, 392 goldentuft, 475 goldfields, 415–416 Goldie’s wood fern, 57 goldthread, 754 Gomphrena, 326 globosa, 326

Goodyera, 203 decipiens, 204 oblongifolia, 204 ophioides, 204 pubescens, 204 repens, 204 ophioides, 204 tesselata, 204 gooseberry, 618–619 goosefoot, 321–325 goosegrass, 252 goose-neck yellow-loosestrife, 674 gopher spurge, 568 gorse, 598 Gossypium, 664 herbaceum, 664 hirsutum hirsutum, 664 punctatum, 664 purpurascens, 664 gourd, 545 goutweed, 337 graceful cinquefoil, 796 graceful sedge, 131 Gracilis, 127 Graham’s rockcress, 477 grama, 234 Grammica campestris, 534 gronovii, 534 indecora, 535 pentagona, 535 granny morning-glory, 536 Granulares, 128 grape, 888–889 grapefern, 64–65 grape honeysuckle, 502 grape-hyacinth, 174 graphephorum, 262 Graphephorum, 262 melicoides, 262 grass-leaved arrowhead, 90 grass-leaved-goldenrod, 401 grass-leaved mud-plantain, 289 grass-leaved rush, 186, 188 grass-leaved stitchwort, 522 grass-like sedge, 137 grass-of-Parnassus, 702 grass-pink, 201 grassy pondweed, 295 grasswort, 343 Gratiola, 710 aurea, 710 obtusa, 710 dubia, 659 neglecta, 710 glaberrima, 710 sphaerocarpa, 710 virginiana virginiana, 710 gray birch, 454 gray club-awn grass, 242 gray dogwood, 539 gray field speedwell, 719 gray goldenrod, 432 gray-headed Mexican-hat, 424 Gray’s flatsedge, 154 Gray’s sedge, 134 gray willow, 845 great burdock, 376

great burnet, 817 great bur-reed, 303 greater bladder sedge, 134 greater bladderwort, 657 greater brown sedge, 127 greater Canada St. John’s-wort, 626 greater celandine, 700 greater creeping rush, 189 greater dodder, 534 greater false leopard’s bane, 394 greater fringed-gentian, 610 greater globe-thistle, 395 greater knapweed, 387 greater masterwort, 339 greater periwinkle, 351 greater purple fringed bogorchid, 209 greater stitchwort, 523 greater straw sedge, 124 greater swine-cress, 488 greater tickseed, 391 greater water dock, 742 greater water-starwort, 708 greater yellow water crowfoot, 758 great lobelia, 496 Great Plains flatsedge, 154 Great Plains wild-rye, 254 great rosebay, 560 great-spurred violet, 886 great St. John’s-wort, 625 great yellow-cress, 490 Grecian foxglove, 710 Greek windflower, 751 green adder’s-mouth, 206 green alder, 451 green amaranth, 316 green arrow-arum, 96 green ash, 678 greenbrier, 301 green buckwheat, 728 green carpetweed, 669 green-dragon, 94 Greene’s rush, 188 green field speedwell, 718 green-flowered shinleaf, 558 green foxtail, 281 green fringed bog-orchid, 209 greenhead-sedge, 163 green-headed coneflower, 425 green-keeled cottonsedge, 162 green milkweed, 350 Green Mountain maidenhair fern, 68 green rockcress, 477 green spleenwort, 52 green-stemmed forsythia, 677 green-violet, 873 greenweed, 581 green-white sedge, 123 Grindelia, 404 hirsutula, 404 squarrosa, 404 integrifolia, 404 quasiperennis, 404 serrulata, 404 Griseae, 128–129 gromwell, 463–465 grooved yellow flax, 658

Grossularia cynosbati, 618 hirtella, 618 missouriensis, 618 reclinata, 619 rotundifolia, 618 Grossulariaceae, 616–619 ground-cedar, 44–45 ground-cherry, 861–863 ground-cherry nightshade, 865 ground-nut, 576 groundsel, 421–422 ground virgin’s-bower, 753 grove blue grass, 275 grove hawthorn, 785 grove-sandwort, 514 Guadalupe waternymph, 177 Guilandina dioica, 582 Guizotia, 404 abyssinica, 404 gum-weed, 404 Gymnadenia, 204 conopsea, 204 Gymnadeniopsis clavellata, 209 Gymnocarpium, 73 ×brittonianum, 73 continentale, 73 dryopteris, 73 jessoense, 73 parvulum, 73 Gymnocladus, 582 dioicus, 582 Gymnosperms, 75–80 Gypsophila, 511 elegans, 511 muralis, 512 paniculata, 512 scorzonerifolia, 512 Gyrostachys beckii, 212 gracilis, 211 ochroleuca, 212 stricta, 212 Gyrotheca tinctoria, 173

H Habenaria blephariglottis, 208 bracteata, 201 ciliaris, 208 clavellata, 209 conopsea, 204 cristata, 209 dilatata, 209 fimbriata, 209 flava, 209 grandiflora, 209 hookeri, 209 huronensis, 209 hyperborea huronensis, 209 lacera, 209 leucophaea, 210 macrophylla, 210 obtusata, 210 orbiculata, 210 macrophylla, 210

i n d e x  9 41

Habenaria, cont. psycodes, 210 rotundifolia, 200 viridis, 201 hackberry, 497 Hackelia, 461 americana, 461 deflexa americana, 461 virginiana, 461 Haemodoraceae, 173 hair-awned muhly, 267 hair grass, 244, 287 hair-like sedge, 118 hair-sedge, 104 hairy Angelica, 338 hairy bedstraw, 826 hairy beggar-ticks, 383 hairy bitter-cress, 482 hairy bush-clover, 587 hairy cat’s-ear, 413 hairy crabgrass, 251 hairy crabweed, 670 hairy crowfoot, 759 hairy eared-rockcress, 475 hairy evening-primrose, 688 hairy false bindweed, 531 hairy forked whitlow-wort, 515 hairy-fruited sedge, 116 hairy goldenrod, 432 hairy ground-cherry, 863 hairy gum-weed, 404 hairy hedge-nettle, 651 hairy honeysuckle, 502 hairy lady’s-mantle, 769 hairy lip fern, 68 hairy mock-vervain, 872 hairy nightshade, 865 hairy pine-sap, 556 hairy pinweed, 528 hairy rosette-panicgrass, 247 hairy rupturewort, 512 hairy sandmat, 568 hairy small-leaved tick-trefoil, 580 hairy smotherweed, 320 hairy Solomon’s-seal, 300 hairy-stemmed gooseberry, 618 hairy sunflower, 407 hairy thoroughwort, 399 hairy tickseed, 391 hairy toad-lily, 195 hairy umbrella-sedge, 163 hairy umbrella-wort, 675 hairy vetch, 600 hairy willow-herb, 683 hairy wood brome, 239 hairy wood-mint, 636 hairy wood rush, 190 halberd-leaved tearthumb, 734 Halenia, 611 deflexa deflexa, 611 heterantha, 611 heterantha, 611 Halerpestes cymbalaria, 758 saximontana, 758 Halesia, 866 carolina, 866 Hall’s bulrush, 166

Haloragaceae, 619–622 Hamamelidaceae, 622 Hamamelis, 622 virginiana, 622 parvifolia, 622 hammer sedge, 116 handsome sedge, 131 Handy’s hawthorn, 784 hard fescue, 260 hard-stemmed bulrush, 166 hare’s-ear-mustard, 483 Harpalyce alba, 419 altissima, 419 Harrimanella, 555 hypnoides, 555 Hasteola suaveolens, 427 hastate-leaved orache, 319 hawkbit, 416 hawk’s-beard, 392–393 hawkweed, 411–413 hawkweed oxtongue, 423 hawthorn, 774–778 Hayden’s sedge, 140 hazel dodder, 534 hazelnut, 454 heart-leaved American-aster, 439 heart-leaved golden Alexanders, 347 heart-leaved paper birch, 453 heart-leaved peppervine, 887 heart-leaved twayblade, 207 heart-leaved umbrella-wort, 676 heart-leaved willow, 846 heart-podded pepperweed, 487 heath, 554 heath American-aster, 439 heather, 527, 553 heath false brome, 235 Hecatonia scelerata, 759 Hedeoma, 638 hispida, 638 pulegioides, 638 Hedera, 342 helix, 342 quinquefolia, 888 hedge false bindweed, 531 hedgehog wood rush, 191 hedge-hyssop, 710 hedge maple, 850 hedge-mustard, 492 hedge-nettle, 651 hedgerow crane’s-bill, 616 hedge-parsley, 346 hedge vetch, 600 Hedyotis caerulea, 828 longifolia, 828 tenuifolia, 828 purpurea longifolia, 828 Hedysarum, 582 alpinum americanum, 582 americanum, 582 canadense, 580

canescens, 580 ciliare, 580 cuspidatum, 580 glabellum, 580 glutinosum, 582 grandiflorum, 580 marilandicum, 581 nudiflorum, 583 obtusum, 581 paniculatum, 581 repens, 588 sessilifolium, 581 violaceum, 588 Helanthium, 88 tenellum, 88 parvulum, 88 Helenium, 404 amarum amarum, 404 autumnale, 404 canaliculatum, 404 parviflorum, 404 canaliculatum, 404 flexuosum, 404 nudiflorum, 404 parviflorum, 404 tenuifolium, 404 Heleochloa schoenoides, 242 Heleoglochin, 129 Helianthemum bicknellii, 526 canadense, 526 sabulonum, 526 dumosum, 526 propinquum, 527 Helianthus, 404 alienus, 407 annuus, 406 lenticularis, 406 macrocarpus, 406 texanus, 406 borealis, 407 cucumerifolius, 406 debilis, 405 cucumerifolius, 406 decapetalus, 407 divaricatus, 407 angustifolius, 407 ×divariserratus, 407 giganteus, 407 alienus, 407 subtuberosus, 407 grosseserratus, 407 hypoleucus, 407 hirsutus, 407 stenophyllus, 407 ×intermedius, 407 ×kellermanii, 407 laetiflorus, 408 rigidus, 407 subrhomboideus, 407 laevis, 383 ×luxurians, 407 maximiliani, 407 microcephalus, 407 mollis, 407 cordatus, 407 occidentalis, 405 occidentalis, 407

pauciflorus, 407 pauciflorus, 408 subrhomboideus, 408 petiolaris, 405 petiolaris, 408 rigidus, 407 salicifolius, 407 strumosus, 408 tomentosus, 408 tracheliifolius, 407 tuberosus, 408 Helichrysum bracteatum, 447 Helictotrichon pubescens, 233 Heliomeris, 408 multiflora multiflora, 408 Heliopsis, 408 helianthoides helianthoides, 409 scabra, 409 helianthoides, 408 heliotrope, 462 Heliotropium, 461 amplexicaule, 462 anchusifolium, 462 curassavicum, 462 europaeum, 462 indicum, 462 hellebore, 198, 203 helleborine, 203 Helleborus trifolius, 754 Helminthotheca, 409 echioides, 409 Helonias graminea, 196 Hemerocallidaceae, 173 Hemerocallis, 173 flava, 173 fulva, 173 japonica, 173 lilioasphodelus, 173 fulvus, 173 Hemicarpha micrantha, 163 Hemizonia fasciculata, 394 ramosissima, 394 hemlock, 80, 340 hemlock-parsley, 340 hemp, 348, 497 hemp dogbane, 348 hemp-nettle, 637 hempvine, 418 henbane, 859 henbit, 638–639 hens-and-chickens, 543 hepatica, 751 Hepatica acutiloba, 751 americana, 751 nobilis acuta, 751 obtusa, 751 Heracleum, 342 lanatum, 342 mantegazzianum, 342 maximum, 342

942   in de x

Heracleum, cont. sphondylium, 342 lanatum, 342 montanum, 342 herbaceous periwinkle, 351 herbaceous sea-blite, 329 Hercules’-club, 339 Herniaria, 512 glabra, 512 hirsuta cinerea, 512 Hesperis, 486 altissima, 492 loeselii, 492 matronalis, 486 Heteranthera, 289 dubia, 289 graminea, 289 reniformis, 289 Heterosperma, 409 pinnatum, 409 Heterotheca, 409 falcata, 423 mariana, 388 subaxillaris latifolia, 409 Heuchera, 853 americana americana, 853 Hibiscus, 664 esculentus, 663 incanus, 664 moscheutos incanus, 664 moscheutos, 664 palustris, 664 oculiroseus, 664 palustris, 664 syriacus, 664 trionum, 665 Hickey’s tree-clubmoss, 44 hickory, 627–629 Hicoria glabra, 628 hirsuta, 628 laciniosa, 628 ovata, 629 tomentosa, 629 hidden dropseed, 284 hidden spikemoss, 49 Hieracium, 409 aurantiacum, 411 caespitosum, 411 canadense, 412 fasciculatum, 412 kalmii, 412 ×fernaldii, 412 flagellare, 411 pilosius, 411 florentinum, 412 ×floribundum, 411 gronovii, 411 kalmii, 412 fasciculatum, 412 lachenalii, 411 maculatum, 412 ×marianum, 412 murorum, 412 officinarum, 412 paniculatum, 412

pilosella, 412 niveum, 412 piloselloides, 412 praealtum, 412 decipiens, 412 pratense, 411 robinsonii, 412 sabaudum, 412 scabriusculum, 413 saximontanum, 413 scabrum, 413 tonsum, 412 umbellatum, 413 ungavense, 412 venosum, 413 Hierochloe alpina, 231 hirta, 231 monticola, 231 odorata, 231 fragrans, 231 orthantha, 231 highbush, 562–563 highbush blueberry, 562–563 highbush-cranberry, 310 highland dog-laurel, 556 highland rush, 189 high mallow, 666 hillside blueberry, 563 Hill’s pondweed, 296 Himalayan blackberry, 813 Himalaya touch-me-not, 448 Hippochaete hyemalis, 60 affinis, 60 scirpoides, 60 variegata, 60 Hipposelinum levisticum, 343 Hippuris, 710 vulgaris, 710 hirsute sedge, 144 Hirtifoliae, 129 hispid crowfoot, 759 hispid hedge-nettle, 651 Hitchcock’s sedge, 129 hoary false alyssum, 476 hoary frostweed, 526 hoary mock-orange, 623 hoary mountain-mint, 646 hoary plantain, 716 hoary sedge, 127 hoary tick-trefoil, 580 hoary vervain, 872 hoary willow, 846 hobblebush, 310 hog-peanut, 575 Holcus, 262 bicolor, 282 halepensis, 282 lanatus, 262 mollis mollis, 262 sorghum, 282 hollow Joe-Pye weed, 401 holly, 351–352 hollyhock, 664 holly-leaved Oregon-grape, 449 Holmes’ hawthorn, 784

Holosteum, 512 umbellatum, 512 Homalocenchrus oryzoides, 264 virginicus, 264 Honckenya, 512 peploides robusta, 512 honesty, 488 honewort, 341 honey-locust, 582 honeysuckle, 500–502 hooded ladies’-tresses, 212 hooded skullcap, 649 hooded windmill-grass, 242 hooked crowfoot, 759 hooked foxtail, 281 Hooker’s bog-orchid, 209 hook-spurred violet, 880 hop, 134–135, 498, 597 hop-hornbeam, 455 hop sedge, 134–135 hoptree, 829 Hordeum, 263 boreale, 263 brachyantherum brachyantherum, 263 distichon, 263 hexastichon, 263 jubatum jubatum, 263 leporinum, 263 murinum jubatum, 263 nodosum, 263 boreale, 263 pusillum, 263 vulgare, 263 trifurcatum, 263 horehound, 635, 639–641 hornbeam, 454 horned bladderwort, 656 horned nightshade, 865 horned-pondweed, 298 Hornemann’s willow-herb, 683 horn-leaved riverweed, 721 horn-poppy, 701 hornwort, 82–83 horsebalm, 636 horse-chestnut, 852 horse-gentian, 504 horse mint, 643 horse-radish, 475 horsetail, 58–60 horsetail spikesedge, 159 horse yellowhead, 413 Hortulan plum, 800 Hosta, 173 japonica, 173 lancifolia, 173 ventricosa, 173 Hostaceae, 173 Hottonia, 746 inflata, 746 hound’s tongue, 460 Houghton’s flatsedge, 154 Houghton’s sedge, 136 Houstonia, 827 caerulea, 828 faxonorum, 828

longifolia, 828 tenuifolia, 828 purpurea calycosa, 828 Houston’s whiteweed, 372 huckleberry, 555 Hudsonia, 527 ericoides, 527 andersonii, 527 intermedia, 527 intermedia, 527 tomentosa, 527 intermedia, 527 Humulus, 498 americanus, 498 japonicus, 498 lupulus, 498 americanus, 498 lupuloides, 498 lupulus, 498 Hungarian lilac, 679 Hungarian milk-vetch, 576 Hungarian vetch, 599 Huperzia, 39 appalachiana, 39 appressa, 39 ×buttersii, 40 josephbeitelii, 39 lucidula, 40 protoporophila, 39 selago, 40 Huperziaceae, 39–40 hyacinth grape, 174 water, 289 Hyacinthaceae, 174 Hyacinthus botryoides, 174 Hybanthus, 873 concolor, 873 Hydatica foliolosa, 853 hydrangea, 623 Hydrangea, 622 arborescens, 623 discolor, 623 oblonga, 623 cinerea, 623 paniculata, 623 quercifolia, 623 Hydrangeaceae, 622–623 Hydrastis, 754 canadensis, 754 Hydrilla, 176 verticillata, 176 Hydrocharis, 176 morsus-ranae, 176 Hydrocharitaceae, 175–177 Hydrocotyle, 342 americana, 342 umbellata, 342 verticillata, 343 Hydrophila aquatica, 541 Hydrophyllum, 462 canadense, 462 linarea, 466 virginianum, 462 atranthum, 462

in d e x  9 43

Hylodesmum, 582 glutinosum, 582 nudiflorum, 583 Hylotelephium, 541 erythrostictum, 541 spectabile, 541 telephium, 541 fabaria, 541 telephium, 541 Hymenatherum tenuiloba, 444 Hymenochlaenae, 130–131 Hymenophyllaceae, 60 Hymenoxys, 413 odorata, 413 Hyoscyamus, 859 niger, 859 Hyoseris amplexicaulis, 414 virginica, 414 Hypericaceae, 624–627 Hypericum, 624 adpressum, 625 ascyron, 625 aureum, 626 bissellii, 626 boreale, 625 canadense, 625 galiiforme, 625 magninsulare, 625 majus, 626 densiflorum, 626 ×dissimulatum, 625 dolabriforme, 626 ellipticum, 626 frondosum, 626 gentianoides, 626 glomeratum, 626 hypericoides multicaule, 626 majus, 626 mutilum, 626 boreale, 625 parviflorum, 626 perforatum, 625 perforatum, 626 prolificum, 626 punctatum, 626 pyramidatum, 625 stragulum, 626 subpetiolatum, 626 virginicum, 627 fraseri, 627 Hypochaeris, 413 glabra, 413 radicata, 413 Hypopitys, 555 americana, 556 lanuginosa, 556 monotropa, 556 Hypoxidaceae, 178 Hypoxis, 178 carolinensis, 178 hirsuta, 178 hyssop, 638 hyssop-leaved fleabane, 396 hyssop-leaved hedge-nettle, 651 hyssop-leaved loosestrife, 661 hyssop-leaved thoroughwort, 399

Hyssopus, 638 anethiodorus, 635 nepetoides, 635 officinalis, 638 scrophulariifolius, 635 Hystrix hystrix, 254 patula bigeloviana, 254

I

Iberis, 486 amara, 486 sempervirens, 486 umbellata, 486 Ibidium cernuum, 211 gracile, 211 plantagineum, 212 strictum, 212 vernale, 212 Ide’s hawthorn, 784 Ilex, 351 ambigua montana, 352 amelanchier monticola, 352 aquifolium, 352 crenata, 352 glabra, 352 laevigata, 352 montana, 352 mucronata, 351 opaca, 352 verticillata, 352 Illecebrum, 512 verticillatum, 512 Illinois pondweed, 296 Ilysanthes attenuata, 659 dubia, 659 inequalis, 659 immigrant pond-lily, 85 Impatiens, 447 biflora, 448 capensis, 448 glandulifera, 448 noli-tangere biflora, 448 nortonii, 448 pallida, 448 roylei, 448 India lovegrass, 258 Indian cucumber root, 194 Indian grass, 281 Indian hedge-mustard, 492 Indian heliotrope, 462 Indian-mallow, 663 Indian-pipe, 557 Indian-plantain, 427 Indian-strawberry, 797 Indian sweet-clover, 591 Indian tobacco, 861 indigo, 577 indigo-bush, 575 inflated narrow-leaved sedge, 129 inland gilia, 722 inland sedge, 147 interior blue grass, 276

intermediate shadbush, 771 interrupted-clubmoss, 48 interrupted fern, 66 Inula, 413 alba, 421 graveolens, 394 helenium, 413 mariana, 388 salicina, 413 Ionactis, 413 linariifolia, 413 Ionoxalis violacea, 697 trichophora, 697 Ioxylon pomiferum, 670 ipecac spurge, 568 Ipomoea, 535 barbigera, 536 coccinea, 536 hederifolia, 536 hederacea, 536 integriuscula, 536 hederifolia, 536 lacunosa, 536 purpurea, 536 quamoclit, 536 tricolor, 536 Ipomopsis, 722 rubra, 722 Iridaceae, 178–181 iris, 179–180 Iris, 178 canadensis, 180 cristata, 179 domestica, 179 ensata, 180 germanica, 180 hookeri, 180 kaempferi, 180 prismatica, 180 pseudacorus, 180 pumila, 180 setosa canadensis, 180 sibirica, 180 tectorum, 180 versicolor, 181 Irish potato, 865 ironweed, 446 ironwort, 650 Ischaemum ophiuroides, 258 Isnardia palustris, 684 Isoetaceae, 2, 40–42 Isoetes, 40 acadiensis, 41 ×brittonii, 41 canadensis, 42 ×dodgei, 41 ×eatonii, 41 echinospora, 40 muricata, 41 ×echtuckerii, 41 engelmannii, 42 ×foveolata, 41 ×harveyi, 42 hieroglyphica, 42 lacustris, 42 hieroglyphica, 42

×novae-angliae, 42 prototypus, 42 riparia, 42 tuckermanii, 42 Isotrema, 82 macrophyllum, 82 tomentosum, 82 Isotria, 204 medeoloides, 205 verticillata, 205 Italian bugloss, 459 Italian rye grass, 265 Italian Timothy, 272 Iva, 414 annua, 414 ciliata, 414 frutescens, 414 oraria, 414 xanthiifolia, 394 ivy, 342, 709 ivy-leaved duckweed, 96 ivy-leaved morning-glory, 536 ivy-leaved speedwell, 719 Ixia chinensis, 179

J Jacea nigra, 387 pratensis, 387 Jack-in-the-pulpit, 94 Jack pine, 79 Jacobaea, 414 vulgaris, 414 Jacob’s ladder, 724–725 jagged-chickweed, 512 Jamaican-forget-me-not, 858 Japanese angelica-tree, 339 Japanese arrow bamboo, 277 Japanese azalea, 560 Japanese barberry, 449 Japanese black pine, 79 Japanese brome, 238 Japanese-clover, 583 Japanese flowering cherry, 801 Japanese flowering crab apple, 793 Japanese flowering-quince, 774 Japanese holly, 352 Japanese honeysuckle, 502 Japanese hop, 498 Japanese knotweed, 730 Japanese maple, 851 Japanese mazus, 704 Japanese meadowsweet, 820 Japanese mountain-spurge, 493 Japanese painted lady fern, 71 Japanese pearlwort, 516 Japanese primrose, 747 Japanese raspberry, 816 Japanese-rose, 792 Japanese sedge, 135 Japanese snowball, 311 Japanese stiltgrass, 265 Japanese sweet-coltsfoot, 423 Japanese tree lilac, 679 Japanese violet, 883 Japanese walnut, 629

944   in de x

Japanese wisteria, 600 Japanese yew, 80 Jasione, 495 montana, 496 Jasmine tobacco, 860 Jerusalem-cherry nightshade, 865 Jerusalem-oak, 326 Jesup’s hawthorn, 784 jetbead, 802 Joe-Pye weed, 401–402 Johnson grass, 282 jointed goat-grass, 226 joint-leaved rush, 186 joyweed, 314 Juglandaceae, 627–629 Juglans, 629 ailantifolia, 629 alba, 629 pro parte, 629 bixbyi, 629 cinerea, 629 cordiformis, 628 laciniosa, 628 nigra, 629 sieboldiana, 629 jumpseed, 736 Juncaceae, 181–189 Juncaginaceae, 192 Juncoides bulbosum, 191 campestre, 191 echinatum, 191 hyperboreum, 191 intermedium, 191 melanocarpus, 191 nemorosum, 191 saltuense, 190 spicatum, 191 Juncus, 182 acuminatus, 186 alpinoarticulatus, 186 americanus, 186 nodulosus, 186 alpinus, 186 ambiguus, 186 anthelatus, 186 arcticus littoralis, 186 articulatus, 186 biflorus, 186 obtusatus, 186 balticus, 182 littoralis, 186 biflorus, 187 brachycarpus, 187 brachycephalus, 187 brevicaudatus, 187 bufonius, 187 halophilus, 186 canadensis, 187 compressus, 187 conglomeratus, 187 debilis, 187 dichotomus, 187 diffusissimus, 187 dudleyi, 187

effusus, 187 conglomeratus, 187 effusus, 188 pylaei, 188 solutus, 188 ensifolius, 188 filiformis, 188 gerardii, 188 greenei, 188 inflexus, 182 marginatus, 188 biflorus, 186 militaris, 188 nodusus, 188 ×oronensis, 189 pelocarpus, 188 crassicaudex, 186 pervetus, 188 platyphyllus, 187 pylaei, 188 secundus, 189 stygius, 184 americanus, 189 subcaudatus, 189 subtilis, 189 tenuis, 189 anthelatus, 186 dichotomus, 187 dudleyi, 187 torreyi, 189 trifidus, 189 vaseyi, 189 xiphioides triandrus, 188 juniper, 75–76 Juniperus, 75 canadensis, 75 communis, 75 depressa, 75 horizontalis, 76 prostrata, 76 repens, 76 virginiana, 75 crebra, 76 prostrata, 76 virginiana, 76 Justicia, 307 americana, 307

K

Kalmia, 556 angustifolia angustifolia, 556 latifolia, 556 polifolia, 556 procumbens, 556 Kalm’s brome, 238 Kalopanax, 343 septemlobus, 343 Kansas hawthorn, 783 Katsura-tree, 525 Keep’s hawthorn, 785 Kenilworth-ivy, 709 Kennedy’s hawthorn, 785 Kentucky blue grass, 276 Kentucky coffee-tree, 582 Kentucky yellow-wood, 578 Kerria, 792 japonica, 792

khaki joyweed, 314 Kickxia, 710 elatine, 710–711 spuria, 710–711 kidney bean, 592 kidney-leaved crowfoot, 757 kidney-leaved mud-plantain, 289 kidney-leaved violet, 885 kidney-vetch, 575 King Solomon’s-seal, 300 knapweed, 386–388 Knautia, 500 arvensis, 500 knawel, 517 Kneiffia arenicola, 686 fruticosa, 686 glauca, 686 latifolia, 686 linearis, 686 perennis, 687 pratensis, 687 pumila, 687 tetragona, 686 knight’s spur, 754 knotroot foxtail, 280 knotted clover, 598 knotted pearlwort, 516 knotted rush, 188 knotweed, 736–740 Kobresia elachycarpa, 147 Kochia, 326 atriplicifolia, 325 hirsuta, 320 hyssopifolia, 320 scoparia, 326 pubescens, 326 Koeleria, 263 cristata, 263 macrantha, 263 Koellia dubia, 646 flexuosa, 647 incana, 646 mutica, 647 pilosa, 647 verticillata, 647 Koelreuteria, 852 paniculata, 852 Kolkwitzia, 500 amabilis, 500 Koniga maritimum, 488 Korea barberry, 449 Korean mountain-ash, 818 kousa big-bracted-dogwood, 537 Kraunhia floribunda, 600 frutescens, 600 macrostachya, 600 Krigia, 414 amplexicaulis, 414 biflora biflora, 414 glandulifera, 414 virginica, 414

kudzu, 592 Krynitzkia californica, 466 Kummerowia, 583 striata, 583 Kyllinga, 163 gracillima, 163

L Labrador-tea, 560 Labrador willow, 844 Laburnum, 583 alpinum, 583 anagyroides, 583 ×watereri, 583 lace lovegrass, 257 Lachnanthes, 173 caroliniana, 173 tinctoria, 173 Lacinaria pycnostachya, 417 scariosa novae-angliae, 417 spicata, 417 Lactuca, 414 biennis, 415 canadensis, 415 integrifolia, 415 hirsuta sanguinea, 415 ×morssii, 415 muralis, 418 pulchella, 418 sativa, 415 scariola, 415 integrata, 415 serriola, 415 spicata integrifolia, 415 lacy scorpion-weed, 466 ladies’s-tresses, 211–212 lady’s-mantle, 769 lady’s slipper, 202–203 lady’s-thumb smartweed, 734–735 Lagenaria, 545 siceraria, 545 Lake Huron green bog-orchid, 209 Lake Mistassini primrose, 747 lake quillwort, 42 lake shore sedge, 140 lakeside sedge, 136 lake yellow-cress, 490 Lamarck’s bedstraw, 825 lamb-succory, 376 Lamiaceae, 630–653 Lamium, 638 album, 639 amplexicaule, 639 dissectum, 639 galeobdolon, 639 hybridum, 639 maculatum, 639 purpureum, 639 incisum, 639 purpureum, 639 Lamprocapnos, 701 spectabilis, 701

i n d e x  9 45

lance-leaved American-aster, 440 lance-leaved arnica, 376 lance-leaved figwort, 856 lance-leaved licorice bedstraw, 825 lance-leaved ragweed, 373 lance-leaved thorough-wax, 340 lance-leaved tickseed, 391 lance-leaved tiger lily, 194 lance-leaved twistedstalk, 195 lance-leaved violet, 883 Landoltia, 95 punctata, 95 Lapathum crispum, 743 minus, 743 Lapland-crowfoot, 754 Lapland rosebay, 560 Laportea, 869 canadensis, 869 Lappa minor, 376 Lappula, 462 americana, 461 deflexa americana, 461 echinata occidentalis, 462 occidentalis occidentalis, 462 squarrosa, 462 redowskii, 462 occidentalis, 462 squarrosa, 462 Lapsana, 415 communis, 415 larch, 77 Lardizabalaceae, 653 Larix, 77 decidua, 77 laricina, 129 large beardtongue, 714 large bluet, 828 large-bracted tick-trefoil, 580 large cranberry, 563 large false ground-cherry, 859 large-flowered bellwort, 99 large-flowered eveningprimrose, 687 large-flowered tickseed, 391 large-fruited amaranth, 316 large-fruited sanicle, 346 large grass-leaved rush, 186 large-leaved avens, 791 large-leaved bog-orchid, 210 large-leaved Dutchman’s pipe, 82 large-leaved goldenrod, 432 large-leaved grove-sandwort, 514 large-leaved linden, 668 large-leaved wood-aster, 400 large-lobed dandelion, 444 large mountain brome, 239 large-seeded false flax, 479 large-seeded hawthorn, 786 large sweet grass, 231 large toothwort, 482 large whorled pogonia, 205

large yellow-loosestrife, 674 large yellow vetch, 599 larkspur, 754 Lasallea concolor, 439 ericoides, 439 novae-angliae, 440 patens, 441 Lasiococcus dumosus, 555 Lasthenia, 415 californica, 416 minor, 416 late lilac, 679 late purple American-aster, 441 late-summer-mint, 637 late thoroughwort, 399 Lathyrus, 583 aphaca, 584 japonicus, 584 maritimus, 584 latifolius, 584 maritimus, 584 myrtifolius, 584 ochroleucus, 584 odoratus, 584 palustris, 584 macranthus, 584 pilosus, 584 pratensis, 584 sativus, 584 sylvestris, 584 tuberosus, 584 Lauraceae, 83 laurel willow, 847 Laurentian fragile fern, 72 Laurus albidus, 83 Lavandula, 639 angustifolia angustifolia, 639 officinalis, 639 spica, 639 vera, 639 Lavatera, 665 trimestris, 665 lavender, 639 lavender-cotton, 426 lavender giant-hyssop, 635 Lawn-daisy, 379 Laxiflorae, 132–133 Layia, 416 platyglossa, 416 leafy annual American-aster, 439 leafy bulrush, 170 leafy bush-clover, 587 leaf-cup, 423 leafy-flowered blackberry, 814 leafy goosefoot, 323 leafy pondweed, 295 leafy spurge, 567, 569 least duckweed, 96 least moonwort, 65 leatherleaf, 553 leatherwood, 867 leathery grapefern, 65 leathery knotweed, 738

Lechea, 527 intermedia, 528 intermedia, 528 juniperina, 528 juniperina, 528 leggettii, 529 moniliformis, 529 ramosissima, 529 maritima, 528 minor, 528 maritima, 528 villosa, 528 moniliformis, 529 mucronata, 528 pulchella, 529 moniliformis, 529 tenuifolia, 529 Le Conte’s violet, 880 Lecticula resupinata, 656 ledge spikemoss, 49 Ledum groenlandicum, 560 leek, 92 Leersia, 264 oryzoides, 264 virginica, 264 ovata, 264 Leggett’s pinweed, 529 Lemna, 95 minima, 96 minor, 96 minuscula, 96 minuta, 96 perpusilla, 96 trinervis, 96 polyrrhiza, 97 torreyi, 96 trisulca, 96 turionifera, 96 valdiviana, 95 minima, 96 lemon-balm, 641 lemon thyme, 653 Lens, 585 culinaris, 585 Lentibularia intermedia, 656 minor, 656 Lentibulariaceae, 654–657 lentil, 585 Leontodon, 416 autumnalis, 426 hastilis vulgaris, 416 hispidus, 416 leysseri, 416 nudicaulis, 416 taraxacoides, 416 saxatilis saxatilis, 416 taraxacoides, 416 Leontodon autumnalis, 426 pratensis, 426 erythrospermum, 444 latiloba, 444 taraxacum, 444 Leonurus, 639 cardiaca, 639 villosus, 639

Lepachys columnifera, 424 Lepidanche adpressa, 534 Lepidanthus suaveolens, 418 Lepidium, 486 campestre, 487 densiflorum densiflorum, 487 typicum, 487 didymum, 487 draba, 487 heterophyllum, 487 latifolium, 487 neglectum, 487 perfoliatum, 487 ruderale, 487 sativum, 488 squamatum, 488 virginicum, 488 virginicum, 488 Lepidotis alopecuroides, 46 inundata, 46–47 Leptamnium virginianum, 692 Leptasea aizoides, 854 Leptilon canadense, 396 pusillum, 396 Leptocephalae, 133 Leptochloa, 264 fascicularis, 264 fusca, 264 fascicularis, 264 uninervia, 264 panicea mucronata, 264 Leptoloma cognatum, 251 Leptorchis liliifolia, 205 loeselii, 205 Lerchenfeldia flexuosa, 245 Lespedeza, 585 ×acuticarpa, 587 angustifolia, 586 brevifolia, 588 bicknellii, 586 bicolor, 585 ×brittonii, 587 capitata, 586 angustifolia, 586 calycina, 587 stenophylla, 586 velutina, 586 cuneata, 586 cyrtobotrya, 587 frutescens, 587 hirta, 587 hirta, 587 intermedia, 588 longifolia, 586 ×nuttallii, 587 procumbens, 587 repens, 588 ×simulata, 586 striata, 583

946  i nd ex

Lespedeza, cont. stuevei, 588 thunbergii, 588 violacea, 588 virginica, 588 lesser bladder sedge, 150 lesser bladderwort, 656 lesser broom-rape, 694 lesser Canada St. John’s-wort, 625 lesser clearweed, 870 lesser hop clover, 597 lesser horse-gentian, 504 lesser periwinkle, 351 lesser pond sedge, 136 lesser purple fringed bogorchid, 210 lesser quickweed, 402 lesser rice-field flatsedge, 154 lesser snakeroot, 372 lesser-snapdragon, 712 lesser swine-cress, 487 lesser tussock sedge, 129 lesser yellow water crowfoot, 759 lettuce, 414–415, 418 Leucacantha cyanus, 387 Leucanthemella, 416 serotina, 416 Leucanthemum, 416 lacustre, 416 nipponicum, 420 ×superbum, 416 vulgare, 416 Leucoglochin, 133 Leucojum, 92 aestivum, 92 Leucophysalis, 859 grandiflora, 859 Leucothoe, 556 axillaris editorum, 556 editorum, 556 elongata, 554 fontanesiana, 556 racemosa, 554 projecta, 554 Levisticum, 343 officinale, 343 Leymus, 264 arenarius mollis, 264 mollis, 264 Liatris, 417 borealis, 417 novae-angliae novae-angliae, 417 pycnostachya pycnostachya, 417 scariosa novae-angliae, 417 spicata, 417 licorice, 582 licorice goldenrod, 432 licorice milk-vetch, 577 Ligurian yarrow, 371 Ligusticum, 343 scoticum, 338, 343 scoticum, 343

Ligustrum, 678 amurense, 678 ibota suave, 678 obtusifolium, 678 leiocalyx, 678 obtusifolium, 678 suave, 678 ovalifolium, 678 sinense, 679 villosum, 679 vulgare, 679 Lilaeopsis, 343 chinensis, 343 Liliaceae, 192–195 lilac, 679 Lilium, 194 canadense, 194 editorum, 194 superbum, 194 lancifolium, 194 philadelphicum, 194 superbum, 194 tigrinum, 194 lily blackberry, 179 lily-leaved wide-lipped orchid, 205 lily-of-the-valley, 299 limestone-meadow sedge, 128 limestone swamp bedstraw, 825 Limnanthaceae, 657 Limnanthemum lacunosum, 669 peltatum, 669 Limnobium, 176 spongiosa, 176 Limnobotrya lacustris, 618 Limodorum tuberosum, 201 Limonium, 721 angustatum, 721 carolinianum, 721 angustatum, 721 nashii, 721 trichogonum, 721 nashii albiflorum, 721 angustatum, 721 trichogonum, 721 Limosae, 133–134 Limosella, 855 aquatica tenuifolia, 855 australis, 855 subulata, 855 Linaceae, 657 Linaria, 711 canadensis, 712 cymbalaria, 709 dalmatica, 712 dalmatica, 712 genistifolia, 712 dalmatica, 712 linaria, 712 maroccana, 712 minor, 708 pinifolia, 712

repens, 712 spartia, 712 striata, 712 linden, 668 linden arrowwood, 310 linden-leaved sage, 648 Lindera, 83 benzoin, 83 pubescens, 83 Lindernia, 659 anagallidea, 659 dubia, 659 anagallidea, 659 Linderniaceae, 659 Lindley’s American-aster, 438 linear-leaved orache, 319 linear-leaved rosettepanicgrass, 249 lined sedge, 133 Linnaea, 500 americana, 500 borealis, 500 americana, 500 Linum, 657 catharticum, 657 intercursum, 658 medium, 658 texanum, 658 perenne, 658 radiola, 658 striatum, 658 sulcatum, 657 sulcatum, 658 usitatissimum, 658 virginianum, 658 texanum, 658 lion’s-foot rattlesnake-root, 420 Liparis, 205 correana, 205 liliifolia, 205 loeselii, 205 Lipocarpha, 163 micrantha, 163 Liquidambar, 311 styraciflua, 311 Liriodendron, 83 tulipifera, 83 Listera auriculata, 206 australis, 206 convallarioides, 206 cordata, 207 Lithospermum, 463 arvense, 459 latifolium, 463 officinale, 463 little barley, 263 little bluestem, 280 little bluet, 828 little club-spur bog-orchid, 209 little evening-primrose, 687 little false cotton-rose, 417 little floating-heart, 669 little green sedge, 117 little hawkbit, 416 little-headed spikesedge, 160 little ladies’-tresses, 212 little-leaved-angelica, 346 little lovegrass, 258 little mallow, 666 little quaking grass, 235

little shinleaf, 558 little skullcap, 650 little yellow-rattle, 695 Littorella, 712 americana, 712 uniflora americana, 712 livid amaranth, 316 livid sedge, 137 lizard’s-tail, 86 Loasaceae, 659 lobelia, 496 Lobelia, 496 bracteata, 496 cardinalis, 496 dortmanna, 496 hirtella, 496 inflata, 496 kalmii, 496 strictiflora, 496 siphilitica siphilitica, 496 spicata, 496 campanulata, 497 hirtella, 497 originalis, 496 spicata, 497 Lobularia, 488 maritima, 488 locust, 593 Loesel’s wide-lipped orchid, 205 Logfia, 417 minima, 417 Loiseleuria procumbens, 556 Lolium, 265 arvense, 265 giganteum, 279 multiflorum, 265 italicum, 265 ramosum, 265 perenne, 265 aristatum, 265 multiflorum, 265 pratense, 279 temulentum arvense, 265 temulentum, 265 Lomatogonium, 611 rotatum, 611 London hedge-mustard, 492 London sycamore, 720 long-beaked beaksedge, 164 long-beaked sedge, 131 long-beaked stork’s-bill, 613 long-beaked willow, 844 long beech fern, 70 long-bracted green orchid, 201 long-bracted spiderwort, 100 long-bracted wild indigo, 577 long-flowered tobacco, 860 long-headed windflower, 752 Long Island Sound hawthorn, 784 long-leaved bluet, 828 long-leaved ground-cherry, 862 long-leaved pondweed, 296 long-leaved redtop-panicgrass, 281

i n d e x  9 47

long-leaved speedwell, 719 long-leaved stitchwort, 523 long-podded poppy, 702 Long’s bitter-cress, 482 long-sepaled beardtongue, 713 long-spined sandbur, 241 long-spurred violet, 885 Long’s sedge, 124 long-stalked crane’s-bill, 616 long-stalked sedge, 118 long-styled sweet-cicely, 344 Long’s woolsedge, 170 long-thorned hawthorn, 785 long-tubercled spikesedge, 161 Lonicera, 500 bella, 502 caerulea villosa, 502 canadensis, 501 dioica, 501 hirsuta, 502 interior, 502 japonica, 502 maackii, 502 morrowi, 501 morrowii, 502 oblongifolia, 502 periclymenum, 502 prolifera, 502 glabra, 502 reticulata, 502 sempervirens, 501 tatarica, 502 villosa, 502 calvescens, 502 fulleri, 502 solonis, 502 tonsa, 502 xylosteum, 502 loose-flowered alpine sedge, 134 loose silky bentgrass, 231 loosestrife, 674–675 Lophotocarpus spathulatus, 90 spongiosus, 90 lop-seed, 705 lopsided oat, 233 lopsided rush, 189 Lorinseria areolata, 53 lotus, 675 Lotus, 588 corniculatus, 588 unifoliatus unifoliatus, 588 lousewort, 694–695 lovage, 343 lovegrass, 258 low agrimony, 768 low baby’s-breath, 512 low frostweed, 527 lowland yellow-loosestrife, 674 low mallow, 666 low nutsedge, 171 Lowrie’s American-aster, 440 low shadbush, 771 low water-milfoil, 621 Lucile’s glory-of-the-snow, 174 Lucy, 461

Ludwigia, 684 alternifolia, 684 linearifolia, 684 pubescens, 684 typica, 684 ×lacustris, 684 palustris, 684 americana, 684 nana, 684 polycarpa, 684 sphaerocarpa, 684 deamii, 684 jungens, 684 macrocarpa, 684 Lunaria, 488 annua, 488 lungwort, 466 lupine, 588–589, 595 Lupinus, 588 pallidipes, 589 perennis, 588 occidentalis, 588 polyphyllus albiflorus, 589 pallidipes, 589 polyphyllus, 589 Lupulinae, 134–135 Luzula, 189 acuminata, 190 acuminata, 190 carolinae, 190 bulbosa, 191 campestris, 191 bulbosa, 191 echinata, 191 frigida, 191 pallescens, 191 carolinae, 190 confusa, 191 echinata, 191 frigida, 191 hyperborea, 191 intermedia, 191 luzuloides, 191 melanocarpa, 191 multiflora, 191 echinata, 191 frigida, 191 fusconigra, 191 multiflora, 191 pallidula, 191 parviflora, 189 melanocarpa, 191 spicata, 191 lychnis, 513 Lychnis, 512 alba, 519 chalcedonica, 513 coeli-rosa, 511 coronaria, 513 dioica, 519 flos-cuculi, 513 githago, 508 saponaria, 517 Lycium, 859 barbarum, 859 chinense, 859 halimifolium, 859 Lycophytes, 39–49 Lycopodiaceae, 42–48

Lycopodiella, 45 alopecuroides, 46 appressa, 46 caroliniana, 47 ×copelandii, 46 ×gilmanii, 46 inundata, 46 ×robusta, 46 Lycopodioides apoda, 49 Lycopodium, 47 alopecuroides, 46 annotinum, 48 acrifolium, 48 montanum, 48 pungens, 48 appressum, 46 canadense, 48 carolinianum, 47 chamaecyparissus, 45 clavatum, 47 subremotum, 47 complanatum, 45 canadense, 45 flabelliforme, 45 dendroideum, 44 digitatum, 45 flabelliforme, 45 hickeyi, 44 inundatum, 46 bigelovii, 46 appressa, 46 lagopus, 47 lucidulum, 40 obscurum, 44 rupestre, 49 sabinifolium sitchense, 45 selaginella spinosa, 49 selaginoides, 49 selago, 40 patens, 40 sitchense, 45 tristachyum, 45 Lycopsis arvensis, 459 Lycopus, 639 americanus, 640 longii, 640 scabrifolius, 640 amplectens, 640 pubens, 640 asper, 640 europaeus, 640 mollis, 640 sinuatus, 640 lucidus americanus, 640 maritimus, 640 membranaceus, 641 parviflorus, 640 pennsylvanicus, 640 pinnatifidus, 640 pubens sessilifolius, 640 rubellus, 640 lanceolatus, 640 sinuatus, 640

uniflorus, 640 velutinus, 640 virginicus, 641 pauciflorus, 640 Lycurus alopecuroides, 267 setosus, 267 Lygodiaceae, 61 Lygodium, 61 palmatum, 61 Lyonia, 557 ligustrina, 557 ligustrina, 557 mariana, 557 lyre-leaved sage, 648 lyre-leaved thale-cress, 475 Lysiella obtusata, 210 orbiculata, 210 Lysimachia, 672 arvensis, 673 borealis, 674 ciliata, 674 clethroides, 674 ×commixta, 674 foemina, 673 hybrida, 674 lanceolata, 674 hybrida, 674 longifolia, 674 maritima, 674 minima, 674 nummularia, 674 ×producta, 674 punctata, 674 verticillata, 674 quadriflora, 674 quadrifolia, 673 revoluta, 674 terrestris, 674 ovata, 674 thyrsiflora, 675 vulgaris, 675 Lythraceae, 659–661 Lythrum, 660 adsurgens, 661 alatum alatum, 661 dacotanum, 661 flexuosum, 661 graefferi, 661 hyssopifolia, 661 junceum, 661 lineare, 661 salicaria, 661 gracilior, 661 tomentosum, 661 verticillatum, 660 virgatum, 661

M Mackay’s fragile fern, 73 Mackenzie’s sedge, 127 Macleaya, 701 cordata, 701 Maclura, 670 pomifera, 670 Macoun’s rabbit-tobacco, 424 Macrocephalae, 135

948   i n d e x

Macrotrys racemosa, 750 madder, 828 mad dog skullcap, 649 Madia, 417 capitata, 417 glomerata, 417 sativa, 417 capitata, 417 congesta, 417 madwort, 459 Magnolia, 84 acuminata, 84 cordata, 84 tripetala, 84 virginiana, 84 tripetala, 84 Magnoliaceae, 83 Magnoliids, 81–86 Mahaleb cherry, 801 Mahonia aquifolia, 449 Maianthemum, 299 canadense, 300 pubescens, 300 racemosum, 300 stellatum, 300 trifolium, 300 maidencane, 230 maidenhair spleenwort, 52 maiden pink, 511 Mairania alpina, 552 Majorana hortensis, 645 majorana, 645 Malaxis, 205 bayardii, 205 brachypoda, 206 liliifolia, 205 monophyllos, 205 brachypoda, 206 unifolia, 206 maleberry, 557 male wood fern, 57 mallow, 666 Maltese knapweed, 387 Malus, 792 ×arnoldiana, 793 baccata, 793 domestica, 793 floribunda, 793 prunifolia, 793 pulcherrima, 793 pumila, 793 sieboldii, 793 ×soulardii, 793 sylvestris, 793 Malva, 665 alcea, 666 crispa, 666 erecta, 666 mauritiana, 666 mendocina, 667 moschata, 666 neglecta, 666 parviflora, 666 pusilla, 666 rotundifolia, 666

sylvestris, 666 mauritiana, 666 verticillata, 666 crispa, 666 Malvaceae, 662–668 Malvastrum, 666 coromandelianum, 666 Manchu tuber gourd, 545 manna grass, 261–262 many-flowered marshpennywort, 342 many-fruited water-primrose, 684 many-headed sedge, 125 manyseed, 516 many-seeded goosefoot, 324 many-seeded plantain, 716 many-spiked flatsedge, 155 maple, 849–851 maple-leaved viburnum, 309 mare’s tail, 710 marginal wood fern, 58 marigold, 384, 394, 442–443 Mariscus cylindricus, 155 mariscoides, 151 retrorsus, 155 schweinitzii, 155 maritime marsh-elder, 414 maritime orache, 319 marjoram, 645 Marrubium, 641 vulgare, 641 marsh arrow-grass, 192 marsh bedstraw, 826 marsh bellflower, 495 marsh-cinquefoil, 774 marsh dandelion, 444 marsh-elder, 414 marsh-felwort, 611 marsh fern, 70 marsh flatsedge, 155 marsh hedge-nettle, 651 marsh horsetail, 60 marsh-mallow, 664 marsh-marigold, 752 marsh meadow-foxtail, 229 marsh mermaid-weed, 621 marsh muhly, 268 marsh-pennywort, 342–343 marsh-St. John’s-wort, 627 marsh straw sedge, 124 marsh thistle, 390 marsh valerian, 504 marsh vetchling, 584 marsh willow-herb, 683 Marsilea, 61 quadrifolia, 61 Marsileaceae, 61 Martynia louisianica, 703 Maruta cotula, 375 Maryland golden-aster, 388 Maryland meadow-beauty, 668 Maryland sanicle, 345 Massachusetts fern, 69 masterwort, 339 mat grama, 234

Matricaria, 418 chamomilla, 418 chamomilla, 418 recutita, 418 discoidea, 418 inodora, 445 maritima, 445 inodora, 445 matricarioides, 418 parthenium, 443 recutita, 418 suaveolens, 418 matrimony-vine, 859 matted muhly, 268 matted sandmat, 568 Matteuccia, 61 pensylvanica, 61 struthiopteris, 61 pensylvanica, 61 Maximilian’s sunflower, 407 May-apple, 449 mayflower, 300 mazus, 704 Mazus, 704 japonicus, 704 miquelii, 704 pumilus, 704 reptans, 704 meadow barley, 263 meadow-beauty, 668 meadow bistort, 728 meadow bottle gentian, 610 meadow brome, 238 meadow crane’s-bill, 616 meadow dropwort, 788 meadow evening-primrose, 687 meadow-foxtail, 229 meadow garlic, 91 meadow goat’s beard, 445 meadow horsetail, 60 meadow-parsnip, 346 meadow-rue, 759–761 meadow rye grass, 279 meadow sage, 648 meadow spikemoss, 49 meadowsweet, 819–820 meadow vetchling, 584 meadow willow, 847 Mead’s sedge, 137 meager sedge, 147 Medeola, 194 virginiana, 194 Medicago, 589 agrestis, 591 arabica, 590 disciformis, 590 falcata, 590 hispida, 591 apiculata, 591 confinis, 591 laciniata, 590 lupulina, 589 cupaniana, 590 glandulosa, 590 minima, 589 compacta, 590 longiseta, 590 monantha, 590 polymorpha, 591 arabica, 591 minima, 591

praecox, 591 rigidula, 589, 591 sativa, 589, 591 sativa falcata, 590 virginica, 588 medicinal agrimony, 768 medick, 589–591 Mediterranean medick, 591 Mediterranean stork’s-bill, 614 Medusa-head, 285 Megalodonta beckii, 382 Meibomia acuminata, 582 canadense, 580 canescens, 580 ciliaris, 580 dillenii, 581 glabella, 580 grandiflora, 580 longifolia, 580 marilandica, 581 michauxii, 581 nudiflora, 583 paniculata, 581 rotundifolia, 581 sessilifolia, 581 Melampodium australe, 371 Melampyrum, 694 latifolium, 694 lineare, 694 latifolium, 694 pectinatum, 694 Melandrium album, 519 dioicum, 519 rubrum, 519 noctiflorum, 520 Melanthiaceae, 195–198 Melanthium hybridum, 198 latifolium, 198 Melastomataceae, 668 Melica purpurascens, 279 Melilotus, 591 albus, 591 indicus, 591 officinalis, 591 Melinum palustre, 288 Melissa, 641 officinalis, 641 melon, 545 memorial rose, 806 Mendoza globe-mallow, 667 Menispermaceae, 669 Menispermum, 669 canadense, 669 Mentha, 641 adspersa, 642 aquatica, 642 aquatica, 642 citrata, 642 arvensis, 641 canadensis, 643 glabrata, 643 haplocalyx, 643

in d e x  9 49

Mentha, cont. parietariaefolia, 642 villosa, 643 canadensis, 643 citrata, 642 gentilis, 642 ×gracilis, 642 hirsuta, 642 longifolia, 643 mollissima, 643 palustris, 642 ×piperita, 642 ×rotundifolia, 643 spicata, 642 longifolia, 643 spicata, 643 suaveolens, 643 sylvestris, 643 variegata, 642 verticillata, 642 ×villosa, 643 alopecuroides, 643 viridis, 643 mentzelia, 659 Mentzelia, 659 oligosperma, 659 Menyanthaceae, 669 Menyanthes, 669 trifoliata, 669 minor, 669 Mercurialis, 569 annua, 569 Mercury, 564–565, 569 Meriolix serrulata, 687 mermaidweed, 621–622, 657 Merritt Fernald’s sedge, 124 Mertensia, 463 maritima maritima, 463 virginica, 463 Mespilus arborea, 770 arbutifolia, 773 melanocarpa, 773 canadensis, 771 rotundifolia, 772 melanocarpa, 773 Mesynium sulcatum, 658 Mexican ground-cherry, 862 Mexican-hat, 424 Mexican lovegrass, 258 Mexican muhly, 268 Mexican prickly-poppy, 699 Mexican-tea, 326 Mibora, 265 minima, 265 Michaux’s sandplant, 514 Michaux’s sedge, 145 Micrampelis lobata, 545 Micranthes, 853 foliolosa, 853 pensylvanica, 853 virginiensis, 853 Micromeles alnifolia, 818 Microseris, 418 douglasii, 418 douglasii, 418

Microstegium, 265 vimineum, 265 Microstylis unifolia, 206 midland sedge, 142 midwestern beggar-ticks, 382 mignonette, 761–762 Mikania, 418 scandens, 418 pubescens, 418 Milium, 265 effusum cisatlanticum, 265 pungens, 273 racemosum, 273 milk-parsley, 344 milk-thistle, 428 milk-vetch, 576–577 milkweed, 349–350 milkwort, 726 milky-bellflower, 495 Millegrana radiola, 658 millet, 265, 270, 280 millet foxtail, 280 millet grass, 265 Mimulus, 704 alatus, 705 brevipes, 705 guttatus, 705 litoralis, 705 micranthus, 705 langsdorfii, 705 guttatus, 705 micranthus, 705 minthodes, 705 moschatus, 705 ringens, 705 colpophilus, 705 minthodes, 705 Mingan moonwort, 65 miniature marigold, 443 mint, 637, 641–643, 645 Minuartia, 513 caroliniana, 513 glabra, 513 groenlandica, 514 glabra, 513 marcescens, 514 michauxii, 514 rubella, 514 Minuopsis caroliniana, 513 michauxii, 514 minute duckweed, 96 Mirabilis, 675 albida, 675 lata, 675 hirsuta, 675 jalapa, 676 lindheimeri, 676 lanceolata, 676 linearis, 675 linearis, 676 subhispida, 676 nyctaginea, 676 Miscanthus, 265 sacchariflorus, 266 sinensis, 266 gracillimus, 266

Misopates, 712 orontium, 712 Miss Jones’ hawthorn, 785 Missouri gooseberry, 618 Missouri ironweed, 446 mistletoe, 887 Mitchella, 828 repens, 828 Mitchell’s sedge, 140 Mitella, 854 diphylla, 854 nuda, 854 prostrata, 854 Mitratae, 135 mixed bladderwort, 656 mockernut hickory, 629 mock-orange, 623 mock-vervain, 871 Modiola, 666 caroliniana, 666 multifida, 666 Moehringia, 514 lateriflora, 514 macrophylla, 514 Mojave suncup, 680 Moldavian dragonhead, 637 Moldavica moldavica, 637 parviflora, 637 thymiflora, 637 Molinia, 266 caerulea, 266 Molluginaceae, 669 Mollugo, 669 verticillata, 669 Momordica, 545 charantia, 545 lanata, 544 Monarda, 643 ciliata, 636 clinopodia, 644 coccinea, 644 didyma, 644 fistulosa, 644 albescens, 644 fistulosa, 644 media, 644 mollis, 644 rubra, 644 hirsuta, 636 media, 644 mollis, 644 punctata, 644 villicaulis, 644 scabra, 644 Moneses, 557 uniflora, 557 Monilophytes, 51–74 monkey-flower, 704–705 monkshood, 749–750 monk’s-wort, 464 Monocots, 87–305 Monolepis, 327 nuttalliana, 327 Monotropa, 557 hypopithys, 556 lanuginosa, 556 uniflora, 557 Monterey spineflower, 728 montia, 745–746

Montia, 745–746 fontana, 745 lamprosperma, 745 lamprosperma, 745 linearis, 746 sibirica, 745 bulbifera, 745 Montiastrum lineare, 746 moorgrass, 266 moor rush, 189 moonseed, 669 moonshine-daisy, 394 moonwort, 62–65 Moraceae, 670 Morella, 671 caroliniensis, 671 heterophylla, 671 pensylvanica, 671 Morison’s spurry, 520 morning-glory, 535–536 Moroccan toadflax, 712 Morrow’s honeysuckle, 502 Morus, 671 alba, 671 tatarica, 671 papyrifera, 670 rubra, 671 tomentosa, 671 tatarica, 671 mosquito bulrush, 170 moss campion, 519 moss phlox, 724 moss-plant, 555 moss stonecrop, 543 motherwort, 639 moth mullein, 857 mountain American-laurel, 556 mountain-ash, 818–819 mountain chestnut oak, 605 mountain cranberry, 563 mountain crane’s-bill, 616 mountain death camas, 196 mountain fetterbush, 557 mountain firmoss, 39 mountain-heath, 557 mountain holly, 352 mountain honeysuckle, 502 mountain ironwort, 650 mountain knapweed, 387 mountain maple, 851 mountain-mint, 646–647 mountain phlox, 724 mountain sandplant, 514 mountain shadbush, 770 mountain snow spurge, 568 mountain-sorrel, 731 mountain spleenwort, 51 mountain-spurge, 493 mountain sweet-cicely, 344 mountain tansy-mustard, 483 mountain tarplant, 417 mountain Timothy, 272 mountain-trumpet, 722 mountain wood fern, 55 mouse barley, 263 mouse-ear chickweed, 509 mouse-ear hawkweed, 412 mouse-ear thale-cress, 475 mousetail, 755 mudflat spikesedge, 160

95 0   i n de x

mud-mat, 704 mud-plantain, 289 mud sedge, 134 mudwort, 855 Mugo pine, 79 Muhlenbergia, 266 alopecuroides, 267 capillaris, 267 ×curtisetosa, 267 erecta, 235 frondosa, 267 glomerata, 268 cinnoides, 268 mexicana, 268 filiformis, 268 phleoides, 267 racemosa, 268 richardsonis, 268 schreberi, 268 palustris, 268 sobolifera, 268 squarrosa, 268 sylvatica, 268 robusta, 268 tenuiflora, 268 uniflora, 268 Muhlenberg’s sedge, 142 muhly, 267–268 mulberry, 670 Mulgedium, 418 pulchellum, 418 spicatum, 415 integrifolium, 415 mullein, 856–857 mullein lychnis, 513 Multiflorae, 135 multi-stemmed St. Andrew’scross, 626 Muscari, 174 atlanticum, 174 botryoides, 174 neglectum, 174 musk mallow, 666 musky monkey-flower, 705 musky stork’s-bill, 614 mustard, 478, 485, 488, 491, 492 Myagrum paniculatum, 488 rugosum, 490 sativum, 479 Mycelis, 418 muralis, 418 Myosotis, 463 arvensis, 464 californica, 466 discolor, 464 laxa, 464 micrantha, 464 palustris, 464 scorpioides, 464 arvensis, 464 stricta, 464 sylvatica, 464 verna, 464 versicolor, 464 virginiana, 461 virginica, 464 Myosoton, 514 aquaticum, 514

Napaea, 666 dioica, 666 hermaphrodita, 667 nap-at-noon, 175 Narcissus, 93 poeticus, 93 pseudonarcissus, 93 Nardosmia arctica, 423 japonica, 423 Nardus, 268 stricta, 268 narrow false oat, 286 narrow-fruited beaksedge, 164 narrow lady fern, 71 narrow-leaved arrowhead, 90 narrow-leaved bitter-cress, 482 narrow-leaved bluecurls, 653 narrow-leaved blue-eyed-grass, 181 narrow-leaved bur-reed, 303 narrow-leaved bush-clover, 586 narrow-leaved cat-tail, 304 narrow-leaved eveningprimrose, 686 narrow-leaved fireweed, 681 narrow-leaved gentian, 610 narrow-leaved glade fern, 73 narrow-leaved goosefoot, 323 narrow-leaved hawk’s-beard, 393 narrow-leaved hawkweed, 413 narrow-leaved montia, 746 narrow-leaved mountain-mint, 647 narrow-leaved mountaintrumpet, 722 narrow-leaved pinweed, 529 narrow-leaved plantain-lily, 173 narrow-leaved sneezeweed, Nabalus, 418 404 albus, 419 narrow-leaved speedwell, 719 altissimus, 419 narrow-leaved thorough-wax, boottii, 419 340 integrifolius, 419 narrow-leaved umbrella-wort, ×mainensis, 419 676 nanus, 420 narrow-leaved vervain, 872 racemosus, 419 narrow-leaved vetchling, 584 serpentarius, 420 narrow-leaved water-plantain, integrifolius, 419 88 trifoliolatus, 420 narrow-leaved white-toppedNahanni oak fern, 73 aster, 427 Najas, 177 narrow-spiked dropseed, 285 caespitosa, 177 narrow triangle moonwort, 64 flexilis, 177 Nartheciaceae, 198 gracillima, 177 Narthecium guadalupensis, 177 glutinosum, 302 guadalupensis, 177 Narukila olivacea, 177 cordata, 289 indica Nasturtium, 488 gracillima, 177 amphibium, 491 minor, 177 austriacum, 491 olivacea, 177 globosum, 491 naked bishop’s-cap, 854 lacustre, 491 naked-bulbil small-floweredmicrophyllum, 488 saxifrage, 853 officinale, 488 naked-stemmed sunflower, 407 microphyllum, 488 naked tick-trefoil, 583 palustre nannyberry, 310 hispidum, 491 Nantucket shadbush, 771

Myosurus , 755 minimus, 755 Myrica, 672 caroliniensis, 671 gale, 672 pennsylvanica, 671 Myricaceae, 671 Myriophyllum, 619 alterniflorum, 620 americanum, 620 ambiguum natans, 621 aquaticum, 620 brasiliense, 620 capillaceum, 621 exalbescens, 621 farwellii, 620 heterophyllum, 620 humile, 621 capillaceum, 621 natans, 621 pinnatum, 621 proserpinacoides, 620 scabratum, 621 sibiricum, 621 spicatum, 621 exalbescens, 621 tenellum, 621 verticillatum, 621 intermedium, 621 pectinatum, 621 Myrrhis, 343 odorata, 343 Myrsinaceae, 672–675 Myrtus chinensis, 866

N

sessiliflorum, 491 ×sterile, 488 sylvestre, 491 Navarretia, 722 leucocephala minima, 722 navel corn-salad, 505 Nazia aliena, 286 racemosa, 286 necklace poplar, 832 necklace sedge, 124 necklace spike sedge, 133 needle beaksedge, 164 needle-leaf, 327 needle spikesedge, 159 needle sprangletop, 264 needle-tipped blue-eyed-grass, 181 neglected reed grass, 240 Negundo aceroides, 850 violaceum, 850 negundo, 850 Nelumbo, 675 lutea, 675 Nelumbonaceae, 675 Nemexia herbacea, 301 Neobeckia aquatica, 491 Neolepia campestris, 487 Neopieris mariana, 557 Neotorularia humilis, 479 Neottia, 206 auriculata, 206 bifolia, 206 convallarioides, 206 cordata, 207 gracilis, 211 lacera, 211 lucida, 212 pubescens, 204 ×veltmanii, 206 Nepalese smartweed, 735 Nepeta, 645 cataria, 645 hederacea, 637 Nephrodium acrostichoides, 58 asplenioides, 71 punctilobulum, 53 nerveless woodland sedge, 133 Nesaea verticillata, 660 Neslia, 488 paniculata, 488 nettle, 869–871 netted chain fern, 53 netted nutsedge, 171 netted popcorn-flower, 466 nettle-leaved bellflower, 495 nettle-leaved goosefoot, 324 Neubeckia cristata, 179 New England American-aster, 440

in d e x  9 5 1

New England groundsel, 422 New England sedge, 114 New England states, 726 New England thoroughwort, 399 New England tuber-bulrush, 103 New England violet, 883 New Jersey highbush blueberry, 562 New Jersey redroot, 762 New York American-aster, 440 New York fern, 69 New York ironweed, 446 New Zealand-spinach, 311 Nezera striata, 658 virginiana, 658 Nicandra, 859 physalodes, 859 Nicotiana, 860 affinis grandiflora, 860 alata, 860 langsdorffii, 860 longiflora, 860 quadrivalvis bigelovii, 861 rustica, 861 ×sanderae, 860 tabacum, 861 Nigella, 755 damascena, 755 sativa, 755 niger-seed, 404 night-flowering campion, 520 nightshade, 681, 863–865 Nikko fir, 77 nimblewill muhly, 268 ninebark, 793 nipplewort, 415 Nipponanthemum, 420 nipponicum, 420 Nippon-daisy, 420 nit grass, 260 noble prince’s-pine, 553 nodding-aster bog, 420 sharp-toothed, 420 nodding beggar-ticks, 382 nodding campion, 520 nodding chickweed, 510 nodding evening-primrose, 687 nodding fescue, 260 nodding ladies’-tresses, 211 nodding saxifrage, 854 nodding sedge, 140 nodding star-of-Bethlehem, 175 nodding stickseed, 461 nodding wakerobin, 197 Nonea, 464 rosea, 464 Norta altissima, 492 North African grass, 287 northeastern beardtongue, 714 northeastern blackberry, 817 northeastern bulrush, 169 northeastern manna grass, 262 northeastern sedge, 117

northern adder’s-tongue fern, 65 northern arrowhead, 90 northern bedstraw, 825 northern beggar-ticks, 383 northern bentgrass, 228 northern blackberry, 814 northern blazing star, 417 northern blueberry, 562 northern bog bedstraw, 825 northern bog-clubmoss, 46 northern bog sedge, 143 northern bog violet, 883 northern catalpa, 455 northern cliff fern, 74 northern cluster sedge, 126 northern crane’s-bill, 616 northern dwarf-gentian, 610 northern eyebright, 693 northern firmoss, 40 northern green rush, 186 northern ground-cedar, 45 northern hawthorn, 783 northern horsebalm, 636 northern interrupted-clubmoss, 48 northern long-awned wood grass, 235 northern long sedge, 145 northern maidenhair fern, 68 northern manna grass, 261 northern marsh violet, 884 northern meadow sedge, 124 northern oak fern, 73 northern painted-cup, 692 northern red oak, 606 northern-rockcress, 479 northern rosette-panicgrass, 248 northern sedge, 114 northern snail-seed pondweed, 297 northern spicebush, 83 northern spikemoss, 49 northern St. John’s-wort, 625 northern sweet-coltsfoot, 423 northern sweet grass, 231 northern tubercled bog-orchid, 209 northern water-horehound, 640 northern water-meal, 97 northern water-milfoil, 621 northern water-plantain, 88 northern white-cedar, 76 northern wild rice, 288 northern wild senna, 594 northern willow, 844 northern wood rush, 191 northern wood sorrel, 697 northern yellow-eyed-grass, 305 north wind bog-orchid, 208 Norway maple, 851 Norway spruce, 78 Norwegian cinquefoil, 797 Nothoholcus lanatus, 262 Nova Scotia agalinis, 690

Nuphar, 85 advena, 85 lutea, 85 variegata, 86 microphylla, 85 minima, 85 rubrodisca, 85 variegata, 86 nut flatsedge, 154 nutsedge, 170–171 Nuttallanthus, 712 canadensis, 712 Nuttall’s alkali grass, 278 Nuttall’s milkwort, 726 Nuttall’s poverty-weed, 327 Nuttall’s reed grass, 240 Nyctaginaceae, 675–676 Nyctelea nyctelea, 461 Nymphaea, 86 advena, 85 fraterna, 86 leibergii, 86 odorata, 86 gigantea, 86 tuberosa, 86 pentaphylla, 675 tetragona, 86 leibergii, 86 tuberosa, 86 Nymphaeaceae, 84–86 Nymphoides, 669 cordata, 669 lacunosa, 669 peltata, 669 Nyssa, 538 sylvatica, 538 caroliniana, 538 dilatata, 538 typica, 538

O oak, 604–606 Oakes’ evening-primrose, 687 Oakes’ eyebright, 693 Oakes’ hawthorn, 786 Oakesiella sessilifolia, 99 Oakes’ pondweed, 296 oak-forest wood rush, 191 oak-leaved goosefoot, 323 oak-leaved hydrangea, 623 oak-leaved mountain-ash, 819 oat, 233, 286 oatgrass, 233, 243–244 obedient false dragonhead, 645 Obione muricata, 320 sibirica, 320 Ocimum, 645 basilicum, 645 crispum, 645 Oclemena, 420 acuminata, 421 ×blakei, 420 nemoralis, 420 Oconee-bells, 546

Odontites, 694 rubra, 694 serotinus, 694 vernus, 694 serotinus, 694 vernus, 694 Oedera trinervia, 402 Oenothera, 684 angustissima, 687 arenicola, 686 austromontana, 687 biennis, 686 austromontana, 687 canescens, 688 centralis, 686 grandiflora, 687 nutans, 687 oakesiana, 687 pycnocarpa, 686 campestris, 680 canovirens, 688 cruciata, 687 sabulonensis, 687 stenopetala, 687 curtiflora, 686 dentata campestris, 680 erythrosepala, 687 filiformis, 686 fruticosa, 686 eamesii, 686 fruticosa, 686 glauca, 686 linearis, 686 microcarpa, 686 gaura, 687 glazioviana, 687 grandiflora, 687 glabra, 687 laciniata, 687 longiflora, 686 muricata, 686 nutans, 687 oakesiana, 687 parviflora, 687 angustissima, 687 lachnocarpa, 686 oakesiana, 687 perennis, 685, 687 pilosella, 685 pilosella, 687 pumila, 687 pycnocarpa, 686 rubricapitata, 687 serrulata, 685 typica, 687 strigosa canovirens, 688 tetragona brevistipata, 686 longistipata, 686 riparia, 686 velutina, 686 villosa, 688 canovirens, 688 Ogden’s pondweed, 297 Ohio buckeye, 851 Oklahoma sedge, 151 okra, 663

95 2   in de x

Oldenlandia purpurea tenuifolia, 828 old-field cinquefoil, 797 oldfield threeawn, 232 oldfield-toadflax, 712 oldman wormwood, 378 Oleaceae, 676–679 Oligoneuron, 420 album, 421 rigidum, 421 Oligosporus campestris, 378 caudatus, 378 dracunculus, 378 glaucus, 378 olive, 548 Omalotheca, 421 supina, 421 sylvatica, 421 Onagraceae, 670–688 one-cone clubmoss, 47 one-flowered broom-rape, 694 one-flowered Indian-pipe, 557 one-flowered-shinleaf, 557 one-glumed spikesedge, 161 one-rowed water-cress, 488 one-seeded burr-cucumber, 545 one-seeded hawthorn, 786 one-sided-shinleaf, 557 onion, 91 Onobrychis, 591 viciifolia, 591 Onoclea, 61 sensibilis, 61 obtusilobata, 61 Onocleaceae, 61 Onopordum, 421 acanthium acanthium, 421 Onosmodium, 464 bejariense occidentale, 464 virginianum, 465 Ontario American-aster, 441 open-field sedge, 129 Ophioglossaceae, 62–65 Ophioglossum, 65 pusillum, 65 vulgatum pseudopodum, 65 Ophrys auriculata, 206 australis, 206 cernua, 211 convallarioides, 206 liliifolia, 205 opium poppy, 702 Opulaster alabamensis, 793 opulifolius, 793 Opuntia, 493 ammophila, 493 calcicola, 493 compressa, 493 humifusa, 493 orache, 319–320 orange day-lily, 173 orange-eye butterfly-bush, 855 orange fringed bog-orchid, 208

orange-fruited horse-gentian, 504 orange-grass St. John’s-wort, 626 orange hawkweed, 411 orange mullein, 857 orange stonecrop, 540 orchard grass, 243 orchid, 200–201, 203–204, 212 bog, 208–210 Orchidaceae, 198–212 Orchis loeselii, 205 rotundifolia, 200 spectabilis, 203 Oregon-grape, 449 Oreosedum album, 543 Oriental knight’s-spur, 754 Oriental lady’s-thumb smartweed, 734 Origanum, 645 majorana, 645 vulgare, 645 Ornithogalum, 174 fistulosum, 193 hirsutum, 178 nutans, 175 umbellatum, 175 Ornithopus, 591 sativus sativus, 591 scorpioides, 578 Orobanchaceae, 688–695 Orobanche, 694 americana, 692 minor, 694 uniflora, 694 purpurea, 694 Orobus myrtifolius, 584 Orono sedge, 124 Orontium, 96 aquaticum, 96 orpine, 541 Orthilia, 557 secunda, 557 obtusata, 557 Oryzopsis, 268 asperifolia, 268 canadensis, 273 pungens, 273 racemosum, 273 Osage-orange, 670 osier willow, 848 Osmorhiza, 343 berteroi, 344 chilensis, 344 claytonii, 344 depauperata, 344 depaupertata, 343 divaricata, 344 longistylis, 344 villicaulis, 344 obtusa, 344 Osmunda, 66 cinnamomea, 66 claytoniana, 66 lunaria, 64 multifida, 65

regalis, 66 spectabilis, 66 ×ruggii, 66 Osmundaceae, 66 Osmundastrum, 66 cinnamomeum, 66 Ostrya, 455 virginiana, 455 Othocallis, 175 siberica, 175 oval-headed sedge, 142 oval sedge, 124 oval-seeded goosefoot, 325 ovoid spikesedge, 160 Oxalidaceae, 696–698 Oxalis, 696 acetosella, 697 montana, 697 brittoniae, 697 corniculata, 697 dillenii, 697 langloisii, 697 cymosa, 697 dillenii, 697 europaea, 697 bushii, 697 filipes, 697 florida, 697 filipes, 697 interior, 697 intermedia, 697 langloisii, 697 montana, 697 repens, 697 stricta, 697 villicaulis, 697 violacea, 697 ox-eye daisy, 416 oxtongue, 423 Oxybaphus hirsutus, 675 nyctagineus, 676 Oxycoccos macrocarpus, 563 oxycoccos, 563 palustris, 563 quadripetalus, 563 Oxyria, 731 digyna, 731 oxytrope, 592 Oxytropis, 592 campestris johannensis, 592 johannensis, 592 Ozark beggar-ticks, 383

P

Pachysandra, 493 terminalis, 493 Pacific silverweed, 772 Packera, 421 aurea, 422 obovata, 422 paupercula, 422 schweinitziana, 422 paddy melon, 545 Paeonia, 697 albiflora, 698 lactiflora, 698 officinalis, 698

Paeoniaceae, 697–698 painted-cup, 692 painted Indian-mallow, 663 paintedtongue, 863 painted wakerobin, 197 palafox, 422 Palafoxia, 422 texana texana, 422 pale alyssum, 474 pale dock, 742 pale duckweed, 96 pale European wood rush, 191 pale false manna grass, 285 pale-leaved rosette-panicgrass, 250 pale-leaved sunflower, 408 pale moonwort, 65 pale purple coneflower, 395 pale sedge, 144 pale-seeded plantain, 716 pale-spiked lobelia, 496 pale St. John’s-wort, 626 pale swallowwort, 350 pale touch-me-not, 448 pale umbrella-wort, 675 palmate hop clover, 597 Palmer’s amaranth, 317 Paludosae, 135 Panax, 344 quinquefolius, 344 trifolius, 344 Paniceae, 136–137 panicgrass, 247–250, 269–271, 281 panicled hawkweed, 412 panicled hydrangea, 623 panicled tick-trefoil, 581 Panicularia acutiflora, 261 borealis, 261 canadensis, 261 grandis, 262 laxa, 262 melicaria, 262 nervata, 262 obtusa, 262 pallida, 285 septentrionalis, 262 Panicum, 269 aculeatum, 250 acuminatum columbianum, 247 villosissimum, 249 wrightianum, 250 addisonii, 249 amarulum, 269 amarulum, 269 amarum, 269 amarulum, 270 amarum, 270 ashei, 248 atlanticum, 249 barbulatum, 249 bicknellii, 248 boreale, 248 boscii, 248 calliphyllum, 248

i n d e x  9 5 3

Panicum, cont. capillare, 251, 269 agreste, 270 campestre, 270 capillare, 270 flexile, 270 clandestinum, 248 clutei, 249 colonum, 252 columbianum, 247 commonsianum, 249 commutatum, 248 ashei, 248 crus-galli, 252 depauperatum, 248 dichotomiflorum, 270 dichotomiflorum, 270 geniculatum, 270 puritanorum, 270 dichotomum, 249 mattamuskeetense, 249 filiforme, 251 flexile, 270 gattingeri, 270 ischaemum, 251 lanuginosum fasciculatum, 248 implicatum, 248 lindheimeri, 247 villosissimum, 249 latifolium, 249 linearifolium, 249 macrocarpon, 249 mattamuskeetense, 249 microcarpon, 249 miliaceum, 270 minutulum, 250 nitidum ramulosum, 249 oligosanthes scribnerianum, 249 oricola, 247 ovale pseudopubescens, 249 philadelphicum, 270 campestre, 270 gattingeri, 270 philadelphicum, 270 tuckermannii, 271 pumilum, 281 sabulorum, 247 sanguinale, 251 scabriusculum, 250 scoparium, 250 sphaerocarpon, 250 spretum, 247 syzigachne, 234 texanum, 287 tuckermannii, 271 verrucosum, 271 verticillatum, 281 villosissimum, 250 virgatum, 269, 271 viride, 281 walteri, 252 wrightianum, 250 xanthophysum, 250 Papaver, 701 dubium, 702 modestum, 702 obtusifolium, 702

rhoeas, 702 setigerum, 702 somniferum, 702 hortense, 702 Papaveraceae, 698–702 paper birch, 453 paper-mulberry, 670 Pappophorum, 271 vaginatum, 271 Papyrius papyriferus, 670 Paraguay starburr, 371 parasol sedge, 114 Parathelypteris, 69 noveboracensis, 69 simulata, 69 Parietaria, 869 judaica, 870 obtusa, 870 occidentalis, 870 pensylvanica, 870 obtusa, 870 Parker’s pipewort, 172 Parlin’s pussytoes, 375 Parnassia, 702 americana, 702 glauca, 702 Parnassiaceae, 702 Paronychia, 515 argyrocoma, 515 albimontana, 515 canadensis, 515 dichotoma, 515 fastigiata, 515 fastigiata, 515 nuttallii, 515 Parosela alopecuroides, 579 leporina, 579 Parrot’s-feather water-milfoil, 620 parsley, 344, 346 parsnip, 344, 346 Parsonsia petiolata, 660 procumbens, 660 Parthenium, 422 hysterophorus, 422 integrifolium integrifolium, 422 lobatum, 422 Parthenocissus, 887 inserta, 888 quinquefolia, 888 tricuspidata, 888 vitacea, 888 partridge-berry, 828 partridge sensitive-pea, 578 Pascopyrum, 271 smithii, 271 Paspalum, 271 ciliatifolium muhlenbergii, 271 circulare, 271 laeve, 271 ciculare, 271 psammophilum, 271 setaceum, 271 muhlenbergii, 272 psammophilum, 271 setaceum, 272

Pastinaca, 344 sativa, 344 pratensis, 344 pasture greenhead-sedge, 163 pasture thistle, 390 path rush, 189 patience dock, 744 Paulownia, 702 tomentosa, 702 Paulowniaceae, 702 peach, 801 peach-leaved bellflower, 495 peach-leaved willow, 844 pea, 592 pea hawthorn, 786 peanut, 576 pear, 801–802 pearlbrush, 788 pear-leaved crab apple, 793 pearlwort, 516 pearl yarrow, 372 pearly everlasting, 373 pea shrub, 577 Peck’s sedge, 114 pectinate-leaved violet, 884 Pectis pinnata, 426 Pedaliaceae, 703 Pedicularis, 694 canadensis canadensis, 695 dobbsii, 695 furbishiae, 695 gladiata, 695 lanceolata, 695 pallida, 695 virginica, 695 Pellaea, 68 atropurpurea, 68 glabella, 68 pellitory, 870 Peltandra, 96 virginica, 96 pendent silver linden, 668 Pennsylvania bitter-cress, 482 Pennsylvania blackberry, 816 Pennsylvania cudweed, 403 Pennsylvania pellitory, 870 Pennsylvania sedge, 114 Pennsylvania smartweed, 735 penny-cress, 492 pennyroyal, 638, 653 pennyroyal bluecurls, 653 pennywort, 342–343 Penstemon, 713 barbatus, 713 calycosus, 713 digitalis, 713 grandiflorus, 714 hirsutus, 714 laevigatus calycosus, 713 digitalis, 713 pallidus, 714 tubiflorus, 714 achoreus, 714 Pentaphylloides floribunda, 788 Penthoraceae, 703 Penthorum, 703 sedoides, 703

peony, 698 Pepo pepo, 545 pepperbush, 530 peppervine, 887 pepperweed, 487–488 Peramium decipiens, 204 ophioides, 204 pubescens, 204 Perezia, 422 multiflora sonchifolia, 422 Perilla, 645 frutescens, 645 crispa, 645 frutescens, 645 ocymoides, 645 crispa, 645 Periploca, 350 graeca, 350 Peritoma, 529 serrulata, 529 periwinkle, 350–351 perennial glasswort, 327 perennial goosefoot, 323 perennial knawel, 517 perennial quaking grass, 235 perennial ragweed, 373 perennial rose-gentian, 612 perennial rye grass, 265 perennial saltmarsh Americanaster, 442 perennial wall-rocket, 484 perennial woolly bean, 594 perezia, 422 perfoliate bellwort, 99 perfoliate-leaved horsegentian, 504 perilla-mint, 645 perplexed tick-trefoil, 581 Persian clover, 598 Persicaria, 731 amphibia, 732 emersa, 734 laevimarginata, 733 stipulacea, 733 arifolia, 734 bicornis, 735 bistorta, 727 caespitosa longiseta, 734 careyi, 734 chinensis, 734 coccinea, 734 hydropiper, 734 hydropiperoides, 732, 734 breviciliata, 734 euronotora, 734 lapathifolia, 734 longiseta, 734 longistyla, 735 maculosa, 735 minor, 735 muehlenbergii, 734 nepalensis, 735 opelousana, 734 orientalis, 735 pensylvanica, 735 perfoliata, 735

95 4   in de x

Persicaria, cont. puncta robustior, 735 punctata, 735 confertiflora, 735 leptostachya, 735 puritanorum, 735 robustior, 735 sagittata, 735 setacea, 735 interjecta, 735 tonsa, 735 tomentosa, 734 virginiana, 736 vivipara, 727 vulgaris, 735 wallichii, 731 wallichii, 736 persimmon, 547 Petalostemum candidum, 579 Petasites, 422 arcticus, 423 aurea, 423 frigidus, 422 arcticus, 423 palmatus, 423 hybridus, 423 japonicus, 423 obovata, 423 officinalis, 423 palmatus, 423 paupercula, 423 schweinitziana, 423 Petrorhagia, 515 prolifera, 515 saxifraga, 515 Petrosedum reflexum, 543 Petroselinum, 344 crispum, 344 petty spurge, 568 petunia, 861 Petunia, 861 axillaris, 861 integrifolia, 861 nyctaginiflora, 861 violacea, 861 Peucedanum, 344 palustre, 344 Phaca robbinsii, 577 Phacelia, 465 bicknellii, 466 brachyloba, 466 cinerea, 466 distans, 466 egena, 466 linearis, 466 magellanica, 465 minor, 466 purshii, 466 tanacetifolia, 466 viscida, 466 whitlavia, 466 Phacocystis, 138–141 Phaestoglochin, 141–143 Phalaris, 272 arundinacea, 272 picta, 272 canariensis, 272

hispida, 233 oryzoides, 264 Phalaroides arundinacea, 272 Pharbitis barbigera, 536 purpurea, 536 Phaseolus, 592 coccineus, 592 leiospermus, 594 max, 582 polystachios, 592 aquilonius, 592 smilacifolius, 592 vulgaris, 592 Phedimus, 542 aizoon, 540 ellacombeanus, 540 spurius, 542 stoloniferus, 542 Phegopteris, 70 connectilis, 70 hexagonoptera, 70 Phellodendron, 829 amurense, 829 sachalinense, 829 japonicum, 829 sachalinense, 829 Philadelphia fleabane, 396 Philadelphia panicgrass, 270 Philadelphus, 623 caucasicus, 623 coronarius, 623 grandiflorus, 623 inodorus, 623 grandiflorus, 623 intectus, 623 latifolius, 623 pubescens, 623 intectus, 623 Phleum, 272 alpinum, 272 commutatum, 272 arenarium, 272 bellardii, 272 commutatum, 272 americanum, 272 nodosum, 272 pratense, 272 nodosum, 272 subulatum, 272 phlox, 723–724 Phlox, 723 bifida, 723 divaricata divaricata, 723 drummondii, 724 latifolia, 724 maculata, 724 odorata, 724 ovata, 724 paniculata, 724 pilosa, 724 virens, 724 ×procumbens, 724 stolonifera, 724 subulata, 724 Photinia floribunda, 773 melanocarpa, 773 pyrifolia, 773

Phragmites, 273 americanus, 273 australis, 273 americanus, 273 communis, 273 Phryma, 705 leptostachya, 705 confertifolia, 705 Phrymaceae, 703–705 Phyllantherum recurvatum, 197 Phyllodoce, 557 caerulea, 557 Phyllostachyae, 143 Phyllostachys, 273 dulcis, 273 Physalis, 861 alkekengi, 862 ambigua, 862 angulata, 862 lanceifolia, 862 pendula, 862 grandiflora, 859 grisea, 862 heterophylla, 862 ambigua, 862 clavipes, 862 heterophylla, 862 nyctaginea, 862 villosa, 862 ixocarpa, 862 longifolia, 862 subglabrata, 862 philadelphica, 862 immaculata, 862 pruinosa, 862 pubescens, 861 grisea, 862 integrifolia, 863 subglabrata, 862 virginiana, 862 subglabrata, 862 virginiana, 863 Physalodes physalodes, 859 Physematium obtusum, 74 Physocarpus, 793 opulifolius, 793 Physoglochin, 143 Physostegia, 645 virginiana elongata, 645 granulosa, 645 speciosa, 645 virginiana, 645 Phytolacca, 705 americana americana, 705 decandra, 705 Phytolaccaceae, 705 Phytosalpinx americanus, 640 aspera, 640 Picea, 77 abies, 78 canadensis, 78 excelsa, 78 glauca, 78 mariana, 78

nigra, 78 rubens, 78 pickerelweed, 289 Pickering’s reed grass, 240 Picradenia odorata, 413 Picris, 423 echiodes, 409 hieracioides, 423 piedmont staggerbush, 557 Pieris, 557 floribunda, 557 pignut hickory, 628 pigweed Russian, 320 Pilea, 870 fontana, 870 opaca, 870 pumila, 870 pumila, 870 Pilosella aurantiaca, 411 caespitosa, 411 flagellaris, 411 novae-angliae, 479 officinarum, 412 piloselloides, 412 praealta, 412 pimpernel, 673 pimpernel willow-herb, 683 Pimpinella, 344 anisum, 345 saxifraga, 345 Pinaceae, 76–80 pin cherry, 801 pincushion, 388 pincushion-plant, 722 pincushions, 503 pine, 79 pine-barren false heather, 527 pine barren flatsedge, 155 pine barren sandplant, 513 pine-drops, 557 pine-needle toadflax, 712 pine-sap, 556 pine-woods coneflower, 425 Pinguicula, 654 vulgaris vulgaris, 654 pink, 202, 391, 482, 511, 558, 560, 709 pink azalea, 560 pink bitter-cress, 482 pink-corydalis, 699 pink cuckoo bitter-cress, 482 pink lady’s-slipper, 202 pinks, 511, 515 pink-shell azalea, 560 pink shinleaf, 558 pink tickseed, 391 pink turtlehead, 709 pinnate hop clover, 597 pin oak, 606 Pinus, 78 balsamea, 77 banksiana, 79 canadensis, 80 divaricata, 79 mugo, 79 nigra, 79 resinosa, 79

i n d e x  9 5 5

Pinus, cont. rigida, 79 strobus, 79 sylvestris, 79 thunbergiana, 79 thunbergii, 79 pinweed, 528–529 pipewort, 172 Piptatherum, 273 canadense, 273 pungens, 273 racemosum, 273 Piptochaetium, 274 avenaceum, 274 Pisum, 592 arvense, 592 sativum, 592 arvense, 592 pitcherplant, 852 pitch pine, 79 pit-seeded goosefoot, 323 Pityopsis, 423 falcata, 423 Plagiobothrys, 466 reticulatus, 466 trachycarpus, 466 plains eryngo, 341 plains lovegrass, 258 Plananthus selago, 40 Plantaginaceae, 705–720 Plantago, 714 altissima, 716 americana, 712 arenaria, 715 aristata, 715 nuttallii, 715 caroliniana, 716 coronopus, 715 gnaphalioides, 716 aristata, 715 halophila, 716 incana, 716 indica, 715 intermedia, 716 juncoides decipiens, 716 glauca, 716 laurentiana, 716 lanceolata, 716 sphaerostachya, 716 major, 715 intermedia, 716 pilgeri, 716 pleiosperma, 716 scopulorum, 716 maritima, 715 borealis, 716 juncoides, 716 media, 716 nuttallii, 715 oliganthos, 716 fallax, 716 patagonica, 716 breviscapa, 716 gnaphalioides, 716 psyllium, 715 purshii, 716 breviscapa, 716 pusilla, 716 major, 716

ramosa, 715 rugelii, 716 asperula, 716 virginica, 716 viridescens, 716 vulgaris, 715 plantain, 716 mud, 289 plantain-leaved pussytoes, 375 plantain-leaved sedge, 116 plantain-lily, 173 plantain viper’s-bugloss, 461 Platanaceae, 720 Platanthera, 207 ×andrewsii, 209 aquilonis, 208 blephariglottis blephariglottis, 208 ciliaris, 208 clavellata, 209 cristata, 209 dilatata dilatata, 209 flava, 209 herbiola, 209 grandiflora, 209 hookeri, 209 huronensis, 209 hyperborea, 209 ×keenanii, 209 lacera, 209 leucophaea, 210 macrophylla, 210 obtusata obtusata, 210 orbiculata, 210 psycodes, 210 Platanus, 720 acerifolia, 720 hybrida, 720 occidentalis, 720 Pleconax conica, 519 Plectoma inflata, 656 Pleopogon setosus, 267 Pleurogyne rotata, 611 Pluchea, 423 odorata succulenta, 423 plum, 798–801 Plumbaginaceae, 721 plumeless-thistle, 384–385 plume-poppy, 701 Plymouth rose-gentian, 612 Pneumaria maritima, 463 Pneumonanthe andrewsii, 610 Poa, 274 agassizensis, 276 alpigena, 276 alsodes, 275 americana, 278 angustifolia, 276 annua, 275 bulbosa, 274 vivipara, 275 chapmaniana, 275

cilianensis, 257 compressa, 275 debilis, 277 glauca, 276 glauca, 276 glaucantha, 276 hypnoides, 258 interior, 276 laxa, 276 fernaldiana, 276 maritima, 278 nemoralis, 274 interior, 276 nevadensis, 277 palustris, 274, 276 pectinacea, 258 pilosa, 258 pratensis, 274 agassizensis, 276 alpigena, 276 angustifolia, 276 pratensis, 276 rigida, 245 saltuensis, 277 languida, 277 saltuensis, 277 secunda, 277 juncifolia, 277 triflora, 276 trivialis, 277 trivialis, 277 Poaceae, 213–288 pod-grass, 301 Podophyllum, 449 peltatum, 449 Podostemaceae, 721 Podostemum, 721 abrotanoides, 721 ceratophyllum, 721 poet’s daffodil, 93 pogonia, 205, 210 Pogonia, 210 affinis, 205 ophioglossoides, 210 trianthophora, 212 pointed broom sedge, 124 pointed-leaved tick-trefoil, 582 poison darnel, 265 poison-hemlock, 340 poison-ivy, 331 poison rubberweed, 413 poison-sumac, 331 poke milkweed, 349 pokeweed, 705 Polanisia, 529 dodecandra dodecandra, 529 trachysperma, 529 graveolens, 529 trachysperma, 529 Polemoniaceae, 721–725 Polemoniella micrantha, 724 Polemonium, 724 caeruleum, 724 vanbruntiae, 725 micranthum, 724 reptans, 725 reptans, 725 rubrum, 722 vanbruntiae, 725

Polycarpon, 516 tetraphyllum tetraphyllum, 516 Polycnemum, 327 majus, 327 Polygala, 725 ambigua, 726 cruciata, 726 aquilonia, 726 nuttallii, 726 paucifolia, 726 polygama, 726 sanguinea, 726 senega, 726 latifolia, 726 verticillata, 726 ambigua, 726 dolichoptera, 726 isocycla, 726 viridescens, 726 Polygalaceae, 725–726 Polygonaceae, 726–744 Polygonatum, 300 biflorum, 300 commutatum, 300 commutatum, 300 hirtum, 300 latifolium, 300 pubescens, 300 Polygonum, 736 achoreum, 738 acre, 735 aequale, 738 oedocarpum, 738 alatum, 735 allocarpum, 739 amphibium coccineum, 734 emersum, 734 laevimarginatum, 733 natans, 733 stipulaceum, 733 arenarium, 738 arenastrum, 738 arifolium, 734 lentiforme, 734 pubescens, 734 articulatum, 738 atlanticum, 739 aubertii, 730 aviculare, 738 angustissimum, 738 arenastrum, 738 aviculare, 738 depressum, 738 erectum, 739 littorale, 739 neglectum, 738 rurivagum, 738 vegetum, 738 baldschuanica, 730 bellardii, 739 bicorne, 735 bistortum, 727 buxifolium, 739 buxiforme, 739 caespitosum longisetum, 734 careyi, 734 chinense, 734

95 6   in de x

Polygonum, cont. cilinode laevigatum, 730 coccineum, 734 rigidulum, 733 terrestre, 734 convolvulus subulatum, 730 cristatum, 730 cuspidatum, 730 douglasii, 739 latifolium, 739 dubium, 735 dumetorum, 730 erectum, 739 achoreum, 738 exsertum, 739 fagopyrum, 727 fowleri fowleri, 739 glaucum, 739 heterophyllum, 738 angustissimum, 738 hydropiperoides, 734 adenocalyx, 734 breviciliatum, 734 bushianum, 734 digitatum, 734 euronotorum, 734 opelousanum, 734 setaceum, 735 hydropiper, 734 lapathifolium, 734 ovatum, 734 prostratum, 734 littorale, 739 longistylum, 735 maritimum, 739 minus, 735 subcontinuum, 735 monspeliense, 738 muehlenbergii, 734 natans, 733 nepalense, 735 orientale, 735 oxyspermum, 739 pensylvanicum, 735 eglandulosum, 735 genuinum, 735 laevigatum, 735 nesophilum, 735 perfoliatum, 735 persicaria, 735 prolificum, 739 punctatum, 735 confertiflorum, 735 leptostachyum, 735 robustius, 735 puritanorum, 735 raii, 739 ramosissimum, 739 prolificum, 739 ramosissimum, 739 robustius, 735 rurivagum, 738 sachalinense, 730 sagittatum, 735 gracilentum, 735 scandens dumetorum, 730

setaceum interjectum, 735 stipulaceum, 733 tataricum, 727 tenue, 736 protrusum, 740 triangulum, 739 virginianum, 736 glaberrimum, 736 viviparum, 727 Polymnia, 423 abyssinica, 404 canadensis, 423 radiata, 423 radiata, 423 Polypodiaceae, 66–67 Polypodium, 66 appalachianum, 67 bulbiferum, 72 fragile, 72 fragrans, 57 ×incognitum, 67 obtusum, 74 virginianum, 67 vulgare virginianum, 67 polypody, 67 Polypogon, 277 monspeliensis, 277 semiverticillatus, 277 viridis, 277 Polystichum, 58 acrostichoides, 58 braunii, 58 ×potteri, 58 pond-lily, 85 pond water-starwort, 708 pondweed, 291–298 Pontederia, 289 cordata, 289 angustifolia, 289 lanceolata, 289 Pontederiaceae, 289 poor-man’s pepperweed, 488 popcorn-flower, 466 poplar, 829–832 poplar hawthorn, 786 poppy, 701–702 Populus, 830 alba, 831 bolleana, 831 angulata, 832 missouriensis, 832 balsamifera, 830 balsamifera, 832 candicans, 832 lanceolata, 832 missouriensis, 832 subcordata, 832 candicans, 832 ×canescens, 831 deltoides, 832 angulata, 832 deltoides, 832 missouriensis, 832 pilosa, 832 grandidentata, 832 ×heimburgeri, 832 heterophylla, 832 italica, 832 ×jackii, 832

nigra, 832 italica, 832 ×rouleauiana, 831 ×smithii, 832 tacamahaca, 832 candicans, 832 lanceolata, 832 tremula, 832 tremuloides, 832 tremuloides, 832 magnifica, 832 porcupine sedge, 149 Porocystis, 144 Porsildia groenlandica, 514 Porteranthus trifoliatus, 792 Portulaca, 746 grandiflora, 746 neglecta, 746 oleracea, 746 retusa, 746 Portulacaceae, 744–746 post oak, 606 Potamogeton, 290 ×absconditus, 297 ×aemulans, 294 alpinus, 294 tenuifolius, 294 americanus, 296 amplexicaulis, 297 amplifolius, 294 angustifolius, 296 ×argutulus, 295 berchtoldii, 294 gemmiparus, 295 bicupulatus, 294 bupleuroides, 297 capillaceus, 294 confervoides, 295 crispus, 295 diversifolius trichophyllus, 294 epihydrus, 295 nuttallii, 295 ramosus, 295 ×faxonii, 296 filiformis, 298 fluitans, 296 foliosus, 291 foliosus, 295 macellus, 295 friesii, 295 gemmiparus, 295 gramineus, 295 maximus, 295 myriophyllus, 295 typicus, 295 ×haynesii, 297 heterophyllus, 296 hillii, 296 illinoensis, 296 lateralis, 298 ×mirabilis, 296 ×mysticus, 294 natans, 296 ×nitens, 296 nodosus, 296 oakesianus, 296 obtusifolius, 296 ogdenii, 297

pectinatus, 298 perfoliatus, 297 bupleuroides, 297 richardsonii, 297 pinnatum, 621 porteri, 296 praelongus, 297 angustifolius, 297 pulcher, 297 pusillus, 297 gemmiparus, 295 minor, 297 mucronatus, 294 tenuissimus, 294 richardsonii, 297 robbinsii, 297 ×scoliophyllus, 294 ×sparganiifolius, 295 ×spathuliformis, 295 spirillus, 297 strictifolius, 297 rutiloides, 297 strictifolius, 298 vaseyi, 298 ×versicolor, 295 zosteriformis, 298 Potamogetonaceae, 290– 298 potato, 536, 865 Potentilla, 794 alba, 796 anglica, 796 anserina, 772 concolor, 772 groenlandica, 772 sericea, 772 argentea, 795 pseudocalabra, 796 arguta, 788 camporum, 797 canadensis, 796 villosissima, 796 canescens, 796 caroliniana, 796 egedii groenlandica, 772 erecta, 796 floribunda, 788 fruticosa floribunda, 788 gracilis, 796 gracilis, 796 pulcherrima, 797 hyparctica, 797 inclinata, 796 indica, 797 intermedia, 797 canescens, 796 litoralis, 797 longipedunculata, 796 macropetala, 796 millegrana, 797 monspeliensis, 797 neumanniana, 798 norvegica, 797 hirsuta, 797 labradorica, 797 monspeliensis, 797 palustris, 774 parvifolia, 774 villosa, 774

in d e x  9 57

Potentilla, cont. pectinata, 797 pensylvanica litoralis, 797 pectinata, 797 virgulata, 797 pentandra, 797 procumbens, 796 pulcherrima, 797 pumila, 796 recta, 797 obscura, 797 pilosa, 797 sulphurea, 797 reptans, 797 rivalis, 797 millegrana, 797 pentandra, 797 robbinsiana, 797 sibbaldii, 818 simplex, 794, 797 argyrisma, 797 calvescens, 797 typica, 797 tabernaemontani, 798 thuringiaca, 798 tridentata, 818 verna, 798 virgulata, 797 Poterium officinale, 817 polygamum, 817 pot-marigold, 384 poverty brome, 239 poverty dropseed, 285 poverty oatgrass, 244 poverty-weed, 327 Powell’s amaranth, 317 prairie-clover, 579 prairie cordgrass, 283 prairie crowfoot, 759 prairie dropseed, 285 prairie dropwort, 788 prairie ironweed, 446 prairie knotweed, 739 prairie Koeler’s grass, 263 prairie moonwort, 64 prairie pepperweed, 487 prairie redroot, 762 prairie rose, 805 prairie sedge, 129 prairie sunflower, 408 prairie wakerobin, 197 prairie wedgescale, 283 prairie willow, 846–847 prairie wormwood, 378 prairie Mexican-hat, 424 Prenanthes alba, 419 nana, 420 altissima, 419 cinnamomea, 419 hispidula, 419 bootii, 419 nana, 420 racemosus, 419 serpentaria, 420 simplicifolia, 420 trifoliolata, 420 nana, 420 pricklegrass, 242

prickly-ash, 829 prickly bog sedge, 146 prickly clover, 597 prickly comfrey, 467 prickly fanpetals, 667 prickly lettuce, 415 prickly-pear, 493 prickly-poppy, 699 prickly quill sedge, 123 prickly saltwort, 328 prickly scorpion’s-tail, 593 prickly sedge, 143 prickly tree-clubmoss, 44 pride-of-Rochester, 622 primrose, 747 primrose-leaved violet, 885 Primula, 747 anisodora, 747 intercedens, 747 japonica, 747 laurentiana, 747 mistassinica, 747 macropoda, 747 noveboracensis, 747 veris, 747 Primulaceae, 746–747 prince’s-feather amaranth, 317 prince’s-feather smartweed, 735 prince’s-pine, 553 Pringle’s hawthorn, 786 privet, 678–679 Proboscidea, 703 louisianica louisianica, 703 professor-weed, 581 proliferous childing-pink, 515 proliferous fescue, 259 Proserpinaca, 621 amblyogona, 621 ×intermedia, 622 palustris, 621 amblyogona, 621 crebra, 621 pectinata, 622 Proso millet, 270 prostrate amaranth, 316 prostrate bishop’s-cap, 854 prostrate sandmat, 568 prostrate vervain, 872 prototype quillwort, 42 Prunella, 645 alba, 646 laciniata, 646 vulgaris, 646 elongata, 646 lanceolata, 646 vulgaris, 646 Prunus, 798 alleghaniensis, 799 americana nigra, 801 avium, 800 cerasus, 800 domestica, 800 insititia, 800 gravesii, 801 hortulana, 800 insititia, 800 mahaleb, 801 maritima, 801

gravesii, 801 maritima, 801 nigra, 801 paniculata, 866 pensylvanica, 799 pensylvanica, 801 persica, 801 pumila, 799 depressa, 801 serotina, 799 serotina, 801 serrulata, 801 spinosa, 801 susquehanae, 801 virginiana, 801 virginiana, 801 Psedera quinquefolia, 888 Pseudabutilon, 667 callimorphum friesii, 667 pedunculatum, 667 stuckertii, 667 Pseudognaphalium, 423 helleri, 424 macounii, 424 micradenium, 424 obtusifolium, 424 Pseudolycopodiella, 47 caroliniana, 47 Pseudolysimachion spurium, 720 Pseudosasa, 277 japonica, 277 Pseudotsuga, 79 menziesii menziesii, 80 taxifolia, 79 Psilocarya nitens, 164 scirpoides, 164 Ptelea, 829 baldwinii, 829 trifoliata trifoliata, 829 Pteridaceae, 67–68 Pteridium, 54 aquilinum, 54 latiusculum, 54 pseudocaudatum, 54 Pteris atropurpurea, 68 Pterospora, 557 andromedea, 557 Ptilimnium, 345 capillaceum, 345 pubescent sedge, 129 Puccinellia, 277 americana, 277 distans, 278 fasciculata, 278 fernaldii, 285 maritima, 277 nuttalliana, 278 pallida, 285 pumila, 278 Pueraria, 592 lobata, 592 montana lobata, 592 thunbergiana, 592

Pulmonaria, 466 saccharata, 466 pumpkin, 545 puncture-vine, 890 Puritan smartweed, 735 purple agalinis, 690 purple amaranth, 316 purple bee-balm, 644 purple chokeberry, 773 purple cliff-brake, 68 purple clover, 598 purple crown-vetch, 593 purple cudweed, 403 purple foxglove, 710 purple giant-hyssop, 635 purple goat’s beard, 445 purplehead, 286 purple-headed sneezeweed, 404 purpleheads, 286 purple Joe-Pye weed, 402 purple knapweed, 386 purple loosestrife, 661 purple lovegrass, 258 purple meadow-parsnip, 346 purple meadow-rue, 760 purple medick, 591 purple milkweed, 349 purple moorgrass, 266 purple mountain-heath, 557 purple mountain saxifrage, 855 purple orpine, 541 purple painted-cup, 692 purple pitcherplant, 852 purple sandgrass, 286 purple scorpion-weed, 466 purple screwstem, 608 purple-stemmed Americanaster, 442 purple-stemmed Angelica, 338 purple-stemmed beggar-ticks, 382 purple threeawn, 233 purple-topped vervain, 872 purpletop tridens, 286 purple virgin’s-bower, 753 purple wood-aster, 400 Pursh’s blue maidencane, 230 Pursh’s plantain, 716 purslane, 746 sea, 311 purslane speedwell, 719 pussytoes, 375 pussy willow, 845 pustulose viper’s-bugloss, 461 putty-root, 200 Pycnanthemum, 646 ×clinopodioides, 646 incanum, 646 incanum, 646 muticum, 647 tenuifolium, 647 torrei, 647 verticillatum, 647 pilosum, 647 verticillatum, 647 virginianum, 647 Pycreus diander, 153 polystachyos, 155

95 8   i n de x

Pyrethrum majus, 379 parthenium, 443 pygmy soapwort, 517 pygmy-weed, 541 Pylae’s soft rush, 188 Pyrola, 558 americana, 558 asarifolia, 558 americana, 558 asarifolia, 558 convoluta, 558 incarnata, 558 ovata, 558 purpurea, 558 chlorantha, 558 convoluta, 558 paucifolia, 558 convoluta, 558 elliptica, 558 maculata, 553 minor, 558 rotundifolia americana, 558 asarifolia, 558 secunda, 557 obtusata, 557 uliginosa, 558 uniflora, 557 virens, 558 convoluta, 558 Pyrus, 801 americana, 818 decora, 819 arbutifolia, 773 glabra, 773 aucuparia, 818 baccata, 793 calleryana, 802 communis, 802 cydonia, 788 decora, 819 groenlandica, 819 floribunda, 773 japonica, 774 melanocarpa, 773 ovalis, 772 pinnatifida, 819 prunifolia, 793 pumila, 793 sanguinea, 772 sieboldii, 793 wangenheimiana, 770

Q quack grass, 285 quaking grass, 235 quaking poplar, 832 Quamoclit quamoclit, 536 vulgaris, 536 Quebec hawthorn, 787 Quercus, 602 acuminata, 605 alba, 604 ×bebbiana, 604 ×benderi, 605 bicolor, 604 ×bimundorum, 604

borealis, 606 maxima, 606 cerris, 605 coccinea, 605 tuberculata, 605 ×faxonii, 604 ×fernaldii, 605 ×fernowii, 604 ×fontana, 605 ×hawkinsii, 606 ilicifolia, 605 imbricaria, 605 ×jackiana, 604 ×leana, 605 macrocarpa, 605 depressa, 605 mandanensis, 605 montana, 605 muehlenbergii, 605 palustris, 606 phellos, 606 prinoides, 606 rufescens, 606 prinoides acuminata, 605 prinus, 605 acuminata, 605 ×rehderii, 605 ×robbinsii, 605 robur, 606 rubra, 606 ambigua, 606 borealis, 606 ×saulii, 604 ×schuettei, 604 stellata, 606 ×stelloides, 606 ×vaga, 606 velutina, 606 quickweed, 402 quill-leaved arrowhead, 90 quill spikesedge, 160 quillwort, 40–42 quince, 774, 788

R rabbit-foot clover, 597 rabbit-tobacco, 424 racemed milkwort, 726 Racemosae, 144–145 Radicula amphibia, 491 armoracia, 475 hispida, 491 palustris, 491 sessiliflora, 491 sylvestris, 491 Radiola linoides, 658 radish, 490 ragged robin lychnis, 513 ragged thoroughwort, 399 ragweed, 373 ragwort, 414, 427 Raimannia laciniata, 687 rain-tree, 852 rambler rose, 806

Ramischia secunda, 557 ram’s-head lady’s-slipper, 202 ram’s-horn, 703 rancher’s fiddleneck, 458 Rand’s eyebright, 693 Rand’s goldenrod, 433 Ranunculaceae, 747–761 Ranunculus, 755 abortivus, 757 acrolasius, 757 indivisus, 757 acris, 757 latisectus, 757 allegheniensis, 757 ambigens, 758 amphibius, 758 aquatilis, 758 capillaceus, 758 diffusus, 758 longirostris, 758 subrigidus, 758 bulbosus, 758 dissectus, 758 caricetorum, 758 cymbalaria, 758 alpinus, 758 saximontanus, 758 delphiniifolius, 758 eucyclus, 757 fascicularis, 758 apricus, 758 ficaria bulbilifer, 754 filiformis ovalis, 758 reptans, 758 flabellaris, 758 flammula, 758 ovalis, 758 reptans, 758 samolifolius, 758 gmelinii, 759 hookeri, 759 prolificus, 759 purshii, 759 terrestris, 759 hispidus, 759 caricetorum, 758 eurylobus, 759 falsus, 759 intermedius, 758 lapponicus, 754 longirostris, 758 micranthus, 759 cymbalistes, 759 delitescens, 759 obtusiusculus, 758 parvulus, 759 pensylvanicus, 759 pseudobulbosus, 759 purshii, 759 recurvatus, 757 repens, 759 degeneratus, 759 erectus, 759 glabratus, 759 pleniflorus, 759 villosus, 759

reptans filiformis, 758 ovalis, 758 rhomboideus, 759 sardous, 759 sceleratus, 757 sceleratus, 759 septentrionalis caricetorum, 758 subrigidus, 758 trichophyllus, 758 confervoides, 758 eradicatus, 758 lutulentus, 758 rape, 478 Raphanus, 489 raphanistrum, 490 raphanistrum, 490 sativus, 490 Rapistrum, 489 rugosum, 490 raspberry, 807–817 Ratibida, 424 columnifera, 424 pinnata, 424 rat-tail six-weeks grass, 288 rattlebox, 578 rattlesnake brome, 238 rattlesnake fern, 65 rattlesnake hawkweed, 413 rattlesnake manna grass, 261 rattlesnake-plantain, 204 rattlesnake-root, 419–420 rayless annual American-aster, 438 rayless chamomile, 418 Ray’s knotweed, 739 Razoumofskya pusilla, 887 red baneberry, 750 red bearberry, 552 redbud, 578 red campion, 519 red chokeberry, 773 red clover, 598 red columbine, 752 red crowberry, 553 reddish pondweed, 294 red elderberry, 308 red false bartsia, 694 red fescue, 259 red-footed spikesedge, 159 red goosefoot, 324 red hemp-nettle, 637 red henbit, 639 red-leaved rose, 806 red lobelia, 496 redmaids, 745 red maple, 851 red morning-glory, 536 red mulberry, 671 red orache, 320 red-osier dogwood, 539 red pine, 79 red raspberry, 815 red-ringed milkweed, 350 redroot, 762 red-rooted amaranth, 317 red-root flatsedge, 153 red sand-spurry, 521 red-seeded dandelion, 444

in d e x  9 5 9

red snowberry, 503 red spruce, 78 red-stemmed gentian, 610 red-stemmed stork’s-bill, 613 red threeawn, 232 redtop bentgrass, 227 redtop-panicgrass, 281 red turtlehead, 709 red wakerobin, 197 red-whiskered clammyweed, 529 reed, 240, 262, 272–273 reed canary grass, 272 reed manna grass, 262 reflexed sedge, 143 Rehsonia sinensis, 600 Reseda, 761 alba, 762 lutea, 762 luteola, 762 odorata, 762 Resedaceae, 761–762 resupinate bladderwort, 656 retrorse sedge, 149 Reynoutria baldschuanica, 730 cilinodis, 730 convolvulus, 730 japonica, 730 sachalinensis, 730 scandens, 730 dumetorum, 730 Reznicek’s sedge, 114 Rhamnaceae, 762–763 Rhamnus, 762 alnifolia, 762 cathartica, 763 citrifolia, 763 davurica, 763 davurica, 763 frangula, 762 utilis, 763 Rheum, 740 rhabarbarum, 740 rhaponticum, 740 Rhexia, 668 lanceolata, 668 mariana exalbida, 668 mariana, 668 stricta, 668 virginica, 668 purshii, 668 Rhinanthus, 695 alectorolophus, 695 borealis, 695 crista-galli, 695 fallax, 695 groenlandicus, 695 major, 695 minor, 695 groenlandicus, 695 minor, 695 oblongifolius, 695 stenophyllus, 695 virginicus, 691 Rhode Island bentgrass, 227 Rhodes windmill-grass, 242

Rhodiola, 542 roanensis, 542 rosea, 542 rhododendron, 560 Rhododendron, 558 calendulaceum, 560 canadense, 560 canescens, 560 carolinianum, 560 catawbiense, 560 groenlandicum, 560 japonicum, 560 lapponicum, 560 maximum, 559 nudiflorum, 560 glandiferum, 560 roseum, 560 periclymenoides, 560 eglandulosum, 560 prinophyllum, 560 roseum, 560 vaseyi, 560 vermiculata, 568 viscosum, 560 rhodora, 560 Rhodora canadensis, 560 Rhodotypos, 802 scandens, 802 rhubarb, 740 Rhus, 330 aromatica aromatica, 330 copallinum latifolium, 330 cotinus, 329 glabra, 330 hirta, 330 ×pulvinata, 330 radicans, 331 rydbergii, 331 typhina, 330 vernix, 331 Rhynchospora, 163 alba, 164 fusca, 164 capillacea, 164 leviseta, 164 capitellata, 164 fusca, 164 glomerata, 164 inundata, 164 macrostachya, 164 inundata, 164 nitens, 164 scirpoides, 164 torreyana, 164 ribbed sedge, 144 ribbon-leaved pondweed, 295 Ribes, 616 americanum, 618 aureum, 617 villosum, 618 cynosbati, 618 atrox, 618 glabratum, 618 inerme, 618 floridum, 618 glandulosum, 618 grossularia, 619

hirtellum, 618 calcicola, 618 saxosum, 618 lacustre, 618 missouriense, 618 nigrum, 618 odoratum, 618 oxyacanthoides calcicola, 618 hirtellum, 618 lacustre, 618 saxosum, 618 prostratum, 618 resinosum, 618 rotundifolium, 618 rubrum, 619 alaskanum, 619 sativum, 619 stamineum, 618 sylvestre, 619 triflorum, 618 triste, 619 albinervium, 619 uva-crispa, 618 reclinatum, 619 sativum, 619 vulgare, 619 rib-seeded sandmat, 568 rice, 264, 268, 288 rice cut grass, 264 rice-field flatsedge, 154 rice grass, 268 Richardson’s pondweed, 297 Richardson’s sedge, 118 rich woods sedge, 129 Ricinus, 569 communis, 569 Ridan alternifolia, 445 ridged yellow flax, 658 rigid sedge, 137 ripgut brome, 239 river birch, 453 river grape, 889 river hawthorn, 784 river-hemp, 594 river horsetail, 59 river tuber-bulrush, 103 riverweed, 721 rivet wheat, 287 roadside agrimony, 768 roadside false alyssum, 476 Robbins’ cinquefoil, 797 Robbin’s milk-vetch, 577 Robbins’ pondweed, 297 Robbins’ spikesedge, 161 Robertiella robertiana, 616 Robinia, 592 fertilis, 593 grandiflora, 593 hispida, 593 fertilis, 593 hispida, 593 pallida, 593 pseudoacacia, 593 speciosa, 593 viscosa viscosa, 593 Robinson’s hawkweed, 412 Robin’s plantain fleabane, 397

rock-brake, 68 rockcress, 475, 477–479 rock elm, 869 rocket candytuft, 486 rock goldentuft, 475 rock muhly, 268 rock polypody, 67 rock scorpion-weed, 465 rock soapwort, 517 rock stonecrop, 543 rock whitlow-mustard, 484 Roman wormwood, 379 rope-root sedge, 117 Rorippa, 489 americana, 490 amphibia, 491 aquatica, 491 armoracia, 475 austriaca, 491 globosa, 491 hispida, 491 islandica, 491 fernaldiana, 491 microphylla, 488 nasturtium-aquaticum, 488 palustris, 490 fernaldiana, 490 glabra, 490 hispida, 490 palustris, 490 williamsii, 490 prostrata, 490 sessiliflora, 491 sylvestris, 491 Rosa, 802 acicularis, 805 bourgeauiana, 805 sayi, 805 arkansana, 805 suffulta, 805 blanda, 805 blanda, 805 glabra, 805 glandulosa, 805 bourgeauiana, 805 canina, 806 carolina, 806 carolina, 806 deamii, 806 glandulosa, 806 lucida, 807 subserrulata, 806 villosa, 806 cinnamomea, 806 eglanteria, 807 floridana, 807 gallica, 806 glauca, 806 johannensis, 805 luciae, 806 lucida, 807 lyonii, 806 majalis, 806 micrantha, 806 mollis, 806 multiflora, 806 nanella, 807 nitida, 804, 807 ×novae-angliae, 806 ×palustriformis, 805

96 0   in de x

Rosa, cont. palustris, 807 pimpinellifolia, 807 rubifolia, 807 rubiginosa, 807 micrantha, 806 rubrifolia, 806 rugosa, 807 sempervirens, 807 setigera, 807 sherardii, 803 spinosissima, 807 subblanda, 805 subserrulata, 806 tomentosa, 806 virginiana, 807 lamprophylla, 807 nanella, 807 wichuraiana, 806 Rosaceae, 763–820 rose, 802–807 rosebay, 560 rose catchfly, 511 rose daphne, 867 rose-gentian, 612 rose loosestrife, 661 rose-mallow, 664–665 rose mock-vervain, 871 rose monk’s-wort, 464 rose pogonia, 210 rosering blanket-flower, 402 roseroot, 542 rosette-panicgrass, 247–250 rosinweed, 428 Rostrales, 145 rosy meadowsweet, 820 rosy sedge, 143 Rotala, 661 ramosior, 661 interior, 661 typica, 661 Roubieva multifida, 326 rough bedstraw, 825 rough bentgrass, 228 rough buttonweed, 822 rough cocklebur, 446 rough dropseed, 284 rough false pennyroyal, 638 rough fleabane, 397 rough-fruited bedstraw, 827 rough-fruited popcorn-flower, 466 rough-fruited water-hemp, 317 rough hawk’s-beard, 393 rough hawkweed, 412 rough-leaved goldenrod, 432 rough marsh-elder, 414 rough-sheathed blue grass, 277 rough wood-aster, 400 round-fruited hedge-hyssop, 710 round-fruited pinweed, 528 round-fruited rosettepanicgrass, 250 round-fruited rush, 187 round-fruited short-scaled sedge, 125 round-headed bush-clover, 586 round-leaved bog-orchid, 210 round-leaved cancerwort, 711

round-leaved dogwood, 539 round-leaved eyebright, 693 round-leaved orchid, 200 round-leaved shadbush, 772 round-leaved sundew, 547 round-leaved thorough-wax, 340 round-leaved thoroughwort, 399 round-leaved trailing ticktrefoil, 581 round-leaved violet, 885 round-pod water-primrose, 684 royal fern, 66 royal hawthorn, 787 royal knight’s-spur, 754 Rubacer columbianum, 816 odoratum, 816 rubberweed, 413 Rubia, 828 tinctoria, 828 Rubiaceae, 820–828 Rubus, 807 abactus, 816 alius, 812 allegheniensis, 811 gravesii, 811 amicalis, 813 amnicola, 816 andrewsianus, 816 arenicola, 812 argutus, 812 randii, 813 arundelanus, 816 ascendens, 817 auroralis, 811 avipes, 816 barbarus, 816 bellobatus, 814 bifrons, 813 bigelovianus, 817 boottianus, 817 brainerdii, 812 brevipedalis, 816 canadensis, 813 chamaemorus, 813 chapmanii, 813 coloniatus, 816 conanicutensis, 816 cubitans, 814 cuneifolius, 813 angustior, 813 subellipticus, 813 curtipes, 812 dalibarda, 813 deaneanus, 817 discolor, 813 elegantulus, 813 enslenii, 814 facetus, 816 felix, 814 fernaldianus, 811 flagellaris, 814 frondisentus, 817 frondosus, 814 geophilus, 814 heterogeneous, 814 hispidoides, 817 hispidus, 814

honorus, 816 idaeus, 815 aculeatissimus, 815 caudatus, 815 egglestonii, 815 eucyclus, 815 heterolasius, 815 idaeus, 815 melanolasius, 815 sachalinensis, 815 strigosis, 815 illecebrosus, 815 insons, 816 insulanus, 814 janssonii, 812 japonicus, 792 jaysmithii, 815 jugosus, 812 junceus, 817 junior, 817 kennedyanus, 813 laciniatus, 816 lawrencei, 817 longissimus, 811 maniseesensis, 814 melanolasius, 815 multiformis, 813 multispinus, 814 ×neglectus, 815 nigricans, 817 nigrobaccus, 811 notatus, 817 obsessus, 812 occidentalis, 816 odoratus, 816 columbianus, 816 malachophyllus, 816 orarius, 816 ortivus, 817 ostryifolius, 816 paludivagus, 812 parviflorus, 816 triphyllus, 816 pauper, 812 pensilvanicus, 816 pergratus, 816 perinvisus, 817 perpauper, 812 pervarius, 814 philadelphicus, 816 phoenicolasius, 816 plicatifolius, 816 positivus, 816 procerus, 813 prosper, 812 pubescens, 816 pilosifolius, 816 pugnax, 811 recurvans, 814 recurvicaulis, 816 repens, 813 rhodinsulanus, 816 saltuensis, 811 sativus, 811 scambens, 815 semisetosus, 817 sempervirens, 814 setosus, 817 significans, 817 singulus, 817 spiculosus, 814

strigosus, 815 canadensis, 815 tardatus, 817 tetricus, 815 triphyllus, 816 udus, 817 vermontanus, 817 Rudbeckia, 424 bicolor, 425 brittonii, 425 fulgida speciosa, 425 hirta, 424 brittonii, 425 hirta, 425 lanceolata, 425 pulcherrima, 425 laciniata, 425 bipinnata, 425 laciniata, 425 pinnata, 424 subtomentosa, 425 triloba triloba, 425 rue, 829 Rufacer carolinianum, 851 rubrum, 851 rufous bulrush, 170 Rugel’s plantain, 716 Rumex, 740 acetosa, 742 pratensis, 742 acetosella, 740 pyrenaicus, 742 alpinus, 742 altissimus, 742 aquaticus fenestratus, 743 occidentalis, 743 aureus, 743 britannica, 742 crispus, 743 crispus, 743 digyna, 731 ×dissimilis, 743 domesticus, 743 fenestratus occidentalis, 743 hastatulus, 743 longifolius, 743 maritimus, 743 obtusifolius, 743 agrestis, 743 obtusifolius, 743 occidentalis, 743 fenestratus, 743 pallidus, 741 patientia, 742 persicarioides, 744 fueginus, 744 persicarioides, 744 ×pratensis, 742 pulcher, 744 triangulivalvis, 744 verticillatus, 744 violascens, 744 running eared-rockcress, 475 running false stonecrop, 542 running groundsel, 422

i n d e x  9 6 1

Ruppia, 299 maritima, 299 longipes, 299 obliqua, 299 Ruppiaceae, 299 rupturewort, 512 Ruscaceae, 299 rush, 181–189 wood, 190–191 rush American-aster, 438 rushfoil, 565 russet sedge, 149 Russia blue lettuce, 418 Russian iris, 180 Russian-olive, 548 Russian-pigweed, 320 rusty cliff fern, 74 rusty flatsedge, 154 Ruta, 829 graveolens, 829 Rutaceae, 828–829 rye cultivated, 280 grass, 265 wild, 254–255 rye brome, 239 ryegrass, 279

S

Sabatia, 611 amoena, 612 angularis, 612 campanulata, 612 gracilis, 612 campestris, 612 chloroides, 612 dodecandra, 611 dodecandra, 612 kennedyana, 612 gracilis, 612 kennedyana, 612 maculata, 612 maritima, 612 nervosa, 612 simulata, 612 stellaris, 612 Sabina virginiana, 76 Sabulina caroliniana, 513 groenlandica, 514 stricta, 514 Saccharodendron nigrum, 850 Sacciolepis, 279 striata, 279 sacred thorn-apple, 859 safflower-thistle, 385 sage, 647–648 sagebrush, 379 sage willow, 845 Sagina, 516 crassicaulis littoralis, 516 decumbens decumbens, 516 smithii, 516 japonica, 516 littoralis maxima, 516

maxima maxima, 516 nodosa, 516 borealis, 516 glandulosa, 516 nodosa, 516 pubescens, 516 procumbens, 517 compacta, 517 Sagittaria, 89 arifolia, 90 calycina spongiosa, 90 cuneata, 90 eatonii, 90 engelmanniana, 90 filiformis, 90 graminea, 90 teres, 90 latifolia, 90 pubescens, 90 longirostra, 90 lorata, 90 montevidensis, 90 planipes, 90 pubescens, 90 rigida, 90 stagnorum, 90 subulata, 90 gracillima, 90 teres, 89 Sago false pondweed, 298 St. Andrew’s-cross, 626 St. John River oxytrope, 592 St. John’s-wort, 625–626 St. Lawrence grapefern, 65 salad burnet, 817 Salicaceae, 829–848 Salicornia, 327 ambigua, 327 bigelovii, 327 depressa, 327 europaea, 327 maritima, 328 mucronata, 327 prostrata, 328 virginica, 327 saline orache, 320 Salix, 833 alba, 843 caerulea, 843 calva, 843 vitellina, 843 amygdaloides, 844 arctica, 844 arctophila, 844 lejocarpa, 844 arguta pallescens, 848 argyrocarpa, 840 denudata, 844 atrocinerea, 845 aurita, 844 balsamifera, 848 lanceolata, 848 ×bebbii, 846 bebbiana, 844 capreifolia, 844 projecta, 844 ×beschelii, 844

candida, 845 denudata, 845 caprea, 845 chlorophylla pellita, 847 cinerea, 845 atrocinerea, 845 cinerea, 845 oleifolia, 845 coactilis, 848 ×conifera, 846 cordata, 846 abrasa, 846 balsamifera, 848 glaucophylla, 847 missouriensis, 846 myricoides, 847 rigida, 846 ×cryptodonta, 844 cutleri, 848 labradorica, 848 denudata, 847 discolor, 845 eriocephala, 846 elaeagnos, 846 eriocephala, 846 eriocephala, 846 eriocephala, 843 exigua, 843 exterior, 846 interior, 846 pedicellata, 846 falcata, 847 fluviatilis sericans, 846 fragilis, 833 glaucophylla, 847 glaucophylla, 847 gracilis, 847 textoris, 847 ×grayii, 844 grisea, 848 herbacea, 846 humilis, 846 angustifolia, 846 microphylla, 847 tristis, 847 incana, 846 interior exterior, 846 longifolia, 846 pedicellata, 846 ×jesupii, 843 longifolia sericans, 846 wheeleri, 846 lucida, 847 angustifolia, 847 intonsa, 847 latifolia, 847 lucida, 847 lucida, 843 serissima, 848 missouriensis, 846 muehlenbergiana, 847 angustifolia, 846 myricoides, 847 myricoides, 847 myrtilloides hypoglauca, 847 pedicellaris, 847

nigra, 847 brevifolia, 847 falcata, 847 occidentalis, 847 ×peasei, 846 pedicellaris, 847 hypoglauca, 847 tenuescens, 847 pellita, 847 ×pendulina, 848 pentandra, 847 lucida, 847 petiolaris, 847 angustifolia, 847 gracilis, 847 grisea, 848 sericea, 848 subsericea, 847 planifolia, 847 planifolia, 847 purpurea, 848 purpureus, 848 pyrifolia, 848 lanceolata, 848 rostrata, 844 capreifolia, 844 projecta, 844 ×rubella, 845 ×sepulcralis, 843 ×sericans, 845 sericea, 848 subsericea, 847 serissima, 848 sitchensis pellita, 847 starkeana bebbiana, 844 triandra, 833 tristis, 847 glabrata, 846 microphylla, 847 uva-ursi, 848 viminalis, 848 vitellina, 843 sallow sedge, 149 Salpiglossis, 863 sinuata, 863 Salsola, 328 collina, 328 kali caroliniana, 328 kali, 328 pontica, 328 platyphylla, 325 tragus, 328 saltgrass, 251 saltmarsh agalinis, 690 saltmarsh alkali grass, 278 saltmarsh arrow-grass, 192 saltmarsh loosestrife, 661 saltmarsh rush, 188 saltmarsh sand-spurry, 521 saltmarsh stitchwort, 523 saltmarsh tuber-bulrush, 103 salt marsh water-hemp, 316 saltmeadow cordgrass, 282 saltwort, 328

962  ind ex

Salvia, 647 azurea, 647 grandiflora, 648 pitcheri, 648 lyrata, 648 nemorosa, 648 officinalis, 648 pratensis, 648 sclarea, 648 splendens, 648 tiliifolia, 648 verticillata, 648 Salvinia, 69 minima, 69 natans, 69 rotundifolia, 69 Salviniaceae, 69 Sambucus, 307 canadensis, 307 nigra, 307 pubens racemosa, 308 racemosa, 308 pubens, 308 Samolus, 866 floribundus, 866 parviflorus, 866 valerandi parviflorus, 866 sandbar lovegrass, 257–258 sandbar willow, 846 sand blackberry, 813 sandbur, 241 sand dropseed, 285 sand-dune wallflower, 485 sand false heather, 527 sandgrass, 265, 286 sandmat, 568 sand phlox, 723 sandplain agalinis, 689 sandplain yellow flax, 658 sandplant, 513–514 sand plantain, 715 sand-spurry, 521 sand Timothy, 272 sandwort, 508, 512, 514 sandy field blackberry, 812 Sanfoin, 591 Sanguinaria, 702 canadensis, 702 rotundifolia, 702 Sanguisorba, 817 canadensis, 817 minor balearica, 817 muricata, 817 officinalis, 817 polygama, 817 polygama, 817 sanicle, 345–346 Sanicula, 345 canadensis, 345 canadensis, 345 grandis, 345 marilandica, 345 gregaria, 346 marilandica, 345 borealis, 345 canadensis, 345 odorata, 346 trifoliata, 346

Santa Maria feverfew, 422 Santolina, 426 chamaecyparissus, 426 Sapindaceae, 848–852 Saponaria, 517 ocymoides, 517 officinalis, 517 pumilio, 517 vaccaria, 523 Sarothra gentianoides, 626 Sarracenia, 852 purpurea purpurea, 852 Sarraceniaceae, 852 sarsaparilla, 339 sassafras, 83 Sassafras, 83 albidum, 83 variifolium, 83 satiny willow, 847 Satureja, 649 acinos, 636 hortensis, 649 vulgaris, 636 neogaea, 636 Satyrium repens, 204 Saururaceae, 86 Saururus, 86 cernuus, 86 Savastana hirta, 231 odorata, 231 savory, 649 Savoy hawkweed, 412 saw-sedge, 151 saw-toothed sunflower, 407 Saxifraga, 854 aizoides, 854 cernua, 854 foliolosa, 853 forbesii, 853 oppositifolia, 854 oppositifolia, 855 paniculata, 855 neogaea, 855 pensylvanica, 853 forbesii, 853 purpuripetala, 853 rivularis, 855 debilis, 855 flexuosa, 855 rivularis, 855 stellaris comosa, 853 virginiensis, 853 Saxifragaceae, 852–855 saxifrage, 345, 853–855 saxifrage childing-pink, 515 Scabiosa, 503 arvensis, 500 australis, 503 columbaria, 503 gigantea, 499 ochroleuca, 503 succisa, 503 scabrous black sedge, 145 scabrous hawthorn, 787

Scandix, 346 cerefolium, 338 odorata, 343 pecten-veneris, 346 scarlet bean, 592 scarlet bee-balm, 644 scarlet hawthorn, 783 scarlet lychnis, 513 scarlet morning-glory, 536 scarlet oak, 605 scarlet painted-cup, 692 scarlet pimpernel, 673 scarlet sage, 648 scarlet smartweed, 734 scentless-chamomile, 445 scentless mock-orange, 623 Sceptridium dissectum, 64 multifidum, 65 oneidense, 65 rugulosum, 65 Schedonorus, 279 arundinaceus, 279 giganteus, 279 pratensis, 279 Schenkia, 612 spicata, 612 Scheuchzeria, 301 americana, 301 palustris, 301 americana, 301 Scheuchzeriaceae, 301 Schizachne, 279 purpurascens, 279 Schizachyrium, 279 littorale, 279 scoparium, 279 scoparium, 280 Schizonotus sorbifolius, 818 Schkuhria, 426 pinnata pinnata, 426 Schoenoplectus, 165 acutus, 166 acutus, 166 americanus, 166 ×contortus, 166 etuberculatus, 166 fluviatilis, 103 glaucus, 103 hallii, 166 heterochaetus, 167 lacustris acutus, 166 tenuiculmis, 167 maritimus, 103 novae-angliae, 103 ×oblongus, 166 occidentalis, 166 pungens pungens, 167 purshianus, 167 purshianus, 167 williamsii, 167 robustus, 103 smithii, 167 levisetus, 167 setosus, 167 smithii, 167 ×steinmetzii, 167

subterminalis, 167 tabernaemontani, 167 torreyi, 168 Schoenus fuscus, 164 Schreber’s wood-aster, 400 Schwalbea, 695 americana, 695 australis, 695 australis, 695 Schweinitz’s flatsedge, 155 Schweinitz’s sedge, 150 Scilla siberica, 175 Scirpinae, 145 Scirpus, 168 acutus, 166 americanus olneyi, 166 ancistrochaetus, 168 atrocinctus, 168 atrovirens, 168 georgianus, 169 autumnalis, 162 capillaris, 104 cespitosus, 171 callosus, 171 delicatulus, 171 clintonii, 171 cylindricus, 103 cyperinus brachypodus, 169 pedicellatus, 170 pelius, 169 debilis, 167 equisetoides, 159 etuberculatus, 166 expansus, 169 fluviatilis, 103 georgianus, 169 glaucus, 103 hallii, 166 hattorianus, 170 hudsonianus, 171 juncoides williamsii, 167 lineatus, 170 longii, 170 maritimus, 103 fluviatilis, 103 macrostachyus, 103 micranthus, 163 microcarpus, 170 rubrotinctus, 170 novae-angliae, 103 obtusa, 160 paludosus, 103 palustris, 160 ×peckii, 169 pedicellatus, 170 pendulus, 170 planifolius, 171 polyphyllus, 170 purshianus, 167 williamsii, 167 quinquefolius, 161 rostellatus, 161 rubrotinctus, 170 smithii, 167 levisetus, 167 setosus, 167

i n d e x  9 63

Scirpus, cont. uniglumis, 161 verecundus, 171 Scleranthus, 517 annuus, 517 perennis, 517 Scleria, 170 flaccida nitida, 171 pauciflora, 170 caroliniana, 171 kansana, 170 pauciflora, 171 reticularis, 171 triglomerata, 171 verticillata, 171 sclerolepis, 426 Sclerolepis, 426 uniflora, 426 verticillata, 426 Scleropoa rigida, 245 scorpion’s tail, 593 scorpion-weed, 465–466 Scorpiurus, 593 muricatus, 593 Scorzoneroides, 426 autumnalis, 426 autumnalis, 426 pratensis, 426 Scotch bellflower, 495 Scotch broom, 579 Scotch cotton-thistle, 421 Scotch elm, 868 Scotch heath, 554 Scotch pine, 79 Scotch rose, 807 Scotch wild lovage, 343 scouring-rush, 60 scratch bedstraw, 824 screwstem, 608–609 Scrophularia, 856 lanceolata, 856 leporella, 856 marilandica, 856 nodosa, 856 pectinata, 856 Scrophulariaceae, 855–857 scrub oak, 605 Scutellaria, 649 altissima, 649 ambigua, 650 australis, 650 churchilliana, 649 epilobiifolia, 649 galericulata, 649 epilobiifolia, 649 pubescens, 649 incana hispida, 649 integrifolia, 649 lateriflora, 649 leonardii, 650 nervosa ambigua, 650 parvula, 650 australis, 650 leonardii, 650 missouriensis, 650 parvula, 650 seabeach dock, 744

seabeach orache, 319 sea-beach sedge, 124 sea-blite, 329 seacliff eyebright, 693 sea coast Angelica, 338 sea-coast tuber-bulrush, 103 sea glasswort, 328 sea-heath, 607 sea-lavender, 721 sea-milkwort, 674 sea-purslane, 311 sea-rocket, 479 seaside alkali grass, 278 sea side amaranth, 317 seaside bluebells, 463 seaside brookweed, 866 seaside crowfoot, 758 seaside goldenrod, 433 seaside heliotrope, 462 seaside knotweed, 739 seaside plantain, 716 seaside sandmat, 568 seaside-sandwort, 512 seaside threeawn, 233 seaside toad rush, 186 Secale, 280 cereale, 280 Securigera, 593 varia, 593 sedge, 113–171 Sedum, 542 acre, 543 aizoon, 540 alboroseum, 541 album, 543 anopetalum, 543 ellacombeanum, 540 erythrostictum, 541 fabaria, 541 hispanicum, 543 kamtschaticum, 540 ellacombeanum, 540 ochroleucum, 543 purpurascens, 541 reflexum, 543 roanense, 542 roseum, 542 roanense, 542 sarmentosum, 543 sexangulare, 543 spectabile, 541 spurium, 542 stoloniferum, 542 telephium, 541 fabaria, 541 ternatum, 543 seep monkey-flower, 705 Selaginella, 48 apoda, 49 eclipes, 49 rupestris, 49 selaginoides, 49 Selaginellaceae, 48–49 selfheal, 646 Selinum, 346 carvifolia, 346 Sempervivum, 543 tectorum, 543 Senebiera didyma, 487 Seneca milkwort, 726

Senecio, 426 aureus, 422 gracilis, 422 intercursus, 422 balsamitae, 422 gaspensis, 422 gracilis, 422 jacobaea, 414 multicapitatus, 427 obovatus, 422 rotundus, 422 pauperculus, 422 balsamitae, 422 praelongus, 422 robbinsii, 422 rotundus, 422 spartioides, 427 multicapitatus, 427 suaveolens, 426 sylvaticus, 427 viscosus, 427 vulgaris, 427 senna, 594 Senna, 593 hebecarpa, 594 obtusifolia, 594 sensitive fern, 61 sensitive-pea, 578 Serapias atrorubens, 203 Sericocarpus, 427 asteroides, 427 linifolius, 427 serpentaria, 81 serpentine sandplant, 514 Serpicula verticillata, 176 Serratula arvensis, 389 sesame, 703 Sesamum, 703 indicum, 703 orientale, 703 Sesban exaltatus, 594 Sesbania, 594 exaltata, 594 herbacea, 594 macrocarpa, 594 sessile-fruited arrowhead, 90 sessile-headed blazing star, 417 sessile-leaved bellwort, 99 sessile-leaved tick-trefoil, 581 Sesuvium, 311 maritimum, 311 pentandrum, 311 Setaria, 280 faberi, 280 geniculata, 280 glauca, 281 italica, 280 parviflora, 280 pumila, 281 verticillata, 281 viridis, 281 breviseta, 281 viridis, 281 weinmannii, 281 setose blackberry, 817 seven-angled pipewort, 172

several-veined sweetflag, 87 Sexilia verticillata, 726 shadbush, 769–772 shaded hawthorn, 788 shagbark hickory, 629 sharp-fruited rush, 186 sharp-leaved cancerwort, 711 sharp-lobed hepatica, 751 sharp-scaled manna grass, 261 sharp-toothed nodding-aster, 420 shaved sedge, 114 sheathed sedge, 137 sheep American-laurel, 556 sheepbit, 495 sheepburr, 462 sheep dock, 742 sheep fescue, 259 shellbark hickory, 628 Shepherdia, 548 canadensis, 548 shepherd’s cress, 492 shepherd’s-needle, 346 shepherd’s-purse, 479 Sherardia, 828 arvensis, 828 Sherard’s downy rose, 807 shingle oak, 605 shining firmoss, 40 shining flatsedge, 153 shining ladies’-tresses, 212 shining rose, 807 shining willow, 847 shinleaf, 557–558 shiny wedgescale, 283 shoreweed, 712 short-awned meadow-foxtail, 229 short-awned mountain-rice grass, 273 short-beaked beaksedge, 164 short-beaked sedge, 123 short-fringed knapweed, 387 Shortia, 546 galacifolia galacifolia, 546 short-spurred corydalis, 700 short-stalked false bindweed, 532 short-tailed rush, 187 showy baby’s-breath, 511 showy blackberry, 813 showy coneflower, 425 showy false goldeneye, 408 showy goldenrod, 434 showy lady’s-slipper, 203 showy mountain-ash, 819 showy orchid, 203 showy orpine, 541 showy tick-trefoil, 580 shrubby-cinquefoil, 788 shrubby rose-mallow, 664 shrubby St. John’s-wort, 626 shrub yellowroot, 761 shy wallflower, 485 sibbaldia, 818 Sibbaldia, 818 procumbens, 818 Sibbaldiopsis, 818 tridentata, 818

96 4  i n de x

Siberian crab apple, 793 Siberian crane’s-bill, 616 Siberian elm, 868 Siberian iris, 180 Siberian orache, 320 Siberian pea shrub, 577 Siberian spring-beauty, 745 Siberian squill, 175 sickle-leaved silk-grass, 423 sicklepod rockcress, 477 sickle-pod wild senna, 594 sickleweed, 341 Sicyos, 545 angulatus, 545 lobata, 545 Sida, 667 callimorpha, 667 hermaphrodita, 667 picta, 663 spinosa, 667 sideoats grama, 234 Sideritis, 650 ciliata, 637 montana, 650 Siebold’s arrowwood, 311 Sigesbeckia, 427 orientalis, 427 Silene, 517 acaulis, 519 arctica, 519 exscapa, 519 alba, 519 anglica, 519 antirrhina, 519 confines, 519 deameana, 519 divaricata, 519 laevigata, 519 armeria, 508 caroliniana, 519 pensylvanica, 519 chalcedonica, 513 coeli-rosa, 511 conica, 519 conica, 519 coronaria, 513 csereii, 519 cucubalus, 520 dichotoma, 518 dioica, 519 flos-cuculi, 513 gallica, 519 hampeana, 519 inflata, 520 latifolia, 519 alba, 519 nivea, 518, 520 noctiflora, 520 nutans, 520 pendula, 520 pensylvanica, 520 pratensis, 520 pumilio, 517 stellata, 520 scabrella, 520 viscaria, 523 vulgaris, 520 vulgaris, 520 silk-grass, 423 silk-tree, 575 silkvine, 350

silky dogwood, 538 silky willow, 848 Silphium, 428 perfoliatum perfoliatum, 428 silverbell, 866 silvergrass, 266 silver-hairgrass, 228 silver-leaved cinquefoil, 796 silver maple, 851 silverpuffs, 418 silverweed, 772 silvery false spleenwort, 73 silvery-flowered sedge, 123 silvery whitlow-wort, 515 Silybum, 428 marianum, 428 Simaroubaceae, 857 simple-stemmed bur-reed, 303 Sinapis, 491 alba, 491 arvensis, 491 juncea, 478 nigra, 478 single-flowered medick, 590 single-veined sweetflag, 87 Sisymbrium, 491 alliaria, 474 altissimum, 492 amphibium, 491 gallicum, 485 humile, 479 irio, 492 loeselii, 492 officinale, 492 leiocarpum, 492 orientale, 492 sylvestre, 491 thalianum, 475 Sisyrinchium, 180 albidum, 181 angustifolium, 181 arenicola, 181 atlanticum, 181 bermudianum albidum, 181 crebrum, 181 fuscatum, 181 graminoides, 181 intermedium, 181 montanum, 181 crebrum, 181 montanum, 181 mucronatum, 181 scabrellum, 181 Sitka ground-cedar, 45 Sium, 346 carsonii, 346 cicutifolium, 346 sisarum, 346 suave, 346 six-angled stonecrop, 543 skullcap, 649–650 skunk-cabbage, 97 skunk currant, 618 skyrocket, 722 sleepy campion, 519 slender amaranth, 317 slender beadgrass, 271 slender bladderwort, 657 slender blue iris, 180

slender bulrush, 167 slender bunch-flower, 198 slender bush-clover, 588 slender cliff-brake, 68 slender cottonsedge, 162 slender cotton-weed, 326 slender crabgrass, 251 slender false bog-clubmoss, 47 slender fimbry, 162 slender goldenrod, 432 slender grama, 234 slender hair grass, 244 slender knotweed, 740 slender ladies’-tresses, 211 slender leafy spurge, 569 slender-leaved agalinis, 690 slender-leaved sundew, 547 slender loose-flowered sedge, 133 slender meadow-foxtail, 229 slender muhly, 268 slender rock-brake, 68 slender rose-gentian, 612 slender saltwort, 328 slender soft brome, 238 slender spikesedge, 161 slender three-seeded-Mercury, 564 slender water-milfoil, 621 slender waternymph, 177 slender wedgescale, 283 slender wild-rye, 255 slender woodland sedge, 116 slender wood-reed, 242 slender yellow-eyed-grass, 305 slender yellow wood sorrel, 697 slim-fruited rush, 187 slim-lobed beggar-ticks, 383 slippery elm, 869 slough grass, 234 small bayberry, 671 small beggar-ticks, 382 small-bristled hound’s-tongue, 460 small bugloss, 459 small carp grass, 233 small cranberry, 563 small dropseed, 285 small enchanter’s-nightshade, 681 smaller forget-me-not, 464 small-flowered agalinis, 690 small-flowered bee-blossom, 686 small-flowered bitter-cress, 482 small-flowered blue-eyed Mary, 709 small-flowered crane’s-bill, 616 small-flowered crowfoot, 759 small-flowered dwarf-bulrush, 163 small-flowered eveningprimrose, 687 small-flowered hairy willowherb, 683 small-flowered-saxifrage, 853 small-flowered sweet-briar rose, 806 small-flowered tamarisk, 866 small-flowered wood rush, 191

small-flowered woolly bean, 594 small-fruited spikesedge, 160 small-headed rush, 187 small-headed sunflower, 407 small-leaved linden, 668 small-leaved pond-lily, 85 small pondweed, 297 small pussytoes, 374 small-seeded false flax, 479 small smartweed, 735 small-spiked false nettle, 869 small sweet grass, 231 Small’s yellow-eyed-grass, 305 small waterwort, 549 small white American-aster, 442 small whorled pogonia, 205 smartweed, 731–736 smartweed dodder, 535 Smilacaceae, 301 Smilacina racemosa, 300 racemosa, 300 stellata, 300 trifolia, 300 Smilax, 301 glauca, 301 leurophylla, 301 herbacea, 301 hispida, 301 tamnoides, 301 rotundifolia, 301 crenulata, 301 quadrangularis, 301 tamnoides, 301 Smith’s blackberry, 815 Smith’s bulrush, 167 smoketree, 329 smooth alder, 451 smooth American-aster, 440 smooth arrowwood, 309 smooth beggar-ticks, 383 smooth blackberry, 813 smooth blackhaw, 311 smooth black sedge, 140 smooth brome, 238 smooth cat’s-ear, 413 smooth cliff fern, 74 smooth cordgrass, 282 smooth crabgrass, 251 smooth Dove’s-foot crane’sbill, 615 smooth false foxglove, 691 smooth forked whitlow-wort, 515 smooth goldenrod, 432 smooth hawk’s-beard, 393 smooth hawthorn, 785 smooth rockcress, 477 smooth rose, 805 smooth rupturewort, 512 smooth saw-sedge, 151 smooth shadbush, 771 smooth-sheathed sedge, 150 smooth small-leaved ticktrefoil, 581 smooth spiderwort, 100 smooth sumac, 330 smooth tick-trefoil, 580 smooth white violet, 884

in d e x  9 6 5

smooth whitlow-mustard, 484 smooth winterberry, 352 smotherweed, 320 snail-seed pondweed, 294, 297 snakeroot, 372 snapdragon, 707–708, 712 sneezeweed, 388, 404 snowball, 311 snow-bed willow, 846 snowberry, 503 snowdrop, 92 snowflake, 92 snow wakerobin, 197 snowy campion, 520 soapwort, 517 soft brome, 238 soft cinquefoil, 797 soft downy rose, 806 soft-haired false gromwell, 464 soft-leaved sedge, 125 soft-stemmed bulrush, 167 Solanaceae, 857–865 Solanum, 863 americanum, 865 carolinense, 863 carolinense, 864 citrullifolium, 863 citrullifolium, 864 dulcamara, 864 dulcamara, 864 villosissimum, 864 lycopersicon, 864 lycopersicon, 865 melongena, 865 nigrum, 864 nigrum, 865 physalifolium, 864 nitidibaccatum, 865 pseudocapsicum, 865 ptycanthum, 865 rostratum, 865 sarrachoides, 865 sisymbriifolium, 865 triflorum, 865 tuberosum, 865 villosum, 865 Solidago, 428 aestivalis, 431 altissima, 431 altissima, 431 procera, 431 arguta, 428 arguta, 431 aspera, 433 asperula, 433 asteroides, 421 axillaris, 431 bicolor, 431 concolor, 432 lantana, 432 caesia, 430 axillaris, 431 caesia, 431 calcicola, 432 canadensis, 431 canadensis, 431 hargeri, 431 scabra, 431 celtidifolia, 433 conferta, 434 cutleri, 432

elliottii, 432 ascendens, 432 pedicellata, 432 erecta, 432 flexicaulis, 432 gigantea, 432 serotina, 432 glutinosa randii, 433 graminifolia major, 401 septentrionalis, 401 hispida, 430 hispida, 432 juncea, 432 neobohemica, 432 ramosa, 432 lanceolata, 401 latifolia, 432 latissimifolia, 432 leiocarpa, 432 macrophylla, 432 thyrsoidea, 432 mexicana, 433 neglecta, 434 nemoralis, 428 haleana, 432 nemoralis, 432 odora, 429 odora, 432 patula, 428 patula, 432 ptarmicoides, 421 puberula, 430, 432 puberula, 432 purshii, 434 racemosa, 433 randii, 433 rigida, 421 rugosa, 433 aspera, 433 celtidifolia, 433 rugosa, 431 sphagnophila, 431 villosa, 433 sempervirens, 433 mexicana, 433 sempervirens, 433 simplex monticola, 433 racemosa, 433 randii, 433 speciosa, 434 erecta, 432 speciosa, 434 squarrosa, 434 suaveolens, 432 tenuifolia, 401 pycnocephala, 401 uliginosa, 434 linoides, 434 neglecta, 434 ulmifolia, 434 ulmifolia, 434 uniligulata, 434 virgaurea alpina, 432 solid-stemmed burnetsaxifrage, 345 Solomon’s seal, 300 somewhat-tailed rush, 189

Sonchus, 434 arvensis, 434 arvensis, 434 glabrescens, 434 asper, 434 oleraceus, 434 asper, 434 Sophora kentukea, 578 Sorbaria, 818 sorbifolia, 818 ×Sorbaronia, 773 fallax, 773 hybrida, 773 Sorbus, 818 alnifolia, 818 americana, 818 arbutifolia, 773 atropurpurea, 773 aucuparia, 818 decora, 819 groenlandica, 819 fennica, 819 hybrida, 819 Sorengia, 281 longifolia, 281 elongata, 281 longifolia, 281 rigidula, 281 Sorghastrum, 281 nutans, 281 sorghum, 282 Sorghum, 281 bicolor, 282 bicolor, 282 drummondii, 282 sudanense, 282 drummondii, 282 halepense, 282 vulgare, 282 technicum, 282 sorrel, 697 sour cherry, 800 South American false velvetleaf, 667 southeastern wild-rye, 254 southern agrimony, 768 southern bellflower, 495 southern blackberry, 812 southern catalpa, 455 southern fragile fern, 73 southern ground-cedar, 45 southern lady fern, 71 southern long-awned wood grass, 235 southern succisella, 503 southern twayblade, 206 southern water-plantain, 88 southern wild rice, 288 southern woodland violet, 883 southern wormwood, 378 southwestern cosmos, 392 sow-thistle, 434 soybean, 582 Spanish chestnut, 601 Spanish needles beggar-ticks, 382 Spanish stonecrop, 543 Sparganium, 302 acaule, 303 americanum, 303

androcladum, 303 angustifolium, 304 chlorocarpum, 303 emersum, 303 acaule, 303 emersum, 303 eurycarpum, 303 greenei, 303 fluctuans, 304 lucidum, 303 minimum, 304 natans, 304 sparse-flowered sedge, 127 Spartina, 282 alterniflora, 282 glabra, 282 ×caespitosa, 283 cynosuroides, 282 polystachya, 282 patens, 282 monogyna, 282 pectinata, 283 suttiei, 283 Spartium scoparium, 579 Spathulata spuria, 542 Spathyema foetidum, 97 Spatularia foliolosa, 853 spatulate-leaved sundew, 547 spearmint, 643 spearscale orache, 319 speckled alder, 451 Specularia perfoliata, 497 speedwell, 716–720 Spergella nodosa, 516 Spergula, 520 arvensis, 520 sativa, 520 morisonii, 520 pentandra, 520 sativum, 520 Spergularia, 520 canadensis, 521 canadensis, 521 leiosperma, 521 marina, 521 media, 521 rubra, 521 perennans, 521 Sphaeralcea, 667 mendocina, 667 munroana, 667 Sphenopholis, 283 intermedia, 283 nitida, 283 obtusata, 283 lobata, 283 pensylvanica, 283 spicebush, 83 spicy-wintergreen, 555 spider-flower, 530 spiderwort, 100 spike bentgrass, 227 spike burgrass, 286 spiked centaury, 612 spiked wood rush, 191

96 6   in d e x

spikemoss, 49 spike muhly, 268 spikenard, 339 spikesedge, 159–161 Spinacea, 328 oleracea, 328 spinach, 328 spindle-tree, 524–525 spineflower, 727 spineless hornwort, 83 spinulose wood fern, 55 Spinulum, 47 annotinum, 48 canadense, 48 spiny amaranth, 317 spiny cocklebur, 446 spiny-leaved sow-thistle, 434 spiny plumeless-thistle, 384 spiny-spored quillwort, 41 spiny threecorner-Jack, 728 spiraea, 818 Spiraea, 819 alba, 819 latifolia, 819 septentrionalis, 819 chamaedryfolia, 820 ulmifolia, 820 japonica, 819 fortunei, 820 latifolia, 819 septentrionalis, 819 opulifolia, 793 prunifolia, 820 plena, 820 septentrionalis, 819 thunbergii, 820 tomentosa, 820 ulmaria, 788 ulmifolia, 820 ×vanhouttei, 820 Spiranthes, 210 casei casei, 211 cernua, 211 ×eamesii, 212 gracilis, 211 ×intermedia, 212 lacera, 211 gracilis, 211 lacera, 211 lucida, 212 ochroleuca, 212 romanzoffiana, 212 tuberosa, 212 vernalis, 212 Spirodela, 97 polyrrhiza, 97 punctata, 95 spleen amaranth, 316 spleenwort, 51–52, 73 split-lipped hemp-nettle, 637 spongeplant, 176 spongy-leaved arrowhead, 90 Sporobolus, 283 asper clandestinus, 284 pilosus, 284 clandestinus, 284

compositus, 284 clandestinus, 284 compositus, 284 drummondii, 284 contractus, 285 cryptandrus, 285 strictus, 285 drummondii, 284 heterolepis, 285 neglectus, 285 serotinus, 268 uniflorus, 268 vaginiflorus, 284 inaequalis, 285 neglectus, 285 vaginiflorus, 285 spot-leaved crowfoot, 759 spotted bee-balm, 644 spotted bellflower, 495 spotted coral-root, 201 spotted crane’s-bill, 616 spotted-duck-meal, 95 spotted hawkweed, 412 spotted henbit, 639 spotted Joe-Pye weed, 402 spotted knapweed, 388 spotted medick, 590 spotted phlox, 724 spotted pondweed, 297 spotted prince’s-pine, 553 spotted sandmat, 568 spotted St. John’s-wort, 626 spotted touch-me-not, 448 spotted water-hemlock, 340 sprangletop, 264 spreading alkali grass, 278 spreading bellflower, 495 spreading cotoneaster, 774 spreading dogbane, 348 spreading Jacob’s-ladder, 725 spreading pellitory, 870 spreading sedge, 116 spreading-sneezeweed, 388 spreading wallflower, 485 spring avens, 792 spring-beauty, 745 spring cinquefoil, 798 spring corydalis, 700 spring forget-me-not, 464 spring ladies’-tresses, 212 spring vetch, 599 spring whitlow-mustard, 484 spruce, 78 spurge, 493, 567–569 spurred-gentian, 611 spurry, 520 square-pod water-primrose, 684 square-stemmed spikesedge, 161 Squarrosae, 145–146 squarrose goldenrod, 434 squarrose sedge, 145 squash, 545 squashberry, 310 squaw-root, 692 squill, 175 squirrel-corn, 701 Stachys, 650 annua, 651 arenicola, 652

arvensis, 651 borealis, 651 brevidens, 652 byzantina, 651 clingmanii, 651 germanica, 651 hispida, 651 homotricha, 651 hyssopifolia, 651 lanata, 651 lythroides, 651 olympica, 651 palustris, 651 arenicola, 652 elliptica, 651 hispida, 651 homotricha, 651 pilosa, 652 puberula, 652 segetum, 651 pilosa, 651 pilosa, 652 arenicola, 652 tenuifolia hispida, 651 staggerbush, 557 staghorn sumac, 330 stalked burgrass, 286 stalked water-horehound, 640 stalked woolsedge, 170 stalkless yellow-cress, 491 standing bugle, 635 standing-cypress skyrocket, 722 Standley’s goosefoot, 324 Staphylea, 865 trifolia, 865 Staphyleaceae, 865 starburr, 371 starch grape-hyacinth, 174 starflower, 674 star-grass, 178 star-like false Solomon’s-seal, 300 star-of-Bethlehem, 175, 193 starry campion, 520 star sedge, 143, 147 starved rosette-panicgrass, 248 Stegia trimestris, 665 Steironema ciliatum, 674 hybridum, 674 lanceolata, 674 longifolium, 674 quadriflorum, 674 revolutum, 674 Stellaria, 521 alsine, 522 apetala, 523 aquatica, 514 borealis, 522 borealis, 522 floribunda, 522 isophylla, 522 calycantha floribunda, 522 laurentiana, 522 corei, 522

graminea, 522 latifolia, 522 holostea, 523 humifusa, 523 longifolia, 522 atrata, 523 eciliata, 523 longifolia, 523 media, 521, 523 procera, 523 pubera sylvatica, 522 sylvatica, 522 tennesseensis, 522 uliginosa, 522 Stellulatae, 146–147 Stenanthium, 196 gramineum, 196 Stenophyllus capillaris, 104 Stephen’s stork’s-bill, 614 stickleaf mentzelia, 659 stickseed, 461 sticky chickweed, 510 sticky false asphodel, 302 sticky nightshade, 865 sticky ragwort, 427 sticky-stemmed pearlwort, 516 stiff-aster, 413 stiff dwarf-gentian, 610 stiff flat-topped-goldenrod, 421 stiff-leaved rosette-panicgrass, 249 stiff sunflower, 407 stiff three-petaled bedstraw, 826 stiff tickseed, 391 stiff tick-trefoil, 581 stiltgrass, 265 stinging nettle, 870 stinking bee-plant, 529 stinking chamomile, 375 stinking hawk’s-beard, 393 stinking hawthorn, 782 stinking lovegrass, 257 stinking pepperweed, 487 stinkwort, 394 Stipa avenacea, 274 canadensis, 273 stitchwort, 522–523 Stomoisia cornuta, 656 stonecrop, 540, 542–543, 703 Stone’s hawthorn, 787 stork’s bill, 613–614 stout dotted smartweed, 735 straggling St. John’s-wort, 626 straight-leaved pondweed, 297 strapwort, 510 strawberry, 789 strawberry-blite, 323 strawberry clover, 597 strawberry ground-cherry, 862 strawberry raspberry, 815 straw-colored flatsedge, 155 strawflower, 447 straw foxglove, 710 straw sedge, 123–124

i n d e x  9 67

Streptopus, 194 amplexifolius, 195 lanceolatus, 195 ×oreopolis, 195 roseus longipes, 195 perspectus, 195 striate campion, 519 strict blue-eyed-grass, 181 strict eyebright, 693 stringy stonecrop, 543 striped bladderwort, 656 striped cream violet, 886 striped maple, 851 striped toadflax, 712 Strobus strobus, 79 Strophocaulos arvensis, 532 Strophostyles, 594 helvola, 594 leiosperma, 594 umbellata, 594 Stuckenia, 298 filiformis, 298 filiformis, 298 occidentalis, 298 pectinata, 298 Styracaceae, 866 Suaeda, 329 americana, 329 calceoliformis, 329 linearis, 329 maritima, 329 americana, 329 maritima, 329 richii, 329 richii, 329 Subularia, 492 aquatica, 492 americana, 492 Succisa, 503 pratensis, 503 succisella, 503 Succisella, 503 inflexa, 503 sugar maple, 851 sulphur cinquefoil, 797 sulphur cosmos, 392 sultan, 372 sumac, 330 summer-cypress, 326 summer grape, 889 summer savory, 649 summer sedge, 130 summer snowflake, 92 suncup, 680 sundew, 547 sundial lupine, 588 sunflower, 406–408 sunflower-everlasting, 408 sun spurge, 568 swallowwort, 350 swamp crowfoot, 758 swamp deciduous dog-laurel, 554 swamp dock, 744 swamp honeysuckle, 502 swamp-loosestrife, 660 swamp lousewort, 695 swamp milkweed, 349

swamp moonwort, 65 swamp poplar, 832 swamp pricklegrass, 242 swamp red currant, 619 swamp rose, 807 swamp rose-mallow, 664 swamp small-floweredsaxifrage, 853 swamp thistle, 390 swamp wedgescale, 283 swamp white oak, 604 swamp wrinkle-leaved goldenrod, 431 swamp yellow-loosestrife, 674 Swan’s sedge, 144 swarthy sedge, 123 sweet-alyssum, 488 sweet basil, 645 sweet-bay, 84 sweet-briar rose, 806–807 sweet cherry, 800 sweet-cicely, 344 sweet-clover, 591 sweet-coltsfoot, 423 sweet coneflower, 425 sweet fennel, 341 sweet-fern, 671 sweetflag, 87 sweetgale, 672 sweet grass, 231 sweet-gum, 311 sweetleaf, 866 sweet marjoram, 645 sweet mock-orange, 623 sweet-pepperbush, 530 sweet-scented bedstraw, 826 sweet-scented camphorweed, 423 sweet-shoot bamboo, 273 sweetshrub, 82 sweet sultan, 372 sweet-vetch, 582 sweet vetchling, 584 sweet white violet, 881 sweet-William-catchfly, 508 sweet William pink, 511 sweet wood-reed, 242 Swertia albicaulis, 609 deflexa, 611 rotata, 611 Swida, 538 alternifolia, 538 amomum, 538 schuetzeana, 539 arnoldiana, 539 racemosa, 539 rugosa, 539 sanguinea, 539 sericea, 539 slavinii, 539 swine-cress, 487–488 swinging sedge, 141 switch panicgrass, 271 swollen-beaked sedge, 150 swollen bladderwort, 656 sword-leaved rush, 188 sword-like bog-mat, 97 sycamore, 720 sycamore maple, 851

Symphoricarpos, 503 albus, 503 albus, 503 laevigatus, 503 orbiculatus, 503 racemosus, 503 Symphyotrichum, 435 amethystinum, 439 anticostense, 438 boreale, 438 ciliatum, 438 ciliolatum, 438 concolor, 437 concolor, 439 cordifolium, 440 dumosum, 439 dodgei, 439 strictior, 439 subulifolium, 439 ericoides, 436 ericoides, 439 firmum, 442 frondosum, 439 laeve, 440 laeve, 440 lanceolatum, 438 interior, 440 lanceolatum, 440 latifolium, 440 lateriflorum, 440 lowrieanum, 440 novae-angliae, 440 novi-belgii, 440 elodes, 441 novi-belgii, 441 villicaule, 441 ontarionis, 441 glabratum, 441 patens, 441 phlogifolium, 441 pilosum, 441 pilosum, 441 pringlei, 441 praealtum, 441 angustior, 441 prenanthoides, 442 puniceum, 436 puniceum, 442 racemosum, 442 simplex, 440 subulatum, 435 subulatum, 442 ×tardiflorum, 439 tenuifolium, 437 tenuifolium, 442 tradescantii, 438 undulatum, 442 urophyllum, 442 Symphytum, 466 asperrimum, 467 asperum, 467 officinale, 467 tanaicense, 467 tuberosum, 467 uliginosum, 467 uplandicum, 467 Symplocaceae, 866 Symplocarpus, 97 foetidus, 97

Symplocos, 866 chinensis, 866 paniculata, 866 Syndesmon thalictroides, 761 Synosma suaveolens, 427 Syrian-mustard, 485 Syringa, 679 amurensis, 679 japonica, 679 japonica, 679 josikaea, 679 ×persica, 679 reticulata, 679 reticulata, 679 villosa, 679 vulgaris, 679 protolaciniata, 679

T tacky scorpion-weed, 466 Taeniatherum, 285 caput-medusae caput-medusae, 285 Taenidia, 346 integerrima, 346 Tagetes, 442 erecta, 443 major, 443 minuta, 443 papposa, 394 patula, 443 tall baby’s-breath, 512 tall beaksedge, 164 tall beggar-ticks, 383 tall blue lettuce, 415 tall bush-clover, 588 tall cottonsedge, 162 tall crowfoot, 757 tall goldenrod, 431 tall hairy lettuce, 415 tall hawkweed, 412 tall hedge-mustard, 492 tall larkspur, 754 tall lettuce, 415 tall meadow-rue, 760 tall oat grass, 233 tall rattlesnake-root, 419 tall river-hemp, 594 tall rye grass, 279 tall scouring-rush, 60 tall skullcap, 649 tall sunflower, 407 tall swamp rosette-panicgrass, 250 tall thistle, 389 tall thoroughwort, 399 tall tickseed, 392 tall white-aster, 394 tall windflower, 752 tall wood-beauty, 788 Tamaricaceae, 866 tamarisk, 866 Tamarix, 866 gallica, 866 parviflora, 866 tetrandra, 866

96 8   in de x

Tanacetum, 443 balsamita, 379 bipinnatum, 443 huronense, 443 huronense, 443 johannense, 443 parthenium, 443 serotinum, 416 suaveolens, 418 vulgare, 443 tansy, 443 tansy-mustard, 483 tansy ragwort, 414 tape-grass, 177 taper-tipped flatsedge, 153 taravine, 307 Taraxacum, 443 erythrospermum, 444 laevigatum, 444 latilobum, 444 officinale, 444 vulgare, 444 palustre, 444 turfosum, 444 vulgare, 444 Tarenaya, 530 hassleriana, 530 tarplant, 417 Tatarian aster, 379 Tatarian honeysuckle, 502 Tatarian maple, 851 Tatarian orache, 320 tawny cottonsedge, 162 Taxaceae, 80 Taxus, 80 baccata, 80 minor, 80 canadensis, 80 canadensis, 80 minor, 80 cuspidata, 80 tea-leaved willow, 847 tearthumb, 734–735 teasel, 500 Tecoma radicans, 455 teel lovegrass, 258 Teesdalia, 492 nudicaulis, 492 Teloxys ambrosioides, 326 botrys, 326 graveolens, 326 multifida, 326 pumilio, 326 Tennessee stitchwort, 522 Tephrosia, 595 latidens, 595 virginiana, 595 glabra, 595 holosericea, 595 terrestrial water-starwort, 708 Tetragonanthus deflexus, 611 Tetragonia, 311 tetragonioides, 311 Teucrium, 652 boreale, 652 botrys, 652

canadense, 652 angustatum, 652 boreale, 652 canadense, 652 littorale, 652 occidentale, 652 virginicum, 652 occidentale, 652 scorodonia, 652 Texas croton, 565 Texas grama, 234 Texas liverseed grass, 287 Texas palafox, 422 thale-cress, 475 Thalesia uniflorum, 694 Thalictrodes racemosum, 750 Thalictrum, 759 confine, 761 dasycarpum, 760 hypoglaucum, 760 dioicum, 760 hypoglaucum, 760 polygamum, 760 hebecarpum, 760 intermedium, 760 pubescens, 760 hebecarpum, 760 revolutum, 760 glandulosior, 760 thalictroides, 761 turneri, 761 venulosum, 761 confine, 761 Thaspium, 346 trifoliolatum, 346 Thelypteridaceae, 69–70 Thelypteris, 70 hexagonoptera, 70 novaboracensis, 69 palustris, 70 pubescens, 70 phegopteris, 70 simulata, 69 Theophrastaceae, 866 Thermopsis, 595 caroliniana, 595 montana, 595 villosa, 595 Thinopyrum, 285 pycnanthum, 285 thistle, 389–390 Thladiantha, 545 dubia, 545 Thlaspi, 492 arvense, 493 bursa-pastoris, 479 campestre, 487 thicket bindweed, 730 thicket-creeper, 888 thicket sedge, 116 thick-spiked blazing star, 417 thin-leaved sedge, 142 thin-leaved sunflower, 407 thorn-apple, 858–859 thorny hawthorn, 785 thorny-olive, 548 thorough-wax, 340 thoroughwort, 399–400 threadleaf, 426

thread-leaved false pondweed, 298 thread-leaved scorpion-weed, 466 thread-leaved sundew, 547 thread-leaved tickseed, 392 thread rush, 188 thread-stalked speedwell, 719 three-angled spikesedge, 161 threeawn, 232–233 three-birds orchid, 212 threecorner-Jack, 728 three-leaved false Solomon’sseal, 300 three-leaved goldthread, 754 three-leaved rattlesnake-root, 420 three-lobed beggar-ticks, 383 three-lobed coneflower, 425 three-lobed false mallow, 666 three-petaled bedstraw, 826–827 three-seeded-Mercury, 564– 565 three-seeded sedge, 127 three-square bulrush, 167 three-toothed-cinquefoil, 818 three-way sedge, 155 Thuja, 76 occidentalis, 76 Thuringiaca, 147 Thunberg’s bush-clover, 588 Thunberg’s meadowsweet, 820 thyme, 652–653 Thymelaeaceae, 866–867 thyme-leaved dragonhead, 637 thyme-leaved pinweed, 528 thyme-leaved sandmat, 568 thyme-leaved sandwort, 508 thyme-leaved speedwell, 719 Thymophylla, 444 tenuiloba tenuiloba, 444 Thymus, 652 chamaedrys, 653 ovatus, 653 pulegioides, 653 serpyllum, 653 vulgaris, 653 Tiarella, 855 cordifolia cordifolia, 855 Tiaridium indicum, 462 tick quack grass, 285 tickseed, 391–392 tick-trefoil, 580–582 tidal spikesedge, 159 tidytips, 416 Tifton medick, 591 Tilia, 667–668 americana, 667 americana, 668 heterophylla, 668 cordata, 668 petiolaris, 668 platyphyllos, 668 ×vulgaris, 668 Tillaea angustifolia, 541 aquatica, 541

Tillaeastrum aquaticum, 541 Timothy, 272 tinged sedge, 125 Tiniaria cilinodis, 730 convolvulus, 730 cristata, 730 dumetorum, 730 tiny mousetail, 755 tiny vetch, 599 Tipularia, 212 discolor, 212 unifolia, 212 Tissa canadensis, 521 media, 521 rubra, 521 Tithymalopsis corollata, 567 helioscopius, 568 ipecacuanhae, 568 lathyris, 568 peplus, 568 platyphyllos, 568 Tithymalus cyparissias, 567 esula, 567 helioscopius, 568 toadflax, 530 toad-lily, 195 toad rush, 186–187 tobacco, 860–861 Tofieldia glutinosa, 302 Tofieldiaceae, 302 tomato, 865 tomentose chickweed, 510 toothcup, 661 toothed flatsedge, 153 toothed medick, 591 toothed spurge, 567 toothed white-topped-aster, 427 toothwort, 481–482 Torilis, 346 japonica, 346 leptophylla, 346 Toringo crab apple, 793 Tormentilla erecta, 796 Torreyochloa, 285 fernaldii, 285 pallida, 285 fernaldii, 285 pallida, 285 Torrey’s beaksedge, 164 Torrey’s bulrush, 168 Torrey’s mountain-mint, 647 Torrey’s rush, 189 Torrey’s thoroughwort, 399 Torularia humilis, 479 Torulinium odoratum, 154 touch-me-not, 448 Tovara virginianum, 736 glaberrima, 736 tower-mustard, 492 town avens, 792

in d e x  9 6 9

Toxicodendron, 330 aboriginum, 331 negundo, 331 radicans, 331 negundo, 331 rydbergii, 331 rydbergii, 331 vernix, 331 Tracaulon sagittatum, 735 gracilentum, 735 Tradescantia, 99 bracteata, 100 canaliculata, 100 incarnata, 100 ohiensis, 100 foliosa, 100 subaspera, 100 virginiana, 100 Tradescant’s American-aster, 442 Tragopogon, 444 dubius, 445 major, 445 major, 445 porrifolius, 445 pratensis, 445 Tragus, 286 berteronianus, 286 racemosus, 286 trailing-arbutus, 554 trailing bush-clover, 587 trailing pearlwort, 516 Trapa, 661 natans, 661 tree-clubmoss, 44 tree-of-heaven, 857 Triadenum, 626 fraseri, 627 virginicum, 627 fraseri, 627 Triantha, 302 glutinosa, 302 Tribulus, 890 terrestris, 890 Trichomanes, 60 intricatum, 60 Trichophorum, 171 alpinum, 171 cespitosum cespitosum, 171 clintonii, 171 planifolium, 171 Trichostema, 653 brachiatum, 653 dichotomum, 653 lineare, 653 puberulum, 653 lineare, 653 setaceum, 653 Triclinium odoratum, 346 Triclisperma paucifolia, 726 tricolor daisy, 403 Tricolpates, 307–890 Tricyrtis, 195 hirta, 195 tridens, 286 Tridens, 286 flavus, 286

Trientalis americana, 674 borealis borealis, 674 Trifolium, 595 agrarium, 597 albopurpureum dichotomum, 597 arvense, 597 aureum, 597 campestre, 597 dalmaticum, 597 dichotomum, 597 dubium, 595 echinatum, 597 elegans, 597 fragiferum, 597 glomeratum, 597 hybridum, 597 elegans, 597 pratense, 597 incarnatum, 598 elatius, 598 macraei dichotomum, 597 medium, 598 pratense, 598 sativum, 598 procumbens, 597 purpureum, 598 repens, 598 resupinatum, 598 striatum, 598 subterraneum, 598 supinum, 597 tomentosum, 598 Triglochin, 192 debilis, 192 gaspensis, 192 maritima, 192 elata, 192 palustre, 192 Trigonella, 598 corniculata, 598 laciniata, 590 monantha, 590 Trillium, 196 cernuum, 197 macranthum, 197 erectum, 197 erythrocarpum, 197 grandiflorum, 197 nivale, 197 purpureum, 197 recurvatum, 197 rhomboideum grandiflorum, 197 undulatum, 197 unguiculatum, 197 Trilobulina fibrosa, 656 striata, 656 Trimorpha acris kamtschaticus, 396 Triodanis, 497 perfoliata, 497 Triodia flava, 286 Trionum trionum, 665

Triosteum, 504 angustifolium, 504 aurantiacum aurantiacum, 504 eamesii, 504 perfoliatum, 504 aurantiacum, 504 Triphora, 212 trianthophora trianthophora, 212 Triplasis, 286 intermedia, 286 purpurea purpurea, 286 Tripleurospermum, 445 inodorum, 445 maritimum maritimum, 445 perforatum, 445 Triplocentron melitense, 387 Tripolium angustum, 438 frondosum, 439 Tripsacum, 286 dactyloides, 286 Triraphis, 286 mollis, 286 Trisetum, 286 flavescens, 286 melicoides, 262 pensylvanicum, 283 purpurascens, 279 spicatum, 286 molle, 286 psilosiglume, 286 triflorum, 286 Triticum, 287 aestivum, 287 cereale, 280 cylindricum, 226 pycnantha, 285 trachycaulum, 255 turgidum, 287 vulgare, 287 Trollius, 761 laxus, 761 Tropaeolaceae, 867 Tropaeolum, 867 majus, 867 tropical whiteweed, 372 Tropidocarpum, 492 gracile, 492 troublesome sedge, 124 trout-lily, 193 Truellum sagittatum, 735 trumpet-creeper, 455 trumpet honeysuckle, 502 Tryphane rubella, 514 Tsuga, 80 canadensis, 80 tube beardtongue, 714 tuber-bulrush, 103 tuberous comfrey, 467 tuberous grass-pink, 201 tuberous sunflower, 408 tuberous water-lily, 86 tuber vetchling, 584 Tuckermann’s panicgrass, 271

Tuckerman’s quillwort, 42 Tuckerman’s sedge, 150 tufted clubsedge, 171 tufted hair grass, 244 tufted hair-sedge, 104 tufted lovegrass, 258 tufted yellow-loosestrife, 675 Tulipa, 195 gesneriana, 193 sylvestris, 193 tulip, 195 tuliptree, 83 tumbleweed amaranth, 316 tumble windmill-grass, 242 tumbling hedge-mustard, 492 tundra alkali grass, 278 Tunica prolifer, 515 saxifraga, 515 tupelo black, 538 turion duckweed, 96 Turkish hawk’s-beard, 393 Turkish warty-cabbage, 479 Turk’s-cap lily, 194 turnip, 478 Turritis, 492 drummondii, 478 glabra, 492 oblongata, 475 turtlehead, 709 Tussilago, 445 farfara, 445 palmata, 423 tussock bellflower, 495 tussock cottonsedge, 162 tussock sedge, 129, 141 twayblade, 206–207 twice-leaved blackberry, 813 twinflower, 500 twining screwstem, 609 twisted sedge, 141 twistedstalk, 195 two-colored bush-clover, 586 two-flowered dwarf-dandelion, 414 two-leaved bishop’s-cap, 854 two-leaved toothwort, 482 two-rowed water-cress, 488 Typha, 304 angustifolia, 304 elongata, 304 glauca, 304 latifolia, 304 Typhaceae, 302–304

U

Ulex, 598 europaeus, 598 Ulmaceae, 867–869 Ulmaria rubra, 788 Ulmus, 867 americana, 868 campestris, 868 fulva, 869 glabra, 868 minor, 868 parvifolia, 868 procera, 868

970 ind ex

Ulmus, cont. pumila, 868 rubra, 869 thomasii, 869 Unamia alba, 421 umbrella flatsedge, 153 umbrella-sedge, 163 umbrella-tree, 84 umbrella-wort, 675–676 Unifolium canadense, 300 racemosa, 300 Uniola spicata, 251 Unisema cordata, 289 upland cotton, 664 upland thoroughwort, 399 upright brome, 238 upright false bindweed, 532 upright knotweed, 739 upright yellow-rocket, 476 upswept moonwort, 64 Urochloa, 287 texana, 287 Urostachys lucidulus, 40 selago, 40 Urtica, 870 canadensis, 869 cylindrica, 869 dioica, 870 angustifolia, 870 gracilis, 870 procera, 870 gracilis, 870 latifolia, 870 procera, 870 pumila, 870 villosa, 670 Urticaceae, 869–871 Utricularia, 654 biflora, 656 ceratophylla, 656 clandestina, 656 cornuta, 656 fibrosa, 656 geminiscapa, 656 gibba, 656 inflata, 656 minor, 656 intermedia, 656 macrorhiza, 655 minor, 656 pumila, 656 purpurea, 656 radiata, 656 resupinata, 656 striata, 656 subulata, 657 vulgaris macrorhiza, 657 Uvularia, 98 grandiflora, 99 perfoliata, 99 sessilifolia, 99

V

Vaccaria, 523 hispanica, 523 pyramidata, 523 vaccaria, 523 vulgaris, 523 Vaccinium, 561 angustifolium, 562 hypolasium, 562 laevifolium, 562 myrtilloides, 563 nigrum, 562 arbuscula, 562 atlanticum, 562 atrococcum, 563 boreale, 562 caesariense, 562 canadense, 563 cespitosum, 562 arbuscula, 562 corymbosum, 563 albiflorum, 563 atrococcum, 563 fuscatum, 563 glabrum, 563 frondosum, 555 fuscatum, 563 hispidulum, 555 macrocarpon, 563 microcarpos, 563 myrtilloides, 563 nigrum, 562 oxycoccos, 563 intermedium, 563 pallidum, 563 pensylvanicum alpinum, 562 angustifolium, 562 stamineum, 563 uliginosum, 563 vacillans, 563 crinitum, 563 vitis-idaea, 561 minus, 563 Vagnera racemosa, 300 stellata, 300 trifolia, 300 Vahlodea, 287 atropurpurea, 287 valerian, 504 Valeriana, 504 officinalis, 504 sitchensis uliginosa, 504 uliginosa, 504 Valerianella, 504 intermedia, 505 locusta, 505 olitoria, 505 patellaria, 505 radiata, 505 fernaldii, 505 missouriensis, 505 stenocarpa parviflora, 505 umbilicata, 505 Validallium tricoccum, 92

Vallisneria, 177 americana, 177 vanilla sweet grass, 231 variable-leaved pepperweed, 487 variable-leaved water-milfoil, 620 variable sedge, 137 variegated scouring-rush, 60 Vasey’s pondweed, 298 Vasey’s rush, 189 veinless medick, 590 veiny-leaved meadow-rue, 761 velvet grass, 262 velvetleaf, 663, 667 velvetleaf Indian-mallow, 663 velvet-leaved blueberry, 563 velvet sedge, 136 velvety rosette-panicgrass, 250 Venice rose-mallow, 665 Ventenata, 287 avenacea, 287 dubia, 287 Venus’-looking-glass, 497 Veratrum, 197 latifolium, 198 luteum, 196 viride, 198 Verbascum, 856 blattaria, 857 densiflorum, 857 lychnitis, 857 nigrum, 857 phlomoides, 857 thapsus, 857 Verbena, 871 angustifolia, 872 ×blanchardii, 872 bonariensis conglomerata, 872 bracteata, 872 bracteosa, 872 canadensis, 871 lambertii, 871 ×engelmanii, 872 hastata hastata, 872 hispida, 872 lambertii, 871 officinalis, 872 ×rydbergii, 872 stricta, 872 urticifolia, 873 leiocarpa, 873 urticifolia, 873 Verbenaceae, 871–873 Verbesina, 445 alternifolia, 445 encelioides, 445 exauriculata, 445 exauriculata, 445 prostrata, 395 Vermont blackberry, 817 vernal sedge, 135 vernal water-starwort, 708 Vernonia, 446 fasciculata, 446 glauca, 446 missurica, 446 noveboracensis, 446

Veronica, 716 agrestis, 718 alpina terra-novae, 720 americana, 718 anagallis-aquatica, 718 arvensis, 718 austriaca, 717 teucrium, 719 beccabunga, 719 americana, 718 catenata, 719 glandulosa, 719 chamaedrys, 719 comosa glaberrima, 719 glandulosa, 719 connata glaberrima, 719 typica, 719 filiformis, 719 glandifera, 718 hederaefolia, 719 humifusa, 719 latifolia, 719 longifolia, 719 officinalis, 719 tournefortii, 719 paniculata, 720 peregrina, 719 peregrina, 719 typica, 719 xalapensis, 719 persica, 719 pocilla polita, 719 polita, 719 scutellata, 719 villosa, 719 serpyllifolia, 719 humifusa, 720 humifusa, 719 serpyllifolia, 720 spuria, 720 teucrium, 719 tournefortii, 719 wormskjoldii, 720 xalapensis, 719 Veronicastrum, 720 serpyllifolium, 719 humifusum, 719 virginicum, 720 vervain, 871–873 vervain mallow, 666 Vesicariae, 147–150 Vesiculina purpurea, 656 vetch, 575, 577–578, 582, 593, 599–600 vetchling, 584 viburnum, 309 Viburnum, 308 acerifolium, 309 glabrescens, 309 ovatum, 309 alnifolium, 310 bushii, 311 cassinoides, 310 angustifolium, 310

i n d e x  971

Viburnum, cont. dentatum, 309 lucidum, 309 venosum, 309 dilatatum, 310 edule, 310 grandifolium, 310 lantana, 310 sphaerocarpum, 310 lantanoides, 310 lentago, 310 nudum, 310 angustifolium, 310 cassinoides, 310 nudum, 310 opulus, 310 americanum, 310 opulus, 310 trilobum, 310 plicatum, 311 tomentosum, 311 prunifolium, 311 rafinesquianum, 311 recognitum, 309 scabrellum venosum, 309 sieboldii, 311 trilobum, 310 venosum, 309 Vicia, 598 angustifolia, 600 segetalis, 600 cracca, 599 cracca, 599 dasycarpa, 600 faba, 599 grandiflora, 599 kitaibeliana, 599 hirsuta, 599 lathyroides, 599 pannonica, 599 sativa, 600 angustifolia, 600 linearis, 600 nigra, 600 segetalis, 600 sepium, 600 montana, 600 sororia affinis, 880 tetrasperma, 600 tenuissima, 600 varia, 600 villosa, 600 varia, 600 villosa, 600 Villarsia cordata, 669 Vinca, 350 herbacea, 351 major, 351 minor, 351 Vincetoxicum medium, 350 nigrum, 350 rossicum, 350 Viola, 873 adunca, 880 minor, 883 aduncoides, 880 affinis, 880

arenaria, 880 arvensis, 881 atlantica, 881 ×bissellii, 882 blanda, 881 brittoniana, 881 canadensis, 880 canadensis, 882 clandestina, 885 concolor, 873 ×conjugens, 884 ×consocia, 881 ×convicta, 886 cucullata, 882 microtitis, 882 ×eclipes, 883 emarginata, 885 acutiloba, 885 eriocarpon, 885 ×fernaldii, 886 ×filicetorum, 881 fimbriatula, 885 ×greenmanii, 882 hirsutula, 883 ×hollickii, 881 incognita, 881 forbesii, 881 ×insessa, 882 ×insolita, 882 japonica, 883 labradorica, 883 labrusca, 889 lanceolata, 877 lanceolata, 883 latiuscula, 880 macloskeyi pallens, 884 ×malteana, 883 ×mulfordae, 882 ×napae, 883 nephrophylla, 883 ×notabilis, 881 novae-angliae, 883 obliqua, 882 odorata, 884 pallens, 884 palmata, 884 dilatata, 884 palustris, 878 palustris, 884 pectinata, 884 pedata, 885 concolor, 885 pedatifida brittoniana, 881 pensylvanica, 885 ×populifolia, 884 ×porteriana, 882 primulifolia, 885 acuta, 885 pubescens, 885 eriocarpon, 885 scabriuscula, 885 renifolia, 885 brainerdii, 885 ×robinsoniana, 884 rostrata, 885 rotundifolia, 885 ×ryoniae, 883 sagittata, 885 subsagittata, 885

selkirkii, 886 sororia, 886 novae-angliae, 883 striata, 886 ×subaffinis, 881 subsinuata, 886 subvestita, 880 tricolor, 886 arvensis, 881 triloba, 884 ×variabilis, 884 Violaceae, 873–886 violet, 873–886 violet bush-clover, 587 violet butterwort, 654 violet crabgrass, 251 violet dock, 744 violet-flowered petunia, 861 violet wood sorrel, 697 viper’s-bugloss, 461 Virginia buttonweed, 822 Virginia chain fern, 53 Virginia dwarf-dandelion, 414 Virginia fanpetals, 667 Virginia ground-cherry, 863 Virginia marsh-St. John’s-wort, 627 Virginia meadow-beauty, 668 Virginia mountain-mint, 647 Virginiana-creeper, 888 Virginia rose, 807 Virginia screwstem, 609 Virginia serpentaria, 81 Virginia spiderwort, 100 Virginia spring-beauty, 745 Virginia stickseed, 461 Virginia three-seeded-Mercury, 565 Virginia virgin’s-bower, 753 Virginia water-horehound, 641 virgin’s bower, 753 Virgulus ericoides, 439 novae-angliae, 440 patens, 441 Viscaceae, 887 Viscaria, 523 vulgaris, 523 Vitaceae, 887–889 Vitis, 888 aestivalis, 889 labrusca, 889 ×novae-angliae, 889 palmata, 889 riparia, 889 vinifera, 889 vulpina, 889 Vulpia, 287 bromoides, 287 dertonensis, 287 megalura, 288 myuros, 288 octoflora, 288 octoflora, 288 tenella, 288 Vulpinae, 150

W wakerobin, 197 Waldsteinia fragarioides, 791 walking spleenwort, 52 wallflower, 485 wall hawkweed, 412 Wallia cinerea, 629 nigra, 629 wall iris, 180 wall-lettuce, 418 wall-rocket, 483–484 wall-rue spleenwort, 52 walnut, 629 Walter’s sedge, 136 wand bush-clover, 588 warty-cabbage, 479 warty panicgrass, 271 Washington hawthorn, 786 Washingtonia claytonii, 344 longistylis, 344 obtusa, 344 water-awlwort, 492 water avens, 792 water bulrush, 167 water-chestnut, 661 water clover, 61 water-cress, 488 water forget-me-not, 464 water-hemlock, 340 water-hemp, 316–317 water-horehound, 640–641 water-hyacinth, 289 waterleaf, 462 water-lily, 86 water lobelia, 496 water manna grass, 261–262 water-marigold, 382 water-meal, 97 watermelon, 544, 864 watermelon nightshade, 864 water-milfoil, 621 water mint, 642 water montia, 745 watermoss, 69 waternymph, 177 water-parsnip, 346 water-pepper smartweed, 734 water-plantain, 88 water-plantain crowfoot, 758 water-primrose, 684 water sedge, 139 water-shield, 84 water smartweed, 733 water speedwell, 718–719 water-starwort, 708 water-thyme, 176 waterweed, 176 water-willow, 307 waterwort, 549 wavy bitter-cress, 482 wavy blue grass, 276 wavy hair grass, 245 wavy-leaved American-aster, 442 wavy waternymph, 177 waxweed, 660 waxy bedstraw, 825

972  inde x

waxy-leaved meadow-rue, 760 wayfaring tree, 310 weak rush, 187 weak spear grass, 277 weak-stalked bulrush, 167 weak stellate sedge, 147 Weatherby’s rabbit-tobacco, 424 wedgeleaf, 384 wedgescale, 283 weeping forsythia, 677 weeping lovegrass, 257 weigela, 505 Weigela, 505 floribunda, 505 western androsace, 746 western bleeding-heart, 701 western dock, 743 western maidenhair fern, 68 western poison-ivy, 331 western rose-gentian, 612 western tansy-mustard, 483 western water-horehound, 640 western wheat grass, 271 West India gherkin, 544 West Indian sage, 648 West Indian wood sorrel, 697 wheat, 287 wheatgrass, 226, 271 wheat sedge, 115 whip hawkweed, 411 whiplash pappus grass, 271 whip nutsedge, 171 white adder’s-mouth, 206 white ash, 677 white-aster, 394 white avens, 791 white baneberry, 750 white beaksedge, 164 white bear sedge, 132 white bedstraw, 824 white beggar-ticks, 382 white blue-eyed-grass, 181 white campion, 519 white-cedar, 76 white charlock, 491 white cinquefoil, 796 white clover, 598 white colic-root, 198 white cuckoo bitter-cress, 482 white cut grass, 264 white dock, 744 white doll’s-daisy, 384 white-edged sedge, 131 white fir, 77 white flat-topped-goldenrod, 421 white flax, 658 white-flowered leaf-cup, 423 white-flowered pincushionplant, 722 white-flowered willow-herb, 683 white fringed bog-orchid, 208, 210 white fringe-tree, 676 white goldenrod, 431 white goosefoot, 323 white-grained rice grass, 268 white henbit, 638 white horehound, 641

white knapweed, 387 white meadowsweet, 819 white morning-glory, 536 White Mountain avens, 792 white mountain saxifrage, 855 white mulberry, 671 white mullein, 857 white northern bog-orchid, 209 white poplar, 831 white prairie-clover, 579 white prickly-poppy, 699 white rattlesnake-root, 419 white-root rush, 187 white snakeroot, 372 white spruce, 78 white-stemmed elkweed, 609 white-stemmed pondweed, 297 white stonecrop, 543 white sweet-clover, 591 white thoroughwort, 399 white-tinged sedge, 113 white-topped-aster, 427 white trout-lily, 193 white turtlehead, 709 white upright mignonette, 761 white vervain, 873 white vetchling, 584 white virgin’s-bower, 753 white wakerobin, 197 white walnut, 629 white water crowfoot, 758 white water-lily, 86 whiteweed, 372 white willow, 843 white wood-aster, 400 white wormwood, 378 whitlow-mustard, 484 whitlow-wort, 515 whorled bedstraw, 826 whorled mallow, 666 whorled marsh-pennywort, 343 whorled milkweed, 350 whorled milkwort, 726 whorled mountain-mint, 647 whorled sage, 648 whorled water-milfoil, 621 whorled yellow-loosestrife, 674 wide-throated monkey-flower, 705 Wiegand’s rush, 186 Wiegand’s sedge, 147 Wiegand’s wild-rye, 256 wig knapweed, 387 wild basil, 636 wild bean, 592 wild bee-balm, 644 wild bishop, 339 wild bleeding-heart, 701 wild blue phlox, 723 wild calla, 95 wild campion, 519 wild carrot, 341 wild chamomile, 418 wild chervil, 338 wild chives, 92 wild cucumber, 545 wild dock, 743 wild feverfew, 422 wild goat’s-rue, 595 wild honeysuckle, 501 wild hound’s-tongue, 460

wild leek, 92 wild marjoram, 645 wild monkshood, 750 wild oat, 233 wild parsnip, 344 wild radish, 489 wild-rye, 256 wild sarsaparilla, 339 wild scorpion-weed, 466 wild sensitive-pea, 578 wild sweet potato morningglory, 536 wild tulip, 195 wild yam, 172 Willdenow’s sedge, 143 willow, 833–848 willow-herb, 682–684 willow-leaved American-aster, 441 willow-leaved yellowhead, 413 willow oak, 606 windflower, 751–752 windmill campion, 519 windmill grass, 242 wine raspberry, 816 winged loosestrife, 661 winged monkey-flower, 704 winged-pigweed, 325 winged spindle-tree, 524 winged sumac, 330 wingpetal, 409 wingstem crownbeard, 445 winter bentgrass, 227 winterberry, 352 wintergreen, 555 winter squash, 545 wire-stemmed muhly, 267 wiry panicgrass, 270 Wissadula callimorpha, 667 wisteria, 600 Wisteria, 600 floribunda, 600 frutescens, 600 sinensis, 600 witch-hazel, 622 witch panicgrass, 270 withe-rod, 310 Wolffia, 97 borealis, 97 brasiliensis, 97 borealis, 97 columbiana, 97 papulifera, 97 punctata, 97 Wolffiella, 97 floridana, 97 gladiata, 97 wolfstail, 267 wood-aster, 399–400 wood-beauty, 788 wood bedstraw, 826 wood blue grass, 276 wood bulrush, 169 wood horsetail, 60 woodland agrimony, 768 woodland arctic-cudweed, 421 woodland figwort, 856 woodland forget-me-not, 464 woodland germander, 652 woodland muhly, 268

woodland ragwort, 427 woodland sage, 648 woodland stonecrop, 543 woodland strawberry, 789 woodland sunflower, 407 woodland yellow flax, 658 wood lily, 194 wood-mint, 636 wood-nettle, 869 wood-reed, 242 woodruff, 821 Woodsia, 74 alpina, 74 glabella, 74 ×gracilis, 74 hyperborea, 74 ilvensis, 74 obtusa, 74 Woodsiaceae, 70–74 wood sorrel, 697 wood violet, 884 Woodwardia, 53 areolata, 53 onocleoides, 53 virginica, 53 wood windflower, 752 woolly bellflower, 495 woolly blue violet, 886 woolly burdock, 376 woolly clover, 598 woolly croton, 565 woolly Dutchman’s pipe, 82 woolly-fruited sedge, 136 woolly hedge-nettle, 651 woolly safflower-thistle, 385 woolly sedge, 136 woolly yarrow, 372 woolsedge, 169–170 wormseed, 326, 485 wormseed wallflower, 485 wormwood, 377–379 Wright’s rosette-panicgrass, 250 Wright’s spikesedge, 159

X Xananthes minor, 656 Xanthium, 446 canadense echinatum, 446 glanduliferum, 446 italicum, 446 oviforme, 446 pensylvanicum, 446 speciosum, 446 spinosum, 446 inerme, 446 strumarium, 446 canadense, 447 glabratum, 447 pensylvanicum, 446 strumarium, 447 Xanthorhiza, 761 apiifolia, 761 simplicissima, 761 Xanthoxalis brittoniae, 697 bushii, 697 corniculata, 697

i n d e x  973

Xanthoxalis, cont. dillenii, 697 filipes, 697 florida, 697 langloisii, 697 repens, 697 Xeranthemum bracteatum, 447 Xerochrysum, 447 bracteatum, 447 Ximenesia encelioides exauriculata, 445 exauriculata, 445 Xolisma ligustrina, 557 mariana, 557 Xylosteum Ciliatum, 501 Xyridaceae, 305 Xyris, 305 bulbosa, 305 caroliniana olneyi, 305 congdonii, 305 difformis, 305 montana, 305 smalliana, 305 olneyi, 305 torta, 305

Y yam, 172 yam-leaved virgin’s-bower, 753 yard dock, 743 yarrow, 371–372 yellow-and-blue forget-menot, 464 yellow avens, 791 yellow ball-mustard, 488 yellow bedstraw, 827 yellow birch, 453 yellow bluebead-lily, 193 yellow chestnut oak, 605 yellow-cress, 490–491 yellow day-lily, 173 yellow evening-primrose, 687 yellow-eyed grass, 305 yellow false crown-vetch, 578 yellow false oat, 286 yellow flatsedge, 154 yellow floating-heart, 669 yellow-flowered knotweed, 739 yellow forest violet, 885 yellow foxglove, 710 yellow foxtail, 281 yellow-fruited sedge, 135 yellow giant-scabious, 499 yellow-green sedge, 117 yellow hawkweed, 411 yellow hawthorn, 783 yellowhead, 413 yellow henbit, 639

yellow horn-poppy, 701 yellow iris, 180 yellow knapweed, 388 yellow ladies’-tresses, 212 yellow lady’s-slipper, 202 yellow-loosestrife, 674–675 yellow medick, 590 yellow mountain saxifrage, 854 yellow-pimpernel, 346 yellow pincushion, 388 yellow pine-sap, 556 yellow-rattle, 695 yellow-rocket, 476 yellowroot, 761 yellow-seeded false pimpernel, 659 yellow spikesedge, 160 yellow sweet-clover, 591 yellow thistle, 390 yellow-throated scorpion-weed, 465 yellowtops, 402 yellow upright mignonette, 761 yellow vetchling, 584 yellow wild indigo, 577 yellow-wood, 578 yew, 80 Yucca, 87 filamentosa, 87 smalliana, 87

Z

Zannichellia, 298 palustris, 298 major, 298 Zanthoxylum, 829 americanum, 829 Zea, 288 mays, 288 Zigadenus elegans glaucus, 196 zigzag clover, 598 zig-zag goldenrod, 432 zigzag hawthorn, 784 Zizania, 288 aquatica angustifolia, 288 aquatica, 288 subbrevis, 288 palustris palustris, 288 Zizia, 347 aptera, 347 aurea, 347 cordata, 347 Zostera, 305 marina, 305 stenophylla, 305 Zosteraceae, 305 Zosterella dubia, 289 Zygophyllaceae, 890

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