Latin American Insects and Entomology 0520078497, 9780520078499

Citation preview

LATIN AMERICAN INSECTS AND ENTOMOLOGY Charles L. Hogue

5>*il^Vc -

C ,A

'

.

.

'

■

>

'

■

Xfti

LATIN AMERICAN INSECTS AND ENTOMOLOGY Charles L. Hogue

Lantern bugs and cicadas, from Madam Merian's Metamorphoses insectorum surinamensium (1705). Note the imagined specimen at the bottom of the plate which combines features of the two insects.

UNIVERSITY OF CALIFORNIA PRESS • Berkeley • Los Angeles • Oxford

!

University of California Press Berkeley and Los Angeles, California University of California Press Oxford, England Copyright © 1993 by The Regents of the University of California Library of Congress Cataloging-in-Publication Data Hogue, Charles Leonard. Latin American insects and entomology / by Charles L. Hogue. p. cm. Includes bibliographical references (p. ) and index. ISBN 0-520-07849-7 (cloth) 1. Insects—Latin America. I. Title. QL476.5.H64 1993 595.7098—dc20 91-48184 CIP Printed in the United States of America 1 2 3 4 5 6 7 8 9 The paper used in this publication meets the mini­ mum requirements of American National Standard for Information Sciences—Permanence of Paper for Printed Library Materials, ANSI Z39.48-1984 ©

IN M E M O R I U M To the great r e g r e t of family, his friends, a n d his colleagues, C h a r l e s H o g u e d i e d s u d d e n l y in m i d - 1 9 9 2 , while the m a n u s c r i p t for this book was b e i n g typeset. T h e Press w o u l d like to t h a n k his son, J a m e s H o g u e , for skillfully a n d meticulously s e e i n g the b o o k t h r o u g h the final stages of p r o o f r e a d i n g a n d p r i n t i n g . His care e n s u r e d that Latin American Insects and En­ tomology, C h a r l e s H o g u e ' s last major work, would a p p e a r w h e n a n d as his father would have wished.

The ubiquity of spirits and the impossibility of killing them seem to personify a feeling of helplessness in the face of an environment so beautiful and so cruel. On the river or work­ ing in a garden the sun hurts, "It is eating," the Sharanahua say, and heads ache for the rest of the day. The incessant gnats feed all day, and, as one lies in a hammock, someone leans over and slaps hard and says, "sandfly," and a black fly, fat with human blood, falls dead. Sundown is a moment of relief which even a hundred mosquitoes cannot mar. In the forest someone shouts to warn of an uula, the huge stinging ants that make one drunk with pain, and, reaching for a handhold on a tree, one must avoid a swarm of red fire ants. Returning, one looks for ticks, huge tapir ticks, gray and voracious, or worse, the almost invisible tiny red ticks that burrow into the skin and hurt for a week. The women dig the egg sacs of chiggers out of toes skillfully so that the sac does not break to leave a budding worm to swell the foot, and they break each and every tiny egg with a needle so that it does not lie in wait for an­ other bare foot. An infected gnat drops a worm's egg into the leg while sucking blood, and two weeks later the pain of the worm turning in the leg is excruciating, and it must be removed by daubing an old, foul, drop of tobacco juice on the skin and slowly winding the worm out on a stick. Women and girls pick lice out of men's hair and their own, break them in their teeth and eat them. When faced by a new animal or insect, I learned to ask both, "Do we eat it?" and "Does it eat us?" Janet Siskind, To Hunt in the Morning

Contents

Preface Acknowledgments Introduction 1 GENERAL ENTOMOLOGY

xi xiii 1 3

7 ECTOPARASITIC ORDERS

206

8 BUGS

216

9 BEETLES

246

2 ECOLOGY

48

10 MOTHS AND BUTTERFLIES

292

3 PRACTICAL ENTOMOLOGY

90

11 FLIES AND MIDGES

360

4 TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

12 SAWFLIES, WASPS, ANTS, 108

AND BEES 13 INSECT STUDY

5 ORTHOPTEROIDS AND OTHER ORDERS

152

6 AQUATIC ORDERS

193

Included Insect and Arthropod Taxa Index

405 470

499 521

IX

Preface

The idea for this book germinated in my mind for many years after my christening in Latin American insect research. As a result of travels to many countries, it became acutely apparent to me that a comprehensive entomological work was sorely needed to serve the many people, both visitors and residents, interested in insects and their other terrestrial arthropod relatives. The curiosity of tourists and general natural historians needed satisfac­ tion, and the more serious minded student and practicing professional needed an upto-date review of the subject. After a long period of note taking and preliminary organization of my then chaotic knowledge of the subject, I resolved to fashion such a volume. Some of my colleagues were incredulous that I could cover such a vast territory. But my experience writing general insect guides told me that with cautious choosing, I could make something really useful, though, of course, far from complete. In fact, writing this book has been an exercise in selectivity, especially with respect to the choice of taxonomic groups to include. I relied on my own experience and the experience of others in this process, and I have tried to give information on the most common, conspicuous, or otherwise nota­ ble (economically or historically important) units, whether species, genera, or higher groups. T h e other topics of discussion, such as ecology and the study of insects,

have also been presented with an eye to the reader's need for an overall understanding of what has transpired and is transpiring in Latin American entomology and to provid­ ing a framework for review and citation of pertinent literature. Some who read this book will feel slighted because of the lack of coverage of topics of particular interest to them, or they may consider that important facts, taxa, or publications have been omitted. I can only ask these readers to remember the vastness of the subject and the necessity for extreme conservatism in choices of matter for inclu­ sion. Of course, I welcome suggestions for additions or changes in emphasis for future editions. I have designed the book to answer questions. In my language and in the selection of taxa/phenomena, and points of information about each, I have been guided by my perception of what most readers want to know rather than a desire to produce an encylopedia of all the facts that might be recorded. The technical litera­ ture, to which I have so freely referred, will serve the latter purpose. Indeed, to present in-depth data, keys to identification, and exhaustive treatments of even the major categories of Latin American insect life would require many volumes and years of effort to produce and would not produce the ready, readable, and portable text that I think is most needed now.

XI

Acknowledgments

For reviewing sections and offering criti­ cisms and information on many topics, I have been fortunate to have the expert assistance of many specialists. I am greatly indebted to the individuals named below for information, identifications, and count­ less improvements in the manuscript (the appearance of their names does not neces­ sarily imply agreement with my final inter­ pretations and statements): J. Richard An­ drews (Náhuatl names), Arthur G. Appel (cockroaches), Phillip A. Adams (Neuroptera), Richard W. Baumann (Plecoptera), Vitor Becker (Lepidoptera), Jackie Belwood (Orthoptera), Harry Brailovsky (bugs), A. Brindle (Dermaptera), Jacob Brodzinsky (fossils), Lincoln P. Brower (but­ terflies), John C. Brown (Lepidoptera), Richard C. Brusca (Crustacea, arthropod anatomy), Gary R. Buckingham (useful insects), James L. Castner (Orthoptera), Gilbert L. Challet (aquatic beetles), John A. Chemsak (Cerambycidae), James C. Cokendolpher (harvestmen), Julian P. Donahue (Lepidoptera), John T. Doyen (darkling beetles), Richard D. Durtsche (study), W. D. Edmonds (dung scarabs), George F. Edmunds, Jr. (mayflies), K. C. Emerson (Mallophaga), Marc E. Epstein (Lepidop­ tera), Terry L. Erwin (beetles), Arthur V. Evans (beetles), Eric M. Fischer (Diptera), Oliver S. Flint, Jr. (Trichoptera), Will Flow­ ers (aquatics), Manfredo A. Fritz (Sphecoidea), Richard C. Froeschner (Hemiptera), David G. Furth (ecology), Rosser W. Garrison (Odonata), P. Genty (palm pol­ linators), Edmund Giesbert (Cerambyci­

dae), Eric E. Grissel (Chalcidoidea), Robert J. Gustafson (botany), Alan R. Hardy (Scarabaeidae), Brian R. Harris (butterflies), Steven Hartman (mantids), Henry A. Hespenheide (beetles), Frank T. Hovore (Cer­ ambycidae), Chistopher A. Ishida (prac­ tical entomology), D. K. McE. Kevan, deceased (orthopteroids), James E. Keirans (ticks), John E. Lattke (general), Claude Lemaire (Saturniidae), Herbert W. Levi (spiders), James E. Lloyd (Lampyridae), Wilson R. Lourenco (scorpions), Richard B. Loomis, deceased (chiggers), Robert J. Lyon (gall wasps), Volker Mahnert (pseudoscorpions), Sergio Martinez (fossils), Mildred E. Mathias (botany), Eustorgio Méndez (medical entomology, parasites), Arnold S. Menke (Cynipidae and other Hymenoptera), Edward L. Mockford (psocids), Jacqueline Y. Miller (Castniidae), Michael J. Nelson (medical entomology), David A. Nickel (orthopteroids), M. W. Nielson (leafhoppers), Lois B. O'Brien (Homoptera), David L. Pearson (tiger beetles), Stewart B. Peck (cave insects), Norman D. Penny (Neuroptera and general), Don R. Perry (canopy insects), Manuel L. Pescador (mayflies), John T. Polhemus (aquatic Hemiptera), Diomedes Quintero Arias, Jr. (whip scorpions), Shivaji Ramalingam (mos­ quitoes), Edward S. Ross (Embiidina), H. F. Rowell (Acrididae), Raymond E. Ryckman (ectoparasitic bugs), Ann L. Rypstra (spi­ ders), Jorge A. Santiago-Blay (scorpions), Jack Schuster (Passalidae), Terry N. Seeno (beetles), Rowland M. Shelley (myriapods), David R. Smith (Symphyta), Roy R. Snell-

XIII

ing (Hymenoptera), Omelio Sosa, Jr. (his­ tory), Paul J. Spangler (aquatics), Lionel A. Stange (Neuroptera), Orley R. Taylor (honeybee), Donald B. Thomas (heteropterans and veterinary entomology), Carlos Trenary (history), Fred S. Truxal (Notonectidae), Alan Watson (Lepidoptera), Thomas K. Wood (Membracidae), A. Willink (history and study), Stephen L. Wood (Scolytidae), and Thomas J. Zavortink (mosquitoes). For suggestions and information on diverse or multiple topics, I also have many other entomologists to thank, including, José Artigas, Tomás Cekalovic, Ana Lia Estévez, Carlos H. W. Flechtmann, Luis F. Jirón, Alberto and Beatriz Larraín, Carlos Machado Allison, and Irene Rut-Wais. There are several colleagues who are responsible for more pervasive involve­ ment in the work as a whole, who have reviewed and contributed to the entire manuscript, to whom I owe a special debt of gratitude: Terry L. Erwin, Arthur V. Evans, Gerardo Lamas, Jack Longino, Scott E. Miller, José Palacios-Vargas, Nel­ son Papavero, and Allen M. Young. For tolerance of my many probings of their knowledge and points of view, I am particu­ larly appreciative of the contributions of Julian P. Donahue, Chris Nagano, Roy R. Snelling, and Fred S. Truxal. I am also indebted to the many local friends and contacts I have made on my Latin American travels—guides, foresters, farmers, Native Americans, and many oth­ ers, too numerous to name, who freely

xiv

ACKNOWLEDGMENTS

shared their firsthand field knowledge of insects and thus contributed to the original­ ity of this book in countless ways. To all who worked with the manuscript primarily in an editorial capacity, I wish to express gratitude, especially to Ernest Callenbach. Several librarians contributed in no small way by finding and interpreting many difficult areas of the literature: Nicole Bouché, Katharine E. S. Dona­ hue, Jennifer Edwards, Donald McNamee, Kathy Showers (Los Angeles County Mu­ seum of Natural History) and Nancy Axelrod (University of California, Berkeley, Entomology Library). The preparation of the ink drawings were greatly assisted by Leland E. Dietz and his Xerox machine. My appreciation is also due to Don Meyers for his careful and considerate handling of my black-and-white photographic needs, as well as to Tracy Robertson and James Robertson for other technical assistance with the figures. I thank James L. Castner, George Dodge, and James N. Hogue for allowing me to use those color photographs bearing their names in the captions. Finally, I acknowledge my wife, Bar­ bara, for her critical role in helping this task to completion, primarily her enor­ mous patience with my needs for time and other of our mutual resources. She also typed most of my original drafts and did a great deal of editing. For these contribu­ tions, not only extended here but lovingly provided in support of most of my entomo­ logical career, I dedicate this book to her.

Introduction

This work provides an introduction to the common and notable insects of Latin Amer­ ica and a foundation for their study. All the countries of the Western Hemisphere south of the U.S.-Mexican border are covered, including the West Indies, the continental islands, and the oceanic islands within 2,500 kilometers of and usually associated with isthmian America and South America (but not Bermuda or Isla de Pascua or Easter Island). In discussing broad geographic regions, I use the terms "Latin America," "New World tropics," and "Neotropics" (and their adjectival forms) as precisely as possible. The first refers to the most inclusive politi­ cal region within the bounds stated above; the second refers to lowland, moist to wet, forested biotypes only; and the third refers to the biogeographic region that includes South America, the West Indies, and tropi­ cal North America. In this volume, I discuss true insects and other kinds of terrestrial arthropods and related creatures (arachnids, millipedes, centipedes, onycophorans, etc.) commonly thought of as "insectlike." In many places, it is overly cumbersome to be exact when referring to these groups, although I have tried to avoid misleading the reader by adding some explanatory phrase, such as "and other terrestrial arthropods," but I have found it difficult to be perfectly consistent in doing so. The meaning of a broader grouping is sometimes implied by the term "insect." For each kind of insect or other ar­ thropod discussed in a separate section in

the systematic chapters, I open with a list of the applicable scientific names of the cate­ gory, including the most commonly used synonyms, and its place in the nomenclatural hierarchy. This is followed by estab­ lished vernacular names in Spanish, Portu­ guese, and other regional languages. I use the widely familiar "Quechua" in place of the more correct "Runasimi." No attempt is made to give a complete synonymy, as this would require a lengthy treatment of its own. The determination of plurals in some antique or indigenous languages is a prob­ lem, and some of those given may not find acceptance by purists. I have replaced a few common names, for example, "wax bugs" for the order Homoptera, "dragon-headed bugs" for Fulgora spp., and "big-legged bugs" for the family Coreidae. I feel that these are more appropriate and correct than previously used names and that they are useful to enhance common parlance about these often-mentioned taxa. There are a few new common names (e.g., "viper worms" for Hemeroplanes sp., Sphingidae; "flag moths" for the subfamily Pericopinae of the Arctiidae; and "shiny scarabs" for the subfamily Rutelinae of the Scarabaeidae. The chapter on study of insects is in­ cluded as information for novices and to enhance the use of the book for teaching and for ready reference for professionals. Much of the information has not been compiled elsewhere. Citations to the literature were chosen according to certain constraints. First, they are always given as authority to, and source of further information on, topics of special

1

interest, statistical statements, historical r e ­ m a r k s , a n d such. I also include basic refer­ ences to g e n e r a l subjects o r sections o r b r o a d categories b u t only t h e most m o d e r n , c o m p r e h e n s i v e , a n d well-referenced, p r e ­ s u m i n g that the r e a d e r will find in its own bibliography all the earlier p e r t i n e n t litera­ t u r e . M i n o r t a x o n o m i c a n d mostly regional p a p e r s a r e e x c l u d e d ; classic o r s t a n d a r d p a p e r s a r e i n c l u d e d , even if short o r out­ d a t e d . References a r e given at t h e e n d of the sections to which they apply, r a t h e r than assembled t o g e t h e r at t h e e n d of t h e book, to facilitate c o m p i l i n g literature by subject. Because I use this format, a few references have been r e p e a t e d , obviating t h e n e e d for cross listings a n d m a k i n g it easier for the r e a d e r to find t h e m . I have seen all refer­ ences except those r a r e ones followed by "[Not seen.]." S o m e n e w observations are included a n d d e n o t e d by "(orig. obs.)." I cover t h e literature p u b l i s h e d t o D e c e m b e r 31, 1991. T h i s book is i n t e n d e d for t h e widest possible a u d i e n c e : s t u d e n t s , lay p e o p l e , travelers, teachers, a n d professional ento­ mologists alike. T h e latter will forgive my use of nontechnical t e r m s w h e n e v e r possi­ ble; b u t generally, I follow v e r n a c u l a r ("wing covers") with technical synonyms ("elytra"). Nevertheless, t h e use of m a n y technical t e r m s has been unavoidable. T h e y a r e e x p l a i n e d in c h a p t e r 1 or a r e available in most any g e n e r a l e n t o m o l o g y textbook.

Illustrations I chose the species for t h e ink d r a w i n g s in the systematic p o r t i o n of t h e book as r e p r e ­ sentative generally of those f o u n d in t h e text. All t h e drawings a r e my own, a n d most a r e based o n m u s e u m specimens entirely o r in part to confirm details p r e s e n t in existing illustrations. I have used t h e latter only casually as a n aid to composition. T h e majority of insects a n d o t h e r types illustrated in the line d r a w i n g s a r e placed in stylized natural settings a n d a r e d e ­ picted as living animals. I have n o t d r a w n figures to scale, except that an a t t e m p t was m a d e to indicate comparative size, that is, larger species a r e larger t h a n smaller, al­ t h o u g h not in p r o p o r t i o n . T h e n e e d occa­ sionally arose to greatly magnify micro­ scopic forms. With t h e exception of those with t h e n a m e s of o t h e r p h o t o g r a p h e r s o r sources noted in t h e captions, all t h e color a n d black-and-white p h o t o g r a p h s of in­ sects a n d p h e n o m e n a a r e m i n e also, con­ verted from 35 m m transparencies taken of live specimens in the field.

Abbreviations T h r o u g h o u t t h e book, only t h e following t h r e e abbreviations a r e used: BL (body length, i.e., front of h e a d to tip of a b d o ­ men); B W L (length of insect from front of h e a d to tips of wings w h e n folded in repose); WS (wingspan).

J

GENERAL ENTOMOLOGY

Entomology (Greek: entomon insect + logos discourse) is the scientific study of insects. A general k n o w l e d g e of insects a n d their a r t h r o p o d relatives is basic to an u n d e r ­ standing of their classification a n d biology in Latin A m e r i c a . T h e r e a r e m a n y good reference works (Parker 1982, K ü k e n t h a l 1923, Grassé 1949) a n d e n t o m o l o g y text­ books (Barth 1972, B o r r o r a n d D e L o n g 1969, B o r r o r et al. 1 9 8 1 , C o r o n a d o et al. 1976, Daly et al. 1978, C a r r e r a 1963, H a y ward 1971, Lara 1977, Richards a n d Davies 1977, 1983) as well as c o m p r e h e n s i v e treat­ ments of anatomy, physiology, classifica­ tion, phylogeny, d e v e l o p m e n t , a n d behav­ ior. Because these works a r e t h o r o u g h a n d widely applicable, it would be r e d u n d a n t and inefficient to r e p e a t their contents in a specialized book such as this. T h e r e m a i n ­ der of this c h a p t e r offers a synopsis of f u n d a m e n t a l topics a n d an explanation of terminology to m a k e t h e book m o r e usable and instructive for g e n e r a l r e a d e r s .

References BARTH, R. 1972. Entomología geral. Fund. Insto. Oswaldo Cruz, Rio de Janeiro. BORROR,

D. J.,

AND D.

M.

DELONG.

1969.

Introducáo ao estudo dos insetos. Ed. Edgard Bluecher, Sao Paulo. Brazilian edition trans­ lated and edited by D. D. Correa et al. BORROR, D. J.,

D.

M.

DELONG,

AND C.

H.

TRIPLEHORN. 1981. An introduction to the

study of insects. 5th ed. Saunders College, Philadelphia. CARRERA, M. 1963. Entomología para vocé. Ed. Univ. Sao Paulo, Sao Paulo. CORONADO, P., R. A. MÁRQUEZ, AND A. MÁR­

QUEZ. 1976. Introducción a la entomología. Ed. Limusa, Mexico City.

2

INTRODUCTION

DALY, H. V., J. T. DOYEN, AND P. R. EHRLICH.

1978. Introduction to insect biology and di­ versity. McGraw Hill, New York. GRASSÉ, P., ed. 1949-. Traite de zoologie. Vols. 6-10. Masson, Paris. HAYWARD, K. J. 1971. Guía para el entomólogo principiante. 2d ed. Univ. Nac. Tucumán, Insto. Miguel Lillo, Misc. no. 37: 1-159. KÜKENTHAL, W. 1923-. Handbuch der zoologie. Vol. 4. Arthropoda. de Gruyter, Berlin. LARA, F. 1977. Principios de entomología. Fac. Cien. Agr. Vet., Univ. Estad. Paulista "Julio de Mesquita Filho," Sao Paulo. PARKER, S. B., ed. 1982. Synopsis and classifica­ tion of living organisms. Vol. 2. McGraw-Hill, New York. RICHARDS, O.

W., AND R.

G.

DAVIES.

1977.

Imms' general textbook of entomology, 10th ed. Vols. 1-2. Chapman and Hall, London. RICHARDS, O. W.,

AND R. G.

DAVIES.

1983.

Tradado de entomología Imms. Vol. 1, Estruc­ tura, fisiología y desarrollo; Vol. 2, Clasifi­ cación y biología. Ed. Omega, Barcelona.

HISTORY OF LATIN AMERICAN ENTOMOLOGY Studies on Latin A m e r i c a n insects a n d related a r t h r o p o d s began late in t h e his­ tory of biology because of t h e belated discovery of t h e New World by E u r o p e a n s a n d its academic isolation for almost two centuries thereafter. T h e history of e n t o ­ mology in the region is best traced as a series of o v e r l a p p i n g accomplishments by different categories of searchers a n d for­ mats of investigation, r a t h e r t h a n as tradi­ tional chronological p e r i o d s that a r e h e r e not readily identifiable. T h e s e categories a r e largely d e t e r m i n e d by t h e kinds of

3

education available or customary at the time. The earliest disciples were broadly trained in philosophy, theology, or medi­ cine; later, the narrower disciplines of natural history, biology, and zoology evolved. Not until very late in the nine­ teenth century did courses in entomology exist and still later full curricula leading to a degree in the subject. No general discussion of Latin Ameri­ can entomology is available, although there are some regional treatments: Lizer y Trelles (1947) recounts the overall scene from South America, as do Lamas (1981) from Peru, Willink (1969) from Argentina, Kevan (1977) from the West Indies, Barrera (1955) from Mexico, Fernández (1978) from Venezuela, and Jirón and Vargas (1986) from Costa Rica. See also Bodenheimer (1929) and Chardon (1949) for many basic notes and Howard (1930) for events in the origins and growth of practical entomology in most parts of Latin America. Gilbert (1977) provides an index to published biographies of deceased ento­ mologists. (In the following sketch, dates of birth and death of major figures are given in brackets []. The titles and publica­ tion dates of historically important works are woven into the text; because they are well known, they are not cited in the references.) Antiquity There are many evidences of pre-Colum­ bian appreciation for insects, arachnids, and myriapods among the classic civiliza­ tions of Mexico, Central America, and northwestern South America. Most refer­ ences are to species that affected human health and welfare. Surviving Mayan (Stempell 1908) and other ancient Mexican murals and codices depict various species of economic and religious importance, in­ cluding stingless bees, scorpions, and but­ terflies (Teotihuacán). Early Mexican pot­ tery, also from the Teotihuacán period

4

GENERAL ENTOMOLOGY

(A.D. 200-800), are adorned with insect designs, and early Mochica pottery from the northern coast of Peru shows human figures clearly engaged in delousing and infested with the chigoe. Other representa­ tions of insect forms appear in sculptures, petroglyphs, and textiles from various cul­ tures (Morge 1973, Tozzer and Allen 1910). Ancient languages and myths con­ tain many entomological allusions, espe­ cially to noxious or ubiquitous species, for example, in Náhuatl, Xochiquetzal, butter­ fly flower goddess (Beutelspacher 1976). Chroniclers With the arrival of Columbus, the insects of the New World became known to West­ ern civilization. One might speculate that the lights seen on the shores of Hispaniola, that night of October 11, 1492, were not native camp fires but glowing Pyrophorns beetles and thus that an insect was the first thing sighted in America: "After the Admi­ ral had spoken he saw the light once or twice and it was like a wax candle rising and falling" {J. First Voyage). Among the conquistadors and colonists who followed were scribes and reporters appointed by the Spanish crown to chroni­ cle the discoveries and bring the influence of Western thinking to the new settle­ ments. Many of the sixteenth-century tech­ nical reports of the natural wonders of the newfound lands contained references to insects. One of the earliest, the Historia General y Natural de las Indias, Islas y Tierrafirme del Mar Océano (first 20 volumes), was written by Gonzalo Oviedo in 1535. It described for the first time such American curiosities as the cucuyo (headlight beetle), chigger, chigoe, cochineal insect, and sting­ less bees. Mentions of the same conspicu­ ous species appeared in other, similar treat­ ments of the period, such as José de Acosta's Historia Natural y Moral de las Indias (1590) and Bernal Diaz's Historia Verdadera de la Conquista de la Nueva España (1568-

1632). Bernabé Cobo [1572-1659] wrote about white butterflies (Ascia monuste) that ittacked crops in Lima in his Historia del Suevo Mundo (1653), the most important work of the period on the natural history of Peru. Fray Bernardino de Sahagún [P-1590] (fig. 1.1) completed his Historia General de las Cosas de Nueva España in 1560, but it was not published until the early nineteenth century. It described many insects, arach­ nids, and myriapods and was later accom­ panied by illustrations originally intended for it, but from which the text was long separated {Codex Florentino, fig. 1.2.). The work explained how the Aztecs treated black widow spider bites and scorpion stings and made special mention of useful

Figure 1.2 Figure from Codex Florentino. The stinging arthropod is described by Sahagún in the early sixteenth century as a "scorpion," but in the figure, it is more similar to the larva of Corydalus cornutus (Megaloptera), called the "water dog" (perro del agua), an insect widely feared as venomous even today in Mexico.

insects such as the maguey worm and the cochineal bug (Curran 1937). Francisco García Hernández [15141578] collected natural objects of medical significance in early colonial Mexico, in­ cluding thirty insects and "worms." His manuscripts were published in various illus­ trated, annotated editions after his death, the best known being Rerum Medicarum Novae Hispaniae Thesaurus sev Planlarum Animalium Mineralium Mexicanorum Historia (published 1648-1651) (d'Ardois 195960, Barrera 1981) in which Tractatus Quarlus, Historia Insectorum Novae Hispaniae, was ostensibly the first unified work on Latin American insects.

Figure 1.1 Fray Bernardino de Sahagún, postconquest chronicler of insect life in the New World. (Frontispiece from Historia General de la Cosas de Nueva España, Edition Pedro Ro­ bredo, Mexico, 1938)

The chroniclers were savants not schooled in biology or in the methods of scientific investigation. Consequently, their statements sometimes contained consider­ able errors, these often the result of believ­ ing too literally the accounts of the Indians. But the firsthand recording of natural history by courtiers, travelers, explorers,

HISTORY OF LATIN AMERICAN ENTOMOLOGY

5

traders, soldiers, missionaries, historians, and adventurers continued in subsequent centuries (e.g., A. de Herrera, Historia Gen­ eral de las Indias Occidentalis [1728]) and remains a tradition even today (e.g., Jacques Cousteau's Amazon Journey [1984]). The creatures described tend to be the same as those described by earlier authors, although stinging ants, large centipedes, Mexican wild silkworms, and tarantulas also appear. Naturalists Following the chroniclers onto the scene were the naturalists, distinguished from the former by possessing some education in the biological sciences. An early example was George Marcgrave [1610—1644], who, during the Dutch invasion of Brazil in 1638—1644, traveled widely and studied insects in that country. An important edi­ tion of his works, citing many indigenous insects, was published in 1648 by one of his traveling companions, Guilielmus Piso, in De Indiae Utriusque re Naturali et Medica Libri XIV Following her ten-year stay (1690-1701) in Surinam, where she collected informa­ tion on insect life histories, Madame Maria Sybilla Merian [1647-1717] (fig. 1.3) pro­ duced her famous Metamorphoses Insectorum Surinamensium (1705), with superbly done color plates (Erlanger 1976). The work contained some errors, including a confu­ sion of the headlight beetles (Pyrophorus), cicadas, and dragon-headed bugs (Fulgora), that engendered misconceptions of the latter's ability to luminesce and stridulate which persist even today (one plate actually shows a mongrel insect, a cicada, bearing the head oí Fulgora) (Frontispiece). Later naturalists, following in this tradi­ tion and notable for significant observa­ tions on Latin American insects, were Hans Sloane, A Voyage to the Islands Madera, Barbadoes, Nieves, St. Christophers and Ja­ maica (1707, 1725); G. Gardner, Travels in

6

GENERAL ENTOMOLOGY

Figure 1.3 Maria Sybilla Merian, famous for her observations of insect natural history in the Guianas in the seventeenth century. (Frontis­ piece from her botanical work, Erucarum hortensis .. ., Amsterdam, 1718)

the Interior of Brazil (1849); Thomas Belt, A Naturalist in Nicaragua (1874); Theodore Roosevelt, Through the Brazilian Wilderness (1914); Konrad Guenther, A Naturalist in Brazil (1931); R. Hingston, A Naturalist in the Guiana Forest (1932); and others. Renaissance Scholars The first works on Latin American insects by those fully qualified as scientists were carried out by established European renais­ sance scholars in absentia. They received specimens and reports from collectors and correspondents on the scene but never set foot in the new lands themselves. Stingless bees were described in Konrad Gesner's encylopedic Historia Animaliurn (1607) but were referenced therein to a work by one Andre Thevet, who "amonst other matters [in the New-found World] reporteth that he

did see a company of flies of Honey-bees about a tree . . . : of which trees there were a great number in a hole that was in a tree, wherein they made Honey and Wax" (Topsel 1967)- In De Animalibus lnsectis (16021618), considered to be the world's first book on entomology, Ulisse Aldrovandi 11522-1605] wrote and figured some Latin American insects, including the cucuyo. This insect, by now famous, also appeared in Thomas Mouffet's Theatrum Insectorum (1634) alongside a rhinoceros beetle (Megasoma). Réné de Reaumur figures and de­ scribes in fine detail a dragon-headed bug (Fulgora) in his Memoir pour servir a I'histoire desinsectes (17M-1742). New World specimens were incorpo­ rated into the rapidly growing European collections of the time. Nehemiah Grew figures many from the cabinets of the Royal Society in England (Museum regalis societatis, 1685). Culminating this phase of historical development were the great taxonomists, Carolus Linnaeus [1707-1778] and J. C. Fabricius [1745-1808], who were able to include a large number of species from the American tropics in their landmark editions of Systema Naturae (1st ed., 1735; 10th ed., 1758) and various Systemas, respectively. Collectors Many of these descriptions were based on specimens provided by a new breed of naturalists to the region, the collectors. Some of the first, who made insectcatching trips to South America in the mid1700s, were Pehr Loefling [1729-1756], Carl Dahlberg [1721-1781], Daniel Ro­ under [1726-1793], and Daniel Solander [1733-1782]. As travel to and conditions in the colo­ nies improved, the number of collectors quickly increased, as did their range (La­ mas 1979, 1981; Papavero 1971, 1973). The majority of these individuals worked

independently, and many paid their ex­ penses through the sale of their collections to museums and private collectors in the United States and Europe. A famous duo of pioneer collectors was Karl von Martius [1794-1868] and Johann von Spix [1781-1826] (Fittkau 1983). In 1817-1820, they traversed much of eastern and Amazonian Brazil, collecting thou­ sands of insects that were studied subse­ quently by European specialists. A later pair were Osbert Salvin [1835-1898] and Fred­ erick Godman [1834-1919] who traveled widely and amassed specimens in Central America and Mexico in the late 1800s. Their extensive collections were assembled in London, systematically worked up by a number of entomologists, and published in a lengthy series of volumes under the title Biología Centrali-Americana (1879-1915), a classic faunal report. Some of the liberes of the infamous French penal colonies in French Guiana from the late 1800s to 1938 made a living by supplying foreign markets with butterflies from the local jungles (Le Moult 1955). Today, many collectors, both commercial and voluntary, continue to pro­ vide material to specialists in their own and other countries. Scientific Expeditions Other collectors and naturalists partici­ pated in or led the organized scientific or biological expeditions that are a very im­ portant part of the growth of entomologi­ cal science in Latin America. These were often sponsored by governments or agen­ cies and included multiple investigators, each with specialized assignments, and were much more elaborate than the simple forays of individuals. Primary examples are numerous and date from the early eighteenth century. Antonio de Ulloa [1716-1795] was a military man appointed as Spanish crown representative to the French Académie de Science Expedition to South America in

HISTORY OF LATIN AMERICAN ENTOMOLOGY

7

1735—1746 with La Condamine to mea­ sure the length of an arc of the meridian at the equator. His Noticias Americanas (1772) contained specific mentions of equatorial insect life, including an account of a locust plague. The monumental expedition of the times, however, must be that of Baron Alexander von Humboldt [1769-1859] and Aimé Bonpland [1773-1858] to ex­ plore northern South America and Mexico in 1799-1804. Their extensive insect collec­ tions were researched by Latreille in Eu­ rope (Papavero 1971, chap. 4). Other exemplary expeditions that fur­ thered entomology in Latin America were several sea voyages with frequent land stops for collecting, such as the expeditions of the French vessel La Coquille (18221823), the Swedish Engentes (1851-1852), and the Austrian Novara (1857-1859). Of special interest also were the Hamburger Südperu Expedition in 1936 (Titschack 1951-1954) and the Machris Brazilian Ex­ pedition of 1956 (entomologist F. Truxal; Delacour 1957). A modern example is the report of entomological results of the 1978-79 Danish Expedition to Patagonia and Tierra del Fuego (Madsen et al. 1980). Later came those expeditions under­ taken by investigators trained in biology or zoology who emphasized work with insects and who conducted studies on their own collections. Three categories of investiga­ tors may be recognized: visiting, expatri­ ate, and native. Visiting Biologists and Zoologists Perhaps the first biologist to produce sig­ nificant entomological results from his own excursions in Latin America was Clau­ dio Gay [1800-1873] (fig. 1.4), an ambi­ tious French traveler who began to work with Chilean insects as early as 1836. Later, he published many research papers, his most important being Historia Física y Po­ lítica de Chile (arthropod portions, 18491852). About the same time, Charles Darwin

8

GENERAL ENTOMOLOGY

Figure 1.4 Claudio Gay, born in France but first trained biologist to make a major contribution to Latin American entomology through his work in Chile. (From portrait in Universidad de Chile; courtesy of José Valencia)

[1809-1882] made his epic global voyage that included major sojourns in South America. He was inclined toward entomol­ ogy and gained some insights into his revo­ lutionary theory of natural selection from observations of South American insects. In the initial sentences of his introduction to the Origin of Species (1859), he states, "When on board H.M.S. 'Beagle,' as naturalist, I was much struck with certain facts in the distribution of the organic beings inhabit­ ing South America . . . [which] throw some light on the origin of species." Some of these facts concerned the distribution of insects and insect examples of sexual se­ lection (e.g., the Chilean stag beetle, Chiasognatkus) much elaborated in his later Descent of Man (1871). Also celebrated among itinerant biolo­ gists of this period was Henry Walter Bates [1825-1892] (fig. 1.5). He spent eleven years on the Amazon and its tributaries

Figure 1.5 Henry Walter Bates, first entomolo­ gist explorer of the Amazon Valley in the mid18005. (Frontispiece from The Naturalist on the River Amazons, John Murray, London, 1892)

(1848-1859) and collected some 14,000 specimens, including 8,000 species new to science. For the first five years of his travels, he was accompanied by Alfred Russel Wallace [1823-1913], who was also an avid collector but who chose to continue his studies in the Malay Archipelago where he produced his own theory of natural selection paralleling Darwin's. Wallace re­ counts his South American experiences in A Narrative of Travels on the Amazon and Rio Negro (1853); Bates recounts his in a later parallel work, The Naturalist on the River Amazons (1892). Bates collected, but he also observed and analyzed, producing many papers on Neotropical Coleóptera. The work that distinguished him as an ento­ mologist was his Contributions to an Insect Fauna of the Amazon Valley, Lepidoptera: Heliconidae, published in 1862 (Moon 1976). The period of the early to mid-1800s was

a time of independence for most of the countries of Latin America and establish­ ment of national universities, museums, and other learned institutions, with depart­ ments paying attention to terrestrial arthro­ pod life forms. Many scholars from Europe emigrated to Latin America. Biology came of age, and considerable progress was made in the academic phases of entomology, pri­ marily insect systematics. But agricultural and medical entomology, knowledge of pes­ ticides, and the role of insects as vectors of disease awaited the threshold of the twenti­ eth century. Other visiting biologists of note were William Beebe [1877-1962], who was gifted with an extraordinary ability of expression and published on many aspects of Neotropical insect biology, for example, High Jungle (1949) (Berra 1977), and A. S. Calvert, who produced works on Costa Rican insects, including the book, written with his wife, A Year of Costa Rican Natural History (1917). Expatriate Biologists A special group of early biologists who worked on insects were expatriates. They were trained in Europe or North America but were drawn to the Neotropics by its exotic and poorly known insect life. They brought with them their education from Western schools and did not merely travel to the New World but spent the rest of their days in their adopted homes. Deserv­ ing special mention in this category is Fritz Müller [1822-1897], born in Germany, who settled in Blumenau, Santa Catarina, Brazil, in 1852. He was a correspondent of Darwin and is best known for his discovery of the type of mimicry named after him. Another outstanding expatriate biologist was German-born Hermann Burmeister [1807-1892]. After a sojourn in Brazil (1850), he settled in Argentina and became the director of the natural history museum in Buenos Aires and published many im­ portant entomological papers. A more mod-

HISTORY OF LATIN AMERICAN ENTOMOLOGY

9

ern example is Felix Woytkowski [18921966], who migrated from his native Po­ land to Peru in 1929. There he collected more than a thousand insect species new to science (Woytkowski 1978). Other notable expatriate entomologists were Hermann von Ihering [1850-1930], who emigrated from Germany to Brazil, Emilio Goeldi [ 1859-1917], Switzerland to Brazil, Adolfo Lutz [1855-1940], Germany to Brazil, Paul Biolley [1862-1909], Switzerland to Costa Rica, and Henri Pittier [1857-1950], Swit­ zerland to Costa Rica. Native Zoologists The first full-fledged biologist specializing in insects who was born in Latin America was Cuban Felipe Poey [1799-1891] (fig. 1.6). He left his birthplace, La Habana, to study in France but returned to produce works in entomology in his own land, especially on Lepidoptera (Centuria de Lepi-

Figure 1.6 Felipe Poey, first native-born Latin American entomologist. (From Memorias de la Sociedad Cubana de Historia Natural, 1, facing p. 8, 1915)

10

GENERAL ENTOMOLOGY

dópleros de Cuba, 1847). Clodomiro Picado [1887-1944] also left for study in France, completing his doctoral dissertation on Neotropical bromeliad communities. He returned to his native Costa Rica to become its most famous biologist. The Argentinian Arribálzaga brothers, Felix [1854-1894] and Enrique [1856-1935], were educated in their homeland where they carried out extensive studies on insect biology and taxonomy, especially on Diptera. The Entomologists By reason of their generalized training, the biologists and zoologists could not be con­ sidered entomologists in the strict sense. But because of their scientific abilities, interest, and emphasis on investigation and publication, the title could be logically bestowed on them. Full curricula in the discipline of ento­ mology were not offered in universities until the very late nineteenth century, so professionals in the study of insect biology are virtually all twentieth-century prod­ ucts. Their numbers now range in the thousands. Who they are and the nature of their accomplishments are best appreci­ ated by reference to the modern literature and bibliographies such as the Zoological Record, Parts lnsecta, Arachnida, and Myriapoda. The amateur entomologist deserves some special recognition here. An active cadre of educated and often highly sophis­ ticated individuals exists who find pleasure in the study of insects. Most are collectors, perhaps the majority working with showy insects like butterflies and beetles, but not always merely for the sake of accumulating specimens. Many take advantage of finan­ cial security acquired in other enterprises to pursue serious questions in entomology. They may even find time to carry out investigations for which the professional finds no support and make valuable contri­ butions directly in their own publications

or in collaboration with professionals. They are therefore distinct from the deal­ ers, whose primary aim in collecting is to profit financially from the sale of their catches. Practical Entomology Mention of diverse pestiferous Latin Amer­ ican insects is common in the earliest chronicles and later works. The first refer­ ence to control was made by Francisco Hernández in his Historia de los Insectos de Nueva España, written in manuscript in about the mid-sixteenth century and stat­ ing that the Mexicans used a concoction of tobacco that they spread over the walls to kill fleas in a house (Hoeppli 1969:177). It may have been Henry Hawks, a Vera Cruz (Mexico) merchant, who provided one of the earliest clues to the connection between mosquitoes and human disease, when he wrote in 1572, "This towne is inclined to many kinde of diseases, by reason of the great heat, and a certeine gnat or flie which they call a musquito, which biteth both men and women in their sleepe. . . . Many there are that die of this annoyance" (Keevil 1957). While evolutionary and taxonomic stud­ ies of insects continued following the birth of scientific entomology and expanded into the early twentieth century, the discovery of arthropod vectors of animal and human disease and the development of chemical control of crop pests fostered increased work in the applied phases of entomology in Latin America. In medical entomology, major strides were made in the battle against yellow fever and malaria because of the newfound knowledge that mosquitoes were the critical link in the spread of these diseases. It was the application of entomo­ logical principles by physicians Carlos Finlay [1833-1915] (fig. 1.7), Walter Reed [1851-1902], and William Gorgas [18541920] which rid La Habana of yellow fever in 1901 and made possible the construction

Figure 1.7 Carlos Finlay, whose ideas led to the mosquito's role in transmission of yellow fever. (From a portrait formerly hung in the Regional Office of the Pan American Health Organization, Washington, D.C.; presently owned by Dr. J. Fermoselle-Bacardi, Coral Gables, Florida. Re­ produced with owner's permission)

of the Panama Canal shortly thereafter (Le Prince et al. 1916, McCullough 1977). In 1909, Carlos Chagas [1879-1934] demon­ strated that a lethal trypanosome parasite of humans (Trypanosoma cruzi) was transmit­ ted by a kissing bug (Panstrongylus megistus). Modern knowledge of agricultural ento­ mology (Doreste et al. 1981, Howard 1930), the identification and control of crop pests, was primarily imported, work­ ers and technology in Europe and North America largely determining the course of events. Although numerous references to pest insects are scattered throughout the literature of the sixteenth, seventeenth, and eighteenth centuries (Kevan 1977), possibly the earliest scientific investigation into what could really be called economic entomology was made in 1801 when a special commission composed of members

HISTORY OF LATIN AMERICAN ENTOMOLOGY

11

of t h e G e n e r a l Assembly of t h e B a h a m a s was sent to t h e West I n d i e s to look into d a m a g e d o n e to cotton by r e d bugs (Dysdercus) a n d t h e chenille (Alabama argillacea). In 1870, B. Pickman M a n n , of Cam­ bridge, was sent to Brazil with a commis­ sion a u t h o r i z e d by D o m P e d r o to study t h e country's n a t u r a l history. H e specialized in coffee a n d maize insects a n d p r e p a r e d r e p o r t s o n each. I n 1897, t h e Comisión para la Extinción la L a n g o s t a (antilocust commission) was established in A r g e n t i n a , the first of m a n y similar agencies, with a p p o i n t e d entomologists, to be f o r m e d in most c o u n t r i e s d u r i n g t h e early p a r t of t h e twentieth century. A m o n g t h e pioneers were W. H . T. T o w n s e n d a n d J o h a n n e s Wille in Peru, G e o r g e Wolcott in P u e r t o Rico, a n d G. E. B o d k i n in British Guiana. T h e earliest book o n agricultural entomol­ ogy was Las Epidemias de las Plantas en la Costa del Perú by M a n u e l García y Merino (1878). Parasitoids a n d p r e d a t o r s of several pests were i n t r o d u c e d into p r o b l e m areas with varying results by t h e 1930s (Myers 1931), a n d several sites b e c a m e t h e scene of i m p o r t a n t e x p e r i m e n t a l trials in biologi­ cal control ( H a g e n a n d F r a n z 1973). H o p e s were realized in t h e Brazilian A m a z o n fly (Melagonislylum mínense, Tachinidae) for control of s u g a r c a n e m o t h s . T h e sterile male t e c h n i q u e for t h e control of screwworm was first tested successfully o n t h e island of C u r a c a o in 1954. Notes o n t h e history of t h e various insects of commercial value in Latin A m e r ­ ica a r e t o be f o u n d in t h e systematic portion of this book (see cochineal insects, silk m o t h , stingless bees, h o n e y b e e , etc.).

References BARRERA, A. 1955. Ensayo sobre el desarrollo histórico de la entomología en México. Rev. Soc. Mex. Entomol. 1: 23-38. BARRERA, A. 1981. Notas sobre la interpretación de los artrópodos citados en el tratado cuarto, Historia de los insectos de Nueva España, de

12

GENERAL ENTOMOLOGY

Francisco Hernández. Fol. Entomol. Mex. 49: 27-34. BERRA, T. M. 1977. William Beebe, an anno­ tated bibliography. Archon, Hamden, Conn. BEUTELSPACHER, C. R. 1976. La diosa Xochiquetzal. Soc. Mex. Lepidop. Bol. Inf. 2: 1-3. BODENHEIMER, F. S. ¡929. Materialien zür Geschichte der Entomologie bis Linné. 1: 1 498; 2: 1-486. Junk, Berlin. CHARDON, C. E. 1949. Los naturalistas en la

América Latina. Sec. Agrie, Pec. Col., Rep. Dominicana, Ciudad Trujillo. CURRAN, C. H. 1937. Insect lore of the Aztecs. Nat. Hist. 39: 196-203. DARDOIS, G. S. ed. 1959-60. Francisco Hernán­ dez, Obras completas. Univ. Nac. México, Mexico City. Vol. 1. 1960, Vida y obra de Francisco Hernández, España y Nueva España en la época de Felipe II by José Miranda. DELACOUR, J. 1957. T h e Machris Brazilian Ex­ pedition: general account. Los Angeles Co. Mus. Contrib. Sci. 1: 1-11. DORESTE, E., F. FERNÁNDEZ, AND P. P PAREDES.

1981. Contribución a la historia de la entomología agrícola en Venezuela. 5th Cong. Venezolano Entomol. (Maracay), Mem. Pp. 29-50. ERLANGER, L. 1976. Maria Sybilla Merian, seventeenth-century entomologist, artist and traveler. Ins. World Dig. 3(1): 12-21. FERNÁNDEZ, F. 1978. Contribución a la historia de la entomología en Venezuela. Red. Fac. Agron. (Maracay) 26: 11-27. FITTKAU, E. J. 1983. Johann Baptist Ritter von Spix, sein Leben und sein wissenschaftliches Werk. Spixiana suppl. 9: 1 1-18. GILBERT, P. 1977. A compendium of the bio­ graphical literature on deceased entomolo­ gists. Brit. Mus. Nat. Hist., London. HAGEN, K. S., ANDJ. M. FRANZ. 1973. A history

of biological control, in R. F. Smith, T. E. Mittler, and C. N. Smith, eds., History of entomology. Annual Reviews, Palo Alto. Pp. 433-476. HOEPPLI, R. 1969. Parasitic diseases in Africa and the Western Hemisphere, early documen­ tation and transmission by the slave trade. Acta Trop. suppl. 10: 1-240. HOWARD, L. O. 1930. A history of applied entomology. Smithsonian Misc. Coll. 84: 1 564. (See Pt. VI, South and Central America and the West Indies, 417-462.) JIRÓN,

L. F., AND R. G.

VARGAS.

1986. La

entomología en Costa Rica: Una reseña his­ tórica. Rev. "Quipu" de Historia de la Ciencia (México) 3(1): 67-77.

KEEVIL, J. J. 1957. Medicine and the Navy 1200-1900. Vol. 1. E. and S. Livingstone, Edinburgh. KLVAN, D. K. McE. 1977. Mid-eighteenth-

century entomology and helminthology in the West Indies: Dr. James Grainger. Soc. Bibliog. Nat. Hist. J. 8: 193-222. LAMAS, G. 1979. Otto Michael (1859-1934), el cazador de mariposas del Amazonas. Col. Suiza Perú Bol. 1979(2): 35-38. LAMAS, G. 1981 [1980]. Introducción a la histo­ ria de la entomología en el Perú. Rev. Peruana Entomol. 23: 17-37. LE MOULT, E. 1955. Mes chasses aux papülons. Ed. Pierre Horay, Paris. LE PRINCE, J. A., A. J. ORENSTEIN, AND L. O.

HOWARD. 1916. Mosquito control in Panama. Putnams, New York.

footed beasts and serpents and insects. Vol. 2. Da Capo, New York. TOZZER, A. M., AND G. M. ALLEN. 1910. Animal

figures in the Maya Codices. Harvard Univ., Peabody Mus. Pap. Amer. Archaeol. Ethnol. 4: 273-372, pis. 1-39. WILLINK, A. 1969. Contribución a la historia de la entomología Argentina. Univ. Nac. Tucumán, Fund. Insto. Miguel Lillo Mise. 28: 1-30. WOYTKOWSKI, F. 1978. Peru, my unpromised land. Smithsonian Inst. and Nati. Sci. Found., Washington, D.C.

LATIN AMERICAN ENTOMOLOGY TODAY

LIZER Y TRELLES, C. A. 1947. Introducción e

historia de la entomología. Argentino Cien. Nat. "Bernardino Rivadavia," Publ. Ext. Cul. Didac. 1 (Curso de entomología) (Buenos Aires), 1-52. (See p. 20ff, Historia de la entomología sudamericana.) MCCULLOUGH, D. 1977. T h e imperturbable Dr.

Gorgas. In D. McCullough, T h e path between the seas: T h e creation of the Panama Canal 1870-1914. Simon & Schuster, New York. Pp. 405-426. MADSEN, H. B., E. S. NIELSEN, AND S. ODUM,

eds. 1980. T h e Danish scientific expedition to Patagonia and Tierra del Fuego 1978-1979. Geogr. Tidssk. 80: 1-28. MOON! H. P. 1976. Henry Walter Bates F.R.S. 1825-1892, Explorer, scientist and Darwin­ ian. Leicestershire Museums, Leicestershire. MORGE, G. 1973. Entomology in the Western world in antiquity and in medieval times. In R. F. Smith, T E. Mittler, and C. N. Smith, eds., History of entomology. Annual Reviews, Palo Alto. Pp. 37-80. MYERS, J. G. 1931. A preliminary report on an investigation into the biological control of West Indian insect pests. Empire Mrkt. Bd. Publ. 42:1-173. PAPAVERO, N. 1971, 1973. Essays on the history of Neotropical dipterology, with special refer­ ence to collectors (1750-1905) 2 vols. Mus. Zool. Univ. Sao Paulo, Sao Paulo. STEMPELL, W. 1908. Die Tierbilder der Mayahandschriften, Zeit. Ethnol. 1908: 704-743. TITSCHACK, E., ed. 1951-1954. Beitrage zür Fauna Perus: Nach der Hamburger SüdperuExpedition 1936, anderer Sammlungen, wie auch auf Grund von Literaturangaben. Vols. 1-4. Fischer, Jena. TOPSEL, E. 1967 [1658]. T h e history of four-

We have seen h o w t h e f o u n d a t i o n s of entomology in Latin America were laid, t h r o u g h four centuries of effort by m a n y types of investigators: chroniclers, general observers, renaissance scholars, collectors, and trained entomologists. By t h e m i d d l e of t h e twentieth century, t h e r e was a firm establishment of t h e t r e n d toward special­ ization, b e g u n first with t h e choice of systematists to study certain limited taxa, followed by t h e separation of t h e applied agricultural a n d m e d i c o v e t e r i n a r y fields a n d m a t u r a t i o n of t h e principles of insect ecology a n d genetics. An i m p o r t a n t m o d e r n specialization has been t h e s t r o n g interest in tropical biology by a large n u m b e r of local s t u d e n t s a n d y o u n g entomologists from N o r t h America a n d E u r o p e . T h e O r g a n i z a t i o n for Tropi­ cal Studies has b e e n a p r i m e m o v e r in this area, principally t h r o u g h t r a i n i n g at field stations in Costa Rica. F u n d a m e n t a l discov­ eries a r e now being m a d e about t h e ecologi­ cal a n d evolutionary strategies of insects in the h u m i d lowland e n v i r o n m e n t s , for ex­ ample, t h e theory of Pleistocene relictual centers of distribution (Brown 1982), t h e theory of island b i o g e o g r a p h y ( M a c A r t h u r and Wilson 1967), a n d t h e organization of insect societies (Wilson 1971). T h e vindication of t h e idea of continen­ tal drift (largely from data collected d u r -

LATIN AMERICAN ENTOMOLOGY TODAY

13

ing t h e I n t e r n a t i o n a l Geophysical Year, 1957—1958) h a s o p e n e d t h e d o o r to u n ­ d e r s t a n d i n g m a n y e n i g m a t i c distributions of Latin A m e r i c a n insect species. T h e s e h a d b e e n e x p l a i n e d by i m p r o b a b l e over­ sea dispersals a n d land b r i d g e s . A m p h i notic p a t t e r n s of austral forms a r e n o w best e x p l a i n e d by their origination o n t h e composite c o n t i n e n t , G o n d w a n a l a n d . A l a n d m a r k study in this a r e a is Lars B r u n d i n ' s , Transantarctic Relationships and Their Significance, as Evidenced by Chironomid Midges ( B r u n d i n 1967). O p e r a t i o n a l i s m (analysis a c c o r d i n g to a prescribed a n d consistent p r o c e d u r e ) h a s now b e c o m e a c a n o n of p h y l o g e n e t i c stud­ ies o n Neotropical g r o u p s , t h a n k s to t h e logic forced into t h e p r o c e d u r e s of t a x o n o mists by t h e a p p r o a c h e s of phenetics, p r o ­ m o t e d in R. Sokal a n d P. H . A. Sneath's, Principles of Numerical Taxonomy (1963), a n d cladistics, f o u n d in W. H e n n i g ' s , Grundziige einer Theorie der Phylogenetischen Systematik (1950). "Trees of d e s c e n t , " o r phyletic dendrograms, now are superimposed on p a t t e r n s of origin a n d d i s p e r s i o n ("area d i a g r a m s " ) . Also of g r e a t p r o m i s e in t h e study of insect systematics, physiology, a n d genetics a r e newly d e v e l o p e d molecular a n d biochemical t e c h n i q u e s for o b t a i n i n g c o m p a r a t i v e d a t a from insect tissues for nucleic acid sequences, metabolic enzymes a n d o t h e r p r o t e i n s , i m m u n o l o g i c a l compati­ bility, a n d g e n e p r o b i n g (Hillis a n d Moritz 1990). T h e m o d e r n m i n i a t u r i z a t i o n a n d i m p r o v e d availability of c o m p u t e r s a n d m a t h e m a t i c a l t e c h n i q u e s for analysis of m o r p h o l o g y a n d distributions h a s h e l p e d immensely in m a k i n g t h e u s e of these logical p r o c e d u r e s feasible. Specific e n t o ­ mological e x p e d i t i o n s t o r e m o t e a n d u n e x ­ plored areas a n d fieldwork by b o t h local a n d foreign specialists a r e m a k i n g possible discoveries in all g e o g r a p h i c a n d study areas (e.g., M a d s e n et al. 1980). Today, e n t o m o l o g y c o n t i n u e s t o b e a rapidly d e v e l o p i n g field of science, seeking to u n d e r s t a n d t h e diversity of insect life in

14

GENERAL ENTOMOLOGY

Latin America a n d its significance to h u ­ m a n k i n d . I n most every country, g o v e r n ­ m e n t s a n d private e n t e r p r i s e a r e recogniz­ ing t h e i m p o r t a n c e of insect forms a n d e m p l o y i n g entomologists. T h e r e h a s b e e n an increase in t h e n u m b e r s of positions filled by local g r a d u a t e s , a l t h o u g h w o r k e r s trained o r i m p o r t e d from o t h e r countries still fill m a n y posts. E d u c a t e d a m a t e u r s also r e m a i n i m p o r t a n t c o n t r i b u t o r s . Gradually, with t h e h e l p of new technologies for acquir­ ing, r e c o r d i n g , a n d d i s p e n s i n g k n o w l e d g e , the f a u n a is b e c o m i n g k n o w n , local biologi­ cal p h e n o m e n a a r e being revealed a n d integrated into universal schemes, losses from destruction of food a n d fiber a n d disease a r e being r e d u c e d , a n d a n a p p r e c i a ­ tion for t h e value of insects a n d their a r t h r o p o d relatives is being realized. T h e need r e m a i n s for m o r e facilities a n d fuller staffing of research institutions a n d g r e a t e r local activity, i n c l u d i n g t h e p o p u l a r i z a t i o n of insect n a t u r a l history t o t h e g e n e r a l public.

References BROWN, JR., K. S. 1982. Paleoecology and re­

gional patterns of evolution in Neotropical forest butterflies. In G. T. Prance, ed., Biologi­ cal diversification in the tropics. Columbia Univ. Press, New York. Pp. 255-308. BRUNDIN, L. 1967. Insects and the problem of austral disjunctive distribution. Ann. Rev. Entomol. 12: 149-168. HENNIG, W. 1950. Grundziige einer Theorie der Phylogenetischen Systematik. Deutcher Zentralverlag, Berlin. HILLIS, D. M., AND C. MORITZ. 1990. Molecular

systematics. Sinauer Associates, Sunderland, Mass. MACARTHUR, R. H., AND E. O. WILSON. 1967.

The theory of island biogeography. Princeton Univ. Press, Princeton. MADSEN, H. B., E. S. NIELSEN, AND S. ODUM,

eds. 1980. T h e Danish scientific expedition to Patagonia and Tierra del Fuego 1978—1979. Geogr. Tidssk. 80: 1-28. SOKAL, R. R., AND P. H. A. SNEATH.

1963.

Principles of numerical taxonomy. Freeman, San Francisco. WILSON, E. O. 1971. T h e insect societies. Har­ vard Univ. Press, Cambridge.

d u r a b l e a n d resistant because of its c o m p o ­ sition of w a t e r p r o o f waxes a n d c o m p l e x molecules of such substances as chitin (a n i t r o g e n o u s polysaccharide) a n d sclerotin (protein). In combination with t h e e p i d e r ­ mis, it forms t h e insect's i n t e g u m e n t ( H e p ­ b u r n 1976). T h e i n t e g u m e n t m a y b e generally thin a n d flexible, as in insect larvae, o r thick a n d rigid, as in most adults a n d larval structures such as t h e h e a d . Rigidity is t h e result of t h e a b u n d a n c e of sclerotin, a n d h a r d areas, o r "sclerites," a r e said to be well sclerotized. Flexibility is allowed by m e m ­ b r a n o u s joints o r articulations b e t w e e n t h e rigid portions. T h u s , t h e i n t e g u m e n t gives the insect its basic form a n d is its p r i m a r y protective system, f o r m i n g a b a r r i e r t o water loss a n d e n t r y of p a t h o g e n i c microor­ ganisms as well as p r o v i d i n g resistance t o physical t r a u m a .

INSECT STRUCTURE AND FUNCTION T h e physical f o r m a n d b o d y workings (Snodgrass 1 9 3 5 , 1 9 5 2 ; W i g g l e s w o r t h 1984; C h a p m a n 1982; K e r k u t a n d Gilbert 1984; King a n d Akai 1 9 8 2 - 1 9 8 4 ; M a n t ó n 1977; Rockstein 1 9 6 4 - 1 9 7 4 , 1978; Smith 1968; T r e h e r n e et al. 1 9 6 3 - 1 9 8 2 ) of insects a n d their terrestrial a r t h r o p o d relatives a r e as r e m a r k a b l e a n d c o m p l e x as those of a n y animal type. N u m e r o u s structural a n d func­ tional systems will be used in t h e text following as organizational topics for a basic review.

References CHAPMAN, R. F. 1982. T h e insects: structure and function. 3d ed. American Elsevier, New York. KERKUT, G. A., AND L. I. GILBERT. 1984. Com­

prehensive insect physiology, biochemistry and pharmacology. Vols. 1 — 13. Pergamon, Elmsford, N.Y KING, R. C , AND H. AKAI, eds.

1982-1984.

Insect ultrastructure. Vols. 1—2. Plenum, New York. MANTÓN, S. M. 1977. T h e Arthropoda, habits, functional morphology, and evolution. Cla­ rendon, Oxford. ROCKSTEIN, M., ed. 1964-1974. T h e physiology of insects. Vols. 1—6. Academic, New York. ROCKSTEIN, M., ed. 1978. Biochemistry of in­

sects. Academic, New York. SMITH, D. S. 1968. Insect cells: Their structure and function. Oliver and Boyd, Edinburgh. SNODGRASS, R. E. 1935. Principles of insect morphology. McGraw-Hill, New York. SNODGRASS, R. E. 1952. A textbook of arthro­ pod anatomy. Comstock, Ithaca. TREHERNE, J. E., M. J. BERRIDGE, AND V B.

WIGGLESWORTH. 1963-1982. Advances in in­ sect physiology. Vols. 1-16. Academic, New York. WIGGLESWORTH, V. B. 1984. Insect physiology. 8th ed. Chapman and Hall, London.

Integument T h e outer, living e p i d e r m i s in insects is a single layer of generally simple, cuboidal cells that secrete a n e x t e r n a l nonliving cuticle. T h e cuticle (Neville 1975) is very

Sclerites may also be s e p a r a t e d by infoldings, k n o w n as a p o d e m e s (if linear, called sutures; if pitlike, apophyses). It is t o t h e internal portions of a p o d e m e s that t h e main muscles of m o t i o n a r e a t t a c h e d , giv­ ing t h e i n t e g u m e n t a s e c o n d a r y function, that of a skeleton (exoskeleton). T h e cuticle derives its color n o t only from its structural c o m p o n e n t s b u t from infusions of p i g m e n t s ( C r o w m a r t i e 1959) a n d microstructural d e v e l o p m e n t s (lamel­ lae, gratings, etc.) that cause scattering, refraction, a n d defraction of light waves striking t h e m , resulting in spectral p h e n o m ­ ena. A m o n g t h e p i g m e n t s a r e c o m m o n colored c o m p o u n d s such as m e l a n i n (black), pterines (white, r e d , yellow), carotenoids ( r e d , b r o w n ) , carminic acid (carmine), a n d flavones (red, yellow). Physi­ cal colors a r e often metallic o r iridescent blues, greens, a n d r e d d i s h h u e s . Many Neotropical butterflies a r e beautifully col­ o r e d from combinations of b o t h pig­ m e n t a r y a n d physical colors localized in t h e wing scales (Ghiradella 1984). Gold a n d silver a r e i n t e r f e r e n c e colors also, b u t u n ­ like t h e o t h e r metallics, which a r e p r o d u c e d

INSECT STRUCTURE AND FUNCTION

15

by pure, narrow wavelengths, these are broad-band reflective mixes of radiation (Neville 1975, 1977). By providing a sur­ face for display of color patterns, the integument serves additional functions— protection by crypsis and mimicry, sexual recognition, and so forth. References CROWMARTIE, R. 1. T. 1959. Insect pigments. Ann. Rev. Entomol. 4: 59-76. GHIRADELLA, H. 1984. Structure of iridescent lepidopteran scales: Variations on several themes. Entomol. Soc. Amer. Ann. 77: 637645. HEPBURN, H. R., ed. 1976. The insect integu­ ment. Elsevier, New York. NEVILLE, A. C. 1975. Biology of the arthropod cuticle. Springer, New York. NEVILLE, A. C. 1977. Metallic gold and silver colours in some insect cuticles. Ins. Physiol. 23:1267-1274. Body Cavity The body cavity of all arthropods is not considered a true coelom, as it lacks a complete mesodermal lining. Morphologists call it a "mixocoel" because of its foi ■ ation embryologically from the fusion of tho blastocoel with parts of the secon­ dary be '- cavity. Because it is filled with blood, emptying into it from an openended circulatory system, it is also known as a hemocoel. Segmentation From their annelid and marine arthropod ancestors, insects and their terrestrial rela­ tives have inherited a segmented body. Between an anteriormost (acron) and pos­ teriormost segment (telson), a varying number of segments are interposed, de­ pending on the group. There were origi­ nally 18 (or 19, if a second antennal segment is recognized) in insects, 19 in chelicerates, and as many as 100 in myriapods. These were more or less equal in form and in the possession of a pair of walking appendages in the first terrestrial

16

GENERAL ENTOMOLOGY

arthropods, much like modern-day centi­ pedes. Evolution eventually favored the fusion of adjoining segments (a process called tagmosis) for various functional pur­ poses (e.g., flight in higher insects), and body regions were formed. Of these, in­ sects display a triple set, including the head (composed of the acron plus four or five highly fused original segments), a thorax (of three segments, pro-, meso-, and metathorax) and abdomen (with eleven segments, the posteriormost being highly modified into genitalia). Arachnids and myriapods show differ­ ent patterns of fusion. In the former, the head is undefined and its segments totally incorporated into the thorax (cephalothorax), which itself may also join into the abdomen, as in mites and ticks. Seg­ mentation in these anterior two regions is concealed by a shield or carapace and is evident ventrally only by the serial set of appendages. The abdomen is either unde­ fined or formed from several segments. Myriapods display only a well-developed head and uniformly segmented thoraxabdomen, with each segment bearing simi­ lar legs. Uniquely, the insect thorax may bear a pair of wings on the meso- or metathorax, or both, but never on the prothorax. Thus, according to the appendages they possess, the three body regions are specialized for separate functions, the head for ingestión and perception, the thorax for locomotion, and the abdomen for metabolic processes and reproduction. The Head and Its Appendages The head (Matsuda 1965) is the most highly modified body region, being a sepa­ rate organ (except in arachnids), in which the primitive segmentation is almost oblit­ erated. It is normally a rigid capsule, containing the main perceptive and integrative neural elements of the animal as well as ingestive organs. The many sensory appendages of the

head include the antennae in insects, centi­ pedes, and millipedes, all with one pair. Arachnids lack antennae, their place usu­ ally being assumed by the pedipalps that have become antennalike. However, in some arachnids, the pedipalps take other forms and functions, as the claws of scorpi­ ons or walking legs in sun spiders. Around the mouth, modified segmented appendages serve as jaws or stylets for chewing or imbibing liquids and bear foodtasting and smelling organs called palpi. In arachnids, these organs are the chelicerae, with the basic scissor form, but they are used directly in feeding by tearing or stab­ bing the food, not chewing. The chelicerae may lose the movable element and become a piercing needle in mites, especially para­ sitic ones, and in spiders they are modified into fangs. Arachnids use the inner portion of the leg coxae to scoop liquid nutrient into the simple mouth. Among insects and milli­ pedes, there are two pairs of jaws, the anterior mandibles and behind them the maxillae; centipedes have two sets of maxil­ lae. In insects, the mandibles and maxillae may retain a primitive toothed or molar form for biting and chewing solid foods, or they may be greatly elongated and bladelike or hypodermiclike for piercing and siphon­ ing. The labium may form only a support­ ive sheath around the latter or be itself spongy and absorptive and act directly in food collection. Flexibility in adaptation of mouth parts has been a major factor in the success of insects as a group, the variety of morphological types making possible an enormous diversity of food niches and feeding strategies. Although notof appendicular origin, the eyes are of major sensory importance to the head capsule (Horridge 1975). There is a pair of larger multifaceted compound eyes in adult insects laterally and usually one to three smaller, single-faceted simple eyes medially on top of the head. In other terrestrial arthropod groups and immature insects, only simple eyes (ocelli) are present,

either in lateral clumps on the sides of the head (millipedes) or on the back of the cephalothorax (arachnids). Eyes are also often absent altogether (many centipedes). The structure and function of the com­ pound eyes are complex. They bulge out on either side of the head to give a wide range of vision in all directions. Each is an aggregation of similar rod-shaped facets called ommatidia, the number of which varies from one per eye in some ants to over 10,000 in dragonflies. Each ommatidium is composed of elongate sensory cells containing light-sensitive pigments, these concentrated toward the center (thus seen as a dark rod, called the rhabdome) and exposed on the exterior through a cap­ ping, duplex lens that gathers and focuses light. There are also cells with diffuse pigment around the lens. The sensory cells are nerve cells and are connected directly to the brain, there being no optic nerve in insects. There are many variations in the de­ tailed structure of the ommatidium, such as the "apposition" versus the "superposi­ tion" types. In the former, the rhabdome is long, and the diffuse pigment cells isolate each ommatidium. In the latter, the rhabdome is short, and the screening pig­ ment moves depending on the amount of light in the environment. Image formation is believed to be basically different in the two types, but little is certain about this aspect of eye function. It is known that insects generally have good visual acuity and light level accommodation. Wave­ length discrimination varies considerably, with a tendency toward the ultraviolet in many species (Silberglied 1979). Many in­ sects, such as bees and butterflies, have good color vision and can orient by polar­ ized light. References HORRIDGE, G. A. 1975. The compound eye and vision of insects. Clarendon, Oxford. MATSUDA, R. 1965. Morphology and evolution

INSECT STRUCTURE AND FUNCTION

17

of the insect head. Amer. Entomol. Inst., Mem. 4: 1-334. SILBERGLIED, R. E. 1979. Communication in the

ultraviolet. Ann. Rev. Ecol. Syst. 10: 3 7 3 398.

The Thorax and Its Appendages T h e t h o r a x ( M a t s u d a 1970), when present as in insects, is a boxlike unit, primarily c o n c e r n e d with locomotion ( H e r r e i d a n d F o u r t n e r 1981). It is t h e site of t h e largest muscles in t h e body, those that m o v e legs a n d , in insects, t h e wings. In all insects a n d a r a c h n i d s a n d s o m e m y r i a p o d s , t h e n u m b e r of a m b u l a t o r y legs is r e d u c e d from o n e pair (primitive) p e r s e g m e n t , a n d t h e legs arise only from t h e thoracic r e g i o n . T h i s a r r a n g e m e n t affords m o r e efficient a n d m o r e r a p i d mobility t h a n t h e original, d i s p e r s e d condition r e ­ tained by c e n t i p e d e s a n d millipedes. A d u l t insects all have t h r e e pairs, o n e pair from each of t h e t h r e e thoracic segments. In a r a c h n i d s , t h e typical n u m b e r of pairs is four, a l t h o u g h t h e p e d i p a l p s p r e c e d i n g these may s o m e t i m e s be involved with locomotion, a n d t h e first t r u e leg pair may substitute as feelers (Amblypygi). T h e insect leg itself is m u l t i s e g m e n t e d a n d typically c o m p o s e d , from base to tip, of a coxa ("hip"), t r o c h a n t e r , f e m u r ("thigh"), tibia ("shin"), a n d tarsus ("foot"). T h e last section h a s o n e to five s e g m e n t s a n d is t i p p e d with g r a s p i n g claws, p a d s , o r both. T h e n u m b e r of leg articles in t h e Myriap o d a a n d Chelicerata differs in t h e various o r d e r s but is always six o r m o r e . Simple, generalized legs a r e for walking a n d r u n ­ n i n g (Wilson 1966). Modifications of form occur in t h e legs of all g r o u p s b u t a r e t h e most diverse in t h e insects, a m o n g which a r e molelike digging legs, j u m p i n g legs with greatly e n l a r g e d , muscle-filled femora, hairy legs for c a r r y i n g pollen, g r a s p i n g a n d clasping legs in ectoparasites, a n d flattened, oarlike s w i m m i n g legs in aquatic insects. I n c e n t i p e d e s , t h e first pair a r e s h a r p fangs associated with poison glands.

18

GENERAL ENTOMOLOGY

References HERREID, C. ¥., II, AND C. R. FOURTNER. 1981.

Locomotion and energetics in arthropods. Plenum, New York. MATSUDA, R. 1970. Morphology and evolution of the insect thorax. Entomol. Soc. Can., Mem. 76: 1-431. WILSON, D. M. 1966. Insect walking. Ann. Rev. Entomol. I I : 103-122. Wings and Flight Insects a r e the only invertebrates with wings. T h e s e u n i q u e s t r u c t u r e s a r e nor­ mally p r e s e n t only in t h e adult stage (may­ flies with winged subimagos being t h e only exception) a n d always arise from t h e mesoa n d metathoracic s e g m e n t s . T h e i r historic origin is still a d e b a t e d question, t h e r e being evidence of i n d e p e n d e n t derivation from t h e body wall a n d their serial homology to t h e legs (Kukalova-Peck 1983) a n d a b d o m i n a l gill plates of aquatic ances­ tors (Kukalova-Peck 1978, M a t s u d a 1981). In t h e most primitive insect o r d e r s , t h e Apterygota, wings never evolved. S o m e g r o u p s , in particular, those a d a p t e d to a parasitic way of life, have lost their wings t h r o u g h r e t r o g r a d e evolution o n o n e o r both segments. T h e D i p t e r a a r e character­ ized by t h e r e p l a c e m e n t of t h e meta­ thoracic pair by nonflight, sensory o r g a n s , the halteres. Anatomically, wings a r e flattened, ex­ pansive o u t g r o w t h s of t h e dorsolateral in­ t e g u m e n t of t h e t h o r a x . T h e y h a v e a n u p p e r a n d lower cuticle, between which r u n nerves a n d blood channels, t h e e p i d e r ­ mis, muscle, a n d o t h e r tissues having been largely obliterated. T h e cuticle is m e m b r a ­ n o u s a n d usually t r a n s p a r e n t , a l t h o u g h it may b e p i g m e n t e d o r covered by d e n s e coverings of hairs o r scales. It is also thickened in a linear p a t t e r n to form veinlike struts for s t r e n g t h . T h e p a t t e r n of the latter is not r a n d o m b u t d e t e r m i n e d phylogenetically a n d is relatively constant a m o n g taxa (Comstock 1918), t h u s provid­ ing useful criteria for identification a n d study of relationships. Each vein root is

n a m e d , a n d its h o m o l o g u e is recognizable in all insects. So a r e t h e b r a n c h e s of some adventitious veins (such as crossveins) a n d cells f o r m e d by closed sets of veins. Slightly different venational plans a r e recognized by various a u t h o r s (see Wootton 1979). T h e p r i m a r y p u r p o s e of wings, of course, is flight, a l t h o u g h o t h e r e n d s may be served. T h e y may be modified as protec­ tive shields (such as t h e elytra of beetles a n d h e m e l y t r a of bugs) o r possess color p a t t e r n s that p r o v i d e protection o r recogni­ tion signals to o t h e r individuals. T h e flight process in insects has been studied extensively a n d f o u n d a e r o d y n a m i cally u n i q u e (Ellington 1984, Goldsworthy and W h e e l e r 1989,Rainey 1976). All move­ m e n t is i m p a r t e d by muscles located within the t h o r a x , n o t in t h e wing itself, t h r o u g h a kind of fulcrum f o r m e d by t h e lateral body wall a n d elastic m e m b r a n e s a n d sclerotic articulations in t h e wing base. T h e s h a p e a n d timing of t h e stroke is also d e t e r m i n e d by these s t r u c t u r e s . It r a n g e s from a slow, simple, u p - a n d - d o w n flapping action as in large butterflies, with a wing beat frequency of only 4 to 5 p e r second, to a c o m p l e x rotational o r twisting m o v e m e n t to a n d fro as well as u p a n d d o w n , with as m a n y as 1,000 beats p e r second (Forcipomyia, D i p ­ tera). Insects in t h e latter category h a v e t h e capacity for e x t r e m e l y agile aerobatics, a n d some can attain flight speeds of nearly 20 kilometers p e r h o u r ( H o c k i n g 1953). Most flight is trivial a n d of short d u r a t i o n , taking insects t h r o u g h their r e g u l a r life r o u t i n e s . S o m e insects such as locusts a n d d r a g o n flies, however, a r e capable of sustained flight over long distances for migration a n d dispersal. Small insects f o u n d h i g h in t h e a t m o s p h e r e a r e m o v e d primarily by wind a n d air drafts a n d form a kind of aerial "plankton." T h e l a n d - b o u n d m y r i a p o d s a n d chelicerates seldom m o v e l o n g distances on their own, a l t h o u g h y o u n g spiders m a y be wafted h u n d r e d s of kilometers, like kites on air c u r r e n t s , by letting o u t long silk

t h r e a d s g r a s p e d by t h e winds (a process called "ballooning").

References COMSTOCK, J. H. 1918. T h e wings of insects. Priv. pub., Ithaca. ELLINGTON, C. P. 1984. T h e aerodynamics of

flapping animal flight. Amer. Zool. 24: 9 5 105. GOLDSWORTHY, G. f., AND C. H. WHEELER, eds.

1989. Insect flight. CRC, Boca Raton. HOCKING, B. 1953. T h e intrinsic range and speed of flight of insects. Royal Entomol. Soc. London, Trans. 104: 223-345, pi. I - V I . KUKALOVA-PECK, J. 1978. Origin and evolution of insect wings and their relation to metamor­ phosis, as documented by the fossil record, f. Morph. 156: 53-126. KUKALOVA-PECK, J. 1983. Origin of the insect wing and wing articulation from the arthro­ pod leg. Can. J. Zool. 61: 1618-1669. MATSUDA, R. 1981. T h e origin of insect wings (Arthropoda: Insecta). Intl. J. Ins. Morph. Embryol. 10: 387-398. RAINEY, R. C , ed. 1976. Insect flight. Blackwell, Oxford. WOOTTON, R. J. 1979. Function, homology and terminology in insect wings. Syst. Entomol. 4: 81-93. The Abdomen and Its Appendages T h e a b d o m e n is generally t h e least m o d i ­ fied of t h e t h r e e body regions ( M a t s u d a 1976). It retains its primitive h o m o m e r o u s segmentation in insects, a l t h o u g h in many, the n u m b e r of s e g m e n t s is r e d u c e d t h r o u g h fusion. I n spiders a n d acarids, only traces of s e g m e n t a t i o n r e m a i n , a n d it is evanescent in o t h e r g r o u p s . M y r i a p o d s have a large n u m b e r of equal s e g m e n t s , but d i p l o p o d s exhibit a fusion of adjoining s e g m e n t pairs to form d u p l e x , secondary segments (hence t h e d o u b l e - p a i r e d legs that give t h e g r o u p its n a m e ) . Fully devel­ o p e d a b d o m i n a l walking legs persist only in this o r d e r a n d t h e C h i l o p o d a . Apterygote insects also possess ventral a p p e n d ­ ages on some basal a b d o m i n a l s e g m e n t s , which r e p r e s e n t vestigial legs. Also found at t h e base of t h e a b d o m e n in varied g r o u p s a r e special modifications

INSECT STRUCTURE AND FUNCTION

19

such as h e a r i n g o r g a n s ( g r a s s h o p p e r s , geom e t r i d moths), copulatory o r g a n s ( O d o nata), o r book lungs (spiders). T h i s region is even intimately fused with a n d function­ ally a p a r t of the t h o r a x in h i g h e r Hymenoptera. T h e t e r m i n u s of the a b d o m e n also has acquired m a n y modifications. In insects, because the sex o p e n i n g ( g o n o p o r e ) exits h e r e , t h e r e a r e c o m p l e x s t r u c t u r e s for copulation a n d oviposition, collectively t e r m e d genitalia ( S c u d d e r 1971). In males, these a r e for sexual p u r p o s e s only a n d are c o m p o s e d basically of p a i r e d lateral grasp­ ing or sensory o r m a n i p u l a t i v e a p p e n d a g e s a n d medial i n t r o m i t t e n t o r g a n s a n d in m a n y cases, p h e r o m o n e dispersing struc­ tures. C o r r e s p o n d i n g female s t r u c t u r e s re­ s p o n d to o r receive t h e male e l e m e n t s . Eggs a r e placed by special a p p e n d a g e s (ovipositors). Because m a n y genitalic struc­ tures a r e often directly involved with the mechanical d e t e r m i n a t i o n of sexual isola­ tion, they p r e s e n t useful c h a r a c t e r s for species discrimination in t a x o n o m y (Tuxen 1970). Modifications of t h e p o s t e r i o r extremity of the a b d o m e n in Chelicerata include single long a n t e n n a l i k e tactile s t r u c t u r e s (Uropygi), the scorpion sting, a n d the spin­ n e r e t s of spiders. C e n t i p e d e s a n d insects also often have p a i r e d , s e g m e n t e d , termi­ nal sensory a p p e n d a g e s (cerci) that o p e r ­ ate like a kind of r e a r set of a n t e n n a e . In noninsect g r o u p s , t h e g o n o p o r e is located ventrally n e a r the base of the a b d o m e n , e l a b o r a t e genitalia a r e absent, a n d copulation is usually r e m o t e from t h e p r i m a r y exits of the r e p r o d u c t i v e system. For e x a m p l e , in spiders, the male fills tubules in the p e d i p a l p s with seminal fluid from its g o n o p o r e a n d transfers the liquid to the female " m a n u a l l y " with these a p ­ p e n d a g e s . Males of o t h e r chelicerates, a p t e r y g o t e insects, a n d chilopods deposit s p e r m packets ( s p e r m a t o p h o r e s ) o n the s u b s t r a t u m to be picked u p by the female. Male millipedes t r a n s f e r t h e s p e r m with

20

GENERAL ENTOMOLOGY

modified legs (gonopods), by contacting the female g o n o p o r e directly or by leaving s p e r m a t o p h o r e s to be picked u p by the female. Miscellaneous terminal a p p e n d a g e s oc­ cur, including large forceps for prey cap­ t u r e a n d defense, which earwigs have, spinnerets for m a n i p u l a t i n g silk in spiders, or even snorkellike b r e a t h i n g a p p a r a t u s e s in many i m m a t u r e aquatic insects. T h e sting of scorpions is the highly modified last a b d o m i n a l segment.

References MATSUDA, R. 1976. Morphology and evolution of the insect abdomen: With special reference to developmental patterns and their bearings upon systematics. Pergamon, New York. SCUDDER, G. G. E. 1971. Comparative morphol­ ogy of insect genitalia. Ann. Rev. Entomol. 16: 379-406. TUXEN, S. L., ed. 1970. Taxonomist's glossary of genitalia in insects. 2d ed. Munksgaard, Copenhagen.

Muscular System Closely tied functionally to the exoskeleton is the main muscular system. All muscles attach to the i n t e g u m e n t internally a n d p r o v i d e motion to the a r t h r o p o d body in all its varied actions. T h e y n e v e r form a body wall plexus but lie in b u n d l e s r u n n i n g between insertions. T h e latter may be b r o a d or a t t e n u a t e d , cover extensive areas o n sclerites, o r fasten to invaginated exten­ sions of the latter, the a p o d e m e s . T h e latter, w h e n long a n d slender, a r e ten­ donlike but are histologically unlike verte­ b r a t e connective tissue, which is virtually absent in insects a n d their a r t h r o p o d rela­ tives. Visceral muscles, as circular, oblique, or longitudinal b a n d s , are confined to the walls of the digestive tract a n d ducts of the r e p r o d u c t i v e system. In insects a n d their relatives, all muscle tissue is striated, w h e t h e r skeletal o r visceral. Physiologically, insect muscle tissue (Ushe r w o o d 1975) is basically the same as that of vertebrates, a l t h o u g h it (apparently) is

capable of slightly m o r e rapid twitches. Very rapid m o v e m e n t s of insects, such as wing beat frequencies of 200 to 300 p e r second, are m a d e possible by vibratory action of elastic p o r t i o n s of the cuticle, the muscles themselves c o n t r a c t i n g n o m o r e rapidly p e r stimulus t h a n those of a bird. T h e t r e m e n d o u s p o w e r p e r body weight and size of m a n y insects (such as the giant h o r n e d scarabs, Dynastes) is also an illusion. Strength results from the e x e r t i o n of short fibers a r r a n g e d along t h e e n t i r e length of leg joint surfaces so that the load is evenly and widely d i s t r i b u t e d . Power o u t p u t a n d metabolic rates of insects, however, a r e much h i g h e r t h a n in v e r t e b r a t e s , the result of a direct a n d c o n t i n u o u s o x y g e n supply via the tracheal system. Gross a n a t o m y of t h e m u s c u l a t u r e is highly c o m p l e x , a n d t h e r e may be h u n ­ d r e d s of discrete muscles in even a small insect. An early anatomist described over 4,000 in the goat m o t h caterpillar, as com­ pared to a m e r e 529 in h u m a n s . T h i s richness of muscles c o m b i n e d with m e ­ chanically diverse articulations p e r m i t s a diverse r e p e r t o i r e of intricate m o v e m e n t s by these animals.

Reference USHERWOOD, P. N. R. 1975. Insect muscles. Academic, London.

Digestive System In all g r o u p s , t h e alimentary canal is a continuous, fairly straight t u b e , with o p e n ­ ings anteriorly via the m o u t h a n d posteri­ orly via t h e a n u s . T h e r e a r e t h r e e regions of the gut, defined by their e m b r y o n i c origins: a f o r e g u t a n d h i n d g u t , both formed by invagination of the blastocoel and lined with e p i d e r m i s a n d cuticle, a n d a mesodermal m i d g u t , lacking a cuticle. T h e r e a r e various diverticula d e p e n d i n g on the g r o u p , most c o m m o n l y a c r o p (temporary s t o r a g e sac leading off t h e esophagus) a n d blind gastric ceca arising

from the m i d g u t o r stomach. Salivary glands empty into the m o u t h cavity or from the tip of a proboscis via long ducts associated with the h y p o p h a r y n x . Most of the digestive enzymes a r e p r o ­ d u c e d by the cells in t h e walls of t h e m i d g u t a n d ceca. T h e n a t u r e of these enzymes varies according to dietary a d a p t a ­ tions, proteases a n d lipases p r e d o m i n a t i n g in carnivores, cellulases a n d related com­ p o u n d s in wood feeders, keratinase a n d collagenase in scavengers of v e r t e b r a t e connective tissues a n d hair, a n d so on. Nutrition and Metabolism Food enters the gut by the m o u t h located o n t h e front o r u n d e r s i d e of t h e h e a d in insects a n d m y r i a p o d s o r the h e a d region in arach­ nids. T h e r e it is mixed with predigestive enzymes from the salivary glands, fangs, or regurgitations. Most digestive processes a r e reserved for the interior of the stomach a n d intestine but in spiders begin p r i o r to swal­ lowing. T h e latter r e g u r g i t a t e o n their prey, causing enzymatic liquification externally. S o m e early c h a n g e s in food in o t h e r types of a r t h r o p o d s may b e w r o u g h t by secre­ tions of the salivary glands. O n its way to t h e stomach via the e s o p h a ­ gus, food may be diverted into a c r o p for storage or g r o u n d u p by a region of the gut set with spines or teeth moved by e x t r a heavy muscles (proventriculus). T h e nutritional r e q u i r e m e n t s of insects, arachnids, a n d so on, a n d their metabolic processes (Gilmour 1961) also vary e n o r ­ mously (Dadd 1973). T h e s a m e essential elements i m p o r t a n t to most animals for e n e r g y (Downer 1981) a n d g r o w t h seem to b e n e e d e d by all, either supplied in the diet or synthesized metabolically or by intesti­ nal symbionts. Nucleic acids a r e synthesized by all in­ sects, as a r e some vitamins. T h e ten essen­ tial a m i n o acids, however, all must be ingested in a p p r o p r i a t e p r o p o r t i o n s to sus­ tain growth. C a r b o h y d r a t e s serve as a m a ­ j o r e n e r g y source, a n d a l t h o u g h they a r e

INSECT STRUCTURE AND FUNCTION

21

often p r e s e n t in t h e diet, they a r e not always essential a n d c a n be c o n v e r t e d from p r o t e i n o r fats. T h e r e a r e considerable differences in t h e ability of different in­ sects to utilize polysaccharides. Wood roaches a n d termites, for e x a m p l e , rely o n p r o t o z o a n s o r bacteria in t h e g u t to b r e a k d o w n cellulose to assimilable sugars. Fats a r e t h e chief f o r m in which e n e r g y is stored, a n d t h e ability t o synthesize t h e m is w i d e s p r e a d . O t h e r lipids such as choles­ terol must b e a c q u i r e d from foodstuffs. Vitamin n e e d s vary considerably, a l t h o u g h generally only water soluble B vitamins must b e p r e s e n t in food. Vertebrate blood is notably lacking in t h e latter, a n d m a n y h e m a t o p h a g o u s insects rely o n symbionts for these c o m p o u n d s . Parasites store B vitamins, ingested by t h e scavenging larvae that feed o n bacteria-rich food, a n d these a r e passed o n t o t h e a d u l t . I n o r g a n i c salts a n d trace minerals a r e n e e d e d m u c h as they a r e in v e r t e b r a t e animals.

References DADD, R. H. 1973. Insect nutrition: Current developments and metabolic implications. Ann. Rev. Entomol. 18: 381-420. DOWNER, R. G. H., ed. 1981. Energy metabo­ lism in insects. Plenum, New York. GILMOUR, D. 1961. T h e metabolism of insects. Freeman, San Francisco.

Growth Food serves n o t only t o p r o v i d e e n e r g y for activity b u t also to build u p stores for long d o r m a n t p e r i o d s , a n d , of course, it is t h e basis for g r o w t h . As a r t h r o p o d s , with a confining, almost n o n e x p a n d a b l e , nonliv­ ing exterior cuticle, insects a n d their kin achieve size increases a n d m a t u r i t y only by periodic s p u r t s of g r o w t h following molt­ ing. T h i s process takes place from a few to m a n y times d u r i n g t h e animal's life, al­ t h o u g h it ceases after a d u l t h o o d in insects. Molting (or ecdysis) is p r e c e d e d by a cessa­ tion of activity a n d catalysis of t h e lower cuticle layers w h e n muscles a n d sense o r ­

22

GENERAL ENTOMOLOGY

gans assume n e w a t t a c h m e n t s . Only t h e old o u t e r cuticular layer is shed, including the linings of t h e larger t r a c h e a e , foregut, and hindgut.

Rhythms and Seasonality Terrestrial a r t h r o p o d s display r h y t h m s in their activity, correlated with a n d a d a p t i n g t h e m generally to changes in their e n v i r o n ­ m e n t . S o m e of these r h y t h m s a r e i n d e p e n ­ d e n t of signals from their s u r r o u n d i n g s ( e n d o g e n o u s o r circadian r h y t h m s ; B r a d y 1974). Examples of such functions a r e daily periods of sleep a l t e r n a t i n g with active locomotion o r feeding a n d p e r i o d s of sing­ ing o r c o u r t i n g . Events in l o n g - t e r m life cycles a r e also cyclic a n d have i n t e r n a l controls interacting with c h a n g e s in ambi­ ent stimuli, day length being a very s t r o n g o n e (Beck 1980). A l t h o u g h t h e physiologi­ cal basis for such functions is still n o t u n d e r s t o o d , a n u n d e r l y i n g "biological clock" m e c h a n i s m is postulated ( S a u n d e r s 1982). L o n g periods of quiescence c o m m o n l y occur in insects a n d relatives, often to carry the animal t h r o u g h a d v e r s e seasons. T h i s is called d i a p a u s e a n d is characteristic of high latitude o r high elevation species in the wintertime (hibernation) o r of desert species d u r i n g d r y p e r i o d s (aestivation). A t these times, growth, d e v e l o p m e n t , a n d activity is a t t e n u a t e d . Finally, d i a p a u s e is b r o k e n with t h e r e t u r n of favorable condi­ tions, a n d e m e r g e n c e occurs. S o m e t i m e s large n u m b e r s m a y r e t u r n to action simul­ taneously, resulting in p o p u l a t i o n explo­ sions. Periods of d o r m a n c y a r e less p r o ­ f o u n d in tropical t h a n t e m p e r a t e insects because of m o r e equable e n v i r o n m e n t a l conditions in t h e lower latitudes (Denlinger 1986).

References BECK, S. D. 1980. Insect photoperiodism. 2d ed. Academic, New York.

BRADY, J. 1974. T h e physiology of inseci circa­ dian rhythms. Adv. Ins. Physiol. 10: 1-115. DENLINCER, D. L. 1986. Dormancy in tropical insects. Ann. Rev. Entomol. 31: 239-264. SAUNDERS, D. S. 1982. Inseci clocks. 2d ed. Pergamon, Oxford.

Luminescence A n o t h e r specialized metabolic j o b to which certain body chemicals a r e p u t is bioluminescence (Harvey 1957). Q u i t e a n u m ­ ber of insects, primarily beetles (glow­ worms, fireflies, h e a d l i g h t beetles, railroad worms) a n d millipedes, h a v e evolved lightproducing o r g a n s (McElroy et al. 1974). T h e m e c h a n i s m of light p r o d u c t i o n is complex (Case a n d Strause 1978) b u t basi­ cally involves t h e oxidation of luciferin in the presence of t h e e n z y m e luciferase. Luciferin is first activated by A T P in t h e presence of m a g n e s i u m , t h e n oxidized to an excited f o r m (adenyl-oxy-luciferin) that decays to a lower e n e r g y f o r m with t h e liberation of light. T h e reaction is cool a n d very efficient, some 9 8 p e r c e n t of t h e energy involved b e i n g released as light.

References CASE, J. F , AND L. G. STRAUSE. 1978. Neurally

controlled luminescent systems. In P. J. Her­ ring, ed., Bioluminescence in action. Aca­ demic, New York. Pp. 331-366. HARVEY, E. N. 1957. A history of luminescence, from the earliest times until 1900. Amer. Phil. Soc, Philadelphia. MCELROY, W. D., H. H. SELIGER, AND M. D E -

LUCA. 1974. Insect bioluminescence. Physiol. Ins. 2 : 4 1 1 - 4 6 0 .

Blood and Circulation All t h e a r t h r o p o d s t h a t a r e t h e subject of this book possess a n o p e n circulatory sys­ tem (Jones 1977). T h a t is, t h e blood moves for t h e most p a r t over a n d a r o u n d t h e tissues a n d o r g a n s , b a t h i n g t h e m a n d ex­ changing molecules with t h e m directly, in a continuous b o d y cavity, t h e h e m o c o e l . I n insects, t h e r e a r e n o blood vessels save t h e main aorta t h a t leads anteriorly, directly

from t h e h e a r t (McCann 1970), a n d e m p ­ ties into sinuses s u r r o u n d i n g t h e brain. I n centipedes, t h e r e a r e short lateral arteries leading t o t h e gut a n d o t h e r m i n o r vessels. T h e r e is a " p u l m o n a r y a r t e r y " to t h e book lungs in spiders as well as s e c o n d a r y vessels to t h e legs, tail, a n d so o n , in o t h e r arach­ nids. T h e heart, which lies dorsally in t h e hemocoel, j u s t b e n e a t h t h e a b d o m i n a l roof, propels t h e blood forward with peri­ staltic contractions. After passing t h r o u g h the body cavity, including t h e legs, a n t e n ­ nae, wings, a n d o t h e r a p p e n d a g e s , a n d often aided by auxiliary, pulsatile o r g a n s at their bases, t h e blood r e e n t e r s t h e h e a r t t h r o u g h lateral pores (ostia). T h e blood itself (or h e m o l y m p h ) con­ sists of a fluid plasma (Florkin a n d J e u n i a u x 1974) in which n u c l e a t e d cells a r e s u s p e n d e d (Crossley 1975). T h e latter a r e of m a n y types b u t normally d o n o t possess h e m o g l o b i n like v e r t e b r a t e corpuscles, t h e oxygen/carbon dioxide t r a n s p o r t function being a s s u m e d by t h e tracheal system (see below). Insect blood is n o t r e d (except in a few specialized types that have h e m o g l o ­ bin, such as blood w o r m s , Chironomus) b u t g r e e n , a m i x t u r e of c a r o t e n o i d a n d bile p i g m e n t ("insectoverdin"), bluish, o r al­ most clear. T h e functions of t h e blood cells include phagocytosis, w o u n d healing, co­ agulation, storage, a n d regulation of inter­ m e d i a t e metabolism. T h e plasma serves primarily as t h e carrier of substances t o tissues a n d also provides a store of n u t r i ­ tive c o m p o u n d s such as sugars a n d p r o ­ teins. Its water acts as a reservoir for t h e m a i n t e n a n c e of cellular fluids.

References CROSSLEY, A. C. 1975. T h e cytophysiology of

insect blood. Adv. Ins. Physiol. 11: 117-221. FLORKIN, M., AND C. JEUNIAUX. 1974. Hemo­

lymph composition. Physiol. Ins. 5: 255-307. JONES, J. C. 1977. T h e circulatory system of insects. C. T. Thomas, Springfield, 111. MCCANN, F. V. 1970. Physiology of inseci hearts. Ann. Rev. Entomol. 15: 173-200.

INSECT STRUCTURE AND FUNCTION

23

Hormones An i m p o r t a n t class of chemicals trans­ p o r t e d by the blood a r e h o r m o n e s (Novak 1975, Sláma et al. 1974). T h e r e are m a n y types, a n d they vary in their effects, even those from a single e n d o c r i n e o r g a n . A few activities m e d i a t e d by h o r m o n e s a r e molt­ ing, m e t a m o r p h o s i s , e g g p r o d u c t i o n , color changes, daily activity r h y t h m s , d o r m a n c y , a n d caste d e t e r m i n a t i o n in social insects. Many of these processes a r e controlled by the balance a n d timely p r o d u c t i o n of only a few basic h o r m o n e s such as the "molting h o r m o n e " (ecdysone) a n d the "juvenile h o r m o n e " (neotenin), often u n d e r the o v e r r i d i n g c o m m a n d of n e u r o s e c r e t o r y hormones. Insect h o r m o n e s a r e c o m p l e x biochemicals p r o d u c e d by two types of e n d o c r i n e o r g a n s . T h e s e a r e the n e u r o s e c r e t o r y cells in the central n e r v o u s system a n d t h e e n d o ­ crine glands which a r e s e p a r a t e masses of tissue specialized for h o r m o n e p r o d u c t i o n . Well-known e x a m p l e s of the latter a r e the c o r p o r a cardiaca, f o r m i n g p a r t of the wall of the aorta, the c o r p o r a allata, situated on either side or s u r r o u n d i n g t h e e s o p h a g u s , a n d the p r o t h o r a c i c g l a n d s , diffuse tissue a g g r e g a t i o n s at the back of t h e h e a d or in the floor of t h e p r o t h o r a x . T h e n e u r o ­ secretory cells send their h o r m o n a l p r o d ­ ucts to t h e target o r g a n s (often glands of the second type) a l o n g t h e a x o n s of n e r v e cells. Secretions from the e n d o c r i n e glands a r e released into t h e blood.

References NOVAK, V. J. A. 1975. Insect hormones. Chap­ man & Hall, London. SLÁMA, K.,

M. ROMANUK, AND F. SORM.

1974.

Insect hormones and bioanalogues. Springer, New York.

Pheromones M u c h like h o r m o n e s (sometimes called " e c t o h o r m o n e s " ) , p h e r o m o n e s (Jacobson 1972) a r e special kinds of biologically ac­ tive substances released by o n e individual

24

GENERAL ENTOMOLOGY

which cause o t h e r individuals of the same species to act in a specific way. T h e s e substances a r e extremely n u m e r o u s in kind a n d influence a m o n g insects a n d their relatives. In fact, entomologists have realized in recent years that the d o m i n a n t m e a n s of c o m m u n i c a t i o n between these c r e a t u r e s is via these m e s s e n g e r substances (Shorey 1976), perceived by olfactory sense o r g a n s , especially on t h e a n t e n n a e , m o u t h p a r t s , a n d tarsi (Lewis 1984). T h e y a r e p r o d u c e d by e c t o d e r m a l (exocrine) glands on the a b d o m e n , wings, or o t h e r parts of the body. S o m e p h e r o m o n a l systems that have been particularly well studied a r e the a p h rodisiacal scents from t h e wings of m a l e butterflies a n d m o t h s or eversible a b d o m i ­ nal glands of the females. T h e s e chemicals serve to d r a w the sexes t o g e t h e r a n d elicit c o u r t s h i p a n d copulatory behavior. T h e trail-marking substances a n d a l a r m chemi­ cals of ants a n d bees that foster a g g r e g a ­ tion are also p h e r o m o n a l , as are the caste a n d activity controlling r e g u l a t o r s in social insect colonies.

References JACOBSON, M. 1972. Insect pheromones. 2d ed. Physiol. Ins. 3: 229-276. LEWIS, T., ed. 1984. Insect communication. Academic, New York. SHOREY, H. H. 1976. Animal communication by pheromones. Academic, New York.

Other External Secretions Allomones a r e c o m p o u n d s p r o d u c e d by insects a n d their relatives that elicit a n t a g o ­ nistic reactions between individuals (Bell a n d C a r d e 1984). T h e y benefit t h e s e n d e r only, usually protecting it by w a r d i n g off an attack by the receiver (Blum 1981). T h e pain-giving (not prey-seducing) v e n o m s of female aculeate H y m e n o p t e r a , r e p u g n a n t o d o r s of m a n y t r u e bugs a n d beetles, a n d emetic body chemicals (cardiac glycocides a n d the like) in a few butterflies a r e of this category. Such also is the function of cant h a r i d i n (Young 1984a, 19846), a t e r p e n o i d

p r o d u c e d by "blister beetles" (Meloidae). When p r o v o k e d , these beetles e x u d e blood containing this substance from the tibiotarsal articulations, a n d they a r e strongly avoided by insectivorous vertebrates a n d carnivorous insects. O t h e r secretions a r e e x t e r n a l but cause no interactive r e s p o n s e in o t h e r or the same species. T h e s e a r e utilitarian sub­ stances involved in the life processes of the producer. E x a m p l e s a r e silk (Denny 1980) for cocoons a n d webs, adhesives to bind e g g s in place, a n d materials such as wax or gums for b u i l d i n g s t r u c t u r e s . V e n o m used by spiders, c e n t i p e d e s , scorpions, a n d oth­ ers to obtain food also b e l o n g in this category. Regardless of function, a r t h r o ­ pod venoms a r e usually c o m p a r e d from chemical or pharmacological s t a n d p o i n t s (Bettini 1978).

References BELL, W.

J.,

AND

R.

T.

CARDE,

eds.

1984.

Chemical ecology of insects. Sinauer, Sunderland, Mass. BETTINI, S., ed. 1978. Arthropod venoms. Springer, Berlin. BLUM, M. S. 1981. Chemical defenses of arthro­ pods. Academic, New York. DENNY, M. W. 1980. Silks—their properties and functions. Soc. Exper. Biol., Symp. 34: 2 4 7 272. YOUNG, D. K. 1984a. Cantharidin and insects: An historical review. Great Lakes Entomol. 17: 187-194. YOUNG, D. K. 19846. Field records and observa­ tions of insects associated with cantharidin. Great Lakes Entomol. 17: 195-199.

Nervous System In insects, as with o t h e r animals, the ner­ vous tissue is c o m p o s e d of n e r v e cells (neurons), which a r e g r o u p e d into linear nerves a n d gangliar masses to form a central n e r v o u s system ( T r e h e r n e 1974, Miller 1979), an a u t o n o m i c (or stomatogastric) system, a n d a p e r i p h e r a l or sen­ sory n e r v e system. T h e first is ventral, lying in the floor of t h e h e m o c o e l , a n d is characterized by a succession of ganglia

interspersed along a p a i r e d , ventral n e r v e cord. T h e n e r v e cell bodies a r e located peripherally in the ganglia, the c e n t e r of which are occupied by a c o m p l e x of n e r v e fibers (the neuropile) that c o n n e c t the ganglia as the n e r v e c o r d . T h e largest a n d most c o m p l e x ganglion is the a n t e r i o r m o s t . It is dorsal, above the p h a r y n x , in the h e a d . T h i s is the brain (Howse 1970), which may actually be com­ posed of two or m o r e fused p r i m a r y gan­ glia. It is the o v e r r i d i n g c e n t e r of n e u r a l integration to which the o t h e r ventral gan­ glia a r e ultimately subjugated, a l t h o u g h each of the latter may have some d e g r e e of autonomy. A b e h e a d e d insect may con­ t i n u e to live a n d exhibit locomotory a n d sensory activity for some time b e f o r e it eventually dies from such injury. T h e major sensory o r g a n s of t h e h e a d , the eyes, a n t e n n a e , a n d palpi, a r e con­ nected by large nerves directly to the brain. T h e brain also contains n e u r o s e c r e t o r y cells a n d functions partly as an e n d o c r i n e o r g a n as explained above. T h e first ventral ganglion is also located in the head region a n d is associated with ingestive processes. T h e r e follows a vary­ ing n u m b e r of s e g m e n t e d ganglia, primi­ tively, o n e p e r s e g m e n t , u p to eleven in insects, a n d m a n y m o r e in m y r i a p o d s , b u t the n u m b e r is often less, d u e to fusion of segments, especially in the t h o r a x . T h e a u t o n o m i c system is closely associ­ ated with the digestive tract a n d consists of a small n u m b e r of small ganglia a n d short fine nerves. Its function is to control vis­ ceral activity. It is also involved with parts of the e n d o c r i n e system. Efferent nerves r u n from the central n e r v o u s system to the muscles in all parts of the body. Afferent nerves lead from t h e sensory system, mainly t h e i n t e g u m e n t a r y sense o r g a n s , to the central n e r v o u s sys­ tem. T h e cell bodies of sensory n e u r o n s a r e located n e a r the sensilla themselves, a n d their axons connect t h e m directly to the ganglia without i n t e r v e n i n g synapses.

INSECT STRUCTURE AND FUNCTION

25

References HOWSE, P. E. 1970. Brain structure and behav­ ior in insects. Ann. Rev. Entomol. 20: 3 5 9 379. MILLER, T. A. 1979. Insect neurophysiological techniques. Springer, New York. TREHERNE, J. E. 1974. Insect neurobiology. North Holland, Amsterdam. Integumentary Sense Organs T h e a r t h r o p o d would b e isolated from its e n v i r o n m e n t by t h e nonliving, encapsulat­ ing cuticle, b u t t h o u s a n d s of s t r u c t u r e s sensitive to e x t e r n a l stimuli (Dethier 1963), collectively called sensilla, cover t h e sur­ face. T h e y a r e especially n u m e r o u s o n t h e a n t e n n a e , tarsal p a d s , a n d palpi a n d com­ m u n i c a t e with t h e n e r v o u s system via nerves of the p e r i p h e r a l system. T h e anat­ omy of sensilla is e x t r e m e l y varied, each type specifically a d a p t e d to t h e p e r c e p t i o n of a certain subset of stimuli i m p o r t a n t to the animal's safety a n d o t h e r life processes. T h e most c o m m o n a n d often most a b u n ­ d a n t sensilla a r e hairlike extensions (setae, chaetae) responsible f o r m e c h a n o r e c e p tion (Mclver 1975). T h e s e may r e s p o n d to touch, stretching, o r b e n d i n g , directly from a n o u t w a r d force o r by p r e s s u r e from a n o t h e r p a r t of t h e body (often they are f o u n d in articulations). T h e hair base may be simple a n d level with the surface o r recessed a n d q u i t e c o m p l e x , as is t h e trichobothria of arachnids. Portions of t h e body wall can b e inner­ vated so that d e f o r m a t i o n s transmit infor­ mation a b o u t mechanical stresses. Even slight vibrations from air c u r r e n t s o r com­ pression waves a r e perceived. For e x a m ­ ple, masses of stretch r e c e p t o r s in t h e swollen, subbasal s e g m e n t of t h e a n t e n n a e of mosquitoes a n d o t h e r flies ( J o h n s t o n ' s o r g a n ) r e s p o n d to deflexions of the flagellum by air m o v e m e n t s , giving these insects an acute sense of h e a r i n g . W h e n the integu­ ment is especially sensitive in this way, a n d t h e r e a r e structural modifications for r e ­ ception such as thin, vibrating m e m b r a n e s , these p o r t i o n s a r e c o n s i d e r e d auditory o r ­

26

GENERAL ENTOMOLOGY

gans (Michelsen 1979). Such a r e t h e t h o ­ racic a n d a b d o m i n a l t y m p a n a of m a n y m o t h s a n d g r a s s h o p p e r s , fore tibial hear­ ing pits of katydids, a n d acoustical win­ dows in the cicada t h o r a x . Sensilla a r e often s t r u c t u r e d for t h e reception of chemicals in air o r liquids. Such c h e m o r e c e p t o r s (Slifer 1970) usually have thin o r p o r o u s walls so t h e molecules may pass t h r o u g h t h e o u t e r p a r t of t h e o r g a n a n d stimulate i n n e r receptive sur­ faces. T h e y m a y be extremely sensitive. Calculations for t h e sex attractant of t h e domestic silk m o t h indicate that a single molecule may elicit a r e s p o n s e . Certain sensilla also react to a m b i e n t t e m p e r a t u r e c h a n g e s , r a d i a n t heat, p r e s ­ sure, humidity, a n d surface m o i s t u r e (Altn e r a n d Loftus 1985). Perception of related factors internally a r e by direct cellular sensitivity.

References ALTNER, H., AND R. LOFTUS. 1985. Ultrastruc-

ture and function of insect thermo- and hygroreceptors. Ann. Rev. Entomol. 30: 273— 295. DETHIER, V. G. 1963. The physiology of insect senses. Methuen, London. MCIVER, S. B. 1975. Structure of cuticular mechanoreceptors of arthropods. Ann. Rev. Entomol. 20: 381-397. MICHELSEN, A. 1979. Insect ears as mechanical systems. Amer. Sci. 67: 696-706. SLIFER, E. H. 1970. The structure of arthropod chemoreceptors. Ann. Rev. Entomol. 15: 121-142. Sound Production C o r r e l a t e d with h e a r i n g in m a n y insects a n d a few a r a c h n i d s is s o u n d p r o d u c t i o n , a n o t h e r m e a n s of c o m m u n i c a t i o n (Haskell 1961). T h e r e a r e a g r e a t variety of m e c h a ­ nisms for m a k i n g s o u n d s that a r e audible to o t h e r insects a n d to t h e h u m a n ear. S o m e , such as those resulting from the vibration of wings in flight, may b e adventitious a n d a p p a r e n t l y have no value to the animal, but most have a specific function a n d originate from u n i q u e , sometimes elaborate struc-

tures. Extraspecific uses usually a r e to star­ tle a n d are protective (Masters 1979); intraspecific functions i n c l u d e t h e calling a n d courtship stimulations between t h e sexes, aggregation, s p r e a d i n g a l a r m , a n d giving the location of o t h e r colony m e m b e r s in social a n d semisocial forms. S o u n d s m a y b e p r o d u c e d as a by­ product of s o m e activity such as feeding o r wing m o v e m e n t , t a p p i n g t h e substrate, a n d ejections of air, b u t the major a n d most effective m e a n s of sonification involve frictional m e c h a n i s m s a n d vibrating m e m ­ branes (tymbals). T h e former, called stridulation, involves two facing surfaces that a r e r o u g h e n e d a n d that, when m o v e d against each other, p r o d u c e a s o u n d . Such a r e t h e narrow s c r a p e r a n d file in t h e base of the fore wings of crickets a n d katydids. Many other insects, beetles, l e p i d o p t e r o u s larvae and p u p a e , a n d so o n , have b r o a d c o r r u ­ gated o r r i d g e d areas that when r u b b e d together, give a variety of g r i n d i n g , hiss­ ing, squeaking, a n d clicking s o u n d s . S o u n d s p r o d u c e d by t h e vibration of a m e m b r a n e d r i v e n by muscles a r e c o m m o n in H o m o p t e r a , H e t e r o p t e r a , a n d some moths b u t a r e best d e v e l o p e d in male cicadas. T h i s s o u n d - p r o d u c i n g o r g a n is located in t h e dorsolateral p a r t of the first abdominal s e g m e n t . S o u n d is m a d e when the tymbal muscle contracts, pulling it back rapidly. Release allows it t o r e t u r n to t h e starting position s u d d e n l y against t h e air, and t h e resulting vibrations set u p highintensity air waves that m a y s o u n d to t h e h u m a n e a r like a d e a f e n i n g screech o r harsh scream.

References HASKELL, P. T. 1961. Insect sounds. Quadrangle Books, Chicago. MASTERS, W. M. 1979. Insect disturbance stridulation: Its defensive role. Behav. Ecol. Sociobiol. 5: 187-200.

Excretion T h e typical insect n e p h r i t i c o r g a n s (Malpighian tubules) a r e long, thin, blindly e n d ­

ing tubes arising from t h e g u t n e a r t h e j u n c t i o n of m i d g u t a n d h i n d g u t a n d e x ­ t e n d i n g freely in t h e body cavity. T h e i r n u m b e r s vary a m o n g different g r o u p s from a few to h u n d r e d s . T h e wall of t h e tubule is o n e cell thick, encircling a lu­ m e n . T h e s e cells extract waste p r o d u c t s of metabolism from t h e blood, n i t r o g e n o u s b y - p r o d u c t s usually in t h e form of uric acid b u t also as u r e a a n d a m m o n i a . Potas­ sium, sodium, a n d o t h e r i n o r g a n i c ions a r e also eliminated, a l o n g with a quantity of water. T h e m a i n t e n a n c e of constant salt levels, water, osmotic p r e s s u r e in the h e m o l y m p h , a n d elimination of n i t r o g e n o u s wastes a r e t h e main excretory tasks (Maddrell 1971) of the Malpighian tubules in insects. T h o s e o r g a n s a r e p r e s e n t in t h e o t h e r g r o u p s , a l t h o u g h they m a y b e replaced by n e phridial glands in a r a c h n i d s a n d some chilopods.

Reference MADDRELL, S. H. P. 1971. T h e mechanisms of

insect excretory systems. Adv. Ins. Physiol. 8: 200-331. Water Relations Terrestrial a r t h r o p o d s a r e subject to water loss ( B a r t o n - B r o w n e 1964, Stobbart a n d Shaw 1974) from excretion, in t h e feces, a n d t h r o u g h t h e cuticle, i n c l u d i n g t h a t lining t h e respiratory system. T h e loss is especially intense in species living in arid e n v i r o n m e n t s . Water is gained primarily in the food b u t also by d r i n k i n g a n d g e n e r a l absorption from h u m i d air. Special o r g a n s of conservation a r e also p r e s e n t in associa­ tion with t h e h i n d g u t , whose n o r m a l func­ tions include r e a b s o r p t i o n o f water from the feces. O n e of these, t h e c r y p t o n e p h r i d ium, incorporates t h e distal e n d s of Malpi­ ghian tubules which loop back o n t o o r into a thickened p o r t i o n of the r e c t u m . W a t e r is recycled from t h e latter back into t h e tubules a n d r e u s e d ; feces from these in­ sects e m e r g e in a very d r y state.

INSECT STRUCTURE AND FUNCTION

27

Aquatic insects have salt a n d water con­ trol p r o b l e m s different from b u t n o less severe t h a n those faced by terrestrial types. Since t h e h e m o l y m p h is h y p e r t o n i c t o t h e outside m e d i u m , t h e r e is a constant ten­ dency for water t o pass into t h e insect t h r o u g h t h e cuticle. T h i s u p t a k e is c o u n t e r ­ balanced by a copious liquid o u t p o u r i n g , which, however, results in a loss of salts. T h i s is c o r r e c t e d by r e a b s o r p t i o n by t h e rectum.

References BARTON-BROWNE, L. B. 1964. Water regulation in insects. Ann. Rev. Entomol. 9: 63-82. STOBBART, R. H., AND J. SHAW. 1974. Salt and

water balance; excretion. Physiol. Ins. 5: 361-446.

Respiration E x c h a n g e of r e s p i r a t o r y gases in insects a n d allied terrestrial a r t h r o p o d s takes a very different f o r m f r o m that f o u n d in o t h e r animals. T h e a n a t o m y of most con­ tains a system of t u b u l e s (tracheal system) d e d i c a t e d directly to t h e tasks of b r i n g i n g oxygen to t h e tissues a n d c a r r y i n g off c a r b o n dioxide a n d o t h e r waste gases. T h e blood plays n o significant role in this process e x c e p t in very small, i m m a t u r e forms that live in d a m p conditions a n d aquatics with blood-filled gills. T h e tra­ cheae o p e n to t h e o u t s i d e t h r o u g h s e g m e n tally a r r a n g e d p o r e s , t h e spiracles, which generally have a closing device to k e e p water loss to a m i n i m u m . L a r g e tubes r u n i n w a r d from the spiracles a n d b r a n c h p r o ­ fusely, often i n t e r c o n n e c t i n g with sacs o r o t h e r tubules a n d t e r m i n a t i n g finally in m i n u t e blind e n d i n g s (tracheoles) directly o n t h e cells. Derived f r o m i n t e g u m e n t a r y e p i d e r m a l cells, t h e e n t i r e system, e x c e p t the tracheoles, is lined with cuticle that has a circular r i n g e d s t r u c t u r e for s t r e n g t h against collapse. T h e rates of diffusion of o x y g e n a n d c a r b o n d i o x i d e are sufficient t o allow these gases to passively r e a c h all tissues, b u t

28

GENERAL ENTOMOLOGY

ventilatory m o v e m e n t s a r e necessary in large a n d very active forms. T h i s is accom­ plished by a b d o m i n a l compression, con­ traction, a n d o t h e r m u s c u l a r m o v e m e n t s . T h e length of the diffusion p a t h , however, is a factor limiting t h e size of insects, in particular, those with t h e bulky muscle masses n e e d e d for flight. Spiders have a tracheal system in t h e a b d o m e n only, including a variety of m o d i ­ fications, a m o n g t h e m "sieve trachea," which a r e large t r u n k s from t h e e n d s of which originate n u m e r o u s individual fine tracheae. Many also possess u n i q u e respira­ tory structures called "book l u n g s , " which are lamellate, trachealike plates e x t e n d i n g into t h e body cavity. Blood flows b e t w e e n the plates, e x c h a n g i n g molecules with t h e c h a m b e r s of t h e tracheoid tubules, t h u s functioning m u c h like a v e r t e b r a t e l u n g . T h e m e c h a n i s m s of e x t e r n a l r e s p i r a t o r y a d a p t a t i o n s in aquatic insects (Miller 1974) go in a great variety of directions. T h e y rely o n t a p p i n g a t m o s p h e r i c air, o r extrac­ tion of dissolved oxygen from t h e sur­ r o u n d i n g liquid, o r combinations of both. A m o n g t h e former, most a r e often associ­ ated air stores of o n e kind o r a n o t h e r . T h e tracheal system itself m a y have sacs o r e n l a r g e m e n t s to a c c o m m o d a t e air s u p ­ plies, o r bubbles m a y b e carried b e n e a t h the wings o r held o n t o t h e g e n e r a l body surface by hairs o r o t h e r extensions of the i n t e g u m e n t . P r e v e n t e d from collapse by these extensions, these air bubbles act as "physical gills," o x y g e n a n d c a r b o n d i o x i d e passing in a n d o u t of t h e m t h r o u g h their surface, which acts like a m e m b r a n e ("plastron respiration"). Spiracles c o m m u ­ nicating with t h e bubbles t a p t h e air store a n d can also function normally s h o u l d t h e water d r y u p o r t h e animal e m e r g e to a s s u m e a terrestrial p h a s e of existence. Species utilizing a t m o s p h e r i c air m u s t come to t h e surface from time to time t o r e s t o r e their gaseous provisions, a l t h o u g h some, such as certain m o s q u i t o larvae, may stay below for very long p e r i o d s of time,

tapping air c a r r i e d in t h e vessels of aquatic plants. Small aquatic insects m a y employ t h e general cuticle as a gill. L a r g e types have other forms of gill s t r u c t u r e s , expansive nlates o r fingerlike extensions filled with blood, o r a rich tracheal n e t w o r k to carry on gas e x c h a n g e . Respiration in m a n y endoparasitic types relies o n similar mecha­ nisms, their lives b e i n g s p e n t in a liquid ambience for long p e r i o d s .

Reference MILLER, P. L. 1974. Respiration: Aquatic insects. 2d ed. Physiol. Ins. 6: 403-467.

Reproduction Insects a n d like a r t h r o p o d s a r e normally bisexual a n d r e q u i r e sexual c o m m u n i o n o r mating (Blum a n d B l u m 1979, T h o r n h i l l and Alcock 1983), with s u b s e q u e n t g a m e t e fusion, for r e p r o d u c t i o n (Davey 1965, E n g l e m a n n 1970). Only in a few cases has p a r t h e n o g e n e s i s — a n d in still fewer cases, hermaphroditism—evolved. T h e produc­ tion of n o r m a l y o u n g by unfertilized fe­ males is p a r t of t h e r e g u l a r r e p r o d u c t i v e process in m a n y H o m o p t e r a , a l t e r n a t i n g with t h e sexual process. Unfertilized eggs may be t h e m e a n s of sex d e t e r m i n a t i o n in others, such as t h e h o n e y b e e , which p r o ­ duces d r o n e s by this m e t h o d . I n t h e cot­ tony cushion scale (Icerya purchasi), both male a n d female g o n a d s d e v e l o p in t h e female, a n d self-fertilization takes place. T h e g o n a d s a n d their i m m e d i a t e ducts are almost always p a i r e d . T h e generative organ m a y b e single o r multiple in myriapods, d e r i v e d from m e s o d e r m a l e m ­ bryonic tissue. T h e g o n o d u c t s join p a i r e d or single e c t o d e r m a l invaginations that lead to the o u t s i d e via t h e g o n o p o r e . T h i s may be located e i t h e r terminally as in most insects o r n e a r the base of the a b d o m e n in arachnids. Male insects a n d m y r i a p o d s usually have a complex set of genitalia s u r r o u n d i n g t h e

g o n o p o r e , a n extension of which t e r m i ­ nates in a n i n t r o m i t t e n t o r g a n o r penis (often called t h e aedeagus). T h e s e geni­ talia, especially t h e claspers of o n e sort o r another, a r e i m p o r t a n t in locking t h e pair securely a n d precisely t o g e t h e r while t h e penis is inserted, f o r m i n g a physical con­ nection that is normally species specific ( E b e r h a r d 1985). T h e y m a y also play a p a r t in physical o r chemical stimulation necessary for successful copulation (their i n n e r surfaces often b e a r sensillar patches) (Alexander 1964). T h e g o n o p o r e is u n elaborated in spiders, t h e function of t h e genitalia being a s s u m e d by t h e p e d i p a l p s . T h e external female genitalia a r e rela­ tively simple c o m p a r e d to t h e male's, b u t some special s t r u c t u r e s (ovipositors) m a y b e p r e s e n t for e g g p l a c e m e n t .

References ALEXANDER, R. D. 1964. The evolution of mat­ ing behaviour in arthropods. Royal Entomol. Soc, Symp. 2: 78-94. BLUM, M. S., AND N. A. BLUM. 1979. Sexual

selection and reproductive competition in insects. Academic, New York. DAVEY, K. G. 1965. Reproduction in the insects. Freeman, San Francisco. EBERHARD, W. G. 1985. Sexual selection and animal genitalia. Harvard Univ. Press, Cam­ bridge. ENGLEMANN, F. 1970. The physiology of insect reproduction. Pergamon, Oxford. THORNHILL,

R., AND J. ALCOCK.

1983. T h e

evolution of insect mating systems. Harvard Univ. Press, Cambridge. Fertilization T h e s p e r m cells p r o d u c e d by the testes a r e i n t r o d u c e d internally into t h e female in most forms, that is, fertilization is internal. T h e y may be first kept in storage in diverticulae of the c o m m o n oviduct, how­ ever, a n d released to fuse with t h e eggs only as they pass, t h e female t h u s control­ ling the time of fertilization. I n t r o d u c t i o n of s p e r m is n o t always directly via the g o n o p o r e . Secondary geni­ talia a r e d e v e l o p e d most notably in O d o -

INSECT STRUCTURE AND FUNCTION

29

nata and spiders. T h e former transfer the sperm from the gonopore to the accessory copulatory organs on the venter of the third abdominal segment; male spiders use syringes in the bulbous apex of the pedipalps for this purpose. Sperm is carried in a liquid medium, or more commonly, com­ pressed into packets (spermatophores) that may be inserted into, or formed, in the common oviduct or its outpocketings (spermathecae), or are placed on the sub­ stratum to be picked up by the female. Size Terrestrial arthropods are subject to size limitations because of the combined restric­ tions of rigidity, lack of permeability, and weight of the cuticle, which becomes too much of an encumbrance to movement in very large forms. Also, the diffusion rates of respiratory gases is insufficient to traverse the distances necessary through prolonged tracheal systems, although this is overcome to some extent by breathing movements. Environmental determinants, such as mois­ ture and food availability, are also impor­ tant (Schoener and Janzen 1968). In spite of these restrictions, some ex­ tremely large insects are found in Latin America, all long lived, herbivorous, forest types. In terms of bulk, the record must be adult males of the large horned scarab, like Megasoma elephas, which may weigh 40 grams or more. Wing expanse is another measure of size and finds its greatest expression in the birdwing moth (Thysania agrippina), with a spread from wing tip to wing tip of up to 30 centimeters. Those with the longest, although slender, bodies are the Neotropical centipede Scolopendra gigantea, which extends 27 centimeters, and walkingsticks, some 26 centimeters {Phi Iba losorna phyllinum) from the head to the tip of the abdomen. Indeed, the wet forests of the Neotropics are traditionally thought to harbor many insect goliaths. While not the largest overall, some that are

30

GENERAL ENTOMOLOGY

the biggest of their category or impressive in any sense are many horned beetles such as Dynastes hércules (17 cm, including horn), morpho butterflies, Morpho hecuba (wingspan 18 cm), tarantulas, Theraphosa lablondi (20 cm leg span), and lubber grasshoppers, Tropidacris (wingspan 25 cm, length to folded wing tips, 13 cm). T h e largest flies in the world are the Neotropical Pantophthalmus (Pantophthalmidae) that measure 4 centimeters in length and weigh over 2.5 grams. At the low end of the size scale are the smallest known insects, parasitic wasps of the genus Alaptus (Myrmaridae) with body lengths of only 0.2 millimeters. Insects and their terrestrial relatives, by and large, are small, the vast majority 6 to 10 millimeters long and 25 to 50 milli­ grams in weight. This is their single most important structural characteristic, en­ abling the exploitation of the infinite num­ ber of small niches of nature. Insects need little space and minimal sustenance to live and hide from predators. Reference SCHOENER, T. W., AND D. H. JANZEN.

1968.

Notes on environmental determinants of tropical versus temperate insect size patterns. Amer. Nat. 102: 207-224.

Genetics and Cytology Insect genetics has been a fruitful field and has contributed a great deal to this field of general science, particularly through stud­ ies on Drosophila. Much of this success is attributable to the ease with which many insects are maintained in the laboratory, their rapid turnover of generations, diver­ sity of phenotypic expressions of gene effects, and in many cases, giant, wellmarked chromosomes. The genetic control of a large number of particular insect characteristics has been elucidated, such as the distribution of dif­ ferent types of hairs, color patterns, resis-

References tance to insecticides, and wing venation. KETTLEWELL, H. B. D. 1973. The evolution of Gross changes in Lepidoptera wing color melanism. Clarendon, Oxford. patterns are known to be determined by LANCÉ, G. 1970. Relations entre le détermisimple gene differences (Robinson 1971). nisme génétique du sexe el la controle hor­ monal de sa differentiation chez les arthroSex in insects is basically determined by podes: Comparaison avec les vertebres. Ann. (he production of different gametes, al­ Biol. 9: 189-230. though epigenetic factors, such as hor­ PAL, R., AND M. J. WHITTEN. 1974. The use of mones, are also important (Langé 1970). genetics in insect control. Elsevier/NorthSex chromosomes may be involved, a vari­ Holland, Amsterdam. ROBINSON, R. 1971. Lepidoptera genetics. Perety of combinations being found. Males gamon, Oxford. heterozygous XY and XO and females homozygous XX is the usual situation. The reverse is true of Lepidoptera and TrichopINSECT BEHAVIOR tera. In Hymenoptera, fertilized eggs de­ velop into females, unfertilized eggs into Insect behavior (Matthews and Matthews males, the latter therefore being haploid 1978) is a rapidly developing field of study individuals. that attempts to explain both the complex Genotype and gene frequencies are anatomical and physiological bases and properties of populations rather than of higher, integrative mechanisms for activity. individual insects. Their behavior is impor­ Only short-term, decisively determined ac­ tant to the understanding of evolutionary tions are recognized in this framework. processes when it is realized that it is shifts Long-lasting, slowly induced actions, such in their frequency, either randomly (ge­ as diapause or maturation, are considered netic drift), by mutation, selection, or exter­ physiologic or developmental phenomena nal events, that lead to speciation and (see other parts of this chapter). higher order phylogenetic changes. A clas­ Physiochemically and anatomically, in­ sic case of the latter is the increase to sects possess the same elements that con­ normalcy of melanism in populations of trol behavior in all animals. Foremost of European moths living in industrial envi­ these is the nervous system (Roeder 1963), ronments where heavy soot pollution dark­ including its sensory component, but the ens their resting substrates (Kettlewell muscular and hormonal components play 1973). No melanics of this type are yet an essential, if secondary, part. It is the known in Latin America. degree of complexity of the first that Mutations are easily induced in insects determines the levels on which lines of by means of radiation and chemicals. T h e action lie. former is even used routinely to create A key element of the nervous system in sterile individuals for mass release in ge­ determining behavior is the associative (ad­ netic control schemes (Pal and Whitten juster, internuncial) neuron, which inter­ 1974). cedes between receptor (efferent) and effec­ The mode of gene operation is also tor (afferent) neurons and has the capacity becoming known in insects. In the giant to redirect and modify otherwise simple chromosomes of fly larvae, characteristic reflex reactions. Large numbers of these swellings, forming after natural hormones form masses (neuropiles) in the brain and contact the cell, appear to indicate activity ventral ganglia and serve as centers of of specific genes. neural integration. These are something Genetic work with other terrestrial ar­ like the cortex or gray matter of the human thropod groups aside from insects has brain and define the overall function of a lagged behind work with insects.

INSECT BEHAVIOR

31

ganglion. They represent the main areas where activities are generated and orga­ nized. A major such center is the corpus pedunculatum ("mushroom-shaped body"), believed to be the site of summation of simultaneous excitation from all sources. It tends to be small in arthropods with simple behavior, large in those with complicated lives, such as the social Hymenoptera. These cells both stimulate and inhibit. Endocrine secretions are not only caused to flow in response to nervous command but are actually part of the nervous system in the form of neurosecretory cells. These cells produce hormones that move along the axons and direct other nerve and endo­ crine tissues to emote. Of course, activity is finally the result of muscular contraction. Insects and their rela­ tives may have very large numbers of dis­ crete muscle bundles that predicate a like­ wise elaborate system of efferent nerves. It is fortunate that a lack of obstructive connec­ tive tissue in these animals makes it possible to dissect and experiment to determine pathways relatively easily. The largest nerves lead to the most active locomotor organs, the wings and legs. Other major efferents control the mouthparts, anten­ nae, cerci, genitalia, and numerous other muscularized structures. The insect behaviorist looks for chains or pathways of stimulation-integrationaction to explain activities (Browne 1974). The latter can be considered to be com­ posed of bits or units that meld together into sequences first, then complexes or systems. The simplest movements have the simplest nerve control and fewest muscles involved. The most complex systems have very large numbers of pathways and pro­ cesses and are so complicated that it is possible to analyze them only in general. An understanding of the way the whole insect acts requires an extension of the rudimentary functioning of the neural, hormonal, and muscular elements. This extension progresses along a scale of in­

32

GENERAL ENTOMOLOGY

creasing complexity, beginning with socalled automatic or instinctive behavior and terminating with learned activity. The simplest instinctive actions are re­ flex arcs, so-called knee-jerk responses, where a part of the body reacts directly to a stimulus without the intercession of an association nerve. An example is the retrac­ tion of the tarsus from a hot surface. A step up from this level occurs when the whole body is coordinated but by nonmodifiable reactions. Where only a single action is identifiable, such as movement away from or toward light or touching or shunning other individuals or objects, the behavior is called a taxis or tropism. Such behavior may be positive or negative. The attraction of moths to artificial light, the catatonic freezing or "death feigning" display many species use to escape harm, and the follow­ ing of odor trails by dung beetles to find food for their young are specific examples. A series of these tropistic elements may be strung together, one triggering the next to form a fixed action pattern. These may take up a sizable part of the behavioral repertoires of most insects. Pupation in giant silk moth larvae offers an appropri­ ate example: changes in photoperiod or some internal stimulus causes them to cease feeding. This initiates defecation and a wandering, searching activity, leading to the discovery of a suitable pupation site. Even if the latter is not found, the larvae will begin to spin silk and form a cocoon of a specific shape in which it finally settles and pupates. This sequence follows the same steps regardless of changes in exter­ nal stimuli (unless acute) and does not vary according to any information learned by the individual. Insects and other terrestrial arthropods are capable of limited learning (Alloway 1972), defined as any relatively permanent change in behavior that results from prac­ tice. Such learning is of a low order and often short lived, but it is often essential to the animal's existence. At least two types

have been seen, classical Pavlovian condi­ tioning and, much more commonly, instru­ mental conditioning, where reinforcement stimuli direct the performance of the in­ sect. The latter is a characteristic especially of social insects, like the honeybee, which can be trained artifically to fly to a colored surface by food offerings. Under natural conditions, this ability is important in re­ cruiting foragers and in efficient utiliza­ tion of a flower nectar food source. Some forms, such as cockroaches and ants, facili­ tate to mazes. The vast majority of these arthropods, however, probably are capable of virtually no learning whatever.

processes are thus served efficiently, al­ though automatically and unswervingly, and have contributed to their success as a group. It is useful to segregate and classify the kinds of motivation driving the insect body because it is often found that single action sequences operate within them. The follow­ ing are only representative, as many exam­ ples fit into the categories given; additional types will appear in the main text of this book.

The complexity of some behavior in insects, particularly social insects, most es­ pecially ants, whose lives parallel our own in some ways, has suggested to some the possibility of the existence of intelligence. As possessed by higher vertebrates, includ­ ing ourselves, no such high degree of learning and reasoning can be truly as­ cribed to these creatures. All activity, re­ gardless of how cunning and comprehend­ ing it seems, can be explained on the basis of fixed action sequences, with very limited learning. T h e nesting of digger wasps (Ammophila) is a classic example: the female wasp first digs a burrow in sandy soil which it then closes over at the mouth. It then leaves to search for prey, captures it, and returns to the location of the burrow. To do this, it has had to learn a few landmarks by which it navigates. Their misplacement, however, may lead the digger wasp to conclude wrongly on the exact location. The nest, when found, is opened and the prey packed within, an egg is laid on it, and the female exits, closes the nest perma­ nently, and leaves to repeat the process elsewhere. All of these are innate, unmodifiable acts.

1. Alimentation. Finding food and feeding involve specific movements, often elabo­ rate. Mosquitoes respond to visual and odor cues to find warm-blooded hosts and then follow tactile stimuli to select a proper station and find a capillary. Inter­ nal pressure from expansion of the stomach causes cessation of feeding and induces flight. 2. Survival. Its host, discovering a mosquito in the act of feeding, will attempt to destroy or remove it. The insect displays flight as a survival act, an extremely common one with winged types. Other survival-related behavior is shelter seek­ ing, catalepsis, and biting. Most protec­ tive coloration is accompanied by pos­ tures that enhance deception or warning patterns. 3. Aggression. Both intra- and interspecific agonistic (fighting) behavior occurs in insects, including male-male competi­ tion for females, as in the horned scar­ abs. Bees may grapple for a nectary or over territory and females. Raiding for food, such as found in many ants, should not be confused with aggression, although the results are the same. The vanquished colony is perceived as food, not as a rival faction.

The remarkable thing about insect be­ havior is that it may be highly complicated, comparable in this respect alone to verte­ brates, yet it is nearly all controlled by instinctive mechanisms. Fundamental life

4. Sex. This essential, overriding drive in all organisms has led to some of the most incredibly complex and even bi­ zarre activities in all groups of terres­ trial arthropods. These are divided into

INSECT BEHAVIOR

33

m a t e finding, c o u r t s h i p , copulation, a n d insemination ( T h o r n h i l l a n d Alcock 1983). 5. Brood care. P a r e n t a l b e h a v i o r occurs in relatively few insects a n d o t h e r terres­ trial a r t h r o p o d s a n d is a p r e c u r s o r to social o r g a n i z a t i o n in g e n e r a l . It greatly increases survivability a n d is necessary for t h e m a i n t e n a n c e of colonies. 6. Inlraspecific communication. T h e ways in which i n f o r m a t i o n is t r a n s m i t t e d be­ tween individuals of the same species a r e t r e m e n d o u s l y varied, e m p l o y i n g vi­ sual, chemical, auditory, tactile, a n d o t h e r m e t h o d s . T h e use of a i r b o r n e p h e r o m o n e s seems to d o m i n a t e , al­ t h o u g h nutritive chemicals, ingested by the receiver (trophallaxis), are transmit­ ted a m o n g m e m b e r s of social insect colonies. S o u n d also ties m a n y nonsocial types together. 7. Tool using. It is an a m a z i n g fact that a few insects actually use tools—in an instinctive way, of c o u r s e . T h e p r i m e e x a m p l e is t h e p e b b l e e m p l o y e d by d i g g e r wasps to t a m p the soil p l u g of their b u r r o w nests. 8. Construction. Many types form structures from a variety of b u i l d i n g materials, both e x t r a n e o u s ( m u d , paper, wood) a n d intrinsic (silk). A r c h i t e c t u r e may be e l a b o r a t e a n d the size a n d s t r e n g t h of m a n y edifices p r o d i g i o u s . A high level of c o o p e r a t i o n can be r e q u i r e d between m e m b e r s of social f o r m s to p u t u p nests. Individual efforts a r e also intricate a n d consistent with r e g a r d to g e o m e t r y a n d engineering. 9. Migration. A l a r g e n u m b e r of species regularly m o v e from o n e territory to a n o t h e r , s o m e even o n long-established a n d precise m i g r a t o r y routes. Unidirec­ tional flight is a c o n s p i c u o u s manifesta­ tion of this behavior, a n d it is most c o n s p i c u o u s in larger, active forms such as butterflies a n d day-flying m o t h s .

34

GENERAL ENTOMOLOGY

S o m e behavioral traits a p p a r e n t l y n o t found in insects a n d their relatives are play, expression of grief or sorrow, a n d h u m o r . T h e s e are characteristics of a v e r t e b r a t e c e r e b r u m a n d set these h i g h e r c r e a t u r e s a p a r t from insects a n d o t h e r a r t h r o p o d s , which function as virtual a u t o m a t o n s .

References ALLOWAY, T. M. 1972. Learning and memory in insects. Ann. Rev. Entomol. 17: 43-56. BROWNE, L. B. 1974. Experimental analysis of insect behavior. Springer, New York. MATTHEWS, R. M., AND J. R. MATTHEWS.

1978.

Insect behavior. Wiley, New York. ROEDER, K. D. 1963. Nerve cells and insect behavior. Harvard Univ. Press, Cambridge. THORNHILL,

R.,

AND J.

ALCOCK.

1983.

The

evolution of insect mating systems. Harvard Univ. Press, Cambridge.

DEVELOPMENT AND LIFE CYCLES Eggs W h e t h e r e x t e r n a l to the female p a r e n t ' s body (oviparity) or temporarily within (viviparity), all insects, spiders, a n d allied terres­ trial a r t h r o p o d s start their lives as eggs (Hinton 1981). Eggs c o m e in a n a m a z i n g variety of shapes a n d sizes. T h e y a r e usually placed singly or in g r o u p s in p r o x i m i t y to the juvenile's food source but may b e scat­ tered indiscriminately only in the g e n e r a l habitat w h e r e d e v e l o p m e n t occurs. Many have elaborate cuticular s c u l p t u r i n g , a n d some possess devices for a t t a c h m e n t to the s u b s t r a t u m or caps (opercula) that o p e n to allow egress of the y o u n g . A n u m b e r of species protect their eggs from m o i s t u r e loss a n d t r a u m a by covering t h e m with froth o r encasing t h e m in o t h e r substances that h a r d e n a r o u n d t h e m (oothecae).

Reference HINTON, H. E.,ed. 1981. Biology of insect eggs. Vols. 1-3. Pergamon, Oxford.

Embryology fust prior to fertilization, insect eggs a r e composed mostly of yolk a n d small islands of cytoplasm s u r r o u n d i n g the female n u ­ cleus on o n e e d g e . W h e n the egg is laid, the nucleus is usually in the m e t a p h a s e of the first meiotic division, in which state it receives t h e s p e r m , o n e of which unites with the oocyte after meiosis is c o m p l e t e . T h e nucleus t h e n m i g r a t e s to t h e c e n t e r of the egg a n d begins to divide mitotically. T h e resulting cells m o v e to the p e r i p h e r y and form the b l a s t o d e r m , or early e m b r y o , which later lodges o n o n e side of the egg. T h e g e r m layers a n d e m b r y o n i c m e m ­ branes soon d e v e l o p , a n d d e t e r m i n a t i o n of segmentation a n d the p r i m a r y o r g a n s a n d tissues e n s u e s . T h e a p p e n d a g e s appear, and after a time, the perfect b o d y of the first juvenile stage is c o m p l e t e . T h i s stage takes different forms d e p e n d i n g o n t h e evolutionary level of the g r o u p . Fairly similar embryological steps a r e followed by other terrestrial a r t h r o p o d s ( J o h a n n s e n and Butt 1941). A major exception a r e the springtails (Collembola), whose eggs u n ­ dergo holoblastic cleavage.

Reference JOHANNSEN, O.

A.,

AND E.

H.

BUTT.

1941.

Embryology of insects and myriapods. Mc­ Graw-Hill, New York.

Development Insects a n d related a r t h r o p o d s must pass through a series of d e v e l o p m e n t a l stages on their way to b e c o m i n g sexually m a t u r e adults (Agrell a n d L u n d q u i s t 1973). T h e s e stages are all the m o r e discrete because of the necessity of m o l t i n g a n d growth in stepwise phases. T h e animal itself between molts is r e f e r r e d to as a n "instar," t h e time period, " s t a d i u m . " In virtually all insects, the first instar possesses the c o m p l e t e n u m ­ ber of s e g m e n t s after h a t c h i n g ; in o t h e r groups, s e g m e n t s a r e a d d e d as develop­ ment p r o c e e d s .

As the animal progresses t o w a r d m a t u ­ rity, it increases in size, a n d c h a n g e s in internal a n d external form a n d p r o p o r ­ tions occur to a g r e a t e r or lesser d e g r e e (Sehnal 1985). In most noninsects a n d primitive a p t e r o u s insects, the i m m a t u r e s are fairly similar to the adults. J u v e n i l e insects of the h i g h e r o r d e r s that possess wings, however, u n d e r g o a fair a m o u n t of body modification, called m e t a m o r p h o s i s , primarily associated with the g r o w t h of the wings a n d exploitation of habitats differ­ ent from the adult. M e t a m o r p h o s i s is said to be " g r a d u a l " (incomplete) in lower winged insects with externally d e v e l o p i n g wing b u d s ; the single j u v e n i l e type is called a n y m p h (or sometimes naiad in aquatics). N y m p h s generally have feeding a n d o t h e r habits similar to the adult; naiads live r a t h e r different lives because of their wa­ ter habitats. M e t a m o r p h o s i s is " c o m p l e t e " in the h i g h e r winged insects. In these, t h e r e a r e two f u n d a m e n t a l j u v e n i l e stages: a larva, which has several instars; it finally molts into a p u p a , which eventually yields the adult. T h e s e early stages look totally unlike a n d live in ways very different from the adult a n d i n d e e d d i v e r g e from t h e m in almost every way. T h i s has c o n t r i b u t e d to the evolutionary success of these insects t h r o u g h the d i c h o t o m o u s specialization of life functions (feeding a n d g r o w t h by im­ m a t u r e s , dispersal a n d r e p r o d u c t i o n by adults). Divergence of body f o r m a n d func­ tion has even taken a f u r t h e r step in m a n y species with varying types of larvae (hyperm e t a m o r p h o s i s ) such as f o u n d in the blis­ ter beetles (Meloidae), chalcidoid wasps, a n d others. I m m a t u r e s of different insect g r o u p s are called by various n a m e s . For e x a m p l e , larvae of L e p i d o p t e r a a r e cater­ pillars; p u p a e of butterflies, chrysalids; larvae of muscoid flies, m a g g o t s ; a n d bee­ tle larvae, grubs. P u p a e generally a r e p r o ­ tected by their location, u n d e r g r o u n d in cells or in wood or o t h e r material or encased in a cocoon of silk s p u n by t h e p r e p u p a l instar.

DEVELOPMENT AND LIFE CYCLES

35

References AGRELL, I. P. S., AND A. M. LUNDQUIST.

1973.

Physiology and biochemical changes during insect development. Physiol. Ins. 1: 159-247. SEHNAL, F. 1985. Morphology of insect develop­ ment. Ann. Rev. Entomol. 30: 89-109.

Maturation T h e adult is t h e sexually capable instar, whose responsibility is to find a m a t e a n d r e p r o d u c e , t h u s p e r p e t u a t i n g t h e species. D e v e l o p m e n t of t h e i n t e r n a l a n d e x t e r n a l sexual o r g a n s c o m p l e t e s g r o w t h a n d usu­ ally molting, a l t h o u g h in some noninsect terrestrial a r t h r o p o d s , m o l t i n g may con­ t i n u e t h r o u g h o u t life. W i n g s in t h e insects also b e c o m e fully g r o w n a n d functional at this time, t h e o n e e x c e p t i o n b e i n g the mayflies, which have a winged instar (subimago) p r e c e d i n g t h e full i m a g o . S o m e a b e r r a n t c o n d i t i o n s occur in in­ sects, such as neoteny, in which t h e a d u l t retains its o u t w a r d larval body form but c o m p l e t e s d e v e l o p m e n t of t h e internal r e ­ p r o d u c t i v e o r g a n s . N e o t e n i c adults m a t e a n d p a r e n t offspring while c o n t i n u i n g to feed a n d live as i m m a t u r e s . T h i s is a c o m m o n condition in railroad w o r m s a n d o t h e r beetles a n d in s o m e primitive flies.

Life Cycles T h e way a n insect o r allied a r t h r o p o d develops in relation to its seasonal environ­ m e n t constitutes its life cycle ( T a u b e r et al. 1985). Life cycles a r e as varied as t h e kinds of animals living t h e m . P e r h a p s a majority of species in arctic o r t e m p e r a t e life zones have a n n u a l g e n e r a t i o n s , that is, o n e com­ plete t u r n o v e r , e g g to e g g p e r year. O t h e r s have b i a n n u a l o r m u l t i a n n u a l cycles. T h e latter implies t h e existence of p r o l o n g e d feeding p e r i o d s , often o n food that is p o o r in n u t r i t i o n (e.g., w o o d - b o r i n g beetle lar­ vae), o r t h e intercession of a p e r i o d of d i a p a u s e . Still o t h e r s a r e s e m i a n n u a l (bivoltine) o r multivoltine, with two to sev­ eral g e n e r a t i o n s p e r year. T h e latter a r e

36

GENERAL ENTOMOLOGY

m o r e typical of tropical o r o t h e r stable e n v i r o n m e n t s w h e r e unfavorable d r o u g h t or cold does not force t e m p o r a r y arrests in d e v e l o p m e n t . Some insects, such as p o m a c e flies (Drosophila), develop very rapidly a n d repeatedly a n d m a y have almost c o n t i n u ­ ous r e p r o d u c t i o n t h r o u g h o u t t h e year. S o m e mosquitoes m a t u r e very quickly in transient water following i n f r e q u e n t r a i n s but r e m a i n d o r m a n t in t h e e g g stage for most of the r e m a i n d e r of t h e year. Certain insect types regularly i n c o r p o ­ rate asexual r e p r o d u c t i o n in their life cy­ cles in addition to sexual r e p r o d u c t i o n . T h i s alternation of g e n e r a t i o n s is typical of a p h i d s , for e x a m p l e . W h e n conditions a r e best for plant g r o w t h a n d t h e r e f o r e feed­ ing, as in t h e b e g i n n i n g of t h e rainy season, emphasis is o n multiplication of n u m b e r s . T h i s is accomplished by t h e "stem m o t h ­ ers" that bear live, sterile, wingless females parthenogenetically a n d as rapidly as possi­ ble. As t h e season favorable for dispersal a p p r o a c h e s , w h e n t h e r e is less o r n o rain a n d winds m a y increase, sexually active, winged males a n d females a p p e a r , to m a t e , mix genes, a n d disperse to n e w localities. T h e females lay eggs that h a t c h into t h e asexual forms once again. P r o d u c t i o n of sexual forms is controlled by c h a n g e s in t e m p e r a t u r e a n d p h o t o p e r i o d ; u n d e r con­ stant tropical conditions, cyclical alterna­ tion of g e n e r a t i o n s m a y n o t occur.

Reference TAUBER, M. J., C. A. TAUBER, AND S. MASAKI.

1985. Seasonal adaptations of insects. Oxford Univ. Press, New York.

EVOLUTION AND CLASSIFICATION T h e reconstruction of t h e historical evolu­ tion a n d d e t e r m i n a t i o n of the interrelation­ ships of the presently e x t a n t o r d e r s of insects a n d o t h e r terrestrial i n v e r t e b r a t e s has n o t b e e n settled by any m e a n s . T h e r e

remain m a n y controversies, even over ma­ jor theses, such as t h e m o n o p h y l y (descent from a single ancestral line) of t h e A r t h r o poda o r of t h e a p t e r y g o t e h e x a p o d s . T h e r e is extensive l i t e r a t u r e o n these dis­ a g r e e m e n t s a n d relevant a r g u m e n t a t i o n (Anderson 1973; B o u d r e a u x 1979; G u p t a 1979; M a n t ó n 1977; Sharov 1966). T h e a r t h r o p o d g r o u p s i n c l u d e d in this book a r e all basically terrestrial, probably by way of several i n d e p e n d e n t , parallel evolutionary pathways, from varied p r e c u r ­ sors a m o n g t h e O n y c o p h o r a , Crustacea (Isopoda), U n i r a m i a ( m y r i a p o d s a n d in­ sects), a n d Chelicerata (arachnids), a n d a r e thus only distantly related ( M a n t ó n 1977: 257-258). T h e o n y c o p h o r a n line seems to attach most closely to t h e m y r i a p o d a n , a n d these animals can n o l o n g e r b e c o n s i d e r e d inter­ mediate phylogenetically between a n n e l i d s and a r t h r o p o d s , t h e latter n o w being recog­ nized as a polyphyletic g r o u p . T h e y a r e not ancestral to e i t h e r that g r o u p o r H e x a p o d a , n o r is t h e latter d e s c e n d e d from t h e M y r i a p o d a . Embryological evi­ dence indicates that all t h r e e have di­ verged i n d e p e n d e n t l y from c o m m o n ances­ tors with u n i r a m o u s (lobate) legs. A l t h o u g h t h e C r u s t a c e a a r e distant, with fundamentally different, b i r a m o u s a p ­ pendages, they s h a r e m a n y features with insects. T h e y a r e almost entirely m a r i n e , with only a few types secondarily a d a p t e d for life in fresh waters a n d o n land. T h e chelicerates, distinguished funda­ mentally by their chelicerate m o u t h p a r t s , are virtually all terrestrial, a l t h o u g h proba­ bly derived from originally m a r i n e ances­ tors. Evolution within t h e s u b p h y l u m is n o t clear. All efforts to subdivide t h e o r d e r s have r e m a i n e d inconclusive, as have associ­ ated phylogenetic speculations. T h o s e with book lungs (Scorpionida, Uropygi, Amblypygi, spiders) p r e s u m a b l y can be g r o u p e d ; scorpions, with their c o m p l e t e s e g m e n t a ­ tion, a r e t h e most primitive. I n body s h a p e and e x t e r n a l genitalia, t h e Opiliones resem­

ble some primitive mites, with which they seem to form a close b r a n c h . T h e o t h e r g r o u p s a r e all isolated. T h e phylogeny of t h e primitively m a n dibulate Uniramia ( a p p e n d a g e s with sin­ gle stem) is fairly well u n d e r s t o o d , at least for t h e insects in g e n e r a l (Kristensen 1981). Myriapods retain h o m o m e r i s m , having only a distinct h e a d , b u t seem to possess the basic b o d y s t r u c t u r e likeliest to p r e c e d e that of insects. T h e ancestors of the insects ( H e n n i g 1981) evolved a t h r e e somite t h o r a x a n d t h r e e pairs of legs at an early time, r e d u c i n g the m a n y equal b o d y parts of t h e m y r i a p o d s . T h i s a r r a n g e m e n t is p r e s e r v e d in all t h e t r u e insects a n d t h r e e primitive o r d e r s ( t h e Parainsecta) that differ from t h e insects in several basic ways, including the m o u t h p a r t s , which a r e not exposed as usual (ectognathy) b u t o v e r g r o w n by cranial folds ( e n t o g n a t h y ) . T h e r e a r e also wingless (Apterygota) p r e ­ decessors in b o d y design to t h e d o m i n a n t insects that evolved wings early in their history (Pterygota) (Kukalová-Peck 1987). At first (Paleoptera), wings were clumsy, outwardly projecting, fixed, flight o r g a n s , as seen in m a n y extinct g r o u p s of t h e Paleozoic (e.g., Palaeodictyoptera) a n d ex­ tant mayflies a n d O d o n a t a , b u t soon ac­ quired improvements, among them the ability to be flexed over t h e b o d y which all t h e h i g h e r o r d e r s h a v e ( N e o p t e r a ) . Even those that have secondarily lost wings alto­ gether, often in association with e c t o p a r a sitism (fleas, lice, b e d b u g s , etc.), retain t h e thoracic s t r u c t u r e of their fully winged ancestors. T h r e e major lines e m e r g e d within t h e h i g h e r winged insects. T h e first, most p r i m ­ itive assemblage (Polyneoptera), which some workers question as m o n o p h y l e t i c , includes the " o r t h o p t e r o i d " g r o u p s , t h e O r t h o p t e r a , Grylloptera, D e r m a p t e r a , a n d o t h e r o r d e r s that display a n e n l a r g e d , fanlike h i n d wing with m a n y longitudinal veins, m u l t i s e g m e n t e d tarsi, a n d m a n y Malpighian tubules a n d ganglia internally.

EVOLUTION AND CLASSIFICATION

37

T h e second, " h e m i p t e r o i d " g r o u p (Par a n e o p t e r a ) is largely c h a r a c t e r i z e d by r e ­ gressive traits, loss o r r e d u c t i o n in t h e n u m b e r of tarsal s e g m e n t s , few Malpighian tubules, a n d ganglia in t h e central n e r v o u s system. T h e m a i n o r d e r s f o u n d h e r e a r e the H e m i p t e r a , Psocoptera, Mallophaga, A n o p l u r a , a n d T h y s a n o p t e r a . B o t h of these major g r o u p s display i n c o m p l e t e metamorphosis. T h e t h i r d a n d t h e most recently derived a n d highly successful g r o u p within t h e N e o p t e r a , a r e those with c o m p l e t e meta­ m o r p h o s i s ( H o l o m e t a b o l a ) . It makes u p m o r e t h a n 80 p e r c e n t of t h e species of living insects. Within t h e g r o u p , various subdivi­ sions can b e r e c o g n i z e d , b u t t h e i r definition a n d t h e o r d e r s to be i n c l u d e d a r e n o t all s u p p o r t e d by incontrovertible evidence. At least " n e u r o p t e r o i d " ( M e g a l o p t e r a , N e u r o p t e r a ) a n d " p a n o r p o i d " (Mecoptera, Trichoptera, Lepidoptera, Díptera, Siphonaptera) lines a r e readily identifiable, b u t t h e Coleóptera and Hymenoptera are indepen­ d e n t a n d stand a p a r t . T h e foregoing can be s u m m a r i z e d in a linear classification such as that following. It s h o u l d be e m p h a s i z e d that such schemes are m a t t e r s of p e r s o n a l p r e f e r e n c e a n d a r e always controversial. T h i s classification is conservative a n d i n t e n d e d to reflect rela­ tionships of b r o a d g r o u p s as well as to p r o v i d e c o n v e n i e n t categories for t h e p r e ­ sentation of t h e systematic material form­ ing t h e major p o r t i o n of this book. (See also taxon table.) T h e r e a d e r interested in alternative ideas s h o u l d consult t h e refer­ ences cited at t h e e n d of this section. C L A S S I F I C A T I O N O F I N S E C T S AND MAJOR GROUPS OF RELATED TERRESTRIAL A R T H R O P O D S

(* = groups not covered in this book; included for reference only.) Phylum Onychophora-onychophorans Phylum Arthropoda-arthropods S u b p h y l u m Biramia Class Crustacea—crustaceans

38

GENERAL ENTOMOLOGY

Subclass Peracarida Order Isopoda-isopods O r d e r Amphipoda— amphipods S u b p h y l u m Chelicerata Class A r a c h n i d a O r d e r *Ricinulei—ricinulids O r d e r Araneae—spiders O r d e r Opiliones— harvestmen S u p e r o r d e r Acari—mites a n d ticks O r d e r *Palpigradi— palpigrades O r d e r *Schizomida—micro whip scorpions Order Uropygi-whip scorpions O r d e r Amblypygi-whipless whip scorpions O r d e r Pseudoscorpionida— pseudoscorpions O r d e r Scorpionida— scorpions O r d e r Solpugida—sunspiders S u b p h y l u m Uniramia Class Myriapoda—myriapods Subclass Chilopoda—centipedes Subclass D i p l o p o d a - m i l l i p e d e s Subclass *Symphyla—symphylans Subclass * P a u r o p o d a — p a u r o p o d s Class Hexapoda—insects (broad sense) Subclass Parainsecta O r d e r *Protura—proturans O r d e r *Diplura—diplurans O r d e r Collembola— springtails Subclass Insecta—true insects Infraclass A p t e r y g o t a O r d e r Thysanura—silverfish a n d bristletails Infraclass Pterygota—winged insects S u p e r o r d e r Paleopteraancient—winged insects O r d e r Ephemeroptera— mayflies

Order Odonata-dragonflies a n d damselflies Superorder Neopteram o d e r n - w i n g e d insects

ORDERS OF UNCERTAIN AFFINITIES

O r d e r Coleóptera—beetles O r d e r Hymenoptera—ants, bees, a n d wasps

"ORTHOPTEROIDS"

O r d e r Plecoptera-stoneflies O r d e r Grylloptera—katydids a n d crickets Order Orthopterag r a s s h o p p e r s a n d allies Order Blattodeacockroaches O r d e r Mantodea—mantids O r d e r Phasmatodea— walkingsticks O r d e r Dermaptera—earwigs O r d e r Isoptera—termites O r d e r Embiidina—web spinners "HEMIPTEROIDS"

O r d e r Psocoptera—psocids O r d e r *Zoraptera— zorapterans Order Mallophaga-biting lice Order Anoplura-sucking lice O r d e r Hemiptera—true bugs (heteropterans and homopterans) Order Thysanoptera-thrips "NEUROPTEROIDS"

O r d e r *Mecoptera— scorpionflies O r d e r Megaloptera— dobsonflies O r d e r Neuroptera—nervewinged insects "PANORPOIDS"

O r d e r Diptera—flies a n d gnats O r d e r Siphonaptera—fleas Order Trichopteracaddisflies O r d e r Lepidoptera— butterflies a n d m o t h s

References ANDERSON, D. T. 1973. Embryology and phylogeny in annelids and arthropods. Pergamon, Oxford. BOUDREAUX, H. B. 1979. Arthropod phylogeny, with special reference to insects. Wiley, New York. GUPTA, A. P., ed. 1979. Arthropod phylogeny. Van Nostrand Reinhold, New York. HENNIG, W. 1981. Insect phylogeny. Wiley, Chinchester, Eng. KRISTENSEN, N. P. 1981. Phylogeny of insect orders. Ann. Rev. Entomol. 26: 135—157. KUKALOVA-PECK, J. 1987. New Carboniferous

Diplura, Monura, and Thysanura, the hexapod ground plan, and the role of thoracic side lobes in the origin of wings. (Insecta). Can. J. Zool. 65: 2327-2345. MANTÓN, S. M. 1977. T h e Arthropoda, habits, functional morphology and evolution. Claren­ don, Oxford. SHAROV, A. G. 1966. Basic arthropodan stock with special reference to insects. Pergamon, Oxford.

FOSSIL INSECTS Known Latin A m e r i c a n fossil insect sites are few, b u t they h a v e p r o d u c e d consider­ able material r e p r e s e n t i n g several types of fossilization. Most r e p r e s e n t relatively r e ­ cent strata (Cenozoic). Impressions in s e d i m e n t a r y rock from the Eocene in South America a r e most significant (Martinez 1982). O n e of t h e best k n o w n beds is found at Sunchal, A r g e n t i n a , in t h e province of Jujuy. Many specimens of weevils a n d o t h e r insects w e r e excavated t h e r e by entomologist T. D. A. Cockerell early in this century. T h e oldest insects from the region a r e of an unidentified o r d e r (Eugeropteron a n d Geropteron) from m i d d l e C a r b o n i f e r o u s beds in t h e Sierra d e los Llanos of t h e province of Rioja, also in A r g e n t i n a . O t h e r i m p o r t a n t sites of insect

FOSSIL INSECTS

39

fossils p r e s e r v e d in s e d i m e n t a r y rocks a r e located in Rio G r a n d e d o Sul, Brazil, a n d at Bajo d e Veliz, in t h e p r o v i n c e of San Luis in A r g e n t i n a ( U p p e r C a r b o n i f e r o u s ) (Marti­ nez pers. c o m m . ) . A n e n o r m o u s therap h o s i n e s p i d e r (Megarachne servinei), mea­ s u r i n g 34 c e n t i m e t e r s from t h e chelicerae to the tip of the a b d o m e n a n d having a leg span of m o r e t h a n 50 centimeters, was discovered in t h e Bajo d e Veliz F o r m a t i o n ( H ü n i c k e n 1980). T h e S a n t a n a F o r m a t i o n n e a r C e a r á , Brazil, h a s yielded fossils of Lower C r e t a c e o u s age (Grimaldi 1990). Two a m b e r deposits h a v e yielded a b u n ­ d a n t specimens, a l t h o u g h of relatively r e ­ cent times. Preservation is exceedingly good. T h e insects w e r e c a u g h t in resinous e x u d a t e s from various trees. T h e viscous substance later h a r d e n e d with burial a n d time a n d r e m a i n e d clear within so t h a t the most m i n u t e s t r u c t u r e s (hairs, genitalia, m o u t h p a r t s ) c a n b e o b s e r v e d today in perfect detail, even o n t h e smallest mites a n d m i d g e s . P r e s e r v a t i o n is so perfect in some cases that a t t e m p t s are b e i n g m a d e to recover a n d replicate fossilized D N A from cells of f u n g u s gnats with large c h r o m o ­ somes (Poinar a n d Hess 1982). A f a m o u s site of late Oligocene to early Miocene A g e is n e a r Simojovel, in the state of C h i a p a s , Mexico ( H u r d et al. 1962). It has b e e n particularly well studied a n d h a s p r o d u c e d a wide variety of taxa (various a u t h o r s 1963, 1971). T h e s e deposits w e r e k n o w n to early p e o p l e s of C e n t r a l A m e r i c a , a n d a m b e r pieces with insect inclusions w e r e fashioned into o r n a m e n t s . T h e origi­ nal resin is believed to h a v e b e e n secreted by a l e g u m i n o u s tree of the g e n u s Hymenaea. Localities in t h e D o m i n i c a n Republic have c o m m a n d e d a t t e n t i o n in r e c e n t years ( B a r o n i - U r b a n i a n d S a u n d e r s 1982) a n d a r e also t h e source of a t h r i v i n g " g e m s t o n e " industry (Rice 1979, Rice a n d Rice 1980). T h e deposits a r e generally accepted as of a b o u t t h e s a m e a g e as t h e C h i a p a s a m b e r a n d likewise derived from Hymenaea. T h e m a n y mines scattered a b o u t t h e c o u n t r y

40

GENERAL ENTOMOLOGY

have p r o d u c e d a m b e r f r a g m e n t s with t h o u ­ sands of specimens of m o r e t h a n o n e h u n ­ d r e d families of insects plus spiders a n d o t h e r a r a c h n i d a ( C o k e n d o l p h e r 1986) a n d invertebrates (Sanderson a n d Farr 1960), including the first fossils of g a r d e n i n g ants (Attini, Trachymyrmex, B a r o n i - U r b a n i 1980). Wilson (1985) f o u n d 37 g e n e r a of ants, of which 34 survive in n e i g h b o r i n g areas of Latin America. I n the Americas, t h e r e a r e also a n u m b e r of o t h e r k n o w n b u t u n e x p l o r e d a m b e r deposits, for e x a m p l e , in Colombia (Cockerell 1923), Brazil (Froes A b r e u 1937), a n d surely o t h e r countries (Poinar a n d A g u d e l o 1980). G o o d preservation is also characteristic of t h e Q u a t e r n a r y A g e r e m a i n s f o u n d in asphalt deposits. S o m e sites in this category a r e located in Trinidad (Blair 1927) a n d at Talara o n t h e n o r t h e r n Peruvian coast ( C h u r c h e r 1966). H e r e , because of t h e stickiness of t h e tarlike m e d i u m a n d t h e attractiveness of t h e surface, which looks like water, asphalt seeps form very efficient small animal traps. T h e most n u m e r o u s kinds of insects f o u n d as fossils in these deposits a r e h a r d - b o d i e d g r o u n d beetles, aquatics, a n d c a r r i o n feeders. Aquatic in­ sects a r e sometimes indicative of the pres­ ence of freshwater pools n e a r t h e asphalt o r overlying it. T h e y w e r e e n t r a p p e d w h e n the water d r i e d u p d u r i n g d r o u g h t peri­ ods. C a r r i o n - f e e d i n g species were c a u g h t along with the carcasses of v e r t e b r a t e s that died in the black q u a g m i r e s . Because of their small size a n d delicate ness, insects a n d their relatives p r o d u c e good fossils only in fine-grained o r h o m o ­ g e n e o u s matrices. T h e f o r e g o i n g a r e of this type. O t h e r m o d e s of fossilization that may b e i m p o r t a n t in Latin America, a n d which have been scarcely investigated, a r e permineralization (such as in m i n e r a l c h a r g e d waters), p e a t a n d soft coal e n c a p ­ sulation, cave sediments (Miller 1986), a n d silicification, especially evident in calcare­ ous nodules. Evidences of feeding, b o r i n g ,

coprolites, a n d trails s h o u l d also be com­ mon in deposits of p l a n t fossils. Insect remains in association with ancient h u m a n remains m a y also b e of considerable ar­ chaeological significance (e.g., W a r n e r a n d Smith 1968).

structure of 40-million-year-old insect tissue. Science 215: 1241-1242. RICE, P. C. 1979. Amber of Santo Domingo— mining in the Dominican Republic. Lapidary J. (Nov. 1979): 1804-1810. RICE, H. E., AND P. C. RICE. 1980. Pepitas de sol

antillano. Americas 32 (10): 3 7 - 4 1 . SANDERSON, M. W., A N D T . H. FARR. 1960. Amber

References BARONI-URBANI, C. B. 1980. First description of fossil gardening ants. Amber collection Stutt­ gart and Natural History Museum Basel: Hymenoptera: Formicidae. I: Attini. Stutt. Beitr. Naturk. Ser. B (Geol. Paleon.) 54: 1-13. BARONI-URBANI, C. B., AND J. B. SAUNDERS.

1982. T h e fauna of the Dominican amber: The present state of knowledge. 9th Carib. Geol. Conf. (Santo Domingo, 1980) Trans. 1: 213-223. BLAIR, K. G. 1927. Insect remains from oil sands in Trinidad. Entomol. Soc. London Trans. 75: 137-141. CHURCHER, C. S. 1966. The insect fauna from the Talara tar-seeps, Peru. Can. J. Zool. 44: 985-993. COCKERELL,T. D. A. 1923. Insects in amber from South America. Amer. J. Sci. 5: 331-333. COKENDOLPHER, J. C. 1986 (1987). A new species of fossil Pellobunus from Dominican Republic amber (Arachnida: Opiliones: Phalangodidae). Carib. J. Sci. 2 2 : 2 0 5 - 2 1 1 . FROES ABREU, S. 1937. Sobre a ocorréncia de

ambur nos arenitos da serie Bahia: Brasil. Inst. Nac. Tech. (Rio de Janeiro) Bol. Inf. 2(4): 8. GRIMALDI, D. A., ed. 1990. Insects from the Santana Formation, Lower Cretaceous, of Brazil. Amer. Mus. Nat. Hist. Bull. 195: 1 191. HÜNICKEN, M. A. 1980. A giant fossil spider (Megarachne servinei) from Bajo de Veliz. Acad. Nac. Cien. Córdoba, Bol. 53: 317-325. HURD, JR., P. D., R. F. SMITH, ANDJ. W. DURHAM.

1962. T h e fossiliferous amber of Chiapas, Mexico. Ciencia 21(3): 107-118, PI. I—II. MARTÍNEZ, S. 1982. Catálogo sistemático de los insectos fósiles de América del Sur. Fac. Hum. Cien. (Univ. Rep., Montevideo) Ser. Cien. Tierra, Rev. 1(2): 2 9 - 8 3 . MILLER, S. E. 1986. Phylum Arthropoda, Class Insecta. In D. W. Steadman, Holocene verte­ brate fossils from Isla Floreana, Galápagos. Smithsonian Contrib. Zool. 413: 1-103. POINAR, JR., G. O., AND F. AGUDELO. 1980. El

ámbar: Oro fósil del nuevo mundo. Americas 32(10): 33-40. POINAR, JR., G. O., AND R. HESS. 1982. Ultra-

with insect and plant inclusions from the Dominican Republic. Science 131: 1 3 1 3 1314. VARIOUS AUTHORS. 1963, 1971. Studies of fos­

siliferous amber arthropods of Chiapas, Mex­ ico, Pts. 1, II. Univ. Calif. Publ. Entomol. 31: 1-53, pis. 1-3; 63: i-vi, 1-106, pis. 1-3. WARNER, R. E., AND G. E. SMITH, JR. 1968. Boll

weevil found in pre-Columbian cotton from Mexico. Science 162(3856): 911-912. WILSON, E. O. 1985. Invasion and extinction in the West Indian ant fauna: Evidence from the Dominican amber. Science 229(4710): 265-267.

INSECT NAMES All k n o w n o r g a n i s m s , i n c l u d i n g insects a n d their relatives, have a scientific n a m e , a n d m a n y also have a c o m m o n n a m e (Goto 1982). Scientific n a m e s a r e applied a c c o r d i n g to r i g o r o u s p r o c e d u r e s (Ride et al. 1985), with consistency, universality, a n d stability as p r i m a r y considerations. Each species m u s t b e a r a u n i q u e Latinized two-part epithet (for subspecies, t h r e e - p a r t ) , consist­ ing of a g e n u s a n d specific n a m e , for e x a m p l e , for t h e c o m m o n housefly, Musca domestica. Species a r e g r o u p e d into a hierar­ chy of categories, t h e most usual b e i n g (in ascending o r d e r ) t h e tribe ( n a m e e n d s in suffix, -irá, e.g., Muscini), subfamily (-inae, Muscinae), family (-idae, Muscidae), a n d o r d e r (no s t a n d a r d suffix, e.g., Diptera). N a m e s a r e given t o species o r h i g h e r taxa as they a r e discovered a n d established by publication. T h e first d e s c r i b e r is enti­ tled to a u t h o r s h i p , a n d all o t h e r s a r e obliged to u s e that n a m e . T h e t e r m "new species" refers to o n e that h a s b e e n so f o u n d for t h e first time, n o t to freshly

INSECT NAMES

41

evolved o n e s ; n o n e of t h e latter h a s yet been observed in n a t u r e . Scientific n a m e s a r e p r o p e r l y p r o ­ n o u n c e d a c c o r d i n g to t h e rules of Latin, but their way of b e i n g s p o k e n usually varies a c c o r d i n g to t h e native accent of t h e speaker. T h i s s h o u l d b o t h e r n o o n e except Latin scholars, as l o n g as t h e n a m e is understood. C o m m o n n a m e s , o r vulgates, a r e a p ­ plied to t h e insects a n d their relatives in all Latin A m e r i c a n c o u n t r i e s . Léxica have b e e n p u b l i s h e d for Chile ( B r ü c h e r 1942, Perez D'Angello 1966), Peru ( D o u r o j e a n n i 1965), Brazil (Baucke 1 9 6 1 , Biezanko a n d Link 1972, M o n t e 1928, d a Silva 1 9 3 0 1934) a n d Haiti ( A u d a n t 1941). Many vulgates a r e a d o p t e d directly from indige­ n o u s l a n g u a g e s , s o m e tribes a n d local cultures b e i n g prolific n o m e n c l a t u r i s t s , especially in Brazil ( M o n t e 1928). T h e s e suffer from f r e q u e n t spelling a n d p r o n u n ­ ciation variations, particularly in Brazil (where, in g e n e r a l , I follow von I h e r i n g ' s [1968] o r t h o g r a p h y ) . At least partial e n t o ­ mological glossaries exist for t h e following native t o n g u e s : M a y a n (Welling 1958), Aztec ( = N á h u a t l , e t c . ; O r d o ñ o 1982), Kunza (= Atacameño; Munizaga and Her­ r e r a 1957), J í v a r o (Guallart 1968), TupíG u a r a n í (Tastevin 1923), a n d Q u e c h u a (García 1976). V e r n a c u l a r n a m e s a p p e a r a c c o r d i n g to n o consistent set of stan­ d a r d s , varying from place to place o r time to time with different origins a n d related to t h e n a t u r e of t h e society e m p l o y i n g t h e m (Stoetzel 1989). P h o n e t i c variations in spelling a r e c o m m o n . Scientists a n d e d u c a t e d p e o p l e often form simple transliterations of technical n a m e s (muscids o r m u s c i d e o s , from Muscidae) o r a c c o m m o d a t e n a m e s of classic origin (scarabs o r escarabajos, from G r e e k karabos). L a y m e n a n d c o u n t r y folk a r e likely to invent q u a i n t , often descriptive appella­ tions that frequently apply to a n insect's b e h a v i o r (saltamonte = "hill j u m p e r " ) o r

42

GENERAL ENTOMOLOGY

stinging abilities (lagarta d e fogo = "fire worm"), a n a t o m y (tijeretas = "scissor bear­ ers"), o r that a r e o n a m a t a p o e t i c (cricket, chicharras), o r that may be without obvious derivation (gallinipper). Sometimes these are literal translations from m o d e r n lan­ guages (scorpions, escorpiones) o r usages (tarantulas) n o t c o m m o n to t h e region. Mixtures of sympatric l a n g u a g e s also o c c u r (sede [Spanish] + ocuilin [Náhuatl] = sedeocuilin = silkworm). T h e only a t t e m p t to s t a n d a r d i z e c o m m o n n a m e s has b e e n m a d e with pest species in English (Stoetzel 1989). Most languages have a b r o a d t e r m for insects a n d like animals, r o u g h l y equiva­ lent to t h e English, for e x a m p l e , " b u g " ("worm" o r " g r u b " ) : bicho (Spanish a n d Portuguese) a n d ocuilin (Náhuatl).

References AUDANT, A. 1941. Identification des insectes d'Haiti par leur nom creóle. Soc. Hist. Geogr. Haiti, Rev. 12(42): 51-55. [Not seen.] BAUCKE, O. 1961. Os nomes comuns dos in­ sectos no Rio Grande do Sul. Sec. Agrie., Porto Alegre. BIEZANKO, C. M., AND D. LINK. 1972. Nomes

populares dos Lepidópteros no Rio Grande do Sul (Segundo Catalogo). Univ. Fed. Santa Maria, Bol. Tec. 4: 3 - 1 5 . BRÜCHER, G. 1942. Lista de algunos nombres vulgares de insectos. Dept. San. Veg. (Min. Agrie, Santiago) Bol. 2(2): 120-125. DA SILVA, B. R. 1930-1934. Nomenclatura popular dos Lepidópteros do Distrito Federal and seus arredores. Vols. 1—5. O Campo, Rio de Janeiro. DOUROJEANNI, M. J. 1965. Denominaciones ver­ naculares de insectos y algunos otros in­ vertebrados en la selva del Perú. Rev. Peruana Entomol. 8: 131-137. GARCÍA, R. J. 1976. Nombre de algunos insectos y otros invertebrados en "Quechua." Rev. Peruana Entomol. 19: 13—16. GOTO, H. E. 1982. Animal taxonomy. Arnold (lnsi. Biol., Stud. Biol. no. 143), London. GUALLART, J. M. 1968. Nomenclature JíbaroAguaruna de la fauna del Alto Marañón (Invertebrados). Biota 7: 195-209. IHERING, R. VON. 1968. Dicionário dos animáis do Brasil. Ed. Univ. Brasilia, Sao Paulo.

MONTE, O. 1928. Os nomes vulgares dos in­ sectos de Brasil. Almanak Agrie. Brasil. 1928: 228-289. MUNIZAGA, C , AND J. HERRERA.

1985. Notas

entomológicas de Socaire (Obtenidas durante la Expedición Chileno-Alemana a Socaire, en mayo de 1957). Notas Centr. Est. Antropol., Univ. Chile, 1: 3 - 1 3 . ORDOÑO, C. M. 1982. Diccionario de zoología Náhuatl. Ed. Innovación, Mexico. PÉREZ D'ANGELLO, V. 1966. Concordancia entre

los nombres vulgares y científicos de los insectos chilenos. Mus. Nac. Hist. Nat. Not. Mens. 10(119): 2 - 7 . RIDE, W. D. L., C. W. SABROSKY, G. BERN ARDÍ,

AND R. V. MELVILLE, eds. 1985. International

code of zoological nomenclature. 3d ed. Intl. Trust Zool. Nomen., London. STOETZEL, M. B. 1989. Common names of insects and related organisms. Entmol. Soc. Amer. Lanham, Md. TASTEVIN, C. 1923. Nomes de plantas e animaes em Lingua Tupy. Rev. Mus. Paulista 13: 6 8 7 763. WELLING, E. C. 1958. Some Mayan names for certain Lepidoptera in the Yucatán penin­ sula. J. Lepidop. Soc. 12: 118.

INSECTS AND HUMAN CULTURE Aside from t h e i r i m p o r t a n c e as pests a n d our academic interest in insects, these crea­ tures, spiders, a n d related a r t h r o p o d s have considerable influence in that p o r t i o n of h u m a n activity that m a y be called t h e h u m a n i t i e s — m u s i c , a r t , literature, lan­ guage, religion, a n d folklore (fig. 1.8). T h e study of these influences is a general area of insect study called cultural e n t o m o l o g y (Hogue 1987). E x a m p l e s a p p e a r a m o n g historical, m o d e r n , a n d i n d i g e n o u s peo­ ples. (Some of t h e m o r e general a r e cited below; m a n y o t h e r specific cases a r e scat­ tered t h r o u g h t h e r e m a i n d e r of this book in t h e sections o n t h e various insects in­ volved; for Mexico, see M a c G r e g o r 1969.) Insects, spiders, c e n t i p e d e s , a n d scorpi­ ons a p p e a r in t h e Mayan Codices (Dresden, Tro-Cortesianus, a n d Peresianus), indicat-

Figure 1.8 Decorative plates from modern Peru prominently featuring ¡mages of the fly (chuspi), revered in Incan times and a design motif in Andean art today. (Original, author's collection) ing an appreciation of their existence a n d their inclusion in cultural events, such as rituals, c e r e m o n i e s , a n d d a n c e s . T h e fa­ m o u s Nasca figures include an i m m e n s e spider (fig. 1.9). Portions of same a r e also stylized as glyphs h a v i n g linguistic signifi­ cance (Tozzer a n d Allen 1910). In t h e e i g h t e e n t h century, it was believed that a small, r e d insect (still unidentified b u t called "coya" in t h e O r i n o c o region) caused severe skin e r u p t i o n s ; its effects could only be r e m e d i e d by ceremoniously passing t h e body t h r o u g h a fire m a d e from a specific grass ("guayacán") ( K a m e n - K a y e 1979). Many such curious accounts of insects fill the accounts of early visitors a n d colonists in t h e New World (Cowan 1865). Insects have lent their n a m e s to m a n y places in Latin America. A m o n g t h e better known a r e C h a p u l t e p e c , t h e "hill of t h e g r a s s h o p p e r s " (chapulín = g r a s s h o p p e r + tepee = hill) w h e r e t h e Aztec E m p e r o r M o n t e z u m a ' s castle was built in what is now p a r t of Mexico City, a n d U r u b a m b a , "plain of t h e insect" (uru = s p i d e r o r

INSECTS AND HUMAN CULTURE

43

evolved ones; n o n e of t h e latter has yet been observed in n a t u r e . Scientific n a m e s a r e properly pro­ n o u n c e d a c c o r d i n g to t h e rules of Latin, but their way of b e i n g s p o k e n usually varies a c c o r d i n g to t h e native accent of the speaker. T h i s should b o t h e r n o o n e except Latin scholars, as long as t h e n a m e is understood. C o m m o n n a m e s , o r vulgates, a r e a p ­ plied to t h e insects a n d their relatives in all Latin A m e r i c a n countries. Léxica have been published for Chile ( B r ü c h e r 1942, Perez D'Angello 1966), Peru (Dourojeanni 1965), Brazil (Baucke 1961, Biezanko a n d Link 1972, M o n t e 1928, d a Silva 1 9 3 0 1934) a n d Haiti ( A u d a n t 1941). Many vulgates a r e a d o p t e d directly from indige­ n o u s languages, s o m e tribes a n d local cultures being prolific nomenclaturists, especially in Brazil ( M o n t e 1928). T h e s e suffer from f r e q u e n t spelling a n d p r o n u n ­ ciation variations, particularly in Brazil (where, in g e n e r a l , I follow von Ihering's [1968] o r t h o g r a p h y ) . At least partial ento­ mological glossaries exist for t h e following native t o n g u e s : Mayan (Welling 1958), Aztec ( = N á h u a t l , etc.; O r d o ñ o 1982), Kunza ( = A t a c a m e ñ o ; Munizaga a n d Her­ rera 1957), J í v a r o (Guallart 1968), TupíG u a r a n í (Tastevin 1923), a n d Q u e c h u a (García 1976). V e r n a c u l a r n a m e s a p p e a r according to n o consistent set of stan­ d a r d s , varying from place to place o r time to time with different origins a n d related to t h e n a t u r e of t h e society e m p l o y i n g t h e m (Stoetzel 1989). P h o n e t i c variations in spelling a r e c o m m o n . Scientists a n d e d u c a t e d people often form simple transliterations of technical n a m e s (muscids o r muscideos, from Muscidae) o r a c c o m m o d a t e n a m e s of classic origin (scarabs or escarabajos, from Greek karabos). L a y m e n a n d c o u n t r y folk a r e likely to invent q u a i n t , often descriptive appella­ tions that frequently apply to an insect's behavior (saltamonte = "hill j u m p e r " ) or

42

GENERAL ENTOMOLOGY

stinging abilities (lagarta d e fogo = "fire worm"), a n a t o m y (tijeretas = "scissor bear­ ers"), or that a r e o n a m a t a p o e t i c (cricket, chicharras), or that may be without obvious derivation (gallinipper). Sometimes these are literal translations from m o d e r n lan­ guages (scorpions, escorpiones) o r usages (tarantulas) n o t c o m m o n to t h e region. Mixtures of sympatric languages also occur (sede [Spanish] + ocuilin [Náhuatl] = sedeocuilin = silkworm). T h e only a t t e m p t to standardize c o m m o n n a m e s has been m a d e with pest species in English (Stoetzel 1989). Most languages have a b r o a d t e r m for insects a n d like animals, roughly equiva­ lent to t h e English, for e x a m p l e , " b u g " ("worm" o r " g r u b " ) : bicho (Spanish a n d Portuguese) a n d ocuilin (Náhuatl).

References AUDANT, A. 1941. Identification des insectes d'Haiti par leur nom creóle. Soc. Hist. Geogr. Haiti, Rev. 12(42): 51-55. [Not seen.] BAUCKE, O. 1961. Os nomes comuns dos in­ sectos no Rio Grande do Sul. Sec. Agrie, Porto Alegre. BIEZANKO, C. M., AND D. LINK. 1972.

Nomes

populares dos Lepidópteros no Rio Grande do Sul (Segundo Catalogo). Univ. Fed. Santa Maria, Bol. Tec. 4: 3-15. BRÜCHER, G. 1942. Lista de algunos nombres vulgares de insectos. Dept. San. Veg. (Min. Agrie, Santiago) Bol. 2(2): 120-125. DA SILVA, B. R. 1930-1934. Nomenclatura popular dos Lepidópteros do Distrito Federal and seus arredores. Vols. 1-5. O Campo, Rio de Janeiro. DOUROJEANNI, M. J. 1965. Denominaciones ver­ naculares de insectos y algunos otros in­ vertebrados en la selva del Perú. Rev. Peruana Entomol. 8: J31-137. GARCÍA, R. J. 1976. Nombre de algunos insectos y otros invertebrados en "Quechua." Rev. Peruana Entomol. 19: 13-16. GOTO, H. E. 1982. Animal taxonomy. Arnold (Inst. Biol., Stud. Biol. no. 143), London. GUALLART, J. M. 1968. Nomenclature JíbaroAguaruna de la fauna del Alto Marañón (Invertebrados). Biota 7: 195-209. IHERING, R. VON. 1968. Dicionário dos animáis do Brasil. Ed. Univ. Brasilia, Sao Paulo.

MONTE. O. 1928. Os nomes vulgares dos in­ sectos de Brasil. Almanak Agrie. Brasil. 1928: 028-289. MrxizAGA, C , AND J. HERRERA. 1985. Notas entomológicas ¿e Socaire (Obtenidas durante la Expedición Chileno-Alemana a Socaire, en mayo de 1957). Notas Centr. Est. Antropol., Univ. Chile, 1: 3 - 1 3 . ORDOÑO, C. M. 1982. Diccionario de zoología Náhuatl. Ed. Innovación, Mexico. PÉREZ D'ANGELLO, V. 1966. Concordancia entre

los nombres vulgares y científicos de los insectos chilenos. Mus. Nac. Hist. Nat. Not. Mens. 10(119): 2 - 7 . RIDE. W. D. 1.., C. W. SABROSKY, G. BERNARDI,

AND R. V. MELVILLE, eds. 1985. International

code of zoological nomenclature. 3d ed. Int). Trust Zool. Nomen., London. STOETZEL, M. B. 1989. Common names of insects and related organisms. Entmol. Soc. Anier. Lanham, Md. TASTEVIN, C. 1923. Nomes de plantas e animaes em Lingua Tupy. Rev. Mus. Paulista 13: 6 8 7 703. WELLING, E. C. 1958. Some Mayan names for certain I.epicloptera in the Yucatán penin­ sula. J. I.epidop. Soc. 12: 118.

INSECTS AND HUMAN CULTURE Aside from t h e i r i m p o r t a n c e as pests a n d our academic interest in insects, these crea­ tures, spiders, a n d related a r t h r o p o d s have considerable influence in that portion of h u m a n activity that m a y be called t h e humanities—music, a r t , literature, lan­ guage, religion, a n d folklore (fig. 1.8). T h e study of these influences is a general area of insect study called cultural entomology (Hogue 1987). E x a m p l e s a p p e a r a m o n g historical, m o d e r n , a n d i n d i g e n o u s peo­ ples. (Some o f t h e m o r e g e n e r a l a r e cited below; many o t h e r specific cases a r e scat­ tered t h r o u g h t h e r e m a i n d e r of this book in the sections on t h e various insects in­ volved; for Mexico, see M a c G r e g o r 1969.) Insects, spiders, c e n t i p e d e s , a n d scorpi­ ons a p p e a r in t h e Mayan Codices (Dresden, fro-Cortesianus, a n d Peresianus), indicat-

Figure 1.8 Decorative plates from modern Peru prominently featuring images of the fly (chuspi), revered in Incan times and a design motif in Andean art today. (Original, author's collection) ing an appreciation of their existence a n d their inclusion in cultural events, such as rituals, ceremonies, a n d dances. T h e fa­ mous Nasca figures include an i m m e n s e spider (fig. 1.9). Portions of same a r e also stylized as glyphs having linguistic signifi­ cance (Tozzer a n d Allen 1910). In the eighteenth century, it was believed that a small, red insect (still unidentified but called "coya" in t h e O r i n o c o region) caused severe skin e r u p t i o n s ; its effects could only be r e m e d i e d by ceremoniously passing t h e body t h r o u g h a fire m a d e from a specific grass ("guayacán") ( K a m e n - K a y e 1979). Many such curious accounts of insects fill the accounts of early visitors a n d colonists in t h e New World (Cowan 1865). Insects have lent their n a m e s to many places in Latin America. A m o n g t h e better known a r e C h a p u l t e p e c , t h e "hill of the g r a s s h o p p e r s " (chapulín = g r a s s h o p p e r + tepee = hill) w h e r e t h e Aztec E m p e r o r Montezuma's castle was built in what is now part of Mexico City, a n d U r u b a m b a , "plain of t h e insect" (uru = s p i d e r or

INSECTS AND HUMAN CULTURE

43

Í x

i

•¿J-U'f'r*

v \ ^ \

iUSPÜi lísl -

/>

caterpillar + pampa = plain), the sacred valley of the Incas near Cuzco in Peru. In modern times, insects symbolize nu­ merous ideas (fig. 1.10), especially in litera­ ture and folklore (Lenko and Papavero 1979). Science fiction and fantasy novels often use the dangerous qualities of many types to instill horror or malevolence. Su­ perstitions and fanciful stories attributing good or bad fortune to many insects, spiders, or the like, are believed by sectors of the population, especially those in re­ mote or primitive areas (Hogue 1985). The cultural use of insects is perhaps best developed among Indian tribes still surviving in many parts of Latin America (Berlin and Prance 1978; Hitchcock 1962; Kevan 1983; Posey 1978, 1983). The study of this aspect of cultural entomology is referred to as "ethnoentomology" (and includes some of the odd practical uses of Figure 1.12 In a variation of the "toucandira ritual" in which giant hunting ants of the genera Dinoponera and Paraponera are used, a mat tied with paper wasps is applied to the chest of this Roucouyenne Indian (French Guiana) to test his courage. (From H. Davis, The Jungle and the Damned 1952, Duell, Sloan and Pearce, New York; reproduced with permission)

Figure 1.9 The spider was an eminent symbol in Peruvian cultures of prehistory. It is displayed on a grand scale among the Nasca figures in the southern desert.

]

:w M»L

.W^~ "

3

• sc C I o

^

km'.

\eifl: . . . .

l

Socti

/'i ^

*

i

Figure 1.10 Political graffiti on wall in San José, Costa Rica, by group using a social insect, the ant, as a symbol of Socialist doctrine; 1978. (Original, author's collection)

Figure 1.11 Image from the Codex Telleriano Remensis of the Aztec deity, Itzpapálotl, in nature represented by wild silk moths of the genus Rothschildia. (Hand copy by Carlos Beutelspacher in Mariposas entre los Antiguos Mexicanos, 1989; reproduced with author's permis­ sion)

insects, such as for food or medicine; see valuable insects, chap. 3). In many Amazonian Indian groups, insects were in the past and are still today venerated religiously, and they play central roles as deities or mythic figures (fig. 1.11). The four guardians of the cardinal points in Warao cosmology are social insects—two wasps, a bee, and a termite (Wilbert 1985). Ritual also incorporates insects, for exam­ ple, the giant hunting ants (Dinoponera) in puberty ceremonies practiced by various Amazonian tribes (Liebrecht 1886; fig. 1.12). Similarly, pain is endured from the stings of wasps whose nests are purposely molested as a part of rites of passage among the Gorotire-Kayapó in Brazil (Posey 1981).

INSECTS AND HUMAN CULTURE

45

n c i r Fihnoentomology of the Kayapó Indi­ e s of central Brazil. J. Ethnobiol. 1: 165-174. 'SFY D. A. 1983. Ethnomethodology as an I'WiV guid e to cultural systems: T h e case of the insects and the Kayapó Indians of Amazonia. Rev Brasil. Zool. 1: 135-144. )ZZER, A. M-, AND G. M. ALLEN. 1910. Animal

figures in the Maya Codices. Harvard Univ, Peabody Mus., Pap., Amer. Archaeol. Ethnol. 4: 273-372, pis. 1-39. WILBERT J. 1985. T h e house of the swallowtailed kite: Warao myth and the art of think­ ing in images. In G. Urton, ed., Animal myths and metaphors. Univ. Utah, Salt Lake City.

Figure 1.13 Modern Peruvian Indian (Yagua) necklace using beetle parts as main decorative element (Original, author's collection) Metallic beetle parts and even galls (Ber­ lin and Prance 1978) are used in body ornamentation by Indians in all parts of the region (fig. 1.13). Insects, especially musical species (crickets and katydids), lu­ minescent forms (headlight beetles), and large beetles, and orthopterans are kept as pets or curiosities. Many species are eaten, both for sustenance and as delicacies (fig. 1.14). References BERLIN, B., AND G. T PRANCE. 1978. Insect galls

and human ornamentation: T h e ethnobotanical significance of a new species of Licania from Amazonas, Peru. Biotropica 10: 81-86. COWAN, F 1865. Curious facts in the history of insects. Lippincott, Philadelphia. HITCHCOCK, S. W. 1962. Insects and Indians of the Americas. Entomol. Soc. Amer. Bull. 8(4): 181-187. HOGUE, C. L. 1985. Amazonian insect myths. Terra 23(6): 10-15. HOGUE, C. L. 1987. Cultural entomology. Ann. Rev. Entomol. 32: 181-199.

46

GENERAL ENTOMOLOGY

Figure 1.14 A bottle of mezcal containing a maguey worm (Comadla redtenbacheri, Cossidae) as an extra treat for the drinker. The beverage was important in ancient and modern Mexican culture. The insect retains today its natural association with the plant and its product. (Los Angeles County Museum of Natural History collection) KAMEN-KAYE, D. 1979. A bug and a bonfire. J.

Ethnopharm. 1: 103-110. KEVAN, Ó. K. MCE. 1983. T h e place of grasshop­ pers and crickets in Amerindian cultures. 2d Trien. Meet. Pan American Acrid. Soc. (Bozeman, Mont., 1979) Proc. P. 8-74c. LENKO, K., AND N. PAPAVERO. 1979. lnsetos no

folclore. Conselho Estad. Artes Cien. Hu­ man., Sao Paulo. LIEBRECHT, F 1886. Tocandyrafestes. Zeit. Ethnol. 18: 350-352. MACGREGOR, R. 1969. La representation des

insectes dans l'ancien Mexique. L'Entomologiste25: 1-8. POSEY, D. A. 1978. Ethnoentomological survey of Amerind groups in lowland Latin Amer­ ica. Fia. Entomol. 61: 225-228. POSEY, D. A. 1981. Wasps, warriors and fearless

INSECTS AND HUMAN CULTURE

47

^

ECOLOGY

As elsewhere, a wide variety of complex, interacting ecological factors d e t e r m i n e the distributions a n d survival a d a p t a t i o n s of insects in Latin A m e r i c a ( H u f f a k e r a n d Rabb 1984, Price 1984). T h e s e may be categorized e i t h e r as g e o g r a p h i c (physical factors) o r biotic (living agents) (Walter 1979). B o t h a r e viewed from historical as well as c o n t e m p o r a r y perspectives.

References HUFFAKER, C. B., AND R. L. RABB, eds.

1984.

Ecological entomology. Wiley, New York. PRICE, P. W. 1984. Insect ecology. 2d ed. Wiley, New York. WALTER, H. 1979. Vegetation of the earth and ecological systems of the geo-sphere. 2d ed. Springer, New York. Translated from 3d rev. German edition.

GEOGRAPHY O f f u n d a m e n t a l i m p o r t a n c e to the occur­ r e n c e of insect life is the surface c h a r a c t e r of t h e land, its configuration, chemistry, a n d relief, o r its physiography, which p r o ­ vides footholds for t h e very existence a n d continuity of individuals. Equally basic are the c o n d i t i o n s of climate, that is, t e m p e r a ­ t u r e , m o i s t u r e , sunlight, as well as the conditions of the m e d i u m (soil, water, o r a t m o s p h e r e ) , which give suitable substance a n d s u p p o r t for life. T h e s e c o m p o n e n t s of g e o g r a p h y w o r k t o g e t h e r to create broad habitats for insects called life zones. From a consideration of the historical g e o g r a p h y of the flora a n d f a u n a of such areas,

48

b i o g e o g r a p h i c provinces may also be de­ fined which contain characteristic g r o u p s of species (faunistics).

Physiography Past a n d present-day s h a p e s a n d positions of landmasses, their elevations, connec­ tions, a n d surface t e x t u r e s , delineate b r o a d areas within Latin A m e r i c a which d e t e r m i n e in a most elemental way the distribution of insects. Physiographic subdi­ visions in Latin America have b e e n out­ lined according to various schemes. A simplified version is p r e s e n t e d below (Fig. 2.1); it is modified from S a u e r (1950) a n d Sick (1969).

References SAUER, C. O. 1950. Geography of South Amer­ ica, Handbk. So. Amer. Indians 6: 319-344. SICK, W. D. 1969. Geographic substance. In E. J. Fittkau, J. lilies, H. Klinge, G. H. Schwabe, and H. Sioli, eds., Biogeography and ecology in South America. 2: 449-474. Junk, T h e Hague.

Climate and Medium Each climatic factor exerts its critical ef­ fects in a variety of ways: t e m p e r a t u r e as freezing point, highs, lows, m e a n s , ranges, heat, cold, daily fluctuations; m o i s t u r e as rainfall, dew, fog, clouds (and, of course, by d e t e r m i n i n g the f u n d a m e n t a l lifes u p p o r t i n g media, aquatic versus terres­ trial); sunlight as day a n d night, s h a d e , illumination, radiation, a n d p h o t o p e r i o d . All act over long time periods as w e a t h e r

Figure 2.1 MAJOR PHYSIOGRAPHIC AREAS OF LATIN AMERICA (from Sauer 1950 and Sick 1969). MIDDLE AMERICA: 1. Mexican Highlands; 2. Isthmian America (lowland Mexico, Central America); 3. West Indies (Greater and Lesser Antillean Islands); 4. Bahamas; SOUTH AMER­ ICA: 5. Pacific Coastal plain; 6. Andes (including Caribbean Borderlands and Bolivian Altiplano); 7. Amazon Basin; 8. Orinoco Basin; 9. Guiana Highlands; 10. Brazilian Highlands (including Brazilian Coastal Mountains) and Mato Grosso, Plateau of Paraná; 11. Llanos de Mamoré; 12. ParanáParaguay Depression; 13. Gran Chaco; 14. Pampas; 15. Patagonia; INSULAR AMERICA: 16. Oceanic islands (Pacific: Galapagos and Revillagigedo Archipelagos, Cocos Island, Easter Island, Juan Fernandez etc. Atlantic: Fernando de Noronha Archipelago, Ascensión Island); 17. Continental islands (Falklands-Malvinas, Tres Marias, Isla de Coiba, Pearl Islands, Magellanic Archipelago).

climate, a n d seasonality a n d , of course, interact with physiography. T h e interplay of physical agents in t h e a t m o s p h e r e cre­ ates a m u l t i t u d e of e n v i r o n m e n t s that a r e also circumscribable a n d c a n even b e m a p p e d . Many schemes have b e e n devised to classify these directly as climatic areas ( K ó p p e n a n d Geiger 1 9 3 1 - 1 9 3 4 ) . Major shifts in t h e e a r t h ' s climatic pat­ tern occur at i r r e g u l a r intervals, c h a n g i n g the rainfall, t e m p e r a t u r e , a n d o t h e r aspects of t h e weather, s o m e t i m e s drastically over wide areas. I n Latin A m e r i c a , t h e bestk n o w n manifestation of such planetaryscale a t m o s p h e r i c fluctuations is t h e El Niño p h e n o m e n o n , itself p a r t of t h e socalled, larger S o u t h e r n Oscillation that in­ volves m u c h of t h e Pacific a n d I n d i a n O c e a n basins of t h e S o u t h e r n H e m i s p h e r e (Philander 1990). At r o u g h l y seven- to tenyear intervals, t h e w a r m e q u a t o r i a l c o u n t e r c u r r e n t in t h e P a n a m a bight shifts strongly to t h e south, u n d u l y h e a t i n g a n d displacing the n o r m a l l y cool, northward-flowing H u m b o l d t C u r r e n t . As a result, heavy rains come t o t h e P e r u v i a n coastal deserts, a n d vegetation flourishes (especially o n t h e lomas; see Special Habitats, below). T h e effects of El N i ñ o o n coastal insect p o p u l a ­ tions h a v e b e e n f o u n d t o be significant in some cases (Beingolea 1987a, 19876). Asso­ ciated w e a t h e r a n o m a l i e s m a y b e felt e v e n far inland o n t h e c o n t i n e n t , for e x a m p l e , p r o l o n g e d d r y n e s s in t h e western A m a z o ­ nian rain forest o r cold snaps in s u b A n d e a n valleys. Brief b u t severe p e r i o d s of cold w e a t h e r fronts, o r i g i n a t i n g in Antarctica, pass across t h e A m a z o n Basin in o d d years, usually d u r i n g early J u l y (Días Frios d e San J u a n , friagem) (Ratisbona 1976: 226, 237). Daily m i n i m u m t e m p e r a t u r e m a y d r o p from a n o r m a l 20° to 8° C. T h e sky becomes heavily c l o u d e d , a s t r o n g wind blows, a n d it fails t o r a i n . Responses of insects to these stresses may b e d r a m a t i c o r subtle b u t a r e as yet virtually u n s t u d i e d in Latin America. T h e

50

ECOLOGY

impact of such s u d d e n w e a t h e r changes on these small, ectothermic creatures surely must be intense, especially o n temperature-sensitive species. Interfer­ ence with r e p r o d u c t i o n a n d p o p u l a t i o n "die-offs" should be expected. T h e effects of climate a r e felt most immediately in t h e a t m o s p h e r e . T h e y a r e manifest equally b u t m o r e slowly in t h e aquatic m e d i u m , in s t a n d i n g (lentic) waters (ponds, lakes, etc.), in m o v i n g (lotic) waters (streams, rivers) (Fittkau 1964; Macan 1962, 1974), a n d in t h e soil (Kühnelt 1961).

References BEINGOLEA, O. D. 1987a. El fenómeno "El Niño" 1982-83 y algunos insectos-plaga en la costa peruana. Rev. Peruana Entomol. 28: 55-57. BEINGOLEA, O. D. 19876. La langosta Schislocerca interrita en la costa norte del Perú, durante 1983. Rev. Peruana Entomol. 28: 35-40. KÓPPEN, W., AND R. GEIGER 1931-1934. Hand-

buch der Klimalologie. Vols. 1 —4. Borntraeger, Berlin. KÜHNELT, W. 1961. Soil biology with special reference to the animal kingdom. Faber 8c Faber, London. FITTKAU, E. J. 1964. Remarks on limnology of central-Amazon rain-forest streams. Int. Verh. Limnol., Verh. 15: 1092-1096. MACAN, T. T. 1962. T h e ecology of aquatic insects. Ann. Rev. Entomol. 7: 261-288. MACAN, T. T. 1974. Freshwater ecology. 2d ed. Wiley, New York. PHILANDER, S. G. H. 1990. El Niño, La Niña and the Southern Oscillation. Academic, San Diego. RATISBONA, L. R. 1976. The climate of Brazil, hi W. Schwerdtfeger, ed., Climates of Central and South America, world survey of climatol­ ogy. 12: 219-293. Elsevier, Amsterdam.

Vegetation Zones Physicochemical e n v i r o n m e n t a l factors not only act directly o n insect life forms b u t influence t h e m indirectly t h r o u g h t h e kinds of plant g r o w t h they allow (biotic factors). G r o u p i n g s of plants a d a p t e d to a particular set of soil a n d climatic conditions delimit b r o a d e n v i r o n m e n t s for insects.

Many types of vegetation a r e p r e s e n t in Latin America, a n d these a r e classified according to various systems (e.g., B e a r d 1944, G r a h a m 1973, H u e c k a n d Seibert 1950, Weber 1969). T h e s e 1 9 7 2 ' Sauer systems a r e only regionally applicable b e ­ cause they often define units in t e r m s of specific plant taxa p r e s e n t . A universal scheme, globally applicable, is H o l d r i d g e ' s (1967, 1982), which c o m b i n e s t h e effects of elevation, latitude, rainfall, a n d t e m p e r a ­ ture to define vegetational formations, in­ d e p e n d e n t of floristic e l e m e n t s . It is very a p p a r e n t that t h e n a t u r e of vegetation plays a p r i m a r y role in d e t e r m i n ­ ing t h e Neotropical insect f a u n a in each physiographic a r e a . T h e special richness of the A m a z o n Basin is a good case in point. Contrary to f o r m e r ideas, t h e region's vege­ tation h a s n o t existed continuously u n ­ changed for tens of millions of years b u t has varied considerably from n e a r d e s e r t t o lush forest in r e c e n t geologic p e r i o d s , par­ ticularly d u r i n g t h e Pleistocene A g e , d u r ­ ing alternating arid a n d h u m i d conditions. In t h e d r i e r phases, m o i s t u r e - r e q u i r i n g vegetation s h r u n k greatly a n d f r a g m e n t e d into forest patches w h e r e rainfall persisted which was a d e q u a t e for their survival ("ref­ uge theory," Haffer 1982; b u t see E n d l e r 1982). This disjunction into forest islands iso­ lated from each o t h e r by grassland o r even desertlike plant cover divided m a n y for­ merly c o n t i n u o u s p o p u l a t i o n s a n d led to their evolution into n e w species. Wet phases, such as t h e world is n o w e x p e r i e n c ­ ing, allowed t h e p a t c h e s to e x p a n d again and t h e gaps b e t w e e n t h e m to close. B u t evidences of t h e f o r m e r islands, o r refugia, are still p r e s e n t as c o n c e n t r a t i o n s of e n ­ demics. A m o n g insects, this is shown espe­ cially well by butterflies (Brown 1982) a n d among a r a c h n i d s by scorpions ( L o u r e n c o 1986). This h e t e r o g e n e i t y in A m a z o n i a n forest and wet forests in o t h e r areas partly ex­ plains t h e latest a n d o n e of t h e most

e x t r e m e forces s h a p i n g t h e m o d e r n e n t o m o f a u n a of t h e basin (Simpson a n d Haffer 1978).

References BEARD, J. S. 1944. Climax vegetation in tropical America. Ecology 25: 127-158. BROWN, JR., K. S. 1982. Paleoecology and re­

gional patterns of evolution in Neotropical forest butterflies. In G. T. Prance, ed., Biologi­ cal diversification in the tropics. Columbia Univ. Press, New York. Pp. 255-308. ENDLER, J. A. 1982. Pleistocene forest refuges: Fact or fancy. In G. T. Prance, ed., Biological diversification in the tropics. Columbia Univ. Press, New York. Pp. 641-657. GRAHAM, A., ed. 1973. Vegetation and vegeta­ tional history of northern Latin America. Elsevier, Amsterdam. HAFFER, J. 1982. General aspects of the Refuge Theory. In G. L Prance, ed., Biological diver­ sification in the tropics. Columbia Univ. Press, New York. Pp. 6-24. HOLDRIDGE, L. R. 1967. Life Zone ecology. Trop. Sci. Ctr., San José, Costa Rica. HOLDRIDGE, L. R. 1982. Ecología basada en zonas de vida. Insto, lnteramer. Coop. Agrie. San José, Costa Rica. HUECK,

K.,

AND P. SEIBERT.

1972.

Vegeta-

tionskarte von Südamerika (Mapa de la vegetación de America del sur). Fischer, Stuttgart. LOURENC.O, W. R. 1986. Diversité de la faune scorpionique de la region amazonienne; cen­ tres d'endémisme; nouvel appui á la théorie des refuges forestiers du Pleistocene. Amazoniana9: 559-580. SAUER, C. O. 1950. Geography of South Amer­ ica. Handbk. So. Amer. Indians 6: 319-344. SIMPSON, B. B., AND J. HAFFER. 1978. Speciation

patterns in the Amazonian forest biota. Ann. Rev. Ecol. Syst. 9 : 4 9 7 - 5 1 8 . WEBER, H. 1969. Zur natürlichen Vegetationsgliederung von Südamerika. In E. J. Fittkau, J. lilies, H. Klinge, G. H. Schwabe, and H. Sioli, eds., Biogeography and ecology in South America. 2:475-518. Junk, T h e Hague.

Artificial Environments T h e foregoing discussion h a s b e e n con­ c e r n e d with n a t u r a l o r original conditions a n d p a t t e r n s of native flora a n d fauna. Since c o m i n g to t h e s o u t h e r n lands of t h e

GEOGRAPHY

51

New World 20,000 to 50,000 years a g o , h u m a n s have modified t h e original life zones to varying d e g r e e s (Kiinkel 1963), even c r e a t i n g l a r g e tracts of essentially new, "artificial life zones." Such a r e t h e cities, farms, a n d vast grasslands for cattle grazing t h a t have c o m e i n t o b e i n g since t h e C o n q u e s t . Even b e f o r e C o l u m b u s , t h e in­ d i g e n o u s p o p u l a t i o n cut a n d b u r n e d siz­ able tracts of forest to s u p p o r t shifting a g r i c u l t u r e , t u r n i n g t h e m into plots of cultivated species that only slowly r e t u r n e d to climax status t h r o u g h successional stages of different vegetation. T h e I n c a n civilization fashioned a n e w A n d e a n land­ scape by their extensive t e r r a c i n g , grazing, irrigation, a n d r o a d b u i l d i n g . I n a d d i t i o n to c h a n g e s contrived for living space a n d a g r i c u l t u r e , t h e r e have also b e e n impacts o n n a t u r a l p o p u l a t i o n s of animals a n d plants t h r o u g h h u n t i n g a n d g a t h e r i n g , u n i n t e n t i o n a l pollution, a n d erosion. Such modifications force adjust­ m e n t s by t h e insect i n h a b i t a n t s . T h i s is most i n t e n s e in t h e u r b a n setting w h e r e new c o m p l e x e s of species a d a p t e d to civili­ zation c o m e into b e i n g . T h e e n t o m o l o g y of cultivated fields o f single o r m i x e d c r o p s also d e p a r t s widely f r o m t h e n o r m in natural environments.

Reference KÜNKEL, G. 1963. Vegetationszerstorung und

Bodenerosion in Lateinamerika. Arch. Naturschutz Landschaft. 3(1): 5 9 - 8 0 .

ECOSYSTEMS Not only t h e vegetation b u t t h e c o m m u ­ nity of all t h e p l a n t s , insects, a n d o t h e r a n i m a l s m u s t be c o n s i d e r e d t o g e t h e r to describe t h e c o n d i t i o n s of a particular habitat. T h e o r g a n i s m s (biocenoses) lo­ cated in a p a r t i c u l a r place (biotope) consti­ tute an ecosystem. W i t h i n t h e ecosystem, each o r g a n i s m h a s a functional role

52

ECOLOGY

(niche). T h e niches of insects in Latin America have b e e n little studied, o t h e r t h a n in agricultural m o n o c u l t u r e s . Habitats a r e recognizable o n different levels. T h e most i m m e d i a t e e n v i r o n m e n t w h e r e t h e d e t e r m i n a n t s of life specifically affect an o r g a n i s m is its microhabitat. Ex­ amples a r e infinite in n u m b e r ; t h e i n t e r i o r of a decaying log o r water-filled c u p of a bromeliad a r e e x a m p l e s . T h e vertical distri­ bution of different species of mosquitoes (Bates 1944) a n d butterflies (Papageorgis 1975) in forests is evidence of t h e effects of subtle e n v i r o n m e n t a l d e t e r m i n a n t s . T h e existence of so m a n y microhabitats partly accounts for t h e vast diversity of t h e in­ sects, c r e a t u r e s so a d e p t at calling h o m e the smallest a n d most d e m a n d i n g of living spaces. Larger, m o r e inclusive areas c o n t a i n i n g microhabitats, such as t h e vegetation zones (e.g., forest canopy) a n d small physio­ g r a p h i c features (caves, lakes), a r e m a c r o habitats. Still g r e a t e r g r o u p i n g s a n d larger expanses of e a r t h f o r m i n g grossly recogniz­ able macrohabitats a r e called life zones or biomes (e.g., tropical forest, deserts). T h e statuses of Latin A m e r i c a n insects in sev­ eral special habitats a r e discussed below.

Special Habitats T h e g r e a t n u m b e r of n a t u r a l Neotropical insect m a c r o - a n d microhabitats m a k e s it impossible to discuss m o r e t h a n a few of t h e most distinctive, peculiar, a n d i m p o r t a n t . T h e y form particular places for insects whose s t r u c t u r e a n d activities m a y be very different from those of t h e o t h e r insects in the s u r r o u n d i n g g e n e r a l e n v i r o n m e n t . Most insects studied in artificial habitats are injurious species a n d a r e discussed in C h a p t e r 3, Practical Entomology. A few investigations have focused o n o t h e r faunal elements of plantations (Young 1986a, 19866), habitations, a n d like a r e a s u n d e r human management.

References BATES, M. 1944. Observations on the distribu­ tion of diurnal mosquitoes in a tropical forest. Ecology 25: 159-170. PAPAGEORGIS, C. 1975. Mimicry in Neotropical butterflies. Amer. Sci. 63: 522-532. YOUNG, A. M. 1986a. Notes on the distribution and abundance of Dermaptera and Staphylinidae (Coleóptera) in some Costa Rican cacao plantations. Entomol. Soc. Wash. Proc. 88: 328-343. YOUNG, A. M. 19866. Notes on the distribution and abundance of ground and arboreal nest­ ing ants (Hymenoptera: Formicidae) in some Costa Rican cacao habitats. Entomol. Soc. Wash. Proc. 88: 550-571. Forest Canopy Until very recently, t h e forest c a n o p y h a s been p e n e t r a t e d by insect researchers only with great difficulty ( H i n g s t o n 1932). T h e earliest w o r k e r s h a d to be c o n t e n t with fortuitous t r e e falls even to gain a glimpse of u p p e r - s t o r y life. T h e c a n o p y could be reached by skilled native climbers, b u t they carried n o scientific expertise aloft, a n d their activities h a d to be directed ineffec­ tively by t h e i r e a r t h b o u n d e m p l o y e r s . Other a p p r o a c h e s , still r e m o t e ones, have been to elevate various kinds of traps to catch some of t h e fauna o r knock it d o w n with quick-acting, b i o d e g r a d a b l e insecti­ cides (Erwin 1983). Greater i m p r o v e m e n t in access c a m e with t h e c o n s t r u c t i o n of a r b o r e a l l a d d e r s and platforms (Porter a n d DeFoliart 1981), towers (a f a m o u s o n e o n B a r r o C o l o r a d o Island, P a n a m a ) , o r elevated causeways, from which observations a n d collections could be m a d e . However, these offer very limited mobility a n d a r e costly to construct and maintain. Lately, s o m e practical m e a n s have been found n o t only to m o v e a b o u t a n d obtain specimens f r o m this c o m p l e x realm b u t even to carry o u t e x t e n d e d studies within it (Mitchell 1982). Even scientists themselves are now able to go into t h e canopy, using m o u n t a i n e e r i n g t e c h n i q u e s (Perry 1980,

1984; Perry a n d Williams 1981). U n f o r t u ­ nately, few entomologists a r e t r a i n e d in b o t h t h e academic a n d athletic aspects of this d e m a n d i n g , a l t h o u g h highly r e w a r d ­ ing, a p p r o a c h . T h e results of these efforts, a l t h o u g h f r a g m e n t e d a n d imperfect, d e m o n s t r a t e the startling fact that t h e c a n o p y has an extremely rich insect c o m p l e m e n t . T h i s s t r a t u m of forests is c o n s i d e r e d to be t h e "last frontier" in tropical e n t o m o l o g y a n d is attracting study from a n u m b e r of view­ points: faunistics, pollination ecology, a n d even populational biology (see below).

References ERWIN, T. L. 1983. Beetles and other insects of tropical forest canopies at Manaus, Brazil, sampled by insecticidal fogging techniques. In S. L. Sutton, T. C. Whitmore, and A. C. Chadwick, eds., Tropical rain forest: Ecology and management. Blackwell, Oxford. Pp. 59-75. HINGSTON, R. W. 1932. A naturalist in the

Guiana forest. Longmans Green, New York. MITCHELL, A. W. 1982. Reaching the rain forest roof (A handbook on techniques of access and study in the canopy). Leeds Phil. Lit. Soc. Leeds, Eng. PERRY, D. R. 1980. 1 probe the jungle's last frontier. Int. Wildlife 10: 5 - 1 1 . PERRY, D. R. 1984. T h e canopy of the tropical rain forest. Sci. Amer. 251: 138-147. PERRY, D.

R.,

AND J.

WILLIAMS.

1981.

The

tropical rain forest canopy: A method provid­ ing total access. Biotropica 13: 283-285. PORTER, C. LL, AND G. R. DEFOLIART. 1981. T h e

man-biting activity of phlebotomine sand flies (Díptera: Psychodidae) in a tropical wet forest environment in Colombia. Arq. Zool. Sao Paulo 30: 81-158. Amazon Inundation Forests Vast expanses b o r d e r i n g t h e A m a z o n a n d its major tributaries a r e annually flooded to a d e p t h of several m e t e r s for p e r i o d s of five to six m o n t h s o r m o r e . T h e s e forests (called igapó locally) h a r b o r an insect f a u n a especially a d a p t e d to t h e stresses o f alter­ nately rising a n d r e c e d i n g waters. T h e terrestrial species that live in these

ECOSYSTEMS

53

forests a r e n u m e r o u s a n d very diverse. T h e y h a v e evolved strategies to c o m p e n ­ sate for the periodic loss of their terrestrial habitat by exercising mobility to escape into t h e c a n o p y o r to o t h e r d r y areas o r acquiring survival a d a p t a t i o n s for c o p i n g with i n u n d a t i o n (Adis 1977, Adis et al. 1988). Spiders a n d m a n y flightless species of g r o u n d beetles, for e x a m p l e s , a r e k n o w n to ascend into the u p p e r forest level (Erwin a n d Adis 1982) d u r i n g flood peri­ ods. M e m b e r s of t h e cockroach genus Epilampra, however, have a c q u i r e d t h e ca­ pacity t o swim a n d h a v e t h e h i n d m o s t spiracles situated o n s h o r t stalks to aid b r e a t h i n g while they a r e in the water. Only minor elements of the g r o u n d fauna a r e p r e s e n t h e r e (e.g., p s e u d o s c o r p i o n s ; Adis a n d M a h n e r t 1985, Adis et al. 1988), t h e r e being n o persistent g r o u n d litter o r stable soil. In r e s p o n s e to t h e d r y i n g p h a s e , aquatic insect i n h a b i t a n t s of these areas r e t u r n to the m a i n c h a n n e l s from which the floodwaters arose. S o m e m a y b e c o m e t r a p p e d in small, d a m m e d d e p r e s s i o n s , b u t normally they e i t h e r take t o t h e a i r a n d fly t o water or b u r r o w into t h e m u d w h e r e they m a y r e m a i n inactive for t h e d u r a t i o n of t h e dry season.

References ADIS, J. 1977. Programa mínimo para analises de ecosistemas: Artrópodos terrestres em florestas inundáveis da Amazonia central. Acta Amazónica 7: 223-229. ADIS, J., AND V. MAHNERT. 1985. On the natural

history and ecology of Pseudoscorpions (Arachnida) from an Amazonian blackwater inundation forest. Amazoniana 9: 297-314. ADIS, J., V. MAHNERT, J. W. DE MORÁIS, AND

J. M. GOMES. 1988. Adaptation of an Amazo­ nian pseudoscorpion (Arachnida) from dry­ land forests to inundation forests. Ecology 69:287-291. ERWIN, T. L., AND J. A D I S . 1982. Amazonian

inundation forests: Their role as short-term refuges and generators of species richness and taxon pulses. In G. T Prance, ed., Biologi­ cal diversification in the tropics. Columbia Univ. Press, New York. Pp. 358-371.

54

ECOLOGY

Soil and Ground Litter T h e soil a n d its overlying layer of o r g a n i c litter constitute t h e habitat of a great m a n y small t o very small a r t h r o p o d s (Kiihnelt 1961). D o m i n a n t a m o n g these are ants, springtails, spiders, psocids, thrips, cryptostigmatic mites, a n d microbeetles (Penny et al. 1978), b u t larger insects a r e sometimes also f o u n d h e r e (Young 1983). It is incredible that such a r t h r o p o d s m a y c o m p r i s e almost 5 0 per­ cent of the total soil a n d litter biota in t h e central A m a z o n forest, ants a n d termites alone f o r m i n g a b o u t 60 p e r c e n t of this (Fittkau a n d Klinge 1973). Most occupy the u p p e r 5 to 8 centimeters of the soil. T h e composition of this f a u n a a n d its ecology have b e e n t h e subject of m a n y studies in the Neotropics, especially in for­ ests (e.g., Williams 1941, Deboutteville a n d Rapoport 1962-1968, Lieberman and Dock 1982) w h e r e m o i s t u r e seems to be the major factor d e t e r m i n i n g the seasonal dis­ tribution of species a n d p o p u l a t i o n fluctua­ tions. A u g m e n t a t i o n s in diversity a n d a b u n ­ d a n c e generally follow increases in water content (Levings a n d W i n d s o r 1984), usu­ ally from rainfall d u r i n g the wet season in all life zones (Dantas a n d S c h u b a r t 1980, L i e b e r m a n a n d Dock 1982, Willis 1976). Strickland (1945) c o n c l u d e d that t h e gen­ eral e n v i r o n m e n t of an a r e a is m o r e i m p o r ­ tant t h a n t h e soil type in d e t e r m i n i n g t h e sizes of g r o u n d a n d litter p o p u l a t i o n s . T h a t a variety of conditions would be d e t e r m i ­ n a n t s was confirmed by studies of forest litter a r t h r o p o d s in s o u t h e a s t e r n Peru (Pear­ son a n d D e r r 1986).

References DANTAS, M., AND H. O. R. SCHUBART. 1980.

Correlacáo dos indices de agregacáo de Acari e Collembola com 4 fatores ambientáis numa pastagem de terra firme da Amazonia. Acta Amazónica 10: 771-774. DEBOUTTEVILLE, C. D., AND E. RAPOPORT, eds.

1962-1968. Biologie de 1'Amérique Australe. Etudes sur la faune de sol; 2: Etudes sur la faune du sol; 3: Etudes sur la faune du sol +

Documents biogéographiques; 4, Documents biog. et ecologiques. Ed. Cen. Nat. Rech. Sci., Paris. FITTKAU,

E. J.,

AND H.

KLINGE.

1973.

On

biomass and trophic structure of the central Amazonian rain forest ecosystem. Biotropica 5:2-14. KÜHNELT, W. 1961. Soil biology with special reference to the animal kingdom. Faber & Faber, London. LEVINGS, S. C , AND D. M. WINDSOR. 1984. Litter

moisture content as a determinant of litter arthropod distribution and abundance dur­ ing the dry season on Barro Colorado Island, Panama. Biotropica 16: 125-131. LIEBERMAN, S., AND C. F DOCK. 1982. Analysis

of the leaf litter arthropod fauna of a lowland tropical evergreen forest site (La Selva, Costa Rica). Rev. Biol. Trop. 30: 27-34. PEARSON, D. L., AND J. A. DERR. 1986. Seasonal

patterns of lowland forest floor arthropod abundance in southeastern Perú. Biotropica 18: 244-256. PENNY, N. D., J.

R. ARIAS, AND H.

O.

R.

SCHUBART. 1978. Tendencias populacionais de fauna de Coleópteros do solo sob floresta de terra firme na Amazonia. Acta Amazónica 8: 259-265. STRICKLAND, A. H. 1945. A survey of the

arthropod soil and litter fauna of some forest reserves and cacao estates in Trinidad, British West Indies. J. Anim. Ecol. 14: 1-11. WILLIAMS, E. C. 1941. An ecological study of the floor fauna of the Panamanian rain forest. Chicago Acad. Sci. Bull. 6: 63-124. WILLIS, E. D. 1976. Seasonal changes in the invertebrate litter fauna on Barro Colorado Island, Panamá. Rev. Brasil. Biol. 36: 6 4 3 657. YOUNG, A. M. 1983. Patterns of distribution and abundance in small samples of litterinhabiting Orthoptera in some Costa Rican cacao plantations. New York Entomol. Soc. J. 91:312-327. Black Water Lakes and Rivers Some lowland tropical river basins contain tributaries a n d landlocked basins (oxbow lakes, cochas) with tea-colored water that in the d e p t h s a p p e a r s black. T h e s e are distin­ guished from so-called white waters n o t only by their color b u t by physical a n d chemical p r o p e r t i e s . W h i t e waters (actually a milky chocolate color) a r e n u t r i e n t rich, neutral to slightly alkaline, a n d turbid.

Black waters usually flow from n u t r i e n t poor, sandy soils a n d t h u s a r e low in minerals b u t a r e acidic a n d m a y contain high concentrations of o r g a n i c c o m p o u n d s (tannic acids, phenolics) leached from vege­ tation a n d toxic to insects. T h e water a n d s u r r o u n d i n g land thus afford unfavorable conditions for insects, a n d such black water basins generally have d e p a u p e r a t e entornofaunas ( J a n z e n 1974). S o m e specially a d a p t e d aquatic types, however, such as certain c h i r o n o m i d m i d g e s (Fittkau 1971) a n d water mites (Tundisi et al. 1979), c a n be very n u m e r o u s , even in the most heavily c h a r g e d water. T h e G u i a n a Shield of n o r t h e r n South A m e r i c a is a large black w a t e r a r e a a n d is notorious for the p o o r n e s s of its productiv­ ity. O t h e r similar such regions a r e f o u n d in the Brazilian H i g h l a n d s a n d o n t h e Yuca­ tán Peninsula. Clear waters (greenish t o clear) a r e also recognized b u t a r e biologically similar t o black water.

References FITTKAU, E. J. 1971. Distribution and ecology of Amazonian chironomids (Diptera). Can. Ento­ mol. 103: 407-413. JANZEN, D. H. 1974. Tropical blackwater rivers, animals and mast fruiting by the Dipterocarpaceae. Biotropica 6: 69—103. TUNDISI, J. G., A. M. P. MARTINS, AND T. MATSU-

MURA. 1979. Estudos ecológicos preliminares em sistemas aquáticos em Aripuaná. Acta Amazónica 9: 311-315. Caves Insects a n d m a n y related terrestrial a r t h r o ­ p o d s of diverse g r o u p s a r e t r u e troglobites (obligate cavernicoles, i.e., animals nar­ rowly a n d specifically a d a p t e d for life d e e p in caves; Culver 1982, H o f f m a n n et al. 1986). However, tropical caves a r e usually d o m i n a t e d by species classed as troglophiles, which also live in n o n c a v e habitats. S o m e a r e omnivores, b u t they a r e m o r e normally specialized for particular foods a n d a r e of two basic types: scavengers a n d

ECOSYSTEMS

55

p r e d a t o r s . T h e f o r m e r feed o n the d r o p ­ pings of bats (Gnaspini 1989), oil birds, a n d o t h e r h i g h e r f a u n a ( g u a n o p h a g e s ) or take n o u r i s h m e n t from o r g a n i c debris such as insect a n d v e r t e b r a t e c a r r i o n a n d plant m a t t e r that washes or falls into the caves or is b r o u g h t in a n d d r o p p e d by o t h e r animals (detritivores). T h o s e a m o n g t h e p r e d a t o r s survive by catching a n d con­ s u m i n g o t h e r live cave dwellers. A wide­ s p r e a d e x a m p l e a r e the long-legged cave crickets of t h e g e n u s Amphiacusta.

1980). Recent work, yet u n p u b l i s h e d , has found over thirty species of eyeless cave a n d soil a r t h r o p o d s in volcanic caves in the Galápagos Islands (Peck pers. c o m m . ) .

T h e scavengers a r e m o r e n u m e r o u s t h a n p r e d a t o r s in kinds a n d n u m b e r s of individ­ uals. A m o n g these a r e the terrestrial isopods (Trichorhina), millipedes (Eurhinocricus), c o c k r o a c h e s {Periplaneta, Blaberus), tineid m o t h s , d u n g beetles (Ataenius), a n d m a n y o t h e r s . T h e y a r e often e x t r e m e l y n u ­ m e r o u s ; d a r k l i n g beetles ( T e n e b r i o n i d a e ) or mites (Acari) m a y c a r p e t p o r t i o n s of the floor of caves to a d e p t h of a c e n t i m e t e r o r more.

DESSEN, E. M. B., V. R. ESTON, M. S. SILVA, M. T. TEMPERINI-BECK, AND E. TRAJANO.

P r e d a t o r s a r e c o n s p i c u o u s for their large size a n d aggressiveness. Spectacular in these respects a r e t h e giant tailless w h i p scorpions (Amblypygi, Phrynus) that prey o n crickets a n d c o c k r o a c h e s . O t h e r s in the category a r e c e n t i p e d e s , m a n y spiders, ants, a n d various beetles, principally r o v e beetles (Staphylinidae) a n d g r o u n d beetles (Carabidae). M a n y soil mites may occupy these niches as well. Cave insects often exhibit m o r p h o l o g i ­ cal a d a p t a t i o n s to life in the d a r k (Dessen et al. 1980), i n c l u d i n g eyes r e d u c e d o r absent, lack of i n t e g u m e n t a r y p i g m e n t a ­ tion, r e d u c t i o n a n d loss of wings, a n d greatly e l o n g a t e d , highly sensitive a p p e n d ­ ages (especially t h e a n t e n n a e ) . T h e e n t o m o l o g y of N e o t r o p i c a l caves has b e e n the subject of m u c h study (Peck 1977, Strinati 1971), b u t m o r e r e m a i n s to b e d o n e . Best k n o w n a r e the cave f a u n a s of Mexico (Reddell 1971), C u b a ( O r g h i d a n et al. 1 9 7 3 - 1 9 8 3 ) , J a m a i c a (Peck 1975), P u e r t o Rico (Peck 1974, 1981a), B a r b a d o s (Peck 19816), a n d Venezuela ( C h a p m a n

56

ECOLOGY

References CHAPMAN, R 1980. The invertebrate fauna of caves of the Serranía de San Luis, Edo. Falcon, Venezuela. Brit. Cave Res. Assoc. Trans. 7: 179-199. CULVER, D. C. 1982. Cave life, evolution and ecology. Harvard Univ. Press, Cambridge. 1980. Levantamento preliminar de fauna de cavernas de algumas regioes do Brasil. Cien. Cult. 32:414-725. GNASPINI, N. R 1989. Análise comparativa da fauna associada a depósitos de guano de morcegos cavernícolas no Brasil: Primeira Approximafáo. Rev. Brasil. Entomol. 32: 183-192. HOFFMANN,

A.,

J.

G.

PALACIOS-VARGAS, AND

J. B. MORALES-MALACARA. 1986. Manual de

biospeleología. México.

Univ.

Nac. Aut.

México,

ORGHIDAN, T , A. NÚÑEZ JIMÉNEZ, V. DECON, S. NEGREA, AND N. V. BAYES. 1973-1983.

Résultats des expeditions biospéleologique cubano-Roumaines á Cuba. Vols. 1-4. Ed. Academiei, Bucharest, Republicii Socialiste Romania. PECK, S. B. 1974. The invertebrate fauna of tropical American caves. Pt. II: Puerto Rico, an ecological and zoogeographic approach. Biotropica 6: 14—31. PECK, S. B. 1975. The invertebrate fauna of tropical American caves. Pt. I l l : Jamaica, an introduction. Int.J. Speleol. 7: 303-326. PECK, S. B. 1977. Recent studies on the inverte­ brate fauna and ecology of sub-tropical and tropical American caves. 6th Intl. Cong. Speleol. Proc 5: 185-193. PECK, S. B. 1981a. Zoogeography of inverte­ brate cave faunas in southwestern Puerto Rico. Nati. Speleol. Soc Bull. 43: 70-79. PECK, S. B. 19816. Community composition and zoogeography of the invertebrate cave fauna of Barbados. Fla. Entomol. 64: 519-527. REDDELL, J. R. 1971. A review of the cavernicole fauna of Mexico, Guatemala, and Belize. Texas Mem. Mus. Austin Bull. 27: 1-327. STRINATI, P. 1971. Recherches biospéleologiques en Amerique du Sud. Ann. Speleol. 26: 439-450.

Lomas T h e low coastal d e s e r t hills of central Peru provide o n e of t h e most u n u s u a l habitats for insects in the N e o t r o p i c (Dogger a n d Risco 1970). It is speculated that, histori­ cally, the lomas w e r e p a r t of a l a r g e r chaparral b i o m e t h a t o n c e e x t e n d e d along the e n t i r e w e s t e r n slopes of the A n d e s (Péfaur 1978). In this zone of e x t r e m e general aridity, wetness in the form of rain comes only at multiyear intervals, decades or m o r e . At these times, explosions of plant life occur o n t h e otherwise p a r c h e d hills, a n d they b e c o m e g r e e n islands in the bleak desert. Normally, only the r e g u l a r annual fogs (garúas, M a y - O c t o b e r ) b r i n g moisture to t h e s e slopes to m a i n t a i n a less plush but m o r e reliable vegetation. Also a d a p t e d to these climatic a n d vegetative cycles a n d living o n the lomas vegetation is a c o m p l e x a n d diverse com­ munity of insects a n d o t h e r terrestrial a r t h r o p o d s , s o m e species of which a r e known from n o w h e r e else (Aguilar 1964). T h e m o r e i m p o r t a n t g r o u p s a r e spiders (Aguilar, Pacheco, a n d Silva 1987), mites, springtails, wax insects, beetles (especially Tenebrionidae), flies, a n d ants (Aguilar 1981), which a r e most a b u n d a n t a n d n u ­ merous in k i n d s in the d a m p e r u p p e r elevations. S o m e special forms a r e wing­ less sticklike f o r m s ("palitos vivientes d e Lima"), i n c l u d i n g two walkingsticks, a j u m p i n g stick (Proscopiidae), a n d assassin bugs (Reduviidae). O n e walkingstick (Libethra minúscula) is o m n i v o r o u s b u t dies with the failing plants at the e n d of the d a m p season. T h e o t h e r (Bostra scabrinota) feeds on one plant b u t can c h a n g e its color to match seasonal c h a n g e s a n d is p r e s e n t year-round (Aguilar 1970). A n u n e x ­ pected e l e m e n t is the water m e a s u r e r (Bacillometra woytkowskii, H y d r o m e t r i d a e ) , a h e t e r o p t e r o u s insect n o r m a l l y associated with bodies of fresh water (Aguilar, Oyeyama, a n d Aguilar 1987). Its p r e s e n c e is associated with the high h u m i d i t y of the lomas in the winter.

References AGUILAR, P. G. 1964. Especies de artrópodos registrados en las lomas de los alrededores de Lima. Rev. Peruana Entomol. 7: 9 3 - 9 5 . AGUILAR, P. G. 1970. Los "palitos vivientes de Lima." 1: Phasmatidae de las lomas. Rev. Peruana Entomol. 13: 1—8. AGUILAR, P. G. 1981. Fauna desértico-costera Peruana. Vil: Apreciaciones sobre diversidad de invertebrados en la costa central. Rev. Peruana Entomol. 24: 127-132. AGUILAR, P. G., F. OYEYAMA, AND Z. P. AGUILAR.

1987. Los "palitos vivientes de Lima." 111: Un Hydrometridae de las lomas costeras. Rev. Peruana Entomol. 28: 89-92. AGUILAR, P. G., V. R. PACHECO, AND R. SILVA.

1987. Fauna desértico-costera peruana. VIII: Arañas de las lomas Zapalla!, Lima (nota preliminar). Rev. Peruana Entomol. 29: 99— 103. DOGGER, J., AND S. H. Risco. 1970. La fauna insectil de las lomas de Trujillo, Estudio del cerro "Campana." Bol. Tec. Circ. Entomol. Norte (Lambayeque) 1(2): 1-5. PÉFAUR, J. E. 1978. Composition and structure of communities in the lomas of southern Peru. Ph.D. diss., Univ. Kansas, Lawrence. The Ocean At first t h o u g h t , it seems i n c o n g r u o u s that the insects, so t r e m e n d o u s l y successful in d o m i n a t i n g the land a n d the inland waters of the world, have failed to c o n q u e r the seas. I n t o l e r a n c e of hypersaline water is certainly not to blame, because m a n y aquatic insects live in highly salinated lakes a n d p o n d s inland a n d at t h e ocean's m a r ­ gin. T h e reasons a p p a r e n t l y lie in t h e fact that they arrived o n t h e scene long after their o t h e r i n v e r t e b r a t e p r e d e c e s s o r s h a d locked u p all the ecological niches. T h e r e a r e a few examples, however, of m a r i n e a d a p t e d g r o u p s ( C h e n g 1976). Insects truly a d a p t e d to life far o u t to sea a r e the fewest a n d consist only of the m a r i n e water striders or "ocean skaters" (several g e n e r a of G e r r i d a e , see w a t e r striders, c h a p . 7). Closer to s h o r e , t h e n u m b e r a n d kinds of m a r i n e insects greatly increases. T h e r e o n e finds the m a r i n e midges ( C h i r o n o m i d a e ) . T h e s e live o n rocky shores a n d have larvae that a r e

ECOSYSTEMS

57

completely at h o m e in t h e salt spray a n d a r e even s u b m e r g e d by seawater d u r i n g high tide. O t h e r intertidal insects a n d a r a c h n i d s a r e f o u n d a m o n g t h e springtails a n d mites (especially H a l a c a r i d a e ) . Several kinds of lice a r e parasitic o n seagoing m a m m a l s , seals a n d sea lions particularly. W h e n t h e host s u b m e r g e s , they escape osmotic dessication by seawater a n d d r o w n i n g by h i d i n g close to t h e skin in air pockets u n d e r t h e host's fur. T h e seashores of t h e c o n t i n e n t s a n d islands of Latin A m e r i c a also s u p p o r t an even m o r e extensive g r o u p of m a r i n e litto­ ral insects (Evans 1968). M a n y of these a r e freshwater g r o u p s , i n c l u d i n g inland h y p e r saline lake types, that h a v e shifted to t h e similar habitats at t h e sea m a r g i n s , espe­ cially salt m a r s h e s a n d m a n g r o v e s . A major­ ity prey o n i n v e r t e b r a t e s , feeding o n o t h e r s h o r e life, such as tiger beetles, rove bee­ tles, p s e u d o s c o r p i o n s , spiders, a n d s h o r e bugs (Saldidae). M a n y of these a r e associated with sea­ weed a n d kelp that a c c u m u l a t e s o n beaches, feeding either directly o n this material (kelp fly larvae, Fucellia; see c h a p . 1 1 ; Coelopidae, s h o r e flies [ E p h y d r i d a e ] ) o r catching a n d d e v o u r i n g t h e wrack scavengers (spi­ d e r s , g r o u n d beetles, r o v e beetles). T h e adults of some, such as salt m a r s h mosqui­ toes a n d tabanids a n d m u d f l a t - b r e e d i n g p u n k i e s , a r e feeders on v e r t e b r a t e blood. T h e i r p o p u l a t i o n explosions can m a k e h u ­ m a n life impossible in seaside areas. Several water b o a t m e n species (especially Trichocorixa reticulata) live in highly saline s h o r e pools (Davis 1966) a n d have even b e e n collected in p l a n k t o n tows in t h e Gulf of California.

References CHENG, L., ed. 1976. Marine insects. NorthHolland, Amsterdam. DAVIS, C. C. 1966. Notes on the ecology and reproduction of Trichocorixa reticulata in a Jamaican salt-water pool. Ecology 47: 850— 852. EVANS, W. G. 1968. Some intertidal insects from

58

ECOLOGY

western Mexico. Pan-Pacific Entomol. 44: 236-241. Torrential Waters S t r e a m waters flowing in t h e r a n g e of 60 to 200 centimeters p e r second o r g r e a t e r a r e considered torrential a n d a r e c o m m o n in the u p l a n d d r a i n a g e s of m o u n t a i n o u s r e ­ gions. O r i g i n a t i n g in countless springs a n d snowfields o n t h e heights of t h e Cordillera, b o u n d i n g over b o u l d e r s a n d c r a s h i n g into foam-covered pools, torrential streams d e ­ scend t h r o u g h gorges a n d n a r r o w can­ yons, over h a r d , rocky beds, before reach­ ing t h e lower, gentle slopes. Fast, cold water offers a refugial niche to m a n y insects, comparatively free from ver­ tebrate p r e d a t i o n because few such large animals can live in such a n inhospitable m e d i u m . Exceptions a r e a few h a r d y insecti­ vorous types, such as t o r r e n t ducks, dip­ pers, a n d a few fish ( i n t r o d u c e d t r o u t a n d possibly some astroblephid catfishes). As a result, some very ancient representatives of several aquatic g r o u p s have persisted for geologic ages as torrenticolous ( r h e o p h i lous) relicts. T h e s e include e n t i r e families, most p r o m i n e n t l y t h e net-winged m i d g e s (Blephariceridae) a n d various beetles (Elmidae, D r y o p i d a e , P s e p h e n i d a e ) . O t h e r simi­ larly a d a p t e d taxa a r e t h e larval stages of lance-winged m o t h flies (Maruina, Psychodidae) a n d blackflies (Simuliidae) a n d m a n y species of T r i c h o p t e r a (especially in the family Hydrobiosidae), H e m i p t e r a [Cryphocricos, N a u c o r i d a e ) , Plecoptera (Araucanioperla, G r i p o p t e r y g i d a e ) , water m i d g e s ( C h i r o n o m i d a e ) , a n d E p h e m e r o p t e r a (Epeorus, H e p t a g e n i i d a e ) . E x t r e m e structural a n d physiological ad­ aptations have evolved in these forms in r e s p o n s e to t h e s t r o n g selective p r e s s u r e s of fast c u r r e n t a n d s m o o t h substrates. T h e s e include holdfast structures (ventral suck­ ers, claws), streamlining, a n d p l a s t r o n respi­ ration. T h e latter makes use of t h e function of air films a n d pockets as "physical gills." A r e q u i r e m e n t for p r o p e r function of this

system is cold, clean, oxygen-rich water. T h e p r e s e n c e of these insects t h e r e f o r e indicates healthy s t r e a m conditions. Food habits for t h e relatively passive grazers a n d p r e d a t o r s d o n o t r e q u i r e r a p i d movement. A d u l t e m e r g e n c e is often "ex­ plosive." To avoid d r o w n i n g , t h e imago rises to t h e surface in a b u b b l e a n d takes wing immediately after contacting t h e air. Torrenticolous insects a r e n o t well known in Latin America. S o m e famous early studies w e r e m a d e o n D i p t e r a in southeastern Brazil by Fritz Müller (1879, 1895) a n d A d o l f o Lutz (1930). New species are c o m m o n l y discovered in t h e habitat, especially in r e m o t e m o u n t a i n s .

References LUTZ, A. 1930. Biología das aguas torrenciais e encachoeiradas. Soc. Biol. Montevideo, Arch. suppl. 1: 114-120. MÜLLER, F. 1879. A metamorphose de uni

insecto díptero. Mus. Nac. Rio de Janeiro, Arch. 5/6: 4 7 - 8 5 , pis. 4 - 7 . MÜLLER, F. 1895. Contributions towards the history of a new form of larvae of Psychodidae (Diptera) from Brazil. Entomol. Soc. London, Trans. 1895: 479-482, pis. 10-11. Tank Plants Some Neotropical plants have parts of their a n a t o m y d e v e l o p e d into c u p - s h a p e d water-holding s t r u c t u r e s (phytotelmata) (Frank a n d L o u n i b o s 1983) a n d a r e r e ­ ferred to as r e s e r v o i r o r t a n k plants. T h e y are of several types. T h e best k n o w n a r e the bromeliads (Bromeliaceae) (Gómez 1977) whose water a c c u m u l a t i o n s p r o v i d e a h o m e for small aquatic animals. T h i s microcosm was first studied c o m p r e h e n ­ sively as a n ecosystem by Picado (1913), who distinguished b e t w e e n t h e aquatic mi­ lieu ("aquarium") a n d terrestrial p o r t i o n ("terrarium"). T h e a q u a r i u m consists of a spacious central c u p a n d e x p a n d e d lateral leaf bases that collect rainwater. L a r g e plants m a y store a liter o r m o r e a n d m a y have a c u p 5 to 10 centimeters, in d i a m e t e r a n d equally

d e e p . T h e s e reservoirs of water p r o v i d e habitats suitable for t h e d e v e l o p m e n t of m a n y aquatic insects, including mosquitoes, especially Wyeomyia a n d o t h e r sabethines a n d Aedes. A n a b u n d a n c e of b r o m e l i a d s in a particular area can foster a substantial mos­ quito p o p u l a t i o n , a good e x a m p l e b e i n g Anopheles darlingi, t h e m a i n malaria vector in Trinidad in t h e 1940s (Downs a n d Pittendrigh 1946). O t h e r r e p r e s e n t a t i v e aquatic inhabitants include water m i d g e s (Chironornus), p u n k i e s (Bezzia), damselflies (Pseudostigmatidae), beetles (Helodidae), a n d water mites. T h e detailed ecology of this fraction has b e e n studied by m a n y entomologists, including Laessle (1961). (See t h e bibliography of b r o m e l i a d a n d pitcher plant reservoir plant e n t o m o l o g y by Fish a n d Beaver [1978].) Detritus collects also in t h e lateral bracts, a n d a special kind of a r b o r e a l soil is cre­ ated which is like a " t e r r a r i u m " to a n o t h e r g r o u p of insects. H e r e a r e f o u n d spiders, carabid beetles, ants, isopods, millipedes, mites, springtails, a n d o t h e r terrestrial forms (Murillo et al. 1983). A few insects actually feed o n t h e leaves of b r o m e l i a d s a n d form yet a n o t h e r guild in association with these plants (Beutelspacher 1972). A n o t h e r category of Neotropical t a n k plants a r e t h e insectivorous " p i t c h e r plants" of t h e family Sarraceniaceae. I n these, d e e p , u r n - s h a p e d leaves have evolved to hold fluids that normally kill a n d digest hapless insects that fall into t h e m . However, i m m a t u r e s of some insect species, mostly mosquitoes of t h e tribe Sabethini, a r e im­ m u n e to t h e corrosive action of these chemi­ cals a n d actually develop normally in this very peculiar aquatic microhabit. I n t h e Neotropics, pitcher plants of only t h e g e n u s Heliamphora a r e known, f o u n d in t h e V e n e z u e l a n - G u i a n a n region. T h e y a r e k n o w n to be occupied by Wyeomyia mosqui­ toes of t h e s u b g e n u s Zinzala (Zavortink 1985), b u t their o t h e r o c c u p a n t s a r e n o t studied. Flower bracts, especially those of t h e

ECOSYSTEMS

59

g e n u s Heliconia (Musaceae), h o l d watery fluids, secreted by t h e p l a n t itself, as well as r a i n w a t e r a n d host a specific microc o m m u n i t y of insects similar to that of b r o m e l i a d s (Seifert 1982). T h e inflores­ cences h a r b o r o n e of two k i n d s of insect g r o u p i n g s d e p e n d i n g o n size a n d s h a p e , a m o u n t of water, a n d a g e . T h e first is c o m p o s e d of fly larvae ( n o n m o s q u i t o ) a n d beetles in older, less v o l u m i n o u s bracts with small a m o u n t s of water. T h e most c o m m o n fly larvae a r e of p o m a c e flies (Drosophilidae), h o v e r flies ( S y r p h i d a e , Copestylum), a n d soldier flies (Stratiomyiidae, Merosargus). T h e beetles a r e primarily scavenging water beetles ( H y d r o p h i l i d a e , Gillisius), leaf beetles ( C h r y s o m e l i d a e , Hisp i n a e , Cephaloleia), a n d rove beetles (Staphylinidae, Odontolinus a n d Quichuana). T h e r e a r e also earwigs ( D e r m a p t e r a , Carcinophora) a n d c o c k r o a c h e s (Blattidae, Litopeltis). I n t h e second g r o u p , m o s q u i t o larvae a r e d o m i n a n t (Seifert 1980), in younger, l a r g e r inflorescences with l a r g e r a m o u n t s of water. At least five g e n e r a a r e r e p r e ­ sented: Wyeomyia, Triehoprosopon, Toxorhynchites, Culex, a n d Sabethes. Various interactions a m o n g t h e insects in these c o m m u n i t i e s h a v e b e e n observed, including p r e d a t i o n , c o m m e n s a l i s m , colo­ nization, a n d c o m p e t i t i o n , a l t h o u g h t h e last seems t o b e weak. T h e clearly defined nature of the communities, the common­ ness of Heliconia plants, a n d t h e ease with which t h e insects m a y b e m a n i p u l a t e d e x p e r i m e n t a l l y m a k e t h e m ideal for investi­ gations of species interactions a n d o t h e r ecological p h e n o m e n a (Seifert 1975, 1 9 8 1 ; Seifert a n d Seifert 1976a, 19766, 1979).

phy of the aquatic fauna inhabiting bromeli­ ads (Bromeliaceae) and pitcher plants (Nepenthaceae and Sarraceniaceae). Florida Antimosq. Assoc, Proc. 49: 11-19. FRANK, J. H., AND L. P. LOUNIBOS, eds. 1983.

Phytotelmata, terrestrial plants as hosts for aquatic insect communities. Plexus, Medford,

N.J. GÓMEZ, L. D. 1977. La biota bromelícola ex­ cepto anfibios y reptiles. Hist. Nat. Costa Rica 1:45-62. LAESSLE, A. M. 1961. A micro-limnological

study of Jamaican bromeliads. Ecology 42: 499-517. MURILLO, R. M.,J. G. PALACIOS, J. M. LABOUGLE, E. M. HENTSCHEL, J. E. LLÓRENTE, K. LUNA, P. ROJAS, AND S. ZAMUDIO. 1983. Variación

estacional de la entomofauna asociada a Tillandsia spp. en una zona de transición biótica. Southwest. Entomol. 8: 292-302. PICADO, C. 1913. Les broméliacées epiphytes considérée comme milieu biologique. Bull. Scient. France Belgique 47: 215-360. SEIFERT, R. P. 1975. Clumps of Heliconia inflorescences as ecological islands. Ecology 56: 1416-1422. SEIFERT, R. P. 1980. Mosquito fauna of Heliconia áurea. J. Anim. Ecol. 49: 687-697. SEIFERT, R. P. 1981. Principal components analy­ sis of biogeographic patterns among Heliconia insect communities. New York Entomol. Soc, J. 89: 109-122. SEIFERT, R. P. 1982. Neotropical Heliconia insect communities. Quart. Rev. Biol. 57: 1-28. SEIFERT, R. P., AND F. H. SEIFERT. 1976a. Natu­

ral History of insects living in inflorescences of two species of Heliconia. New York Entomol. Soc.,J. 84: 233-242. SEIFERT, R. P., AND F. H. SEIFERT. 19766. A

community matrix analysis oí Heliconia insect communities. Amer. Nat. 110: 461-483. SEIFERT, R. P., AND F. H. SEIFERT.

1979. A

Heliconia insect community in a Venezuelan cloud forest. Ecology 60: 462-467. ZAVORTINK, T. J. 1985. Zinzala, a new subgenus of Wyeomyia with two new species from pitcher-plants in Venezuela (Diptera, Culicidae, Sabethini). Wasmann J. Biol. 43: 46-59.

References BEUTELSPACHER, C. R. 1972. Some observations on the Lepidoptera of bromeliads. J. Lepidop. Soc. 26: 133-137. DOWNS, W. G., AND C. S. PITTENDRIGH. 1946.

Bromeliad malaria in Trinidad, British West Indies. Amer. J. Trop. Med. Hyg. 26: 46-66. FISH, D., AND R. A. BEAVER. 1978. A bibliogra­

60

ECOLOGY

Miscellaneous Special Habitats A few additional i m p o r t a n t special habitats that have b e e n i g n o r e d o r received mini­ mal attention with r e g a r d to their e n t o m o faunal aspects in Latin A m e r i c a a r e t h e following.

1. Hot, mineral springs. D e Oliveira (1954) noted e p h y d r i d flies in t h e h o t effluent of a geyser at El Tatio in Chile. S o m e general collections from T e r m a s d e Chillan in t h e same c o u n t r y have b e e n identified by Ruiz a n d S t u a r d o (1935). 2. Inland salt lakes and playas, such as t h e i m m e n s e salares of t h e Bolivian alti­ plano (Salar d e Coipasa, Lago d e Poopó, Salar d e Uyuni). A few insects of hypersaline waters have b e e n d e ­ scribed, notably mosquitoes (Bachm a n n a n d Casal 1963) a n d C o r i x i d a e of the g e n u s Trichocorixa from A r g e n t i n a ( B a c h m a n n 1979: 326f.) a n d t h e Mexi­ can lakes, Texcoco a n d A h u a t l e (see water b o a t m e n , c h a p . 8). E p h y d r i d shore flies of t h e g e n u s Dimecoenia (eco­ logically a n a l o g o u s to t h e Mexican a n d N o r t h A m e r i c a n Ephydra) a r e a b u n d a n t in these waters as well ( d e Oliveira 1958). 3. Streams. Entomological studies of lode waters a r e n u m e r o u s b u t n o t c o m p r e ­ hensive a n d deal mostly with h i g h l a n d , h a r d - b o t t o m s t r e a m s (lilies 1964; C a m ­ pos et al. 1984; Patrick a n d collabs. 1966; T u r c o t t e a n d H a r p e r 1982a, 19826). A d a p t i v e strategies were stud­ ied o n s t r e a m invertebrates, mostly in­ sects in t h e A r g e n t i n e Rio N e g r o , by Wais(1985). 4. Alpine ponds and lakes. Available a r e only t h e r e p o r t s of Robeck et al. (1980) for miscellaneous A n d e a n habitats a n d Gilson (1939) for Lake Titicaca. 5. Bogs. N o p u b l i s h e d observations. 6. Deserts, b o t h local a n d large, as t h e great S o n o r a n o r A t a c a m a . N o a p p a r ­ ent c o m p r e h e n s i v e observations. 7. Sand dunes, b o t h coastal a n d inland desert. N o a p p a r e n t observations. 8. Inland freshwater observations.

m o u n t a i n o u s f a u n a s (Palacios-Vargas 1985); dispersal a n d recolonization of new t e r r a i n will b e d e m o n s t r a t e d by insects a n d terrestrial arthropods found o n new volcanoes, b o t h at sea a n d o n t h e c o n t i n e n t s . I n b o t h cases, the principles of island b i o g e o g r a p h y a r e applicable. 10. Guianan tepuis. T h e flat-topped j u n g l e mesas of t h e G u i a n a Shield a r e ac­ knowledged islands s u p p o r t i n g m a n y e n d e m i c biotic e l e m e n t s . T h e y have been studied mostly by botanists, b u t some u n i q u e insects have b e e n col­ lected a t o p M o u n t R o r a i m a a n d C e r r o Neblina (Spangler 1 9 8 1 , 1985; Waterh o u s e 1900). 11. Lakes and rivers. Limnological studies, particularly of bodies of fresh water larger t h a n streams, a r e n u m e r o u s , b u t in these, m o r e attention is usually paid to t h e m a c r o f a u n a t h a n to insects. Some works have c o n c e n t r a t e d o n t h e latter (Wais 1984). 12. Land crab burrows. T h e b u r r o w s of tropical coastal land crabs s u p p o r t a complex ecosystem, including m a n y insects a n d related a r t h r o p o d s (Bright a n d H o g u e 1972), most of which have aquatic life stages in t h e water that accumulates within t h e m from rainfall a n d g r o u n d w a t e r seepage. 13. Lava tubes. N o a p p a r e n t observations. Also r e m a i n i n g to b e elucidated a r e t h e entomological characteristics of t h e g r e a t n u m b e r of vegetation types t h r o u g h o u t the region, for e x a m p l e , c e r r a d o , p a l m forests, caatinga, cloud forest, m a n g r o v e s , grassland, p u n a , p á r a m o (Bernal 1985), alpine, a n d savanna. T h e foregoing a r e all fruitful fields of investigation for f u t u r e s t u d e n t s of ecologi­ cal a n d b i o g e o g r a p h i c entomology.

swamps. N o a p p a r e n t

9. Volcanoes. O l d volcanoes may provide in­ formation o n t h e effects of isolation o n

References BACHMANN, A. O. 1979. Notas para una monografía de las Corixidae Argentinas (In-

ECOSYSTEMS

61

secta, Heteroptera). Acta Zool. Lilloana 35: 305-350. Mosquitos Argentinos que crian en aguas salobres y saladas. Soc. Entomol. Argentina, Rev. 25: 21-27. BERNAL, A. 1985. Estudio comparativo de la artropofauna de un bosque alto-andino y de un pajonal paramuno en la región de Monserrate (Cund.). In H. Sturm and O. Rangel, Ecología de los paramos andinos: Una visión preliminar integrada. Univ. Nac. Colombia (lnst. Cien. Nat., Mus. Nat. Hist., Bibl. José Jerónimo Triana no. 9). Bogotá. Pp. 225-260.

plano expedition. Pt. 1, Aquatic insects except Diptera. Acad. Nat. Sci. Philadelphia Proc 132: 176-217. Ruiz, E, AND C. STUARDO O. 1935. Insectos colectados en las Termas de Chillan. Rev. Chilena Hist. Nat. 39: 313-322. SPANGLER, P. J. 1981. New and interesting water beetles from Mt. Roraima and Ptari-tepui, Venezuela (Coleóptera: Dytiscidae and Hydrophilidae). Aquat. Ins. 3: 1-11. SPANGLER, P. J. 1985. A new genus and species of riffle beetle, Neblinagena prima, from the Venezuelan tepui, Cerro de la Neblina (Coleóptera, Elmidae, Larinae). Entomol. Soc. Wash., Proc. 87: 538-544.

BRIGHT, D. B., AND C. L. HOGUE.

TURCOTTE, P., AND P. P. HARPER. 1982a. T h e

BACHMANN, A. O., AND O. H. CASAL. 1963.

1972. A

synopsis of the burrowing land crabs of the world and list of their arthropod symbionts and burrow associates. Los Angeles Co. Mus. Nat. Hist., Contrib. Sci. 220: 1-58. CAMPOS, H., J. ARENAS, C. JARA, T. GONSER, AND

R. PRINS. 1984. Macrozoobentos y fauna lótica de las aguas limnéticas de Chiloé y Aysén continentales (Chile). Medio Ambiente 7: 52-64. DE OLIVEIRA, S. J. 1954. Contribuicáo para o conhecimento do género "Dimecoeniá" Cresson, 1916. 1: "Dimecoenia lenti" sp. n. encon­ trado no Chile (Díptera, Ephydridae). Rev. Brasil. Biol. 14: 187-194. DE OLIVEIRA, S. J. 1958. Contribuicáo para o conhecimento do género "Dimenia" Cresson, 1916. IV: Descricáo da larva e do pupário de "Dimecoenia grumanni" Oliveira, 1954 (Diptera, Ephydridae). Rev. Brasil. Biol. 18: 1 6 7 169. GILSON, H. C. 1939. T h e Percy Sladen Trust Expedition to Lake Titicaca in 1937. I: De­ scription of the expedition. Linnean Soc. London, Trans. 1: 1—20. ILLIES, J. 1964. T h e invertebrate fauna of the Huallaga, a Peruvian River, from the sources down to Tingo María. Intl. Verein Theor. Angew. Limnol., Verh. 15: 1077-1083. PALACIOS-VARGAS, J. 1985. Microartrópodos del

Popocatepetl (aspectos ecológicos y biogeográficos de los ácaros oribátidos e insectos colémbolos). Ph.D. diss., Univ. Nac. Aut. Méx­ ico, Mexico City. PATRICK, R., AND COLLABORATORS. 1966. T h e

Catherwood Foundation Peruvian-Amazon Expedition: Limnological and systematic stud­ ies. Acad. Nat. Sci. Philadelphia, Monogr. 14: 1-495. ROBECK, S. S., L. BERNER, O. S. FLINT, J R . , N. NIESER, AND P. J. SPANGLER. 1980. Results

of the Catherwood Bolivian-Peruvian alti­

62

ECOLOGY

macroinvertebrate fauna of a small Andean stream. Freshwater Biol. 12: 411-419. TURCOTTE, P., AND P. P. HARPER. 19826. Drift

patterns in a high andean stream. Hydrobiologia89: 141-151. WAIS, 1. R. 1984. Two Patagonian basins— Negro (Argentina) and Valdivia (Chile)—as habitats for Plecoptera. Ann. Limnol. 20: 115-122. WAIS, I. R. 1985. Strategies adaptatives aux eaux courantes des invertébrés du bassin du fleuve Negro, Patagonia, Argentina. Int. Verein. Limnol., Verh. 22: 2167-2172. WATERHOUSE, C. O. 1900. Report on a collec­ tion made by Messrs. E V McConnell and J. J. Quelch at Mt. Roraima in British Guyana. Linnean Soc. London, Trans. 8: 74—76.

HISTORICAL BIOGEOGRAPHY T h e historical perspective in zoogeog­ r a p h y provides insights into t h e origins of species a n d faunas a n d their c h a n g e s t h r o u g h time relative to geologic events (Fittkau et al. 1969). Insects m a k e excellent material for such studies because of t h e very long history of m a n y taxa a n d t h e variety of types available t o test hypotheses of all kinds (Gressitt 1974, M u n r o e 1965). A g r o u p m a y b e f o u n d which will fulfill almost any given set of conditions for any geologic p e r i o d . For e x a m p l e , u p l a n d , sed­ e n t a r y types with limited vagility (such as narrowly a d a p t e d , torrenticolous Diptera) are particularly useful for t r a c i n g past

m o u n t a i n o u s land c o n n e c t i o n s as far back as t h e late Paleozoic. T h e existence of two, contrasting means of p o p u l a t i o n d i s r u p t i o n , that is, dispersal to isolated areas versus fragmention in place, is t h e basis for t h e "dispersalist" a n d "vicariance" schools of biogeographic t h o u g h t (Ferris 1980) to explain t h e speciation process in evolving organisms. Actually, b o t h m e c h a n i s m s may cause t h e b r a n c h i n g of phyletic lines and a r e p a r t of a m o d e r n , unified theory of b i o g e o g r a p h y (Pielou 1979, B r o w n a n d Gibson 1983). Because insects have b e e n o n e a r t h for a very long time, at least since t h e m i d d l e of the Paleozoic e r a , c o n t i n e n t a l c o n n e c t i o n s and disjunctions (tectonics) (Dietz a n d Holden 1970, Marvin 1973, Smith et al. 1981) a r e major vicariant events that have affected their evolution a n d dispersal (Carbonell 1977: 1 5 5 - 1 6 1 ) . I n Latin America, the tectonic d e v e l o p m e n t of t h e C a r i b b e a n and isthmian r e g i o n s seems to be m u c h more complicated (Bonini et al. 1984, D u r ­ ham 1985, Rosen 1985) t h a n that of t h e South A m e r i c a n p o r t i o n s (Jenks 1956, Harrington 1962) with c o n s e q u e n t p r o b ­ lems in e x p l a i n i n g t h e origins of o r g a n i s m s there ( L i e b h e r r 1988, W o o d s 1989). T h e origin of m a n y g r o u p s o n G o n d wanaland, t h e g r e a t s o u t h e r n c o n t i n e n t that was c o m p o s e d of w h a t is n o w S o u t h America, Africa, Antarctica, Australia, a n d India, is still evident in t h e restricted occur­ rences of their d e s c e n d a n t s in those areas and in t h e s o u t h e r n m o s t p o r t i o n s of S o u t h America today (e.g., water m i d g e s [ B r u n din 1966, 1967]; see Keast 1973 for o t h e r insect examples of these so-called amphinotic o r austral disjunctive distribu­ tions). Close affinities of s o m e eastern Brazilian insects with West African species, such as a m o n g Schistocerca g r a s s h o p p e r s (Carbonell 1977:169), t h e amblypygid ge­ nus Phrynus (= Tarantula) (Quintero 1983), t h e psocid g e n e r a Belaphapsocus a n d Notiopsocus (New 1987), a n d t h e t e r m i t e

g e n u s Mimeutermes ( E m e r s o n 1955), a r e c u r r e n t evidence of past u n i o n of t h e two continents at midlatitudes. T h e formation of t h e A m a z o n Basin is a direct result of events b r o u g h t into play by the break in t h e South America—Africa connection a b o u t 90 million years a g o . A c c o r d i n g to o n e theory (Putzer 1984), t h e A m a z o n River system was p r o b a b l y con­ t i n u o u s d u r i n g t h e early Mesozoic with t h e Niger River, a n d t h e main flow was west­ ward to t h e Pacific O c e a n . B u t with t h e relatively recent (Miocene) uplift o f t h e A n d e s , t h e flow was d a m m e d a n d a n e n o r m o u s lake f o r m e d at their foot. As t h e plates s e p a r a t e d , t h e western p o r t i o n of t h e river r e v e r s e d its flow a n d c a m e t o e m p t y into t h e Atlantic. A n extensive coastal plain was also c r e a t e d along t h e west side of t h e Cordillera from ejection of volcanic material a n d pluvial o u t w a s h . T h e s e vast p h y s i o g r a p h i c c h a n g e s have created a m i x e d heritage for t h e basin's present-day insect p o p u l a t i o n s . S o m e a r e derived from t h e O l d World, o t h e r s a r e derived from h i g h l a n d s t o t h e s o u t h a n d n o r t h , a n d still o t h e r s have evolved in situ after long p e r i o d s of isolation by river a n d climatic barriers. T h i s variety of causes is o n e reason for t h e basin's incredible spe­ cies richness. O n a m u c h m o r e limited scale, local n a t u r a l disasters, such as volcanic e r u p t i o n s a n d h u r r i c a n e s , take a toll o n insect life a n d may actually cause t h e extinction of small populations o r even very regional species a n d constitute vicariant events, a l t h o u g h such consequences have yet to b e docu­ m e n t e d , especially those of low-density for­ est species (Elton 1975). C h a n g e s in t h e c o u r s e of rivers, a n especially c o m m o n o c c u r r e n c e in m e a n d e r i n g lowland d r a i n ­ ages, such as t h e A m a z o n , c a n b r e a k u p c o n t i n u o u s p o p u l a t i o n s a n d halt g e n e flow sufficiently t o create new entities. Dispersalist e x a m p l e s o f t h e effects of geology o n t h e history of insect life c o m e from t h e oceanic islands scattered o n t h e

HISTORICAL BIOGEOGRAPHY

63

as physical and biotic factors. All tropical deserts, for example, will not have the same species or genera of darkling beetles or all high mountains the same butterfly groups. Past dispersals and evolution of organisms will result in different assem­ blages of taxa in geographic areas regard­ less of similarities they may have in their environment (Mooney 1977). These assem­ blages, particularly the existence of en­ demics, give taxonomic character to any region from which it may be recognized as a distinct biogeographic unit with biotic uniformity. The Neotropical Region itself is one of the earth's broadest and most distinctive of such categories. T h e reason for this seems to be due primarily to the fact that South America and Africa, before their disjunc­ tion, constituted the largest block of land (western Gondwanaland) in the tropics and was therefore a separate evolutionary center (Raven and Axelrod 1975). Subse­ quent long isolation permitted undis­ turbed and independent development of an already rich biota (Fittkau 1969). The composition and spatial arrange­ ment of the world's biogeographic regions have changed through time. Past evolution­ ary patterns have been postulated for some other organisms (Hallam 1973) but are not known for insects in Latin America, mainly because of their poor fossil record there. The origins of these regions and contempo­ rary insect geography are determinable, nevertheless, by combining knowledge of a group's morphology, mobility, geographic and ecological dispersal opportunities in the past, and historical disruptions in its range. There are relatively few such stud­ ies on Neotropical insects (Halffter 1975, 1976, 1987), but it is common for taxonomists to treat the zoogeography of their groups routinely in systematic papers (e.g., Nielsen and Robinson 1983). In the Neotropics, different systems of classification of today's biogeographic subregions have been applied by authors choos­

66

ECOLOGY

ing varied organisms as indicators and more or less inclusive amounts of land area. For all or most of Latin America, few are based on the entire flora (Gentry 1982, Cabrera and Willink 1973) or fauna (Gilmore 1950, Goldman and Moore 1946, Fittkau 1969), fewer on both (Darlington 1965, Udvardy 1975). Some are derived from an analysis of diverse entomofaunal elements (Halffter 1964, 1974, 1975, 1987; O'Brien 1971) or specific taxa (Porter 1980, Ichneumonidae; Kuschel 1969, beetles; Halffter 1975, dung beetles). Most, how­ ever, are limited to much narrower taxo­ nomic groups and restrictive areas (Peña 1966, Chilean darkling beetles; Scott 1972, Antillean butterflies; de Armas 1982, Antillean scorpions; Lane 1943, sabethine mos­ quitoes). Some of the latter have produced complex results, such as Lamas's (1982) recognition of forty-eight biotic regions for Peru, based on the country's butterfly fauna alone. The aquatic medium has been treated to a limited degree. Stream insects fall into two main groups, the cool-adapted (oligostenothermic) types found in certain mountain­ ous areas (Andes, especially southern Andes, Guianas, southeastern Brazil), and warm-adapted (polystenothermic) types spread almost everywhere else in the low­ lands (lilies 1965, 1969). For entomological purposes, in Latin America, the simple classification of biogeo­ graphic regions shown in figure 2.2 is useful (although a much more detailed system is proposed by Udvardy [1975]). References CABRERA,

A.

L.,

AND A.

WILLINK.

1973.

Biogeografía de América Latina. Org. Est. Americanos, Ser. Biol., Monogr. 13: 1-122. DARLINGTON, JR., P. J. 1965. Biogeography of

the southern end of the world. McGraw-Hill, New York. DE ARMAS, L. F. 1982. Algunos aspectos zoogeográficos de la escorpionfauna antillana. Poeyana238: 1-17. FITTKAU, E. J. 1969. The fauna of South Amer-

Figure 2.2. MAJOR NEOTROPICAL BIOGEOGRAPHIC REGIONS (modified and expanded from Fittkau 1969). MIDDLE AMERICA: 1. Mexican Highlands (part of Nearctic Region); 2. Mexican Low­ lands and Central America; 3. West Indies (Antilles); SOUTH AMERICA Guiana-Brazil; 4. Caquetio (Orinoco and south); 5. Hylaea (Amazonia); 6. Bororó (Brazilian plateau); 7. Cariri (northeastern Brazil); 8. Tupi; 9. Guaraní (southern Brazil); 10. Incasia (northern Andes) Andes-Patagonia; 11. Pampas; 12. Patagonia (includes Juan Fernández, Falklands, etc.); 13. Subandes (eastern Andean foot ranges); 14. Chile; 15. Andes (central and southern).

ica. In E. J. Fittkau, J. lilies, H. Klinge, G. H. Schwabe, and H. Sioli, eds., Biogeography and ecology in South America. 2:624—658. Junk, T h e Hague. GENTRY, A. H. 1982. Neotropical floristic diver­ sity: Phytogeographical connections between Central and South America, Pleistocene clima­ tic fluctuations, or an accident of the Andean orogeny? Missouri Bot. Gardens Ann. 69: 557-593. GILMORE, R. M. 1950. Fauna and ethnozoology of South America. Handbk. So. Amer. Indi­ ans 6: 345-464. GOLDMAN, E. A., AND R. T. MOORE. 1946.

The

biotic provinces of Mexico. J. Mammal. 26: 347-360. HALFFTER, G. 1964. La entomofauna Ameri­ cana, ideas acerca de su origen y distribución. Fol. Entomol. Mex. 6: 1-108. HALFFTER, G. 1974. Elements anciens de l'entomofaune néotropicale: Ses implications biogéographiques. Quaest. Entomol. 10: 2 2 3 262. HALFFTER, G. 1975. Elements anciens de l'entomofaune néotropicale: Ses implications biogéographiques. Mus. Nat. Hist. Natur. (Paris) Mem. Ser. A. (Zool.) 2: 114-145. HALFFTER, G. 1976. Distribución de los insectos en la zona de transición Mexicana: Relaciones con la entomofauna de norteamérica. Fol. Entomol. Mex. 35: 1-64. HALFFTER, G. 1987. Biogeography of the mon­ tane entomofauna of Mexico and Central America. Ann. Rev. Entomol. 32: 95-114. HALLAM, A. 1973. Atlas of palaeobiogeography. Elsevier, Amsterdam. ILLIES, J. 1965. Phylogeny and zoogeography of the Plecoptera. Ann. Rev. Entomol. 10: 1 1 7 140. ILLIES, J. 1969. Biogeography and ecology of Neotropical freshwater insects, especially those from running waters. In E. J. Fittkau, J. Illies, H. Klinge, G. H. Schwabe, and H. Sioli, eds., Biogeography and ecology in South America. 2: 685-708. Junk, T h e Hague. KUSCHEL, G. 1969. Biogeography and ecology of South American Coleóptera. In E. J. Fittkau, J. lilies, H. Klinge, G. H. Schwabe, and H. Sioli, eds., Biogeography and ecology in South America. Vol. 2. Junk, T h e Hague. LAMAS, G. 1982. A preliminary zoogeographical division of Peru, based on butterfly distribu­ tions (Lepidoptera, Papilionoidea). In G. T Prance, ed., Biological diversification in the tropics. Columbia Univ. Press, New York. Pp. 336-357.

68

ECOLOGY

LANE, J. 1943. The geographic distribution of Sabethini (Dipt., Culicidae). Rev. Entomol. Sao Paulo 14: 409-429. MOONEY, H. A., ed. 1977. Convergent evolution in Chile and California, Mediterranean cli­ mate ecosystems. Dowden, Hutchinson and Ross, Stroudsburg, Penn. NIELSEN, E. S., AND G.

S. ROBINSON.

1983.

Ghost moths of southern South America (Lepidoptera: Hepialidae). Entomonograph 4: 1-192. O'BRIEN, C. W. 1971. T h e biogeography of

Chile through entomofaunal regions. Ento­ mol. News 82: 197-207. PEÑA, L. E. 1966. A preliminary attempt to divide Chile into entomofaunal regions based on the Tenebrionidae (Coleóptera). Postilla 97: 1-17. PORTER, C. 1980. Zoogeografía de las lchneumonidae Latino-Americanas (Hymenoptera). Acta Zool. Lilloana 36: 1-52. RAVEN, P. H., AND D. 1. AXELROD. 1975. History

of the flora and fauna of Latin America. Amer. Sci. 63: 420-429. SCOTT, J. A. 1972. Biogeography of Antillean butterflies. Biotropica 4: 32-45. UDVARDY, M. D. F 1975. A classification of the

biogeographical provinces IUCN, Occ. Pap. 18: 1-48.

of the world.

Faunistics T h e present-day composition of species in a particular g e o g r a p h i c a r e a c o m p r i s e s its fauna. As shown above, t h e n a t u r e of faunas, t h e relative n u m b e r of different ecological types, a n d t h e t a x o n o m i c spec­ t r u m offer clues to t h e geologic history of the a r e a a n d a r e i m p o r t a n t to ecologists as indicators of c o m m u n i t i e s a n d ecosystems. T h e e x t e n t of t h e entire Latin A m e r i c a n e n t o m o f a u n a is n o t yet k n o w n . Estimates go as high as 20 million species. Attempts have b e e n m a d e to identify t h e insects and related a r t h r o p o d s of m a n y p o r t i o n s of the region, b u t n o n e can be c o n s i d e r e d com­ plete. It involves a g r e a t deal of effort and taxonomic expertise even to compile a g e n e r a l list of all t h e speices in a small area. Only a few g e n e r a l lists a n d c u r s o r y faunal reviews have b e e n published; consult the Faunal Surveys in c h a p t e r 13 for a listing of these.

POPULATION BIOLOGY T h e size, s t r u c t u r e , energetics, composi­ tion, a n d mobility of insect p o p u l a t i o n s in various life zones, m a c r o h a b i t a t s , a n d microhabitats a r e highly significant, a n d a very large b o d y of literature on b o t h theo­ retical a n d a p p l i e d insect d e m o g r a p h y exists (Young 1982). Such investigations always r e q u i r e quantitative analysis of population p a r a m e t e r s , a n d particular sam­ pling t e c h n i q u e s a r e n e e d e d in each spe­ cialty a r e a ( S o u t h w o o d 1980, Wolda 1984). Most works deal with specific insects (Ehrlich a n d Gilbert 1973), often of eco­ nomically i m p o r t a n t types, limited taxa (Torres 1984), o r guilds ( u n r e l a t e d species with similar niches) ( H e i t h a u s 1979) within particular e n v i r o n m e n t s . S o m e of this re­ search may i n c l u d e c o m p a r i s o n s between different habitat types. Investigations of whole c o m m u n i t i e s a r e very few owing to their complexity a n d their multifold inter­ actions with t h e e n v i r o n m e n t . T h e lower layers of tropical forests (low­ land, m i d l a t i t u d e moist to rain forests) have attracted t h e most attention from insect p o p u l a t i o n biologists in Latin A m e r ­ ica. S a m p l i n g is direct; specimens can be observed directly o r c a p t u r e d fairly easily with nets, t h e " s w e e p i n g " t e c h n i q u e often being used for soft foliage, h e r b s , or grasses (Allan et al. 1973, J a n z e n 1973). A n intensive t h i r t e e n - m o n t h survey of Amazo­ nian forest was u n d e r t a k e n in 1 9 7 7 - 7 8 in the Ducke Forest Preserve n e a r M a n a u s (Penny a n d Arias 1982), for which passive Malaise traps c a u g h t most of t h e samples. Long-term studies a r e r a r e (Wolda 1983). O v e r a p e r i o d of several years, Elton (1973) s a m p l e d t h e forest "field layer" (15 c m to 1.8 m above t h e g r o u n d ) in a variety of localities a n d with different methods a n d c o n c l u d e d that insect life generally exists in low n u m b e r s a n d that the sizes of most a r e small b u t that t h e species richness is very high. T h e most common f o r m s f o u n d were ants, spiders,

a n d o r t h o p t e r o i d s . T h e very diverse noc­ t u r n a l fauna of flying insects is c o m p o s e d of m u c h larger insects. Using sweep n e t samples to study diver­ sity a n d distribution, various a u t h o r s (e.g., J a n z e n 1973) have discovered p r o f o u n d differences along elevational transects, for e x a m p l e , in Costa Rica a n d t h e Venezuelan A n d e s ( J a n z e n et al. 1976). Latitudinal effects ( J a n z e n a n d Pond 1975) as well as seasonal effects ( J a n z e n a n d S c h o e n e r 1968, T a n a k a a n d T a n a k a 1982) have also been included. Light t r a p p i n g h a s revealed s o m e p o p u lational traits of Neotropical insects (Ricklefs 1975; Wolda 1978a, 19786). I n lowland forest, t h e seasonality of flight activity o f nocturnal forms is p r o n o u n c e d . Generally, n u m b e r s a r e d e p r e s s e d in t h e m i d - d r y period a n d maximally e x p a n d e d in t h e early weeks o f wetness in both m a i n l a n d (Wolda 1980) a n d island (Tanaka a n d Ta­ n a k a 1982) areas. T h e suppressive effect of m o o n l i g h t o n t h e activity of night-flying insects is also often n o t e d (Wolda a n d Flowers 1985). T h e insects of t h e forest canopy, al­ t h o u g h now accessible by special tech­ niques, resist quantitative study because of the difficulties r e m a i n i n g in taking sizable samples a n d covering a significant a r e a of habitat. Some advances have been m a d e in the study of c a n o p y b e e biology by climb­ i n g entomologists (Perry 1983). T r a p s (Mal­ aise, "photocollectors") have also b e e n raised to u p p e r levels for r e m o t e assess­ m e n t . In this way, t h r e e responses by arboreal insects in A m a z o n i n u n d a t i o n forests have b e e n detected d u r i n g t h e flood season: t e m p o r a r y immigration, survival in place, a n d dying o u t (Adis 1977). A fruitful a p p r o a c h for obtaining statisti­ cally a d e q u a t e data from t h e c a n o p y h a s c o m e from t h e insecticide fogging tech­ nique. Rapid-acting p y r e t h r o i d s a r e blown into t h e trees. ( T h e s e mildly toxic chemi­ cals rapidly d e g r a d e in t h e e n v i r o n m e n t

POPULATION BIOLOGY

69

a n d d o not h a r m n o n t a r g e t organisms.) All the e x p o s e d insects a r e s t u n n e d a n d fall to the g r o u n d , w h e r e they a r e collected, iden­ tified, a n d c o u n t e d . I n this way, almost c o m p l e t e samples c a n b e taken. Cock­ roaches (Fisk 1982), wax bugs (Wolda 1980), g r a s s h o p p e r s (Roberts 1973), a n d carabid beetles (Erwin 1983) have b e e n analyzed. S o m e c o m p a r i s o n s o f species richness of t e m p e r a t e r h e o p h i l i c (Stout a n d V a n d e r m e e r 1975) forms to those in Costa Rican a n d A n d e a n (Patrick 1966) s t r e a m s have b e e n m a d e . Results generally show that m i d l a t i t u d e s t r e a m f a u n a s a r e significantly m o r e diverse t h a n those in c o m p a r a b l e , high-latitude streams. Soil a n d litter types a r e studied by m e a n s of t h e Berlese (Tullgren) funnel (Beebe 1916). Samples m a y also b e t a k e n with a c o r e a u g e r a n d s e p a r a t i o n by flota­ tion in liquid s u s p e n s i o n s . Substrata m a y be e x p e r i m e n t a l l y m a n i p u l a t e d , which al­ lows t h e investigator to m e a s u r e m i n u t e habitat characteristics with g r e a t accuracy a n d c o m p a r e t h e m to n a t u r a l plots (Stanton 1979). F r o m these various p o p u l a t i o n studies, it is generally c o n c e d e d t h a t in tropical for­ ests, insect a b u n d a n c e is low b u t the diver­ sity very high. It seems also that t h e d i s p e r ­ sion of insect p o p u l a t i o n s over a particular a r e a is s e l d o m even b u t usually very patchy. T h i s may b e caused by t h e i r r e g u l a r distri­ b u t i o n of habitats (Wiens 1976), b u t it can be so even if e n v i r o n m e n t a l n e e d s a p p e a r continuously a n d a b u n d a n t l y available. Moisture c o n t e n t of litter a n d soil may b e very u n e v e n l y d i s t r i b u t e d , for e x a m p l e , a n d a major factor affecting the p r e s e n c e or absence of ants, springtails, mites, a n d o t h e r a r t h r o p o d s (Levings a n d W i n d s o r 1984). However, in spite of t h e e n o r m o u s n u m b e r s of leaves available o n a single tree or in a forest, only a few a r e n o r m a l l y utilized by h e r b i v o r e s at o n e time (notwith­ s t a n d i n g p o p u l a t i o n explosions w h e n defo­ liation m a y occur).

70

ECOLOGY

References ADIS, J. 1977. Programa mínimo para análises de ecosistemas: Artrópodos terrestres em florestas inundáveis da Amazonia central. Acta Amazónica 7: 223-229. ALLAN, J. D., L. W. BARNHOUSE, R. H. PRESTBYE,

AND D. R. STRONG. 1973. On foliage arthro­ pod communities of Puerto Rican second growth vegetation. Ecology 54: 628—632. BEEBE, W. 1916. Fauna of four square feet of jungle debris. Zoológica 2: 107-119. EHRLICH,

P. R., AND L. E. GILBERT.

1973.

Population structure and dynamics of the tropical butterfly Heliconius ethilla. Biotropica 5: 69-82. ELTON, C. S. 1973. T h e structure of inverte­ brate populations inside Neotropical rain for­ est. J. Anim. Ecol. 42: 55-104. ERWIN, T L. 1983. Beetles and other insects of tropical forest canopies at Manaus, Brazil, sampled by insecticidal fogging techniques. In S. L. Sutton, T C. Whitmore, and A. C. Chadwick, eds., Tropical rain forests: Ecology and management. Blackwell, Oxford. Pp. 59-75. FISK, F. W. 1982. Abundance and diversity of arboreal Blattaria in moist tropical forest of the Panama Canal area and Costa Rica. Amer. Entomol. Soc, Trans. 108: 479-489. HF.ITHAUS, E. R. 1979. Community structure of Neotropical flower visiting bees and wasps: Di­ versity and phenology. Ecology 60: 190—202. JANZEN, D. H. 1973. Sweep samples of tropical foliage insects: Effects of seasons, vegetation types, elevation, time of day and insularity. Ecology 54: 687-708. JANZEN, D. H., M. ATTAROFF, M. FARIÑAS, S. REYES, N. RINCÓN, A. SOLER, P. SORIANO,

AND M. VERA. 1976. Changes in the arthro­ pod community along an elevational transect in the Venezuelan Andes. Biotropica 8: 1 9 3 203. JANZEN,

D. H., AND C. M. POND.

1975. A

comparison, by sweep sampling, of the arthro­ pod fauna of vegetation in Michigan, Englz d and Costa Rica. Royal Entomol. Soc. London, Trans. 127: 30-50. JANZEN, D. H., AND T

W. SCHOENER.

1968.

Differences in insect abundance and diversity between wetter and drier sites during a tropi­ cal dry season. Ecology 49: 96-110. LEVINGS, S. C , AND D. M. WINDSOR 1984. Litter

moisture content as a determinant of litter arthropod distribution and abundance dur­ ing the dry season on Barro Colorado Island, Panama. Biotropica 16: 125—131. PATRICK, R. 1966. The Catherwood Foundation

Peruvian Amazon Expedition: Limnological and systematic studies. Monogr. Acad. Nat. Sci. Phil. 14: 1-495. PENNY, N. D., AND J. R. ARIAS. 1982. Insects of

an Amazon forest. Columbia Univ. Press, New York. PERRV, D. R. 1983. Access methods, observa­ tions, pollination biology, bee foraging behav­ ior, and bee community structure within a Neotropical wet forest canopy. Ph.D. diss., Univ. Calif, Los Angeles. RICKLEFS, R. E. 1975. Seasonal occurrence of night-flying insects on Barro Colorado Is­ land, Panama Canal Zone. New York Ento­ mol. Soc, J. 83: 19-32. ROBERTS, H. R. 1973. Arboreal Orthoptera in the rain forests of Costa Rica collected with insecticide: A report on the grasshoppers (Acrididae), including new species. Acad. Nat. Sci. Philadelphia Proc. 125: 49-66. SOUTHWOOD, T R. E. 1980. Ecological methods, with particular reference to the study of populations. 2d ed. Chapman & Hall, Lon­ don. STANTON, N. L. 1979. Abundance and diversity of Homoptera in the canopy of a tropical forest. Ecol. Entomol. 4: 181-190. STOUT, J., AND J. VENDERMEER 1975. Compari­

son of species richness for stream-inhabiting insects in tropical and mid-latitude streams. Amer. Nat. 109: 263-280. TANAKA, L. K., AND S. K. TANAKA. 1982. Rainfall

and seasonal changes in arthropod abun­ dance on a tropical oceanic island. Biotropica 14: 114-123. TORRES, J. A. 1984. Niches and coexistence of ant communities in Puerto Rico: Repeated patterns. Biotropica 16: 284-295. WIENS, J. A. 1976. Population responses to patchy environments. Ann. Rev. Ecol. Syst. 7: 81-120. WOLDA, H. 1978a. Fluctuations in abundance of tropical insects. Amer. Nat. 112: 1017-1045. WOLDA, H. 19786. Seasonal fluctuations in rain­ fall, food and abundance of tropical insects. J. Anim. Ecol. 47: 369-381. WOLDA, H. 1980. Seasonality of tropical insects. I. Leafhoppers (Homoptera) in Las Cumbres, Panama.]. Anim. Ecol. 49: 277-290. WOLDA, H. 1983. "Long-term" stability of tropi­ cal insect populations. Res. Pop. Ecol. Suppl. 3:112-126. WOLDA, H. 1984. Diversidad de la entomofauna y como medirla. 9th Cong. Latinoamericano Zool. (Arequipa), Inf. Final. Pp. 181-186. WOLDA, H., AND R. W. FLOWERS. 1985. Sea­

sonality

and diversity

of mayfly

adults

(Ephemeroptera) in a "nonseasonal" environ­ ment. Biotropica 17: 330-335. YOUNG, A. M. 1982. Population biology of tropical insects. Plenum, New York.

FOOD RELATIONS I m p o r t a n t to their ecological roles a r e t h e kinds of food a n d feeding habits exhibited by insects (Brues 1946, C u m m i n s 1973). A majority, herbivores (or p h y t o p h a g e s ) , act as p r i m a r y r e d u c e r s of plant life (d'Araújo y Silva et al. 1 9 6 7 - 6 8 , G u a g l i u m i 1966, Martell 1974, Martorell 1976), including a relatively few s a p r o p h a g e s that live only o n d e a d plant material such as d e a d w o o d . N e c r o p h a g e s specialize o n animal c a r r i o n , a n d c o p r o p h a g e s utilize animal feces. H i g h e r o n the p y r a m i d is a n o t h e r large a n d highly competitive g r o u p , t h e p r e d a t o r s , which catch, kill, a n d c o n s u m e o t h e r living animals, most often o t h e r insects (Clausen 1962) b u t sometimes even healthy verte­ brates (Formanowicz et al. 1981, Hayes 1983). (Body toxins sometimes p r o t e c t a n u r a n s from p r e d a t i o n by spiders; t h e latter may even react negatively t o aposematic coloration of this potential p r e y [Szelistowski 1985].) Modifying t h e strategy of p r e d a t i o n are the parasites, which r o b food (including blood) from hosts o n o r within whose bodies they intimately live without causing their d e a t h , a n d parasitoids, which live like parasites b u t eventually kill t h e host. T h e latter two types a r e n o t always clearly s e p a r a t e d from each o t h e r (Askew 1971, Price 1975). T h e r e are also a variety of o t h e r n a r r o w feeding specialists o n o d d n u t r i e n t sources, such as hair follicle secre­ tions (hair follicle mites), beeswax (wax moths), c u l t u r e d fungi ( g a r d e n i n g ants), a n d m a n y others. W h o l e guilds may b e tied to a particular v e r t e b r a t e host, a good e x a m p l e being the a r t h r o p o d associates of sloths (Waage a n d Best 1985). Kleptoparasites a n d social parasites steal u n ­ g u a r d e d prey from s p i d e r webs o r wasp

FOOD RELATIONS

71

food caches. Many insects c o m b i n e such habits a n d a r e c o n s i d e r e d o m n i v o r e s . Herbivory a n d s a p r o p h a g y a r e vastly c o m p l e x feeding practices because of t h e variety of plant types a n d anatomical parts available as food for insects, including leaves (Ernest 1989), stems, b u d s , seeds, a n d fruits (Strong et al. 1984). Species r a n g e from highly restricted in their choice of species o r p a r t (host specific) to catholic in their tastes (generalists). T h e r e a r e in­ sects that graze o n d i a t o m s a n d algae from films o n rocks in s t r e a m s , lichen browsers, a n d fern feeders ( F o r n o a n d B o u r n e 1984, H e n d r i x 1980), in a d d i t i o n to those that eat a n g i o s p e r m s generally, both in aquatic ( C u m m i n s 1973) a n d terrestrial (Zimmer­ m a n et al. 1979) e n v i r o n m e n t s . Living, d e a d , a n d e v e n d e c o m p o s i n g ( W i n d e r 1977) m a t t e r is utilized as food. T h e greatest availability to herbivores is found in forests, especially tropical rain forests, w h e r e t h e n u m b e r of plant species is p r o d i g i o u s . S o m e g e n e r a l studies in this v e n u e a r e those c o m p a r i n g h e r b i v o r e d a m ­ age in r i p a r i a n versus d r y forests (Stanton 1975), t h e effects of seed p r e d a t i o n in d e t e r m i n i n g tree distribution (Janzen 1970), a n d host specificity (Pipkin et al. 1966). T h e i m p o r t a n c e of insect r e d u c e r s of d e a d w o o d a n d its c o n v e r s i o n t o soil in forests c a n n o t be overstated ( M o r ó n 1985). T h e s u p e r a b u n d a n c e of plants for h e r b i ­ vores would seem to allow e n o r m o u s explo­ sions of insect n u m b e r s , b u t t h e latter a r e r a r e events that a r e often associated with h u m a n p e r t u r b a t i o n s , such as misuse of insecticides a n d m o n o c u l t u r e . N a t u r a l fac­ tors limiting e x p a n s i o n s of plant-eating insects a r e parasitism, p r e d a t i o n , a n d u n palatability (from both physical a n d chemi­ cal d e t e r r e n t s ) , l e a d i n g to intraspecific com­ petition. T h e r e is e v i d e n c e that t h e latter may be increased by t h e plant in direct r e s p o n s e t o insect attack (Wratten et al. 1988). A special category of h e r b i v o r e s a r e t h e gall m a k e r s . M a n y flies, wasps, a n d mites,

72

ECOLOGY

including entire families in these o r d e r s , the Cecidomyiidae, Cynipidae, a n d Eriophyidae, respectively, stimulate their plant hosts to form a b n o r m a l neoplastic growths called galls ( A n a n t h a k r i s h n a n 1984). T h e s e take many forms a n d occur o n all p a r t s of the plant a n d a r e often h a r m f u l (Fernandes 1987). T h e y serve t h e d e v e l o p i n g stage of the insect as a nutrient-rich, a b u n d a n t , a n d reliable food supply. T h e m e c h a n i s m of gall formation is not fully u n d e r s t o o d a n d var­ ies a m o n g t h e g r o u p s involved. Very little is k n o w n of t h e galls of Latin America (Occhioni 1979). Leaf m i n e r s ( H e r i n g 1951) form a n o t h e r specialized g r o u p of herbivores. It is t h e larvae of several families of Diptera (Agromyzidae, T e p h r i t i d a e , A n t h o m y i i d a e ) a n d L e p i d o p t e r a (Nepticulidae, Tischeriidae, Lyonetiidae, etc.), in particular, t h a t have a d a p t e d to t h e confined microhabitat be­ tween t h e u p p e r a n d lower e p i d e r m a l tis­ sues of leaves. Understandably, their chief morphological characteristics a r e small size a n d a compressed s h a p e . A p p e n d a g e s a r e r e d u c e d o r absent, a l t h o u g h t h e m o u t h parts a r e well developed. T h e same is t r u e of b o r e r s , in wood a n d o t h e r tissues, found a m o n g t h e C o l e ó p t e r a (Cerambycidae, Buprestidae) a n d L e p i d o p t e r a (Pyralidae, Castniidae, Cossidae). N e c r o p h a g o u s insects occur chiefly in the orders Coleóptera a n d Diptera (Jirón a n d Cartin 1981). Several beetle families feed entirely o n d e a d v e r t e b r a t e animals o r their skins a n d hair, such as t h e carrion beetles (Silphidae) a n d d e r m e s t i d s (Dermestidae). I m p o r t a n t carrion fly g r o u p s a r e the blowflies (Calliphoridae) a n d flesh flies (Sarcophagidae). T h e succession of com­ munities of such insects in h u m a n cadavers is t h e key to their use in d e t e r m i n i n g time of d e a t h as sometimes e m p l o y e d in forensic medicine (see Valuable Insects, c h a p . 3). Dead insect carcasses also p r o v i d e a form of carrion for n e c r o p h a g o u s species (Young 1986). T h e s e insects a r e i m p o r t a n t in nutri­ e n t t u r n o v e r a n d , from t h e s t a n d p o i n t of

environmental hygiene, fortunately, a r e a b u n d a n t a n d w i d e s p r e a d in n a t u r e (Mo­ rón a n d T e r r ó n 1984). A major g r o u p of t h e ecologically i m p o r ­ tant c o p r o p h a g e s , insects that feed in o n e stage o r a n o t h e r o n t h e feces of o t h e r animals, a r e t h e P h a n a e i n e a n d C o p r i n e d u n g scarabs (Scarabaeidae) (Peck a n d Howden 1984). S o m e g r o u p s a r e highly specific, for instance, larval sloth m o t h s (Pyralidae, see Sloth Moths, c h a p . 10) a n d the scarab beetle, Uroxys gorgon, o n sloth d u n g (Young 1981). A g r e a t m a n y insects with sucking m o u t h p a r t s survive entirely o n , o r fre­ quently s u p p l e m e n t their diets with, liquid foods. T h e variety a n d quality of n u t r i e n t s available in solution is great. C a r b o h y ­ drates, a m i n o acids, minerals, vitamins, and even fats a r e dissolved in such diverse and unlikely fluids as blood a n d l y m p h from animals, s a p , r a i n w a t e r solutions, nectar, a n i m a l a n d plant secretions (sweat, milk, tears), honeydew, d e c o m p o s i t i o n products, w e e p i n g w o u n d s , decay juices, plant juices, fecal matter, a n d u r i n e . W h a t e v e r t h e food source, its n u t r i e n t content m u s t be a p p r o p r i a t e for each spe­ cies (Dadd 1973). Insects generally r e ­ quire t h e s a m e basic classes of dietary substances—minerals, calories, p r o t e i n , carbohydrates, fats, vitamins—as verte­ brates, b u t m a n y peculiar o r restricted fac­ tors a r e n e e d e d to fulfill t h e metabolic quirks of idiosyncratic species. S o m e a r e incapable e v e n of digesting their food without t h e i n t e r v e n t i o n of symbiotic mi­ croorganisms in their guts (e.g., primitive termites a n d s o m e cockroaches), while oth­ ers m u s t cultivate their food, t h e best ex­ ample of which a r e t h e g a r d e n i n g ants (Attinae).

References ANANTHAKRISHNAN, T. N. 1984. Biology of gall insects. Edward Arnold, London. DARAÚJO Y SILVA, A. G., C. R. GONCALVES, D. M. GALVÁO, A. J. LOBO GONCALVES,

J. GOMES,

M. DO NASCIMENTO

SILVA, AND

L. DE SIMONI. 1967-68. Quarto catálogo dos insetos que vivem ñas plantas do Brasil. Pts. 1-2. Min. Agrie, Dept. Def. Insp. Agropec, Rio de Janeiro. ASKEW, R. R. 1971. Parasitic insects. American Elsevier, New York. BRUES, C. T. 1946. Insect dietary. An account of the food habits of insects. Harvard Univ. Press, Cambridge. CLAUSEN, C. P. 1962. Entomophagous insects. Hafner, New York. CUMMINS, K. W. 1973. Trophic relations of aquatic insects. Ann. Rev. Entomol. 18: 183— 206. DADD, R. H. 1973. Insect nutrition: Current developments and metabolic implications. Ann. Rev. Entomol. 18: 381-420. ERNEST, K. A. 1989. Insect herbivory on a tropical understory tree: Effects of leaf age and habitat. Biotropica 21: 194-199. FERNANDES, G. W. 1987. Gall forming insects: Their economic importance and control. Rev. Brasil. Entomol. 31: 379-398. FORMANOWICZ, J R . , D. R., M. M. STEWART, K . TOWNSEND, F. H . POUGH, AND P. F.

BRUSSARD. 1981. Predation by giant crab spiders on the Puerto Rican frog Eleutherodactylus coqui. Herpetologica 37: 125-129. FORNO, I. W., AND A. S. BOURNE. 1984. Studies

in South America of arthropods on the Salvinia auriculata complex of floating ferns and their effects on 5. molesta. Bull. Entomol. Res. 74: 609-621. GUAGLIUMI, P. 1966. Insetti e aracnidi delle pianti comuni del Venezuela segnalati nel periodo 1938-1963. Rel. Mono. Agrar. Subtrop. Trop. (Nov. Ser. 86): 1-391. HAYES, M. P. 1983. Predation on the adults and prehatching stages of glass frogs (Centrolenidae). Biotropica 15: 74-76. HENDRIX, S. D. 1980. An evolutionary and

ecological perspective of the insect fauna of ferns. Amer. Nat. 115: 171-196. HERING, M. 1951. Biology of the leaf miners. Junk, T h e Hague. JANZEN, D. H. 1970. Herbivores and the num­ ber of tree species in tropical forests, Amer. Nat. 104: 501-528. JIRÓN, L. F , AND V. M. CARTÍN. 1981. Insect

succession in the decomposition of a mammal in Costa Rica. New York Entomol. Soc. J. 89: 158-165. MARTELL, C. 1974. Primer catálogo de los insectos fitófagos de México. Fitófilo 27: 1 176. MARTORELL, L. F. 1976. Annotated food plant

FOOD RELATIONS

73

catalog of the insects of Puerto Rico. Agrie. Expt. Sta., Univ. Puerto Rico, Puerto Rico. MORÓN, M. A. 1985. Los insectos degradadores, un factor poco estudiado en los bosques de México. Fol. Entorno!. Mexicana 65: 131-137. MORÓN,

M. A.,

AND R.

A.

TERRÓN.

1984.

Distribución altitudinal y estacional de los insectos necrófilos en la Sierra Norte de Hildago, México. Acta Zool. Mexicana (n.s.) 3: 1-47. OCCHIONI, P. 1979. "Galhas," "cecídeas" ou "tumores vegetáis" em plantas nativas da flora do Brasil. Leandra 8—9: 5—35. PECK, S. B., AND H. F. HOWDEN. 1984. Response

of a dung beetle guild to different sizes of dung bait in a Panamanian rainforest. Biotropica 16: 235-238. PIPKIN, S. B., R. L. RODRIGUEZ, AND J. LEON.

1966. Plant host specificity among flower feeding Neotropical Drosophüa (Díptera: Drosophilidae). Amer. Nat. 100: 135-156. PRICE, P. W., ed. 1975. Evolutionary strategies of parasitic insects and mites. Plenum, New York. STANTON, N. 1975. Herbivore pressure on two types of tropical forests. Biotropica 7: 8—11. STRONG, D. R., J. H. LAWTON, AND R. SOUTH-

WOOD. 1984. Insects on plants, community patterns and mechanisms. Harvard Univ. Press, Cambridge. SZELISTOWSKI, W. A. 1985. Unpalatability of the

poison arrow frog Dendrobates pumilio to the ctenid spider Cupiennius coccineus. Biotropica 17: 345-346. WAAGE, J. K., AND R. C. BEST. 1985. Arthropod

associates of sloths. In G. G. Montgomery, ed., The evolution and ecology of armadillos, sloths, and vermilinguas. Smithsonian Inst., Washington, D.C. Pp. 297-311. WINDER, J. A. 1977. Some organic substrates which serve as insect breeding sites in Bahian cocoa plantations. Rev. Brasil. Biol. 37: 351 — 356. WRATTEN, S. D., P. J. EDWARDS, AND L. WINDER.

1988. Insect herbivory in relation to dynamic changes in host plant quality. Biol. J. Linnean Soc. 35: 339-350. YOUNG, A. M. 1986. Carcass-scavenging by

Taeniopoda reticulata (Orthoptera: Acrididae) in Costa Rica. Entomol. News 97: 175-176. YOUNG, O. P. 1981. T h e utilization of sloth dung in a Neotropical forest. Coleop. Bull. 35: 427-430. ZIMMERMAN,

H. G.,

H.

E.

E R B , AND R. E.

MCFADYEN. 1979. Annotated list of some cactus-feeding insects of South America. Acta Zool. Lilloana33: 101-112.

74

ECOLOGY

DIVERSITY T h e best recent c o u n t of t h e total n u m b e r of n a m e d insect species in t h e world is 1,111,225 (Arnett 1983), a l t h o u g h m a n y textbooks still place t h e figure at 650,000 to 700,000. Plausible a r g u m e n t s a r e of­ fered by a u t h o r s for m a n y m o r e , 3 to 5 million o r 10 million (May 1988) existing species. By liberal e x t r a p o l a t i o n of some data, t h e r e a r e indications of even as m a n y as 3 0 million actual living species of tropi­ cal forest a r t h r o p o d s alone (Erwin 1982) or possibly several times that a m o u n t (Stork 1988). W h a t e v e r figure is accurate, most of t h e total could b e e x p e c t e d to occupy Latin America, because of t h e r e ­ gion's very rich flora, large area, a n d complexity of physical a n d biotic habitats. An e d u c a t e d guess of 4 0 p e r c e n t of t h e total world's biota could b e m a d e for t h e n u m b e r of species ultimately discoverable in t h e region, based o n available insect host diversity a n d land habitat area p r e s e n t c o m p a r e d to that of t h e o t h e r zoogeog r a p h i c regions. Most of these species occur in lowland forests. T h e reasons for t h e especially high di­ versity in t h e tropical p o r t i o n s a r e n o t fully u n d e r s t o o d , a l t h o u g h various theories have b e e n p r o p o s e d (Dobzhansky 1950, Pianka 1966). N o d o u b t , t h e inclusion of vast areas of wet lowland forest that pro­ vide a b u n d a n t niches a n d a stable climate, at least in refuges, h a s m u c h to d o with it. T h e r e is little d o u b t that t h e r e a r e m o r e species of insects of most major g r o u p s in the tropics t h a n in t e m p e r a t e areas, with the possible exception of some parasitic H y m e n o p t e r a a n d bees (Wolda 1983), a p h i d s (Dixon et al. 1987), t h e Plecoptera, a n d a few o t h e r s . A c c o r d i n g to J a n z e n (1976), t h e e x p l a n a t i o n lies chiefly in t h e a m o u n t h e r e of "harvestable productivity" a r r a n g e d in a sufficiently h e t e r o g e n e o u s m a n n e r . T h i s allows utilization by a vast c o m p l e x of small a n d very a d a p t a b l e reduc­ ing animals, most fittingly, t h e insects.

H e t e r o g e n e i t y is p r o v i d e d by climatic and g e o g r a p h i c variation over wide areas for long p e r i o d s of time. T h e best e x a m p l e is p r o v i d e d by t h e A m a z o n Basin, w h e r e insect diversity is very high as a result of habitat d i s t u r b a n c e s — f r o m a constantly changing vegetation, dissection of t h e land­ scape by rivers, a l t e r n a t i o n of i n u n d a t e d dry forest types, a n d possibly o t h e r p h e n o m e n a — f o r at least t h e d u r a t i o n of the P l i o c e n e - P l e i s t o c e n e e r a (Erwin a n d Adis 1982). T h e stability of t h e wet lowland tropics gives t h e insects that live t h e r e a h e a d start on speciation a n d allows t h e m to a c c u m u ­ late, b u t because of competition, n o single area becomes greatly e n r i c h e d . T h e fact that t h e r e a r e a g r e a t n u m b e r of species but relatively few individuals is t h o u g h t by some t o b e largely d u e t o t h e high p r o d u c ­ tivity a n d availability of r e s o u r c e s in such forests a n d t h e r e d u c e d seasonality that makes it possible for insects to occupy marginal niches ( M a c A r t h u r 1969).

References ARNETT, JR., R. H. 1983. Status of the taxonomy of the insects of America north of Mexico: A preliminary report prepared for the subcom­ mittee for the insect fauna of North America Project. For Stand. Comm. System Res., Entomol. Soc. America. DIXON, A. F. G., P. KINDLMANN, L. LEPS, AND

J. HOLMAN. 1987. Why there are so few spe­ cies of aphids, especially in the tropics. Amer. Nat. 129: 580-592. DOBZHANSKY, T. 1950. Evolution in the tropics. Amer. Sci. 38: 209-221. ERWIN, T. L. 1982. Tropical forests: Their richness in Coleóptera and other arthropod species. Coleop. Bull. 36: 74-75. ERWIN, T. L., AND J. ADIS. 1982. Amazonian

inundation forests: Their role as short-term refuges and generators of species richness and taxon pulses. In G. T. Prance, ed., Biologi­ cal diversification in the tropics. Columbia Univ. Press, New York. Pp. 358-371. JANZEN, D. H. 1976. Why are there so many species of insects? 15th Int. Congr. Entomol. Proc. Pp. 84-94. MACARTHUR, R. H. 1969. Patterns of communi­

ties in the tropics. Biol. J. Linnean Soc. 1: 19— 30. MAY, R. M. 1988. How many species are there on earth? Science 341: 1441-1449. PIANKA, E. R. 1966. Latitudinal gradients in species diversity: A review of concepts. Amer. Nat. 100:33-44. STORK, N. E. 1988. Insect diversity: Facts, fic­ tion and speculation, Biol. J. Linnean Soc. 35: 321-337. WOLDA, H. 1983. Diversity, diversity indices and tropical cockroaches. Oecologia 58: 290-298.

ENEMIES Insects suffer diseases from microbial p a t h o g e n s , as d o h i g h e r o r g a n i s m s (Cantwell 1974, Steinhaus 1963). N o species is i m m u n e from infections by a host of vi­ ruses, rickettsias, bacteria, n e m a t o d e s (Nickle a n d Welch 1984), a n d p r o t o z o a n s . Some of these have practical application in the control of insect pests (Roberts a n d Castillo 1980). Inocula of several types of Bacillus thuringiensis already a r e p r o d u c e d commercially a n d a r e very successful in t h e battle against m a n y caterpillar pests a n d blackfly larvae (see C o n t r o l of Insect Pests, c h a p . 3). Insects a r e also afflicted with m a n y fun­ gal infections (Madelin 1966). T h e s e a r e especially well d e v e l o p e d in t h e m o r e moist portions of t h e Latin A m e r i c a n tropics. Species of t h e g e n u s Cordyceps (Ascomycotina: Clavicipitales) a r e t h e most com­ m o n . Dead specimens of m a n y kinds of insects a r e often f o u n d with t h e fruiting bodies of these fungi projecting from t h e m . I n parts of t h e Peruvian rain forest, they a r e well k n o w n to t h e natives as tamshi a n d a r e t h o u g h t by t h e naive to b e meta­ m o r p h o s i n g plants (see Giant Solitary H u n t i n g A n t , c h a p . 12). Such e n t o m o p h a g o u s fungi a r e u n d o u b t e d l y i m p o r t a n t in t h e regulation of a r t h r o p o d p o p u l a t i o n s a n d h e l p to maintain stability in rain forest ecosystems (Evans 1982). Insects bear t h e b u r d e n of b e i n g t h e

ENEMIES

75

principal food of a g r e a t m a n y types of v e r t e b r a t e a n i m a l s . U n d o u b t e d l y , t h e larg­ est single g r o u p of insectivores a r e birds, the majority of which r e q u i r e insects in their diet at s o m e time of t h e i r lives. M a n y eat insects exclusively, such as flycatchers (Sherry 1984), swifts a n d swallows (Collins 1968, H e s p e n h e i d e 1975), a n d w o o d p e c k ­ ers. S o m e show specific f o r a g i n g a d a p t a ­ tions, even selecting u p p e r versus lower leaf surfaces ( G r e e n b e r g a n d G r a d w o h l 1980). A n t b i r d s specialize in eating insects forced to e x p o s e themselves while e s c a p i n g a r m y ant f e e d i n g swarms. T h e r e a r e m a n y insect-feeding t e r r e s ­ trial m a m m a l s , primarily o p o s s u m s , e d e n ­ tates ( a n t e a t e r s a n d armadillos), a n d pri­ mates. A n t e a t e r s c o n s u m e g r e a t n u m b e r s of t e r m i t e s a n d ants in tropical a r e a s a n d are s t r o n g d e t e r m i n a n t s of t h e latter's distribution a n d a d a p t a t i o n s for survival ( L u b i n e t a l . 1977). E n o r m o u s quantities of night-flying in­ sects, mostly m o t h s , beetles, a n d gnats, a r e c o n s u m e d by bats. A p p r o x i m a t e l y 70 per­ cent of living species a r e insectivores, eat­ ing o n t h e a v e r a g e a q u a r t e r to a half of their b o d y weight in insects nightly (Hill a n d Smith 1984: 63). Reptiles a n d a m p h i b i a n s a r e heavy in­ sect p r e d a t o r s ( L i e b e r m a n 1986). Most lizards (Lescure a n d Fretey 1977) a n d a few kinds of snakes a n d small crocodilians (Seijas a n d R a m o s 1980) rely o n a n insect diet a n d m a y b e s t r o n g selective agents in insect a d a p t a t i o n ( H u n t 1983). Legless liz­ a r d s living in r o t t i n g w o o d a p p e a r to p r e y almost entirely o n t e r m i t e s . A m p h i b i a n s , particularly frogs, a r e also e n t o m o p h a g e s . A r b o r e a l species t e n d to specialize in ants a n d mites (Formanowicz et al. 1981), while litter-dwelling f o r m s h a v e a m o r e varied insect diet (Toft 1981). S a l a m a n d e r s a r e significant only from t h e n o r t h e r n A n d e s n o r t h w a r d . Fossorial caecilians m a y also take s o m e soil insects in a d d i t i o n t o their p r i m a r y e a r t h w o r m diet, a l t h o u g h t h e aquatic species may b e m o r e insectivorous.

76

ECOLOGY

C a r n i v o r o u s plants a r e highly a d a p t e d for t r a p p i n g a n d digesting insect p r e y (Lloyd 1931). T h e r e a r e several categories r e p r e s e n t e d in Latin America, differing in m e t h o d of c a p t u r e a n d form of t h e t r a p ­ ping device. T h o s e with pitfalls, t h e socalled pitcher plants, all belong to t h e g e n u s Heliamphora (Sarraceniaceae) a n d a r e f o u n d only in t h e G u i a n a a r e a a n d Venezuela. T h e y g r o w in d e n s e clusters in bogs. T h e leaves a r e modified into u r n s h a p e d structures, filled with a secretion that is corrosive t o most insects that fall into it. S o m e aquatic insects have evolved special m e c h a n i s m s t o protect t h e m from this chemical action a n d p o p u l a t e this gen­ erally unsuitable microhabitat (see T a n k Plants, above). T h e s u n d e w s (Drosera, Droseraceae) em­ ploy a totally different m e a n s of e n t r a p ­ m e n t . T h e plant consists of a basal rosette of leaves, covered with glands raised o n elongated tentaclelike stalks, each covered with a sticky, extremely viscid mucilage. Unwary insects a r e first s n a r e d by t h e glands; t h e n the stalks a n d the leaves b e n d inward, completely i m p r i s o n i n g t h e prey, which is t h e n digested by secretions within. S u n d e w s grow w h e r e t h e soil is p o o r in n u t r i e n t s , in swamps, in bogs, a n d o n r o t t i n g logs a n d d e c o m p o s i n g vegetation. B l a d d e r w o r t s (the w i d e s p r e a d Utricularia a n d C u b a n Biovularia, Lentibulariaceae) are mostly rootless, floating o r epiphytic plants. T h e most c o m m o n species live freely s u s p e n d e d in p o n d s a n d slow water­ courses, often backwater lakes a n d sluggish j u n g l e streams. T h e traps a r e attached to the plant by a stalk a n d a r e b o r n e o n t h e roots a n d have t h e form of a small, flat­ t e n e d , hollow, saclike body. T h e o p e n i n g or d o o r is set with n u m e r o u s g l a n d u l a r hairs. Small aquatic insects, such as mosquito larvae, water fleas, a n d m i d g e larvae, be­ c o m e c a u g h t by these hairs a n d e n g u l f e d by the bladder, in which they a r e finally di­ gested. Some

bromeliads

apparently

have

evolved the insectivorous habit (Frank a n d O'Meara 1984). C o m p e t i t i o n exists between insects a n d o t h e r o r g a n i s m s . S o m e t i m e s this is active, as in t h e case of h u m m i n g b i r d s driving butterflies away from c o m m o n nectar sources ( T h o m a s et al. 1986), b u t m o r e often it is passive, for e x a m p l e , u s u r p a t i o n of nectar sources by Africanized bees lead­ ing to loss of food to o t h e r h o n e y b e e strains a n d native bees (Roubik 1979).

References CANTWELL, G. E., ed. 1974. Insect diseases. Vol. 1. Marcel Dekker, New York. COLLINS, C. T. 1968. The comparative biology of two species of swifts in Trinidad, West Indies. Fla. St. Mus. Bull. 11: 257-320. EVANS, H. C. 1982. Entomogenous fungi in tropical forest ecosystems: An appraisal. Ecol. Entomol. 7: 47-60. FORMANOWICZ, J R . , D. R., M. M. STEWART, K. TOWNSEND, F. H . POUCH, AND P. F.

BRUSSARD. 1981. Predation by giant crab spiders on the Puerto Rican frog Eleutherodactylus coqui. Herpetologica 37: 125-129. FRANK, J. H., AND G. F O'MEARA.

1984. T h e

bromeliad Catopsis berteroniana traps terres­ trial arthropods but harbors Wyeomyia larvae (Diptera: Culicidae). Fla. Entomol. 67: 4 1 8 424. GREENBERG, R., AND J. GRADWOHL. 1980. Leaf

surface specializations of birds and arthro­ pods in a Panamanian forest. Oecologia 46: 115-124. HESPENHEIDE, H. A. 1975. Selective predation by two swifts and a swallow in Central Amer­ ica. Ibis 117: 82-99. HILL, J. E., AND J. D. SMITH.

1984. Bats, a

natural history. Univ. Texas Press, Austin. HUNT, J. H. 1983. Foraging and morphology in ants: T h e role of vertebrate predators as agents of natural selection. In P. Jaisson, Social insects in the tropics. 1st Int. Symp., Int. Union Stud. Soc. Ins. and Soc. Mexicana Entomol. (Cocoyoc 1980) Proc. Pp. 83-104. LESCURE, J., AND J. FRETEY. 1977. Alimentation

du lézard Anolis rnarmoratus speciosus Garman (Iguanidae) en Guyane francaise. Mus. Nat. Hist. Natur. (Paris) Bull. (Ecol. Gen.) 35: 4 5 52. LIEBERMAN, S. S. 1986. Ecology of the leaf litter herpetofauna of a Neotropical rain forest: La Selva, Costa Rica. Acta Zool. Mexicana (n.s.) 15: 1-72.

LLOYD, F. E. 1931. T h e carnivorous plants. Chronica Botánica, Waltham, Mass. LUBIN, Y. D., G. G. MONTGOMERY, AND O. P.

YOUNG. 1977. Food resources of anteaters (Edentata: Myrmecophagidae). I. A year's census of arboreal nest of ants and termites on Barro Colorado Island, Panama Canal Zone. Biotropica 9: 26-34. MADELIN, M. F. 1966. Fungal parasites of in­ sects. Ann. Rev. Entomol. 11: 423-448. NICKLE, W. R., AND H. E. WELCH. 1984. History,

development, and importance of insect nematology. In W. R. Nickle, ed., Plant and insect nematodes. Marcel Dekker, New York. Pp. 627-653. ROBERTS, D. W., AND J. M. CASTILLO.

1980.

Bibliography on pathogens of medically im­ portant arthropods: 1980. World Health Org. Bull. 50 (suppl.): 1-197. ROUBIK, D. W. 1979. Africanized honeybees, stingless bees and the structure of tropical plant-pollinator communities. Proc. 4th Int. Symp. Pollination, Maryland Agrie. Exper. Sta., Spec. Misc. Publ. 2: 403-417. SEIJAS, A. E., AND S. RAMOS.

1980. Caracte­

rísticas de la dieta de la baba (Caiman crocodilus) durante la estación seca en las sabanas moduladas del Estado Apure, Venezuela. Acta Biol. Venezolana 10: 373-389. SHERRY, T. W. 1984. Comparative dietary ecol­ ogy of sympatric, insectivorous Neotropical flycatchers (Tyrannidae). Ecol. Monogr. 54: 313-338. STEINHAUS, E. A., ed. 1963. Insect pathology. 2 vols. Academic, New York. THOMAS, C. D., P. M. LACKIE, M. J. BRISCO, AND

D. N. HEPPER. 1986. Interactions between hummingbirds and butterflies at a Hamelia patens bush. Biotropica 18: 161 — 165. TOFT, C. A. 1981. Feeding ecology of Panaman­ ian litter anurans: Patterns in diet and forag­ ing mode. J. Herpetol. 15: 139-144.

PROTECTION FROM ENEMIES Insects employ a vast a r r a y of direct d e ­ vices to protect t h e m from their e n e m i e s . T h e most elemental is flight t o escape danger. Species with wings use t h e m often for this p u r p o s e , some with incredible swiftness a n d agility, such as s k i p p e r butter­ flies. O t h e r s may r u n to safety, like g r o u n d beetles that scurry into soil cracks o r b u r -

PROTECTION FROM ENEMIES

77

row a m o n g g r o u n d litter w h e n t h r e a t e n e d . Insects resting o n leaves o r tree t r u n k s often d r o p to t h e g r o u n d (weevils), j u m p o r h o p (leafhoppers), o r s n a p (click bee­ tles) from t h e i r p e r c h e s to d i s a p p e a r a m o n g debris o r vegetation o n t h e soil surface. T h e r e they r e m a i n motionless, playing d e a d (catalepsy) until t h e t h r e a t passes. L a r g e size a n d a n a r m o r e d , often s p i n e d , b o d y is a n o t h e r way m a n y insects, especially s o m e of t h e gigantic h o r n e d scarabs (Dynastes, Megasoma), avoid h a r m . T h e y a r e simply too m u c h of a m o u t h f u l for insectivorous p r e d a t o r s . A g r e a t m a n y insects a r e biochemically u n p a l a t a b l e , possessing substances in their blood a n d tissues r e n d e r i n g t h e m bitter o r otherwise o b n o x i o u s to t h e palates of p r e d ­ ators. T h e most c o m m o n c o m p o u n d s in this category a r e c a r d e n o l i d e s , t e r p e n o i d s , alkaloids, a n d a m i n e s , which in m a n y cases, tint t h e b o d y fluids yellow (leaf beetles, fireflies). S o m e of these com­ p o u n d s a r e p r o d u c e d by t h e insects' own metabolic processes, b u t m a n y a r e seques­ tered from t h e plants o n which insects feed a n d a r e s t o r e d in b o d y tissues a n d blood (Rothschild 1973). Such chemicals a r e also ejected from t h e body as sprays (winged p h a s m i d s , milli­ pedes), injected in stings o r bites (vespoid wasps), secreted o n t o t h e b o d y surface (wax bugs), o r e x p o s e d in o t h e r ways to h u r t , frighten, o r disgust e n e m i e s . T h e purely physical effects of bites can also d i s c o u r a g e , as a n y o n e c a n attest w h o has felt t h e closing j a w s of a l a r g e l o n g - h o r n e d beetle o r giant l u b b e r g r a s s h o p p e r . Highly c o m p l e x d e s i g n s of color a n d form, as well as behavior, p r o v i d e t h e insect with a r e s e m b l a n c e to s o m e object in its e n v i r o n m e n t which e i t h e r possesses o n e of its o w n direct protective m e c h a n i s m s o r is s o m e h o w u n i n t e r e s t i n g to a potential p r e d a t o r ( B e r n a r d i 1985). Such d e c e p t i o n s a r e e x t r e m e l y well d e v e l o p e d a m o n g in­ sects, especially in t h e rich Neotropical

78

ECOLOGY

e n t o m o f a u n a , a n d a r e e x a m p l e s of t h e well-known p h e n o m e n o n of mimicry (Pas­ t e u r 1982, R e t t e n m e y e r 1970), directed toward t h e visual sense of v e r t e b r a t e p r e d a ­ tors (Robinson 1982). T h e c o n c e p t of m i m ­ icry historically originated in classical stud­ ies o n Neotropical butterflies by H e n r y Walter Bates (1862) (Stearn 1981), Alfred Russell Wallace (1870), a n d Fritz Müller (1879) a n d is still studied actively in this g r o u p of insects (Williamson a n d Nelson 1972). All mimetic systems have t h r e e c o m p o ­ n e n t s : a m o d e l , o r object having s o m e direct protective capability; a mimic, t h e species obtaining protection by r e s e m b l i n g the m o d e l ; a n d a d u p e , t h e o p e r a t o r or e n e m y that is fooled by t h e r e s e m b l a n c e ( T u r n e r 1977). Gaudy, noticeable colors a n d p a t t e r n s often p r o v i d e ways of e n h a n c ­ ing t h e efficacy of these direct protective devices a n d of otherwise deceiving o r con­ fusing e n e m i e s (protective coloration). Bright color a n d / o r c o n s p i c u o u s m a r k i n g s can advertise unpalatability, r e m i n d i n g a p r e d a t o r of a past u n p l e a s a n t e x p e r i e n c e a n d k e e p i n g it from a d v a n c i n g toward o r injuring t h e insect even with an explor­ atory bite (Rothschild 1973). T h e bright r e d s a n d yellows f o u n d o n tiger m o t h s a n d l u b b e r g r a s s h o p p e r s a r e for this p u r p o s e . Designs may be startling, such as t h e bright bars o r dazzlingly bright color fields o n the u p p e r wing surfaces of m a n y butterflies (e.g., Morpho). T h e s e confuse the senses of a p u r s u e r , which sees c o n s p i c u o u s color in contrast to inconspicuous a n d fails to recog­ nize o r to be able to follow t h e m o v e m e n t s of t h e insect. Two distinct types of mimicry a r e recog­ nized, o n e in which t h e m o d e l is some i n n o c u o u s , i n a n i m a t e object like a stone, a leaf, a twig, o r wood. T h i s is crypsis a n d is exemplified by the walkingstick's similarity to a piece of wood o r t h e katydid's imita­ tion of a leaf. W h e n t h e m o d e l is an animal actively avoided by t h e p r e d a t o r as noxious, unpal-

atable, o r d a n g e r o u s , o r even j u s t difficult to catch ( H e s p e n h e i d e 1973), t r u e mimicry is in effect, a large n u m b e r of variations of which a r e possible a n d a r e practiced by insects in n a t u r e (Vane-Wright 1976). Nu­ m e r o u s similarly a p p e a r i n g species exhibit a c o m m o n d e f e n s e signal (pattern) that advertises t h e i r universal unacceptability. This is t h e classic situation r e f e r r e d to as Miillerian mimicry. M o r e simply, o n e o r m o r e edible mimics m a y simulate a n o t h e r organism in s o m e way repellent to t h e d u p e , so escaping attention, an a r r a n g e ­ ment k n o w n as Batesian mimicry. Rarely, the d u p e m a y itself also be t h e m o d e l ( H o g u e 1984: 149).

practice simple cryptic r e s e m b l a n c e of leaves o r sticks. It is i m p o r t a n t to recognize also that mimetic similarities e x t e n d to b e h a v i o r a n d s t r u c t u r e as well as color a n d p a t t e r n . T h e attine j u m p i n g spiders (Salticidae, Attinae) would be only p o o r a n t simulators by their d a r k colors alone, if their attitudes (fore­ legs held forward like a n t e n n a e ) a n d m o r ­ phology ( n a r r o w e d base of a b d o m e n ) d i d not also follow their model's physical a p ­ p e a r a n c e a n d c o m p o r t m e n t . A single spe­ cies usually relies o n a variety of protective abilities, not only o n e (Kettlewell 1959).

Mimicry is a n i m p o r t a n t a r e a of study in the Neotropics w h e r e spectacular e x a m ­ ples a b o u n d (e.g., Chai 1988, Poole 1970, Young 1971). T h e best-known Miillerian mimicry series (pi. 3f) a r e those involving c o m m o n p a t t e r n s a m o n g various l e p i d o p teran families (see Butterflies, c h a p . 10). In some instances, it is n o t yet clear w h e t h e r a mimetic g r o u p is Miillerian o r Batesian, as in those D í p t e r a (Blephariceridae, Bibionidae, T i p u l i d a e ) a n d L e p i d o p t e r a (Zygaenidae, Pyralidae, C t e n u c h i n a e ) with a black body a n d legs, d a r k e n e d wing m e m b r a n e , and golden yellow s c u t u m ( H o g u e 1981). Research includes e x p e r i m e n t s to m e a s u r e the actual survival value of p r e s u m e d mim­ icry systems ( B r o w e r et al. 1963, 1967).

BATES, H. W. 1862. Contributions to an insect fauna of the Amazon valley, Lepidoptera: Heliconidae. Linnean Soc. London Trans. 23: 495-566, PI. LV-LV1. BERNARDI, G., ed. 1985. Camouflage et mimetisme. Soc. Entomol. France Bull. 90: 10041103.

Mimicry m a y h a v e aggressive as well as purely defensive p u r p o s e s . H e r e the mimic simulates a n o r g a n i s m with which the d u p e n o r m a l l y interacts. T h e m o d e l takes a d v a n t a g e of t h e d u p e to p r e y o n o r parasitize it. E x a m p l e s a r e female Photuris fireflies that l u r e males of different species to their d e a t h by s e n d i n g signal flashes like those of their prey species females (see fireflies, c h a p . 9). T h e type of mimicry e m p l o y e d by a species may vary a c c o r d i n g to t h e life stage involved ( d e v e l o p m e n t a l mimicry). T h u s , •he n y m p h s of certain m a n t i d s may resem­ ble stinging ants, while t h e adults may

References

BROWER, L. P., J.

V. Z. BROWER, AND C.

T.

COLLINS. 1963. Experimental studies of mim­ icry, 7. Relative palatability and Miillerian mimicry among Neotropical butterflies of the subfamily Heliconiinae. Zoológica 48: 65-84. BROWER, L. P., L. M. COOK, AND H. J. CROZE.

1967. Predator responses to artificial Bates­ ian mimics released in a Neotropical environ­ ment. Evolution 21: 11-23. CHAI, R 1988. Wing coloration of free-flying Neotropical butterflies as a signal learned by a specialized avian predator. Biotropica 20: 20-30. HESPENHEIDE, H. A. 1973. A novel mimicry complex: Beetles and flies. J. Entomol. 48: 49-56. HOGUE, C. L. 1981. Blephariceridae. In S. H.

Hurlbert, G. Rodriguez, and N. Dias de Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 285-287. HOGUE, C. L. 1984. Observations on the plant hosts and possible mimicry models of "lan­ tern bugs" (Fulgora spp.) (Homoptera: Fulgoridae). Rev. Biol. Trop. 32: 145-150. KETTLEWELL, H. B. D. 1959. Brazilian insect adaptations. Endeavour 18: 200-210. MÜLLER, F. 1879. Ituna and Thyridia; a remark­ able case of mimicry in butterflies. Translated by R. Meldola. Entomol. Soc. London Proc. 1879: 20-29.

PROTECTION FROM ENEMIES

79

PASTEUR, G. 1982. A classificatory review of mimicry systems. Ann. Rev. Ecol. Syst. 13: 169-199. POOLE, R. W. 1970. Habitat preferences of some species of a Müllerian-mimicry complex in northern Venezuela, and their effects on evolution of mimic wing pattern. New York Entomol. Soc.J. 78: 121-129. RETTENMEYER, C. W. 1970. Insect mimicry. Ann. Rev. Entomol. 15: 43-74. ROBINSON, M. H. 1982. Defensa contra depreda­ dores que cazan por medios visuales. In G. A. de Alba and R. W. Rubinoff, eds., Evolución en los trópicos. Smithsonian Trop. Res. Inst., Panama. Pp. 57-76. ROTHSCHILD, M. 1973. Secondary plant sub­ stances and warning coloration in insects. Royal Entomol. Soc. London Symp. 6: 5 9 - 8 3 . STEARN, W. T. 1981. Henry Walter Bates (18251892), discoverer of Batesian mimicry. Biol. J. Linnean Soc. 6: 5—7. TURNER, J. R. G. 1977. Mimicry: A study in behaviour, genetics, ecology and biochemis­ try. Ann. Rev. Ecol. Syst. 13: 169-199. VANE-WRIGHT, R. I. 1976. A unified classifica­ tion of mimetic resemblances. Biol. J. Lin­ nean Soc. 8:25—56. WALLACE, A. R. 1870. Mimicry, and other protective resemblances among animals. In A. R. Wallace, Contributions to the theory of natural selection: A series of essays. Macmillan, London. Pp. 45-129. WILLIAMSON, G. B., AND C. E. NELSON.

1972.

Fitness set analysis of mimetic adaptive strate­ gies. Amer. Nat. 106: 525-537. YOUNG, A. M. 1971. Mimetic associations in natural populations of tropical papilionid butterflies (Lepidoptera: Papilionidae). New York Entomol. Soc.J. 79: 210-224.

SOCIAL INSECTS Evolution has c a r r i e d several g r o u p s of insects to a r e m a r k a b l y h i g h level of g r o u p o r g a n i z a t i o n . T h e s e a r e t h e social insects, the termites, a n t s , a n d certain categories of wasps a n d bees (Jaisson 1983, Richards 1953, Wilson 1971). E a c h r e p r e s e n t s an i n d e p e n d e n t lineage of social d e v e l o p m e n t from different, nonsocial ancestors. T r u e sociality (eusociality) is character­ ized by several essential e l e m e n t s : (1) m e m ­ bers of t h e colony a r e all siblings directly

80

ECOLOGY

related to the f o u n d i n g p a r e n t s , n o t indi­ viduals of diverse p a r e n t a g e t h a t have c o m e to live t o g e t h e r secondarily; (2) m e m ­ bers c o o p e r a t e in the c a r e of y o u n g a n d defense of t h e colony ( H e r m a n n 1984); (3) a division of labor exists b e t w e e n m e m b e r s , often correlated with differences in b o d y form; (4) g e n e r a t i o n s o v e r l a p so that the offspring aid t h e p a r e n t s in the work of the colony; a n d (5) colonies usually m a k e a n d maintain s o m e sort of nest s t r u c t u r e . Ex­ c h a n g e of social m e s s e n g e r chemicals also often occurs (trophallaxis), which acts to bind a n d control g r o u p behavior. Castes are d e t e r m i n e d in several ways, n u t r i t i o n a n d p h e r o m o n e s b e i n g primarily i m p o r ­ tant (Lüscher 1977, O s t e r a n d Wilson 1979, Watson et al. 1985). It is interesting that n o g r o u p of insects with sucking m o u t h p a r t s has evolved sociality; m a n d i ­ bles seem to be necessary in m a n i p u l a t i n g nest material a n d n u r s i n g duties. So-called altruism (kin selection) is p r e s e n t also a m o n g social insects, that is, n o r e p r o d u c ­ tion in favor of a n o t h e r individual, a n d may also involve self-sacrifice in defense of the colony. T h e p a t h to t h e origin of sociality seems to be in increasing interaction of adults with their offspring, coupled with in­ creased longevity of the p a r e n t s (Andersson 1984) a n d acquisition of g r o u p defen­ sive specializations such as the sting (Starr 1985). Several logical stages in the p r o g r e s ­ sion from completely solitary lives to eusocial insects may be recognized, al­ t h o u g h they have not necessarily o c c u r r e d in the history of all g r o u p s ( A l e x a n d e r 1974, H a r o 1982, Lin a n d M i c h e n e r 1972, Michener 1958, Wilson 1975). Simple p a r e n t a l care is the first step. T h e m o t h e r r e m a i n s with a n d protects the y o u n g , sometimes for a considerable time, k e e p i n g t h e m from h a r m a n d sometimes p r o v i d i n g occasional food a n d shelter. This is f o u n d in such d i s p a r a t e g r o u p s as the earwigs, t r e e h o p p e r s , a n d cockroaches. W h e n the m o t h e r constructs nest cham-

bers in which she places provisions of food especially for t h e y o u n g , called mass provi­ sioning, a m o r e c o m p l e x level of interac­ tion has b e e n r e a c h e d , such as that prac­ ticed by t h r e a d - w a i s t e d wasps (Ammophila). This advances to a still h i g h e r level of organization w h e n provisioning is p r o g r e s ­ sive, t h a t is, fresh food m a d e available as d e v e l o p m e n t c o n t i n u e s , t h e habit of sand wasps (Bembix). O t h e r subsocial or semisocial stages a r e d e m o n s t r a t e d by c o m m u ­ nal nesting of female bees or wasps, even by g r o u p s of spiders o c c u p y i n g a single web, a l t h o u g h truly social spiders have n o t been discovered. Because of their s e p a r a t e origins, the various g r o u p s of truly social insects dis­ play some f u n d a m e n t a l differences in biol­ ogy. Termites, because they retain g r a d u a l m e t a m o r p h o s i s , r e q u i r e less care of the young, which a r e mobile n y m p h s quite capable of f e e d i n g themselves. T h e y d o not have a d o r m a n t p u p a l p e r i o d requir­ ing speical p r o t e c t i o n a n d care, a n d they have m o r e flexible control over the redirec­ tion of d e v e l o p m e n t to d e t e r m i n e castes than social insects with c o m p l e t e m e t a m o r ­ phosis. T h e r e p r o d u c t i v e m a l e r e m a i n s with the " q u e e n " m o t h e r , continuously in­ seminating h e r d u r i n g h e r long life, in contrast to his usual early d e m i s e after o n e mating in o t h e r social species. Alate r e p r o ductives leave t h e nest, pair off a n d mate, and f o u n d a n e w colony together. A m o n g the offspring, w o r k e r a n d soldier castes may develop. T h e f o r m e r have n o r m a l chewing m a n d i b l e s . T h e ants a r e a n o t h e r major g r o u p , like termites, to h a v e evolved social behavior. New colonies a r e normally f o u n d e d m u c h like termites, l a r g e s w a r m s of flying males attracting females, b u t the female begins the b r o o d a l o n e , h e r m a t e having died soon after m a t i n g with her. Castes are similar also to those of termites, the work­ ers having u n m o d i f i e d m a n d i b u l a r tools and soldiers having outsized h e a d s a n d jaws used as w e a p o n s for colony defense.

T h e eusocial wasps a r e the h o r n e t s a n d p a p e r wasps, all in the family Vespidae. O n e social g e n u s {Microstigmus) of S p h e c i d a e is also known. I n social wasps a n d bees, t h e w o r k e r caste is little differentiated a n a t o m i ­ cally from the q u e e n , usually only smaller. Behaviorally, however, they are quite dis­ tinct, being sterile a n d responsible p r i m a r ­ ily for the g a t h e r i n g of food, nest construc­ tion, a n d n u r s i n g the larvae. New colonies a r e f o r m e d by splitting of old colonies, some nonsterile female offspring leaving the nest a n d m a t i n g with males of o t h e r colonies. As with ants, the males die, a n d the females build a new h o m e a n d start a family whose m e m b e r s quickly take o n tasks, releasing the m o t h e r for r e p r o d u c ­ tive activities only. She may live several years, b e c o m i n g d o r m a n t d u r i n g cold sea­ sons in t e m p e r a t e areas or ovipositing con­ tinuously in tropical regions. T h e r e a r e several g r o u p s of eusocial b e e s — h o n e y b e e s , stingless bees, b u m b l e ­ bees, a n d certain types of c a r p e n t e r bees a n d sweat bees. T h e y a r e m u c h like social wasps in their biology b u t are never wholly carnivorous. T h e i r food is normally of plant origin, consisting of n e c t a r a n d pol­ len. T h e y are, t h e r e f o r e , m u c h m o r e inti­ mately associated with flowering plants t h a n the wasps, which mainly take animal prey. T h e study of social insects has attracted a g r e a t m a n y entomologists a n d behaviorists. T h e popularity of the subject partly derives from the similarities of insect soci­ eties to the h u m a n condition, a n d m a n y scientists believe that k n o w l e d g e g a i n e d from the workings of the insect may have direct b e a r i n g o n some aspects of o u r o w n lives (Wilson 1975). T h i s is particularly t r u e in the area of c o m m u n i c a t i o n a n d n e u r a l integration. T h e brains of the so­ cial g r o u p s are the largest a n d most com­ plex of insects a n d can serve as simplified models of integrative systems involving primitive l e a r n i n g as well as instinctive activity.

SOCIAL INSECTS

81

References ALEXANDER, R. D. 1974. T h e evolution of social behavior. Ann. Rev. Ecol. Syst. 5: 325-383. ANDERSSON, H. 1984. T h e evolution of eusociality. Ann. Rev. Ecol. Sys. 15: 165-189. HARO, A. DE. 1982. Algunas consideraciones sobre el origen y evolución de las sociedades de insectos. In P. Jaisson, Social insects in the tropics. 1st Int. Symp. (Cocoyoc 1980), Int. Union Stud. Soc. Ins. and Soc. Mexicana Entomol. Proc. 1: 6 5 - 7 1 . HERMANN, H. R., ed. 1984. Defensive mecha­ nisms in social insects. Praeger, New York. JAISSON, P. 1983. Social insects in the tropics. 1st Int. Symp. (Cocoyoc 1980), Int. Union Stud. Soc. Ins. and Soc. Mexicana Entomol. Proc. Vols. 1-2. LIN, N., AND C. D. MICHENER 1972. Evolution

of sociality in insects. Quart. Rev. Biol. 46: 131-159. LÜSCHER, M. 1977. Phase and caste determina­ tion in insects. Pergamon, Oxford. MICHENER, C. D. 1958. T h e evolution of social behavior in bees. 10th Int. Cong. Entomol. (Montreal) Proc. 2: 441-447. OSTER, G. F., AND E. O. WILSON. 1979. Caste

and ecology in social insects. Princeton Univ. Press, Princeton. RICHARDS, O. W. 1953. T h e social insects. Macdonald, London. STARR, C. K. 1985. Enabling mechanisms in the origin of sociality in the Hymenoptera—The sting's the thing. Entomol. Soc. Amer. Ann. 78: 836-840. WATSON, J. A. L., B. M. OKOT-KOTBER, AND

C. NOIROT, eds. 1985. Caste differentiation in social insects. Pergamon, Oxford. WILSON, E. O. 1971. T h e insect societies. Har­ vard Univ. Press, Cambridge. WILSON, E. O. 1975. Sociobiology. Harvard Univ. Press, Cambridge.

SYMBIOSIS Two o r m o r e kinds of o r g a n i s m s living in close association to t h e benefit of o n e (commensalism) o r all p a r t n e r s (mutualism) ( B o u c h e r 1982) a r e said to exhibit symbio­ sis. ("Symbiosis" is u s e d in t h e b r o a d sense h e r e to i n c l u d e all f o r m s of intimate asso­ ciations, n o t j u s t t h o s e of m u t u a l benefit for which t h e t e r m is u s e d in a n a r r o w sense.) T h e N e o t r o p i c s p r o v i d e countless,

82

ECOLOGY

fascinating e x a m p l e s of this p h e n o m e n o n involving insects. S o m e of t h e best-known e x a m p l e s occur between insects a n d h i g h e r plants (Bernays 1989). Most often, t h e f o r m e r a r e ants, a n d major e x a m p l e s a r e discussed in c h a p t e r 12, o n H y m e n o p t e r a (see ants a n d plants). Many symbiotic relationships exist between insects a n d lower plants, especially fungi (Lichtwardt 1986, W h e e l e r a n d Blackwell 1984). Nest s h a r i n g , o r inquilinism, is a type of symbiosis wherein both p a r t n e r s a r e in­ sects (Kistner 1982). E x a m p l e s a r e certain silverfish, millipedes, beetles, mites, a n d cockroaches, called m y r m e c o p h i l e s , that s h a r e a c o m m o n dwelling with ants. O t h e r social insects a r e likewise visited by termitophiles, melittophiles, a n d sphecophiles, de­ p e n d i n g o n w h e t h e r they a r e termites, bees, o r wasps, respectively. T h e r e a r e m a n y variations in t h e d e g r e e of closeness in t h e association a n d its specific n a t u r e (Kistner 1979). I n t h e Neotropics, army ants a n d leaf c u t t e r ants a r e f r e q u e n t hosts (see c h a p . 12). T h e visitors live o n the debris, scraps of food, a n d even corpses of ants. T h i s is mutualistic coexistence, t h e ants profiting by t h e nest debris removal, the guests by a reliable food source.

A related p h e n o m e n o n is exhibited by ants a n d o t h e r insects t h a t h a v e a particu­ lar f o n d n e s s f o r t h e h o n e y d e w secretions p r o d u c e d by several g r o u p s of sap-sucking insects. H o n e y d e w is a sugar-rich solution excreted from t h e digestive tract (aphids) or from special i n t e g u m e n t a r y glands (scale insects, m e t a l m a r k a n d hairstreak butterfly larvae). T h e solution is a food in r e t u r n for which t h e h o n e y d e w p r o d u c e r receives protective, dispersal, cleaning, and o t h e r t e n d i n g favors from t h e feeding insects. (See wax b u g s , a p h i d s , c h a p . 8; metalmarks, hairstreaks, c h a p . 10.) Phoresy is yet a n o t h e r symbiotic relation­ ship b e t w e e n insects. T h e t e r m is used for the t e m p o r a r y a t t a c h m e n t of a m u c h smaller, relatively s e d e n t a r y form to a much larger, m o r e vagile form. T h e former receives t r a n s p o r t a t i o n a n d is able to disperse far m o r e widely t h a n it could on its o w n . M a n y o f t h e mites often f o u n d living o n beetles a r e practicing this habit. This is also t h e case with t h e p s e u d o scorpions so c o m m o n u n d e r t h e elytra of large l o n g - h o r n e d beetles, especially t h e harlequin beetle (see h a r l e q u i n beetle, chap. 9). P h o r e s y is often a m e a n s for parasites a n d p r e d a t o r s to find access to their prey (Clausen 1976).

T h e invasive species finds its way into the host's colony a n d is a c c e p t e d into its society by a variety of physical a n d chemical decep­ tions (Hólldobler 1971). Staphylinid beetles may actually resemble a n d b e h a v e like their hosts ( W a s m a n n i a n mimicry) (Akre a n d R e t t e n m e y e r 1966). T h e t r a i l - m a r k i n g sub­ stances secreted by t h e ants m a y serve as attractants to inquilines (Moser 1964). T h e latter may also p r o d u c e "tranquilizing" chemicals that pacify t h e ants o r mimic the food offerings of w o r k e r ants.

T h e above e x a m p l e s a r e d e s i g n a t e d "ectosymbiosis" (Hartzill 1967) in contrast to "endosymbiosis" (Koch 1967). T h e latter refers to t h e habitation of m a n y microor­ ganisms in t h e g u t , o t h e r b o d y cavities, a n d tissues of insects (Boush a n d C o p p e l 1974). T h e best-known cases a r e t h e flagellate protozoans of primitive termites a n d trichomycete fungi associated with n u m e r o u s insects (Lichtwardt 1986).

T h e strategies of inquilines vary; some prey directly on t h e host, o t h e r s o n the i m m a t u r e s of t h e latter o r o n accumulated nest refuse. O t h e r s a r e a p p a r e n t l y totally n e u t r a l , t h e p u r p o s e s of their invasions remaining unknown.

References AKRE, R. D., AND C. W. RETTENMEYER.

1966.

Behavior of Staphylinidae associated with army ants (Formicidae: Ecitonini). Kans. Entorno!. Soc. J. 39: 745-796. BERNAYS, E. A., ed. 1989. Insect-plant interac­ tions. Vol. I. CRC, Boca Raton, Fla.

BOUCHER, D. H. 1982. T h e ecology of mu­

tualism. Ann. Rev. Ecol. Syst. 13: 315-347. BOUSH,

G.

M.,

AND H.

C.

COPPEL.

1974.

Symbiology: Mutualism between arthropods and microorganisms. In G. E Cantwell, ed., Insect diseases. 1: 301-326. Marcel Dekker, New York. CLAUSEN, C. P. 1976. Phoresy among entomophagous insects. Ann. Rev. Entomol. 2 1 : 343-368. HARTZILL, A. 1967. Insect ectosymbiosis. In S. M. Henry, ed., Symbiosis. 2: 107-140. Academic, New York. HÓLLDOBLER, B. 1971. Communication between ants and their guests. Sci. Amer. 224(3): 8 6 93. KISTNER, D. H. 1979. Social and evolutionary significance of social insect symbionts. In H. R. Hermann, ed., Social insects. 1:339413. Academic, New York. KISTNER, D. H. 1982. T h e social insects bestiary. In H. R. Hermann, ed., Social insects. 3: 1 24. Academic, New York. KOCH, A. 1967. Insects and their endosymbionts. In S. M. Henry, ed.. Symbiosis. 2: 1-10. Academic, New York. LICHTWARDT, R. W. 1986. T h e Trichomycetes

(fungal associates of arthropoda). Springer, Berlin. MOSER, J. C. 1964. Inquiline roach responds to trail-making substance of leaf-cutting ants. Science 143: 1048-1049. WHEELER, Q., AND M. BLACKWELL, eds.

1984.

Fungus-insect relationships: Perspectives in ecology and evolution. Columbia Univ. Press, New York.

POLLINATION T h e majority of flowering plants a r e d e ­ p e n d e n t on insects as vehicles f o r t h e transfer of their pollen from a n t h e r to stigma (Meeuse a n d Morris 1984, Faegri a n d Van d e r Pijl 1979, Real 1983, Richards 1978). T h e flower itself, by color a n d o d o r attractants (Yeo 1973) a n d nectar a n d pol­ len r e w a r d s , is t h e p r i m e m e c h a n i s m for involving insects in pollination (Bacior 1974). Plants also may entice insects to visit a n d pollinate t h e m by p r o v i d i n g wax a n d resins o r false r e w a r d s such as t h e p r o m i s e of sex or food, t h e latter usually in t h e form of carrion o r feces.

POLLINATION

83

Many insects, in turn, require plants, their leaves, stems, wood, and so on, or floral products to sustain their lives and have a stake in pollination equal to that of the plant. Fruit setting in figs (Ficus), for example, is entirely dependent on minute wasps whose whole lives are likewise depen­ dent on the tree (see fig wasps, chap. 12). Although grasses are generally regarded as wind-pollinated plants, in the still atmo­ sphere of tropical forests, insects may still be necessary to effect pollen transfer (Soderstrom and Calderón 1971). Pollination biology of insects and plants in the tropical portions of Latin America is an active field of study (Heinrich and Raven 1972, Jones and Little 1983). Most work is concerned with the mechanisms of pollen acquisition and deposition (see ex­ amples below); few deal with the important topics of transport during foraging activity and with interflower relationships (Frankie and Baker 1974). Pollination of trees in isolated remnants of tropical forests has also been studied (Raw 1989). Pollinating insects are of two general types: (1) large, powerful, long-distance fliers, including sphinx moths (Cruden et al. 1976, Haber and Frankie 1989, Linhart and Mendenhall 1977), large bees (Sazima and Sazima 1989), and butterflies, and (2) comparatively weak-flying, short-distance fliers, such as small flower and bee flies (Syrphidae and Bombyliidae), moth flies (Psychodidae), punkies (Ceratopogonidae), chalcidoid wasps, minute bees, and some beetles (Meloidae, Oedemeridae, Scarabaeidae). In addition to flight, many have elon­ gate, sucking mouthparts and possess spe­ cialized anatomical structures for carrying or storing pollen and nectar. Most also have good color vision for locating day-blooming flowers and excellent olfactory senses to find plants with nocturnal blossoms. Flowers assume a variety of shapes and special adaptations to enhance or ensure visits and their pollination by coadapted

84

ECOLOGY

insects. Many, for example, have spots or color marking near the nectaries ("nectar guides") to direct the insect's attentions toward them. These may be visible only to pollinators with ultraviolet vision, as many insects have. Other flowers have very long corollas (such as Posoqueria, Calliandra) with deep nectaries that only the long tongues of sphinx moths can reach. Some plants produce flowers without nectaries but which so resemble others that do give nectar reward that they are visited by pollinators nevertheless, a form of floral mimicry (Haber 1984). Some woody tropical forest plants prac­ tice "mass flowering." Individuals bloom profusely and synchronously during a short period (often during the dry season) and attract large numbers and diverse kinds of insect pollinators, especially bees. These insects tend to range freely between individuals of such plants, ensuring crosspollination. Gentry (1978) suggested that insects may be induced to move between plants by insectivorous birds chasing them. Other trees are slightly asynchronous bloomers, having different individuals with short overlapping flowering periods. This is thought to be a strategy to foster pollinator movement between trees (Perry 1980). Many orchid flowers exhibit floral adap­ tations for pollination to an extreme de­ gree (Darwin 1890, Van der Pijl and Dodson 1966, Williams 1982). The lip petal of the Colombian orchid Masdevallia bella has a gill-like, fleshy appearance exactly like the underside of a mushroom and is polli­ nated by fungus gnats. The latter ostensi­ bly visit the flower as they would in search of an oviposition site in their normal mush­ room hosts. Several orchid species attract male euglossine bees, providing them with oily, fragrant substances used in the bee's courtship, in return for securing their services as pollinators (see orchid bees, chap. 12). Coryanthes, or "bucket orchid" flowers,

all found in tropical Central and South America, are complexly formed to ensure pollination by bees. The lower part is a cuplike container that holds a sweet fluid accumulated from fluid-producing glands above on the tip of the flower's stalk. Male euglossine bees visit the flowers to collect substances from the petals and in doing so often lose their footing and fall into the wet, sticky pool below. Unable to use their wings and fly out, they are forced to escape the bucket by crawling through an opening in its side, in one wall of which are the pollen packets. Because of the tight squeeze, the bee brushes past the packets, which detach and adhere to it. The next flower visited receives the packets, which are scraped off onto the stigma and polli­ nate it. Scents emitted by flowers are not always sweet. Many members of the orchid subtribe Pleurothallideae (Orchidaceae) and the genera Dracontium (Araceae), Sterculia, and Herrania (Sterculiaceae) at­ tract carrion flies with putrescent odors. A curious collection of plants utilize "trap flowers" to ensure pollination. The principle is to detain the pollinator for a short time, to ensure that it will pick up and deposit pollen. It is released hours or days later to repeat the process in the flowers of other individuals of the same species. The South American bladder-flower plant (Araujia sericofera) is an example. Moths may visit their first bladder flower without being detained, merely picking up pollen packets (pollinia) on their tongues and carrying them off. When the moth feeds from a second flower, these pollinia become wedged into a slitlike structure on the petals where they come in contact with the receptive female organs. Larger, stronger moths, like sphingids, can pull away from this snare, but smaller, weaker species remain caught permanently. Phenylacetaldehyde emitted from the flowers is the active chemical agent that these plants

use initially to attract the moths (Cántelo andjacobson 1979). Another trap flower is that of the aristolochia vine (Aristolochia spp.). The complex chambered blossom attracts in­ sects with its bright outer lip and carrionmimicking odors. Visiting insects push their way down the dark tubular part into the base of the flower, or "prison," where the receptive young stigma is located. Their advance is prompted by a lighter colored or semitransparent area in the chamber, and their exit is prevented by backward-pointing hairs in the narrowed funnel region. By this mechanism they are trapped for some days until the stamens mature and shed their pollen, which dusts the insects. Then the hairs wither and the insects can leave, to enter another young flower and go through the trial again but pollinating the young stigma with the grains they have picked up on their previ­ ous captive experience. The flower buds of the giant Amazon water lily (Victoria amazónica) open at night and emit a strong fruity odor that attracts scarab beetles of the genus Cyclocephala (Lovejoy 1978; pi. 3h). They close by the next morning, trapping the beetles in a deep chamber inside for a day where they feed on sterile inner anthers, depositing pollen on the style. They are released in the afternoon when the flowers open again, picking up pollen from the outer ripe anthers. Cyclocephala also pollinate Cyclanthus (Cyclanthaceae) in a similar manner: they spend twenty-four hours in the flower spathe but are not trapped (Beach 1982). Cyclocephala are known to be pollinators of aroid's, such as Dieffenbachia (H. Young 1986), and beetles generally are important in reproduction of a variety of Neotropical plant families. Flowering in many beetlepollinated plants is accompanied by an increase in metabolic rate that leads to the production of heat. This causes the volatil­ ization of various odors from the flower that attract the beetles (Meeuse 1975).

POLLINATION

85

Insect pollination is very i m p o r t a n t in a g r i c u l t u r e (Free 1970). T o set fruit a d e ­ quately for a profitable c r o p , m a n y culti­ vated plants r e q u i r e pollination by insects, whose p r e s e n c e in p l a n t a t i o n s is e n c o u r ­ aged by g r o w e r s . H o n e y b e e s have long b e e n r e c o g n i z e d a s valuable i n this way, a n d hives a r e p u r p o s e l y placed in fields a n d o r c h a r d s t o increase seed a n d fruit set (Martin a n d M c G r e g o r 1973). T h e cacao tree is pollinated chiefly by p u n k i e flies of t h e g e n u s Forcipomyia (A. Young 1986; see p u n k i e s , c h a p . 11) a n d adult gall m i d g e s (A. Y o u n g 1985). Be­ cause m a n y of these a r e b r o m e l i a d tank b r e e d e r s , t h e p r o x i m i t y of these plants is essential to successful cacao fruiting (Privat 1979). A l t h o u g h c o m m o n o n t h e plants, ants a n d H o m o p t e r a a n d o t h e r insects a r e probably n o t i m p o r t a n t ( W i n d e r 1978). T h e i n t r o d u c e d oil p a l m (Elaeis guineensis) has b e e n found to d e p e n d o n certain fruit beetles (Nitidulidae—Mystrops, Haptoncus) for pollination. F o r m e r l y t h o u g h t to be pestiferous, they a r e n o w welcomed in areas w h e r e this valuable tree is cultivated (Genty et al. 1986). Nitidulid a n d scarab beetles, weevils, a n d o t h e r insect pollinators are equally i m p o r t a n t to native palms (Barfod et al. 1987, Beach 1984), which a r e a s s u m i n g u s e as oil p r o d u c e r s in some areas. T h e Brazil n u t tree (Bertholletia excelsa) a p p a r e n t l y d e p e n d s o n euglossine bees for pollination. O n l y large species a r e capable of u n c u r l i n g t h e floral a n d r o e c i u m (protec­ tive h o o d a r o u n d t h e a n t h e r s ) (Nelson et al. 1985).

References ARMBRUSTER, W. S., AND G. L. WEBSTER 1979.

1987. A note on the pollination of Phytelephas microcarpa (Palmae). Biotropica 19: 191 — 192. BEACH, J. H. 1982. Beetle pollination of Cyclanthus bipartitus (Cyclanthaceae). Amer. J. Bot. 69: 1074-1081. BEACH, J. H. 1984. The reproductive biology of the peach of "Pejibayé" palm (Bactris gasipaes) and a wild cogener (B. porschiana) in the Atlantic lowlands of Gosta Rica. Principes 28: 107-119. CÁNTELO,

W. W., AND M. JACOBSON.

1979.

Phenylacetaldehyde attracts moths to bladder flower and to blacklight traps. Envir. Entomol. 8: 444-447. CRUDEN, R. W., S. KINSMAN, R. E. STOCKHOUSE

II, AND Y. B. LINHART.

1976. Pollination,

fecundity, and the distribution of mothflowered plants. Biotropica 8: 204-210. DARWIN, C. 1890. The various contrivances by which orchids are fertilized by insects. 2d ed. John Murray, London. FAEGRI, K., AND L. VAN DER PIJL. 1979. T h e

principles of pollination ecology. 3d ed. Pergamon, Elmsford, N.Y. FRANKIE, G. W., AND H. G. BAKER. 1974. T h e

importance of pollinator behavior in the reproductive biology of tropical trees. Inst. Biol. Univ. Nat. Autón. México, Ser. Bot., Ann. 45(1): 1-10. FREE, J. B. 1970. Insect pollination of crop plants. Academic, London. GENTRY, A. H. 1978. Anti-pollinators for massflowering plants? Biotropica 10: 668-669. GENTY, P., A. GARZÓN, AND E LUCCHINI. 1986.

Polinización entórnenla de la palma Africana en América tropical. Oléagineaux 4 1 : 99— 112. HABER, W. A. 1984. Pollination by deceit in a mass-flowering tropical tree Plumería rubra L. (Apocynaceae). Biotropica 16: 269-275. HABER, W. A., AND G. W. FRANKIE.

1989. A

tropical hawkmoth community: Costa Rican dry forest Sphingidae. Biotropica 21: 155— 172. HEINRICH, B., AND P. H. RAVEN. 1972. Ener­

getics and pollination ecology. Science 176: 597-602. JONES, C. E., AND R. T

LITTLE,

eds. 1983.

Handbook of experimental pollination biol­ ogy. Van Nostrand Reinhold, New York.

Pollination of two species of Dalechampia (Euphorbiaceae) in Mexico by euglossine bees. Biotropica 11: 278-283. BACIOR, L. W. 1974. Behavioral aspects of coadaptations between flowers and insect pollinators. Missouri Bot. Garden Ann. 61: 760-769.

Pollen dispersal by hawk moths in a Lindenia rivalis Benth. population in Belize. Biotropica 9: 143. LOVEJOY, T. E. 1978. Royal water lilies: Truly Amazonian. Smithsonian 7: 77-82.

BARFOD, A., A. HENDERSON, AND H. BALSLEV.

MARTIN, E. C , AND S. E. MCGREGOR.

86

ECOLOGY

LINHART, Y. B., AND J. A. MENDENHALL. 1977.

1973.

Changing trends in insect pollination of com­ mercial crops. Ann. Rev. Entomol. 18: 2 0 7 226. MEEUSE, B. J. D. 1975. Thermogenic respira­ tion in aroids. Ann. Rev. Plant. Physiol. 26: 117-126.

cocoa tree flowering, fruit set, and pollinator availability in Costa Rica. J. Trop. Ecol. 2: 163-186. YOUNG, H. J. 1986. Beetle pollination of Dieffenbachia longispatha (Araceae). Amer. J. Bot. 73: 931-944.

MEEUSE, B., AND S. MORRIS. 1984. T h e sex life

of flowers. Facts on File, New York. NELSON, B. W., M. L. ABSY, E. M. BARBOSA, AND

G. T. PRANCE. 1985. Observations on flower visitors to Bertholletia excelsa H.B.K. and Couratari tenuicarpa A.C. Sm. (Lecythidaceae). Acta Amazónica Suppl. 15: 225-234. PERRY, D. R. 1980. T h e pollination ecology and blooming strategy of a Neotropical emergent tree Dipteryx panameñas. Biotropica 12: 3 0 7 313. PRIVAT, F. 1979. Les bromeliacees, lieu de developpment de quelques insectes pollinisateurs de fleurs de cacao. Brenesia 16: 197— 212. RAW, A. 1989. T h e dispersal of euglossine bees between isolated patches of eastern Brazilian wet forest. Rev. Brasil. Entomol. 33: 1 0 3 107. REAL, L., ed. 1983. Pollination biology. Aca­ demic, New York. RICHARDS, A. J., ed. 1978. T h e pollination of

flowers by insects. Linnean Soc. Symp. 6: 1 214. SAZINA, I., AND M. SAZIMA. 1989. Mamangavas e

irapuás (Hymenoptera, Apoidea): Visitas, interacáes e conseqüécias para polinizacáo do maracujá (Passifloraceae). Rev. Brasil. Ento­ mol. 33: 109-118. SODERSTROM, T . R . , AND C . E . CALDERÓN. 1 9 7 1 .

Insect pollination in tropical rain grasses. Biotropica 3: 1-16. VAN DER PIJL, L., AND C. H. DODSON.

forest 1966.

Orchid flowers: Their pollination and evolu­ tion. Univ. Miami, Coral Gables. WILLIAMS, N. G. 1982. T h e biology of orchids and euglossine bees. In J. Arditti, ed., Orchid biology. II: 119-171. Cornell Univ. Press, Ithaca. WINDER, J. A. 1978. T h e role of non-dipterous insects in the pollination of cocoa in Brazil. Bull. Entomol. Res. 68: 559-574. YEO, P. F. 1973. Floral allurements for pollinat­ ing insects. Royal Entomol. Soc. London Symp. 6: 151-157. YOUNG, A. M. 1985. Studies of cecidomyiid midges (Diptera: Cecidomyiidae) as cocoa pollinators (Theobroma cacao L.) in Central America. Entomol. Soc. Wash. Proc. 87: 4 9 79. YOUNG, A. M. 1986. Habitat differences in

INSECT CONSERVATION T h e u n n a t u r a l despoliation of wildlife in Latin America, as in most parts of t h e world today, is p r o c e e d i n g at a n alarmingly rapid pace. Habitat destruction is t h e pri­ mary cause of insect extinction t h r o u g h deforestation, erosion, a n d land alteration for agriculture, animal h u s b a n d r y , h y d r o ­ electric e n e r g y a n d r a w material p r o d u c ­ tion, mining, a n d u r b a n i z a t i o n . Insect life suffers also from misplaced insecticides, i m p o r t a t i o n of alien o r g a n i s m s , pollution, fires, a n d direct commercial exploitation (pets a n d decorative uses) (Faria 1940), b u t these a r e far less significant t h a n loss of habitat. T h e r e is some effect from scien­ tific a n d hobby collectors, b u t this is also of little h a r m a n d may actually have benefi­ cial results t h r o u g h advances in t h e knowl­ e d g e gained t h r o u g h these activities. Unfortunately, very little direct effort is being e x p e n d e d to c o u n t e r these negative t r e n d s . Economic d e v e l o p m e n t e v e r y w h e r e takes p r e c e d e n c e over protection of n a t u r e , especially insects, a life form against which t h e r e is almost universal enmity. T h e idea of purposeful c o n c e r n over t h e fate of insects in t h e e n v i r o n m e n t is a new a n d little a p p r e c i a t e d concept, in spite of the fact that insects a r e a form of wildlife ecologically essential to m a n k i n d in so m a n y ways, including their direct value as pollinators, r e d u c e r s of organic matter, m a i n t e n a n c e of e n v i r o n m e n t a l cleanliness, a n d food for o t h e r "desirable" animals (Pyle et al. 1981). T h e signing by most Latin A m e r i c a n countries of t h e C o n v e n t i o n o n I n t e r n a ­ tional T r a d e in E n d a n g e r e d Species of F a u n a a n d Flora (CITES) establishes con-

INSECT CONSERVATION

87

trols o n a n d m o n i t o r s i m p o r t a t i o n a n d e x p o r t of certain species (Fuller a n d Smith 1984); presently, t h e only specific terres­ trial a r t h r o p o d f r o m these c o u n t r i e s which is listed is t h e Mexican r e d - k n e e d t a r a n t u l a (Hemley 1986). Most c o u n t r i e s have laws r e g u l a t i n g t h e collecting of insects for scientific a n d c o m m e r c i a l p u r p o s e s , b u t these have virtually n o effect o n ameliorat­ ing t h e g r e a t e r causes of h a r m . Only Brazil ( O t e r o a n d B r o w n 1986) a n d Mexico have e n a c t e d legislation p r o h i b i t i n g t h e t a k i n g of specific insects: a swallowtail butterfly (Parides ascanius) is given special p r o t e c t i o n in Brazil, a n d t h e m o n a r c h butterfly is p r o t e c t e d in its very localized overwinter­ ing sites in t h e m o u n t a i n s of central Mex­ ico. T h e Invertebrate Red Data Book (Wells et al. 1983) of t h e I n t e r n a t i o n a l Union for t h e C o n s e r v a t i o n of N a t u r e a n d N a t u r a l Re­ sources lists m a n y N e o t r o p i c a l insect species as v u n e r a b l e , r a r e , t h r e a t e n e d , a n d e n d a n g e r e d , b u t n o legal r e g u l a t i o n s yet control their r e m o v a l f r o m t h e e n v i r o n ­ ment. O n e solution to o v e r e x p l o i t a t i o n of s o m e species by c o m m e r c i a l collectors m a y be insect " f a r m i n g " o r " r a n c h i n g , " a tech­ n i q u e f o u n d successful in P a p u a N e w G u i n e a with s o m e o f t h e large bird-wing butterflies (Ornithoptera). A few such e n t e r ­ prises a r e n o w in o p e r a t i o n in Latin A m e r ­ ica (e.g., Brazil; Kesselring p e r s . c o m m . ) . T h e idea h a s b e e n s u g g e s t e d generally ( C r a n e a n d F l e m i n g 1 9 5 3 , National Re­ search Council 1983). Indirectly, c o n s i d e r a b l e p r o g r e s s has b e e n a c c o m p l i s h e d by t h o s e c o u n t r i e s that h a v e set aside habitat as national p a r k s a n d n a t u r e p r e s e r v e s . M a n y o f t h e latter h a v e b e e n m a d e possible by private indi­ viduals, consortia, a n d agencies such as the Xerces Society, T h e N a t u r e C o n ­ servancy, a n d World Wildlife Fund-U.S. I n t e r n a t i o n a l p r o g r a m s also foster conser­ vation m e a s u r e s that h e l p in t h e preserva­ tion a n d c o n s e r v a t i o n o f insects t h r o u g h t h e U.S. Agency for I n t e r n a t i o n a l Devel-

88

ECOLOGY

o p m e n t (USAID), t h e United Nations E d u ­ cational a n d Social C o o p e r a t i o n O r g a n i z a ­ tion ( U N E S C O ) , a n d t h e U.S. Peace Corps. A critical consideration in t h e establish­ m e n t of preserves is t h e "minimal size factor." Little is k n o w n r e g a r d i n g t h e a m o u n t of habitat that n e e d s to b e p r o ­ tected in o r d e r for t h e insect f a u n a to c o n t i n u e t o thrive. A major baseline study directed at this question, a n d including s o m e insects, for A m a z o n i a n forest is t h e cooperative World Wildlife F u n d - U . S . a n d Brazilian National Research I n s t i t u t e proj­ ect n o r t h of M a n a u s o n various-sized, mea­ s u r e d plots of isolated forest ( " M i n i m u m Critical Size of Ecosystems Project," Lovejoy 1980). W h a t t h e f u t u r e holds for t h e insects, a r a c h n i d s , a n d o t h e r similar terrestrial ar­ t h r o p o d s of Latin A m e r i c a is u n f o r e s e e ­ able, b u t it seems realistic t o look f o r w a r d with pessimism. B u t for a very few excep­ tions (Lamas 1974), t h e d i s a p p e a r a n c e of species goes almost entirely u n d o c u ­ m e n t e d . I n t h e forests alone, species a r e probably b e c o m i n g extinct b e f o r e they a r e discovered. T h i s r e d u c t i o n of insect diver­ sity is certainly o c c u r r i n g , however, a n d will accelerate at a r a p i d pace because of deforestation. T h e t r e n d is e x a c e r b a t e d by t h e low p o p u l a t i o n densities of inverte­ brates in this habitat type (Elton 1975). Unquestionably, a g r e a t e r a p p r e c i a t i o n of the benefits of p r o t e c t i n g these life forms is n e e d e d , a n d m o r e action to prevent wholesale ruination of their living space is called for. Laws alone a r e n o t t h e answer. Public awareness a n d u n d e r s t a n d i n g of the issue is critical to all p r o g r e s s t o w a r d ensur­ ing t h e survival of this p a r t of t h e Earth's h e r i t a g e (Lamas 1978).

population density of invertebrates inside Neotropical rain forest. Biol. Conserv. 7: 3 15. FARIA, A. 1940. Ca$a e commercio de borboletas e outros insetos ornamentaes. Rev. Entomol 11:607-608.

NATIONAL RESEARCH COUNCIL (U.S.A). 1983.

FULLER,

OTERO, L. S., AND K. S. BROWN, JR. 1986.

K.

S., AND G.

SMITH.

1984.

Latin

American wildlife trade laws. World Wildlife Fund, Washington, D.C. HEMLEY, G. 1986. Spotlight on the red-kneed tarantula trade. Traffic (U.S.A.) 6(4): 16-17. LAMAS, G. 1974. Supuesta extinción de una mariposa en Lima, Perú (Lepidoptera, Rhopalocera). Rev. Peruana Entomol. 17: 119-120. LAMAS, G. 1978. Mariposas y conservación de la naturaleza en el Perú. Col. Suiza Perú Bol. (July): 6 1 - 6 6 . LOVEJOY, T. E. 1980. Discontinuous wilderness:

Minimum areas for conservation. Parks 5(3): 13-15. Butterfly farming in Papua New Guinea. Managing Animal Resources Series. National Academy Press, Washington, D.C. Biology and ecology of Parides ascanius (Cra­ mer, 1775) (Lep., Papilionidae), a primitive butterfly threatened with extinction. Atala 10-12: 2 - 1 6 . PYLE, R. M., M. BENTZIEN, AND P. OPLER. 1981.

Insect conservation. Ann. Rev. Entomol. 26: 233-258. WELLS, S. M., R. M. PYLE, AND N. M. COLLINS.

1983. T h e IUCN invertebrate red data book. Int. Union Conserv. Nat. Res., Gland, Switzerland.

References CRANE, J., AND H. FLEMING. 1953. Construction

and operation of butterfly insectaries in the tropics. Zoológica 38: 161-172. ELTON, C. S. 1975. Conservation and the low

INSECT CONSERVATION

89

Ó

1962. Destructive and useful insects. 4th ed. McGraw-Hill, New York. PELUFFO, A. T. 1942. Insectos y otros parásitos de la agricultura y sus productos en el Uru­ guay. Univ. Rep., Fac. Agron., Montevideo.

PRACTICAL ENTOMOLOGY

PFADT, R. D., ed. 1978. Fundamentals of ap­

For t h e h u m a n p o p u l a t i o n of Latin A m e r ­ ica, as e l s e w h e r e in t h e world, c o m p e t i t i o n from insects for food a n d fiber a n d inter­ ference with health a n d welfare a r e in­ tense. Insects a r e injurious in several ways. Each is t h e subject of formal fields of study, r e p r e s e n t i n g c o m p a r t m e n t s of t h e vast a n d c o m p l e x subject of "practical" ("economic" or "applied") entomology. Insects a r e h a r m ­ ful by d a m a g i n g g r o w i n g crops a n d o t h e r useful plants (agricultural a n d forest e n t o ­ mology), by a n n o y i n g a n d inflicting dis­ ease o n h u m a n s (medical e n t o m o l o g y ) a n d domesticated animals (veterinary e n t o m o l ­ ogy), a n d by d e s t r o y i n g useful p r o d u c t s (stored p r o d u c t a n d s t r u c t u r a l e n t o m o l ­ ogy). T h e relationships of insects to m a n in p o p u l a t i o n c e n t e r s f o r m a n o t h e r special topic ( u r b a n e n t o m o l o g y ) .

lists a n d cursory reviews o f p r o b l e m species are available for specific areas. T h e s e in­ clude Bolivia (Squire 1972), Colombia (Gallego 1967, Posada 1976), El Salvador (Berry 1959), French G u i a n a (Remillet 1988), G u a t e m a l a (Alvarado 1939), H o n d u ­ ras (Koone a n d Bañegas 1958), U r u g u a y (Peluffo 1942), t h e West Indies (Wolcott 1933), a n d t h e Lesser Antilles (Ballou 1912). Insects cause direct d a m a g e to h u m a n s t h r o u g h their feeding o r by biting a n d stinging. T h e i r m e r e p r e s e n c e may also be h a r m f u l , b u t they a r e m o r e serious as vectors of organisms pathological to plants as well as to m a n a n d his d o m e s t i c a t e d animals.

T h e e c o n o m i c d a m a g e caused in these ways is incalculable in m o n e t a r y t e r m s a n d h u m a n suffering. Most entomologists in Latin A m e r i c a a r e e m p l o y e d to c o m b a t h a r m f u l species. T h e r e s e a r c h l i t e r a t u r e in t h e field is i m m e n s e . Because of t h e g r e a t size of t h e subject, only a brief review is possible h e r e . A l t h o u g h most publications in practical e n t o m o l o g y deal with t h e t e m p e r a t e parts of t h e world ( N o r t h A m e r i c a a n d E u r o p e [Metcalf et al. 1962, Pfadt 1978]), c o m p r e ­ hensive research o n tropical pests has devel­ o p e d in r e c e n t years ( L a m b 1974). T h e lat­ ter includes large p a r t s o f Latin America, yet references giving detailed i n f o r m a t i o n o n regional e c o n o m i c e n t o m o l o g y r e m a i n sparse a n d in m a n y cases, o u t d a t e d . S o m e

ALVARADO, J. A. 1939. Los insectos dañinos y los insectos auxiliares de la agricultura en Guate­ mala. Published by author, Guatemala City. BALLOU, H. A. 1912. Insect pests of the Lesser Antilles. Comm. Agrie, Imp. Dept. Agrie. West. Ind. Pamph. Ser. 71: 1-210. BERRY, P. 1959. Entomología económica de El Salvador. Min. Agrie. Gan. (Santa Tecla, El Salvador) Bol. Tec. 24: 1-255. GALLEGO, F. L. 1967. Lista preliminar de insectos de importancia económica y secundarios, que afectan los principales cultivos, animales do­ mésticos y al hombre, en Colombia. Fac. Nac. Agron. (Medellín) Rev. 26(65): 32-66.

90

References

KOONE,

H.

D.,

AND A.

D.

BAÑEGAS.

1958.

Entomología económica hondurena. Min. Re­ curs. Nat. (Tegucigalpa) Bol. Teen. 6: 1 — 139. [Not seen.] LAMB, K. P. 1974. Economic entomology in the tropics. Academic, London. METCALF, C. L., W. P. FLINT, AND R. L. METCALF.

plied entomology. 3d ed. Macmillan, New York. POSADA, L. 1976. Lista de insectos dañinos y otras plagas en Colombia. Insto. Colombiano Agropec, Bogotá. REMILLET, M. 1988. Catalogue des insectes ravageurs des cultures en Guyane francaise. ORSTRM, Cayenne. SQUIRE, F. A. 1972. Entomological problems in Bolivia. Pest Art. News Sum. 18: 239-268. WOLCOTT, G. N. 1933. An economic entomol­ ogy of the West Indies. Entomol. Soc. Puerto Rico, San Juan.

AGRICULTURAL ENTOMOLOGY Insects d a m a g e c r o p plants a n d r e d u c e t h e value of agricultural p r o d u c e by eating t h e vegetative p a r t s a n d fruits, by acting as vec­ tors of diseases ( C a r t e r 1973), o r by contami­ nation with t h e i r p r e s e n c e . D a m a g e may be reflected in a variety of ways: wilted leaves, dead stems, discolored o r spoiled fruit, gall formation ( F e r n a n d e s 1987), a n d often t h e death of t h e plant. Loss of food, textile fi­ ber, a n d o r n a m e n t a l plant p r o d u c e in Latin America c a n n o t be calculated with accuracy but surely r u n s into equivalents of billions of dollars annually. No c o u n t r y o r any plant of the g r e a t spec­ trum of Neotropical cultigens is i m m u n e from t h e d e p r e d a t i o n s of injurious insects and mites; t h e p r o b l e m s a r e universal, as evident from several g e n e r a l publications (Caswell 1962; Frohlich a n d Rodewald 1970; Gallo 1988; Hill 1983; K r a n z et al. 1979; Ebeling 1959; F l e c h t m a n n 1983). A variety of reviews o r catalogs of agricultural pests describe local situations: A r g e n t i n a (Rizzo 1977, Molinari 1948), Brazil (Bondar 1913), C e n t r a l A m e r i c a (King a n d Saunders 1984, S a u n d e r s et al. 1983), Co­

lombia ( A n o n y m o u s 1968), C u b a ( B r u n e r et al. 1975), Dominican Republic (Sontoro 1960), G u a t e m a l a a n d El Salvador (Bates 1932), H o n d u r a s (Passoa 1983), t h e Lesser Antilles (Fennah 1947), Mexico (MacG r e g o r a n d G u t i é r r e z 1983, M o r ó n a n d T e r r ó n 1988), Peru (Wille 1952), P u e r t o Rico (Chiesa Molinari 1942), S u r i n a m (van D i n t h e r 1960), a n d U r u g u a y (Ruffinelli a n d Carbonell 1954). Most injurious species a r e a d a p t e d to a particular host, b u t a few attack almost any plant, t h e best e x a m p l e s being migra­ tory locusts (Schistocerca) a n d leaf c u t t e r ants (Acromyrmex, Atta; C h e r r e t t a n d Pere­ g r i n e 1976). T h e following a r e some of t h e m o r e i m p o r t a n t pests of widely g r o w n c r o p species e n c o u n t e r e d by Latin A m e r i ­ can agronomists. T h e most serious enemies of t h e cacao tree (Leston 1970, Entwhistle 1972) a r e scale insects {Pseudococcus, Planococcus, Dysmicoccus) that act as vectors of viruses a n d fungal diseases, causing dieback a n d signifi­ cant fruit r e d u c t i o n o n affected plants. T h e cacao thrips (Selemnothrips rubrocinctus) a n d a p h i d s (e.g., Toxoptera aurantii) cause similar d a m a g e . Coffee trees serve as hosts to over two h u n d r e d insect species (Le Pelley 1968, 1973). T h e t r u n k s a n d stems a r e suscepti­ ble to t h e larvae of a variety of b o r i n g beetles in Latin America, especially t h e coffee berry b o r e r (Hypothenemus hampei, Scolytidae), which causes t h e d r o p p i n g a n d decaying of berries, a n d "black b o r e r s " (Apate, Bostrichidae) that hollow t h e stems. T h e coffee leaf m i n e r (Leucoptera coffeella, Lyonetiidae) d a m a g e s leaves to such a d e g r e e that they fall off t h e plant. Leaf curl, b u r n i n g , a n d s t u n t i n g a r e c o m m o n l y caused by various mealybugs (Pseudococcus) a n d scale insects. T h e attacks of several kinds of l e p i d o p terous stem b o r e r s can cause t h e virtual loss of entire crops of rice (Cheaney a n d J e n n i n g s 1975, Grist a n d Lever 1969). I n Latin America, t h e chief o f f e n d e r s in this

AGRICULTURAL ENTOMOLOGY

91

category a r e Diatraea s p p . a n d o t h e r Pyralidae ( K a p u r 1964). C o r n (maize) is native t o t h e New World, b u t this gives it n o i m m u n i t y to m a n y i n t r o d u c e d pests such as e a r w o r m s (Heli­ coverpa s p p . , Noctuidae) that d e s t r o y t h e kernals a n d stem b o r e r s , mainly pyralid m o t h s of t h e g e n u s Diatraea, that may cause the e n t i r e plant to d r o o p a n d d i e . O t h e r pests i n c l u d e leaf d e s t r o y e r s , m a n y leaf beetles ( C h r y s o m e l i d a e ) , a n d a host of o t h e r noctuid leaf w o r m s a n d c u t w o r m s . A l t h o u g h m a n i o c ( m a n d i o c a , cassava, yuca, tapioca) suffers from relatively few a r t h r o p o d pests, they c a n cause extensive d a m a g e to this i m p o r t a n t food c r o p (Bellotti a n d van S c h o o n h o v e n 1978, Samways a n d Ciociola 1980). T h e cassava shoot-fly, Neosilva perezi (Lonchaeidae; sometimes e r r o n e o u s l y cited as Silba o r Lonchaea chalybea), is w i d e s p r e a d a n d does major d a m a g e . Y o u n g larvae m i n e in grow­ ing shoots a n d m a y cause t h e e n t i r e s h r u b to d i e . T h e m a n i o c gall m i d g e (Latrophobia [= Autodiplosis, Eudiplosis] brasiliensis, Cecidiomyiidae) is well k n o w n because it causes an obvious d e f o r m a t i o n , r e d galls, o n t h e leaves of plants it attacks. T h e s e galls r e ­ d u c e photosynthesis b u t rarely b r i n g a b o u t loss of vitality of t h e whole plant. I n parts of Brazil, t h e galls a r e k n o w n as mamica de rama o r veruga da mandioca. S p i d e r mites s h o u l d also be c o u n t e d a m o n g t h e m o r e serious pests, especially Tetranychus a n d Mononychellus s p p . T h e larvae of t h e ashy s p h i n x , Erinnyis ello ( W i n d e r 1976), a n d pyralid m o t h Chilozela (Becker 1986) feed directly o n m a n i o c leaves. T h e vegetative p a r t s of t h e b a n a n a plant a r e attacked by t h e m e a l y b u g Pseudococcus comstocki ( O s t m a r k 1974). A d u l t b a n a n a fruit-scarring beetles (Colaspis hypochlora) eat t h e y o u n g , u n f u r l e d leaves a n d stems plus t h e fruit, causing scars that m a k e t h e latter unsalable a n d allowing t h e e n t r y of decay microbes. T h e b a n a n a t h r i p s (Hercinothrips bicinctus) feeds o n t h e fruit, discol­ o r i n g it a n d r e d u c i n g its m a r k e t value, b u t

92

PRACTICAL ENTOMOLOGY

it does n o t cause systemic effects o n t h e plant. Plants a r e killed in m a n y areas from the borings of b a n a n a weevil (Cosmopolites sordidus) larvae in t h e r h i z o m e a n d p s e u d o stem at g r o u n d level. T h e pink bollworm (Pectinophora gossypiella) is t h e most i m p o r t a n t pest of cotton a n d occurs in nearly all g r o w i n g areas. Heavy feeding o n t h e bolls by t h e larvae leads to fiber loss a n d seed d e s t r u c t i o n . T h e cotton bollworm (Helicoverpa zea) a n d the famous cotton boll weevil (Anthonomus grandis) ( B u r k e et al. 1986) both d o similar d a m a g e . Red spider mites, especially (Tetra­ nychus cinnabarinus), cause t h e leaves of cotton plants to wither a n d d r o p off. T h e cotton a p h i d (Aphis gossypii) also often in­ fests t h e foliage. P i n e a p p l e is d a m a g e d directly by t h e root-feeding p i n e a p p l e m e a l y b u g (Dysmicoccus brevipes, Pseudococcidae) b u t also suf­ fers from an associated fungus wilt disease (Phytophthora) that causes leaf d e g e n e r a t i o n a n d small fruit. Citrus is impossible to grow profitably in m a n y areas because of t h e attacks of n u ­ m e r o u s insects (Ebeling 1959). Most seri­ ous a r e t h e citrus s p i d e r mites, including Metatetranychus citri, a n d t h e six-spotted mite (Eotetranychus sexmaculatus). Consider­ able d a m a g e also results from overwhelm­ ing p o p u l a t i o n s of citrus white flies (Trialeurodes, Dialeurodes, Aleyrodidae) a n d scale insects (California r e d scale, Aonidiella aurantii; West I n d i a n r e d scale, Sclenaspidus articulatus; a n d citrus mussel scale o r pur­ ple scale, Lepidosaphes beckii). T h e maggots of t h e fruit flies (Rhagoletis, Anastrepha), especially t h e M e d i t e r r a n e a n fruit fly (Ceratitis capitata, T e p h r i t i d a e ) , m a y ac­ c o u n t for almost total fruit losses in heavily infested areas. Mealybugs (e.g., Planococcus citri) a r e also a major p r o b l e m . S u g a r c a n e also comes u n d e r attack from a large n u m b e r of insects a n d mites (Long a n d Hensley 1972, G u a g l i u m i 1 9 7 2 - 7 3 , Williams e t a l . 1969). F r o g h o p p e r s , particu­ larly t h e s u g a r c a n e f r o g h o p p e r (Aeneolamia

varia saccharina, C e r c o p i d a e ) , a r e this crop's most serious pest in m a n y countries. Sap feeding by t h e n y m p h s o n t h e roots results in w i t h e r i n g of t h e leaves a n d stunted stalks. T h e c a n e l e a f h o p p e r (Saccharosydne saccharivora) is a t r o u b l e s o m e species in J a m a i c a a n d elsewhere. Many s u b t e r r a n e a n , r o o t - f e e d i n g scale insects, mealybugs, cicadas, a n d scarab beetle lar­ vae ("white grubs") a r e also prevalent in various a r e a s . Several stalk-boring larvae of t h e pyralid m o t h g e n u s Diatraea (espe­ cially t h e s u g a r c a n e borer, D. saccharalis) are recognized as p r i m a r y pests as well. Aphids (Aphis sacchari, Sipha flava) serve as vectors of virus a n d fungal diseases a n d injure t h e plants directly by sap removal. O t h e r c r o p s a n d their major pests in La­ tin A m e r i c a a r e avocado (fruit flies, Tephritidae); alfalfa (spotted alfalfa a p h i d , Therioaphis maculata)\ beans (bean a p h i d , Aphisfabae); c o c o n u t p a l m (Lever 1969) (co­ conut scale, Aspidiotus destructor; coconut mite, Eriophyes guerreronis; planthopper, Myndus crudus, vector of lethal yellowing disease) ( H o w a r d et al. 1983); guayaba (fruit flies) (Espinoza 1972); m a g u e y (fruit flies, Euxesta); m a n g o (fruit flies); potatoes (potato t u b e r w o r m , Phthorimaea operculella); papaya (fruit flies); s o r g h u m (sugar­ cane borer, Diatraea saccharalis) (Young a n d Teetes 1977); tobacco (tobacco a n d t o m a t o hornworms, Manduca sexta a n d M. quinquemaculata); a n d w h e a t (wheat thrips, Frankliniella tritici).

References ANONYMOUS. 1968. Catálogo de insectos en culti­ vos de importancia económica en Colombia. Assoc. Latinoamer. Entomol. Publ. 1: 1-156. BATES, M. 1932. Insectos nocivos: Estudio de las principales plagas guatemaltecas con algunos datos de Honduras y El Salvador. Serv. Tec. Coop. Agrie, Unit. Fruil Co., Guatemala City. BECKER, V. O. 1986. Correct name for the species of Chilozela (Lepidoptera: Pyralidae) whose caterpillars damage cassava in South America. Bull. Entomol. Res. 76: 195-198. BELLOTTI, K. A., AND A. VAN SCHOONHOVEN.

1978. Mite and insect pests of cassava. Ann. Rev. Entomol. 23: 39-67. BONDAR, G. 1913. Os insectos daninhos na agricultura. Sec Agrie. Indus. Comm., Sao Paulo. BRUNER, S. C , L. C. SCARAMUZZA, AND A. R.

OTERO. 1975. Catálogo de los insectos que atacan a las plantas económicas de Cuba. 2d ed. Acad. Cien. Cuba, Insto. Zool., La Habana. BURKE, H. R., W. E. CLARK, J. R. CATE, AND P. A.

FRYXELL. 1986. Origin and dispersal of the boll weevil. Entomol. Soc. Amer. Bull. 32: 228-238. CARTER, W. 1973. Insects in relation to plant disease. 2d ed. Wiley, New York. CASWELL, G. H. 1962. Agricultural entomology in the tropics. Arnold, London. CHEANEY, R. L., AND P. R. JENNINGS.

1975.

Problemas en cultivos de arroz en América Latina. Centr. Int. Agrie. Trop. (Ser. GS-15), Cali. CHERRETT, J. M., AND D. J. PEREGRINE. 1976. A

review of the status of leaf-cutting ants and their control. Ann. Appl. Biol. 84: 128-133. CHIESA MOLINARI, O. 1942. Entomología agrí­

cola, identificación y control de insectos y otros animales dañinos o útiles a las plantas. Talleres Gráficos Accurziol, San Juan. CHIESA MOLINARI, O. 1948. Las plagas de la

huerta y el jardín y modo de combatirlas. Ed. Bell (Bibl. Pampa Argentina, Rev. Min. Agrie. Gan. Int. Gen. Serv. Pais), Buenos Aires. EBELING, W. 1959. Subtropical fruit pests. Univ. Calif. Press, Berkeley. ENTWHISTLE, P. F. 1972. Pests of cocoa. Long­ mans, London. ESPINOZA, W. O. 1972. Control fitosanitario en plantaciones de guayaba. Univ. Indus. Santan­ der, Bucaramanga, Colombia. FENNAH, R. G. 1947. T h e insect pests of food crops in the Lesser Antilles. Dept. Agrie. Antigua, British West Indies. [Not seen.J FERNANDES, G. W. 1987. Gall forming insects: Their economic importance and control. Rev. Brasil. Entomol. 31: 379-398. FLECHTMANN, C. H. W. 1983. Acaros de im­

portancia agrícola. Liv. Nobel, Sao Paulo. FRÓHLICH, G., AND W. RODEWALD. 1970. Pests

and diseases of tropical crops and their con­ trol. Pergamon, Oxford. GALLO, D. 1988. Manual de entomología agrícola. 2d ed. Ed. Agron. Ceres Ltda., Sao Paulo. [Not seen.] GRIST, D. H., AND R. J. A. W. LEVER. 1969. Pests

of rice. Longmans, London. GUAGLIUMI, P. 1972—73. Pragas de cana de

AGRICULTURAL ENTOMOLOGY

93

acucar no nordeste do Brasil. MIC, Insto., Acucar e do álcool, Dr. Ad., Ser. Doc. Colecáo Canavieira 10, Rio de Janeiro. HILL, D. S. 1983. Agricultural insect pests of the tropics and their control. 2d ed. Cambridge Univ. Press, Cambridge. HOWARD,

F. W., R. C.

NORRIS,

AND D. L.

THOMAS. 1983. Evidence of transmission of palm lethal yellowing agent by a planthopper, Myndus crudus (Homoptera: Cixiidae). Trop. Agrie. 60: 168-171. KAPUR, A. P. 1964. Taxonomy of the rice stem borers. In Int. Rice Res. Inst., T h e major insect pests of the rice plant. Johns Hopkins Univ. Press, Baltimore. Pp. 3 - 4 3 . KING, A. B. S., AND J. L. SAUNDERS. 1984. T h e

invertebrate pests of annual food crops in Central America. Trop. Devel. Res. Inst., Overseas Devel. Adm., London. KRANZ, J., H. SCHMÜTTERER, AND W. KOCH.

1979. Diseases, pests, and weeds in tropical crops. Wiley New York. LE PELLEY, R. H. 1968. Pests of coffee. Long­ mans, London. LE PELLEY, R. H. 1973. Coffee insects. Ann. Rev. Entomol. 18: 121-142. LESTON, D. 1970. Entomology of the cocoa farm. Ann. Rev. Entomol. 15: 273-294. LEVER, R. J. A. W. 1969. Pests of the coconut palm. United Nations Food Agrie. Org. Agrie. Ser. 77: 1-190. LONG, W. H., AND S. D. HENSLEY. 1972. Insect

pests of sugarcane. Ann. Rev. Entomol. 17: 149-176. MACGREGOR, R., AND O. GUTIÉRREZ. 1983. Guía

de insectos nocivos para la agricultura en México. Ed. Alhambra Mexicana, Mexico City MORÓN,

M.

A.,

AND R.

A.

TERRÓN.

1988.

Entomología practica. Insto. Ecol., México. OSTMARK, H. E. 1974. Economic insect pests of bananas. Ann. Rev. Entomol. 19: 161-176. PASSOA, S. 1983. Lista de los insectos asociados con los granos básicos y otros cultivos selectos en Honduras. Ceiba 25: 1-97. Rizzo, H. F. E. 1977. Catálogo de insectos perjudiciales en cultivos de la Argentina. 4th ed. Hemisferio Sur, Buenos Aires. RUFFINELLI, A., AND C. S. CARBONELL M. 1954.

Segunda lista de insectos y otros artrópodos de importancia económica en el Uruguay. Assoc. Ing. Agron. Montevideo Rev. 94: 33-82. SAMWAYS, M. J., AND A. I. CIOCIOLA. 1980. O

complexo de artrópodos da mandioca (Mani­ hot esculenta) Crantz em Lavras, Minas Gerais, Brasil. Soc. Entomol. Brasil An. 9: 3-10.

94

PRACTICAL ENTOMOLOGY

SAUNDERS, J. L., A. B. S. KING, AND C. L.

VARGAS. 1983. Plagas de cultivos en América Central. Centro. Agron. Trop. Inves. Ens., Turrialba, Costa Rica. SONTORO, R. 1960. Notas de entomología agríco­ la dominicana. Sec. Estad. Agrie. Comer., Rep. Dominicana, Santo Domingo. [Not seen.] VAN DINTHER, J. B. M. 1960. Insect pests of cultivated plants in Surinam. Landbouwproefstation Suriname Bull. 76: 1 — 159. WILLE, J. E. 1952. Entomología agrícola del Perú. 2d ed. Min. Agrie, Lima. WILLIAMS, J. R., J. R. METCALF, R. W. MONTGOM­

ERY, AND R. MATHES, eds. 1969. Pests of

sugarcane. Elsevier, Amsterdam. WINDER, J. A. 1976. Ecology and control of Erinnyis ello and E. alope, important insects in the New World. Pest Art. News Sum. 22: 4 4 9 466. YOUNG, W. R., AND G. L. TEETES. 1977. Sor­

ghum entomology. Ann. Rev. Entomol. 22: 193-218.

FOREST ENTOMOLOGY A special b r a n c h of agricultural entomol­ ogy deals with forest pests (Dourojeanni Ricordi 1963, Gray 1972). T h e t r e m e n d o u s value of wood p r o d u c t s makes this o n e of the most i m p o r t a n t fields economically but o n e s o m e w h a t neglected in Latin America. For this reason a n d because t h e r e a r e a large n u m b e r of commercial t i m b e r species a n d forest types, it is difficult to generalize a b o u t forest pests in this p a r t of the world. Only a few a r e a studies o r surveys have b e e n c o n d u c t e d (Martorell 1945). Investi­ gations o n eucalyptus, s n a p d r a g o n tree (Gmelina arbórea), a n d p i n e (Pinns caribea) pests m a y b e c o m e m o r e a p p r o p r i a t e as these exotic timber types replace native Neotropical h a r d w o o d species. T h e r e has been a tendency to regard insect c o m m u n i t i e s in mixed tropical for­ ests as relatively stable, that is, subject to only small p o p u l a t i o n fluctuations, com­ p a r e d to t e m p e r a t e forests. T h u s , the likeli­ h o o d of severe o u t b r e a k s a r e t h o u g h t to be

r e m o t e . However, as various a u t h o r s have r e p o r t e d a n u m b e r of localized p o p u l a t i o n explosions in O l d World tropical forests similar to those e n c o u n t e r e d in t e m p e r a t e regions, t h e possibility r e m a i n s for similar occurrences in Neotropical forests. W o o d - b o r i n g beetles a r e t h e most com­ mon a n d serious t i m b e r pests. T h e i r larvae molest all p a r t s of the y o u n g , m a t u r e , a n d harvested t r e e . Most b e l o n g to the families Cerambycidae, Scolytidae, C u r c u l i o n i d a e , Platypodidae, a n d B u p r e s t i d a e . D a m a g e t o standing t i m b e r is almost wholly d u e to termites of t h e family T e r m i t i d a e , in par­ ticular, m e m b e r s of t h e g e n u s Coptotermes (Harris 1966). I n their r e s i n - g a t h e r i n g ac­ tivities, stingless bees may d a m a g e n u r s e r y seedlings by b o r i n g into a n d g o u g i n g t h e stems of new plantings (Gara 1970). Many l e p i d o p t e r o u s species a r e n o doubt injurious to t i m b e r trees in Latin America as they a r e t o those of t e m p e r a t e forests, b u t little is k n o w n of the economic impact of t h e n u m e r o u s leaf-feeding a n d wood-boring types. T h e m a h o g a n y w e b worm (Macalla thyrsisalis, Pyralidae) is o n e recognized pest species of m a h o g a n y (How­ ard a n d Solis, 1989).

References DOUROJEANNI RICORDI, M. 1963. Introducción

al estudio d e los insectos que afectan la explotación forestal en la selva Peruana. Rev. Peruana Entomol. Agrie. 6(1): 27-38. GARA, R. I. 1970. Report of forest entomology consultant. Inter-American Inst. Agr. Sci., Org. Amer. States, Turrialba, Costa Rica, UNDP Project 80: 1-21. GRAY, B. 1972. Economic tropical forest ento­ mology. Ann. Rev. Entomol. 17: 313-354. HARRIS, W. V. 1966. T h e role of termites in tropical forestry. Ins. Soc. 13: 255-266. HOWARD, F. W., AND M. A. S o n s . 1989. Distribu­ tion, life history, and host plant relationships of mahogany web worm, Macalla thyrsisalis (Lepidoptera: Pyralidae). Florida Entomol. 72: 469-479. MARTORELL, L. F. 1945. A survey of the forest insects of Puerto Rico. Univ. Puerto Rico J. Agrie. 29: 69-608.

MEDICAL ENTOMOLOGY Many insects, spiders, mites, m y r i a p o d s , a n d o t h e r a r t h r o p o d s a r e medically i m p o r ­ tant, acting either as agents of h a r m to h u m a n s o r as vectors of p a t h o g e n i c micro­ organisms. T h i s is such an i m p o r t a n t aspect of o u r existence that several textbooks treat the subject from a n overall perspective in considerable detail (Faust et al. 1962; F l e c h t m a n n 1973; Horsfall 1962; J a m e s a n d H a r w o o d 1969; Kettle 1984; Smith 1973). Regional discussions a r e also avail­ able for A r g e n t i n a (del Ponte 1958), Brazil (Pinto 1930), Central America (Baerg 1929), P a n a m a (Méndez a n d Chaniotis 1987), a n d South America (Biicherl 1969). Because they inject o r dispense v e n o m s , m e m b e r s of m a n y g r o u p s (Biicherl a n d Buckley 1971) a r e serious agents of medical p r o b l e m s t h r o u g h o u t Latin America. T h e most i m p o r t a n t o f f e n d e r s a r e scorpions (Tityus, Centruroid.es), spiders (Latrodectus, Loxosceles, a n d Phoneutria), a n d stinging Hym e n o p t e r a (Apis, ants, a n d wasps) (Akre a n d Davis 1978). Poisons that act topically (vesicants) are p r o d u c e d by millipedes, blis­ ter beetles, fire beetles (Paederus), a n d oth­ ers (Hoffman 1927). Nettling hairs o r spines, such as a d o r n m a n y caterpillars (Saturniidae, L i m a c o d i d a e , Megalopygidae, etc.), also i m p l a n t toxins (urtication, erucism). Reactions to toxic substances (Tu 1984) may b e slight to severe, even fatal in r a r e cases. Such effects occur either t h r o u g h direct toxification o r by eliciting allergic responses t h r o u g h a n t i g e n s (Frazier 1969), superficially (Orkin a n d Maibach 1985) o r systemically. Any p r o t e i n derived from t h e insect's body may cause a h a r m f u l reaction if it comes in contact with tissue topically o r by injection. A c o m m o n m e a n s of injection is t h r o u g h the bite of blood-feeding forms, that is, mosquitoes, ticks, mites, a n d sand flies (Feingold et al. 1968). T h e antigen is contained in t h e saliva a n d e n t e r s t h e

MEDICAL ENTOMOLOGY

95

b l o o d s t r e a m directly o r via t h e lymphatics. Hypersensitive individuals exhibit varied s y m p t o m s , r a n g i n g f r o m mild d e r m a t o s i s to anaphylactic shock, which c a n b e fatal. Many p e o p l e h a v e e x t r e m e fears o r p h o b i a s of insects a n d similar c r e a t u r e s , leading to p s y c h o n e u r o s e s . A c o m m o n form is d e l u s o r y parasitosis (Waldron 1963), t h e u n s h a k a b l e belief that one's skin a n d orifices a r e infested with m i n u t e , barely visible insects, mites, o r o t h e r ver­ min. T h i s c o n d i t i o n a p p e a r s to b e a symp­ tom of a variety of o r g a n i c a n d m e n t a l d i s o r d e r s . E x t r e m e p h o b i a s also affect m a n y p e r s o n s , especially against large, very hairy, d a r k , o r noisy species. T h e s e e m o t i o n a l c o m p l a i n t s a r e n o t as well d o c u ­ m e n t e d in Latin A m e r i c a as in o t h e r parts of t h e world b u t a r e surely as w i d e s p r e a d . H u m a n myiasis is a n o t h e r major p r o b ­ lem caused directly by larval D i p t e r a , espe­ cially of blowflies a n d a few special types like t h e h u m a n botfly (Dermatobia). T h e b o d y m a y b e i n v a d e d by m a g g o t s , leading to a variety of s y m p t o m s , m a n y highly r e p u g n a n t psychologically as well as physi­ cally (Beesley 1974; a n d see Myiasis, c h a p . 11).

s p r e a d in this way, particularly u n d e r very u n s a n i t a r y conditions w h e n poverty o r so­ cial d i s r u p t i o n , such as w a r o r n a t u r a l disasters, prevails in a h u m a n p o p u l a t i o n . Poliomyelitis, typhoid fevers, cholera, lep­ rosy, a n d o t h e r diseases m a y also find new h u m a n hosts in this m a n n e r . Research indicates that t h e A I D S virus is n o t transmit­ ted by insects, blood feeding o r otherwise. Biological vectors a r e f o u n d entirely a m o n g blood-feeding types, especially mos­ quitoes a n d o t h e r biting m i d g e s a n d flies, a l t h o u g h ticks a n d mites a r e also signifi­ cant. T h e i r ecology is a principal factor de­ t e r m i n i n g t h e effectiveness of these insects as vectors ( M u i r h e a d - T h o m s o n 1968). T h e o r g a n i s m s of over a d o z e n major types of h u m a n diseases a r e t r a n s m i t t e d by a r t h r o ­ p o d s in Latin America. T h e most notorious a n d w i d e s p r e a d of these is malaria, which is caused by four species of plasmodial proto­ zoans. T h e s e o r g a n i s m s invade various or­ gans a n d destroy r e d blood cells, releasing toxins into t h e circulation which cause rack­ ing chills a n d fever. Vectors a r e several spe­ cies of mosquitoes in t h e g e n u s Anopheles.

Insects a n d their relatives a r e highly effi­ cient a n d diverse as t r a n s m i t t e r s of o t h e r p a t h o g e n i c o r g a n i s m s . Disease m i c r o o r g a n ­ isms m a y b e c a r r i e d by t h e insect passively (mechanically) o r m a y pass t h r o u g h certain of its d e v e l o p m e n t a l stages in t h e a r t h r o ­ p o d host, which is t h e n c o n s i d e r e d a n obli­ gatory o r "biological" vector. Several major g r o u p s a r e vectors of h u m a n a n d animal p a t h o g e n s , i n c l u d i n g biting flies, fleas (Bibikova 1977), kissing b u g s , bloodfeeding mites, a n d ticks.

Leishmaniasis is a n affliction also caused by protozoans, at least t h r e e species of flagellates in t h e g e n u s Leishmania. Sand flies (Lutzomyia, Psychodidae) carry these agents that invade a n d chemically destroy both d e r m a l a n d internal tissues of vital o r g a n s . A n o t h e r flagellate p r o t o z o a n , Trypanosoma cruzi, that develops in kissing bugs (Reduviidae, Triatominae) brings o n a seri­ ous ailment called Chagas's disease in many parts of Latin America. Visceral o r g a n s suffer c h r o n i c d a m a g e , which leads ultimately to d e a t h in m a n y u n t r e a t e d cases.

Many viruses, bacteria, a n d a m o e b i c o r w o r m cysts a r e mechanically t r a n s m i t t e d . T h e y a r e c a r r i e d o n t h e bodies, o n t h e m o u t h p a r t s , a n d in t h e intestines of filth flies, c o c k r o a c h e s , a n d o t h e r insects that f r e q u e n t c o n t a m i n a t e d m a t t e r a n d food e a t e n later by h u m a n s . Many dysenteries, t a p e w o r m , a n d n e m a t o d e diseases a r e

Various parasitic n e m a t o d e w o r m s intro­ d u c e d from t h e O l d World have become established in certain areas a n d cause a variety of filarial infections. T h e s e include Wuchereria bancrofti (Bancroftian filariasis), Onchocerca volvulus (onchocerciasis), and Dirofilaria imitis ( d o g h e a r t w o r m ) . While seldom fatal, they wreak considerable dam-

96

PRACTICAL ENTOMOLOGY

age by i n v a d i n g t h e tissues, p r o d u c i n g inflammation, e n l a r g e m e n t , a n d destruc­ tion. W h e n essential o r g a n s such as t h e eye or brain a r e involved, critical functions of the senses m a y b e i m p a i r e d . Vectors a r e mostly mosquitoes, b u t blackflies, p u n k i e s , and tabanids also serve as carriers. A large a n d g r o w i n g n u m b e r of viruses ("arboviruses") a r e being discovered which require biting fly, mite, a n d tick vectors. T h e worst of these historically has b e e n t h e yellow fever virus, t r a n s m i t t e d by mosqui­ toes, especially t h e yellow fever mosquito, Aedes aegypti. Several kinds of encephalitides, h e m o r r h a g i c fever, a n d d e n g u e fe­ ver a r e also in this category. T h e viral genus Phlebovirus, t r a n s m i t t e d by phlebotomine sand flies (Lutzomyia) a n d mosquitoes, contains m a n y species of h u m a n p a t h o ­ gens causing intense flulike diseases (Tesh 1988).

are again causing major p r o b l e m s in areas formerly t h o u g h t free of t h e m .

Epidemic (Rickettsia prowazekii) a n d e n ­ demic (R. mooseri) t y p h u s o r g a n i s m s pass to humans from t h e bodies of lice a n d fleas. A third rickettsia (R. rickettsii), that of Rocky M o u n t a i n spotted fever, is b o r n e by hard ticks. T h e s e microbes i n v a d e a n d destroy t h e i n n e r lining of small blood vessels. H i g h fevers, often followed by death, occur. Similar s y m p t o m s follow in­ fection by t h e spirochetes of relapsing fevers (Borrelia) t r a n s m i t t e d t h r o u g h t h e bite o r body secretions of ticks a n d lice.

ous animals and their venoms. 3. Venomous invertebrates. Academic, New York. DEL PONTE, E. 1958. Manual de entomología médica y veterinaria Argentinas. Ed. Lib. Colegio, Buenos Aires.

T h e i n f a m o u s p l a g u e bacillus (Yersinia pestis) still resides in animal reservoirs in parts of Latin A m e r i c a a n d sometimes makes its way t o t h e h u m a n p o p u l a t i o n via flea bites. Fortunately, epidemics like those of the past in E u r o p e a n d elsewhere have not occurred in recent times. Most of these diseases have b e e n con­ trolled by m o d e r n insecticides applied against t h e carriers a n d by d r u g s that kill the p a t h o g e n s . However, as a result of relaxation of a b a t e m e n t c a m p a i g n s a n d development of chemical resistance by both insects a n d m i c r o o r g a n i s m s , some diseases a r e e x p e r i e n c i n g a r e s u r g e n c e a n d

References AKRE, R. D., AND H. G. DAVIS. 1978. Biology

and pest status of venomous wasps. Ann. Rev. Entomol. 23: 215-238. BAERG, W. J. 1929. Some poisonous arthropods of North and Central America. 4th Int. Cong. Entomol. (Ithaca, 1928) Trans. 2: 418-438. BEESLEY, W. N. 1974. Arthropods—Oestridae,

myiases and acariñes. In E. J. L. Southby, Parasitic zoonoses. Academic, New York. Pp. 349-368. BIBIKOVA, V. A. 1977. Contemporary views on the interrelationships between fleas and the pathogens of human and animal diseases. Ann. Rev. Entomol. 22: 23-32. BOCHERL, W. 1969. Giftige arthropoden. In E. J. Fittkau, J. lilies, H. Klinge, G. H. Schwabe, and H. Sioli, Biogeography and ecology in South America. 2: 764-793. Junk, T h e Hague. BÜCHERL, W., AND E. E. BUCKLEY. 1971. Venom-

FAUST, E. C , P C. BEAVER, AND R. C. JUNG.

1962. Animal agents and vectors of human disease. Lea and Febiger, Philadelphia. FEINGOLD, B. E, E. BENJAMINI, AND D. M I -

CHAELI. 1968. T h e allergic responses to insect bites. Ann. Rev. Entomol. 13: 137-158. FLECHTMANN, C. H. W. 1973. Acaros de im­

portancia médico veterinaria. Liv. Nobel, Sao Paulo. FRAZIER, C. A. 1969. Insect allergy, allergic and toxic reactions to insects and other arthro­ pods. Green, St. Louis. HOFFMAN, W. A. 1927. Irritation due to insect secretion. Amer. Med. Assoc. J. 88: 145 — 146. HORSFALL, W. R. 1962. Medical entomology, arthropods and human disease. Ronald, New York. JAMES,

M. T.,

AND R. F. HARWOOD.

1969.

Herm's medical entomology. 6th ed. MacMillan, London. KETTLE, D. S. 1984. Medical and veterinary entomology. Wiley, Somerset, N.J. MÉNDEZ, E., AND B. CHANIOTIS. 1987. Reseña

de las principales enfermedades transmitidas por garrapatas en Panamá. Rev. Med. Pan­ amá 12: 217-223. MÉNDEZ, E., AND B. CHANIOTIS. 1987. Reseña

MEDICAL ENTOMOLOGY

97

de las principales enfermedades transmitadas por insectos en Panamá. Rev. Med. Panamá 12: 205-216. MUIRHEAD-THOMSON, R. C. 1968. Ecology of in­

sect vector populations. Academic, London. ORKIN, M., AND H. I. MAIBACH. 1985.

Cutane­

ous infestations and insect bites. Marcel Dekker, New York. PINTO, C. 1930. Arthrópodes parásitos e transmissores de doencas. Vols. 1—2. Pimento de Mello, Rio de Janeiro. SMITH, K. G. V., ed. 1973. Insects and other arthropods of medical importance. Trus. Brit. Mus. Nat. Hist., London. TESH, R. B. 1988. The genus Phlebovius and its vectors. Ann. Rev. Entomol. 33: 169-181. Tu, A. T 1984. Handbook of natural toxins: Insect poisons, allergens, and other inverte­ brate venoms. Vol. 2. Marcel Dekker, New York. WALDRON, W. 1963. Psychiatric and entomologi­ cal aspects of delusory parasitosis. Amer. Med. Assoc. J. 186: 213-214.

VETERINARY ENTOMOLOGY T h e study of a r t h r o p o d a g e n t s a n d vectors of diseases of d o m e s t i c a t e d animals, pets, g a m e , a n d wildlife constitutes the field of veterinary e n t o m o l o g y (Kettle 1984, Southby 1982, Williams 1985). O n e of the most serious afflictions suf­ fered by cattle a n d o t h e r livestock in Latin America is babesiosis o r cattle fever (caused by Babesia bigemina). It is p r e s e n t t h r o u g h o u t the region a n d is t r a n s m i t t e d by ticks, mainly Boophilus rnicroplus. N u m e r o u s viruses s p r e a d by m o s q u i t o vectors infect horses, o t h e r q u a d r u p e d s , a n d poultry. T h e s e a r e mainly t h e e n c e p h a litides, such as V e n e z u e l a n e q u i n e a n d eastern e q u i n e types that can rapidly deci­ m a t e h e r d s o r flocks a n d a r e also transmis­ sible to h u m a n s . T h e h e a r t w o r m (Dirofilaria imitis) is fairly c o m m o n in the m o r e tropical por­ tions of Latin A m e r i c a . Its insect hosts a r e p u n k i e s a n d mosquitoes. W o r m s living in the h e a r t i m p a i r its function in dogs, which a r e particularly susceptible to this filarial parasite.

98

PRACTICAL ENTOMOLOGY

A great n u m b e r of e x t e r n a l parasites, including ticks of all types, m a n g e mites, keds (Hippoboscidae), fleas, a n d biting a n d sucking lice, infest every kind of do­ mestic animal (Steelman 1976). T h e feed­ ing a n d allergenic effects of these pests cause considerable a n n o y a n c e to their hosts a n d greatly diminish their g r o w t h a n d vitality. Similarly, the attacks of biting flies, mostly blackflies, mosquitoes, horse­ flies a n d deerflies, a n d h e m a t o p h a g o u s Muscidae, k e e p animals o n e d g e a n d nega­ tively affect their general health. T h e sites of bites may become septic, a n d d e a t h may e n s u e from loss of blood. Particularly h a r m f u l are the h o r n fly (Haematobia irritans), whose constant p e s t e r i n g can cause significant weight loss in cattle, a n d the stable fly (Stomoxys calcitrans), which can drive q u a d r u p e d s to fits. T h e f o r m e r spe­ cies has invaded South A m e r i c a as far south as n o r t h e r n Brazil b u t seems not to be a major p r o b l e m because of its p o o r a d a p t a t i o n to tropical climates. It may b e c o m e a serious pest if it reaches t e m p e r ­ ate regions farther south ( T h o m a s pers. comm.). As with h u m a n s , myiasis is a p r o b l e m — b u t to a m u c h g r e a t e r d e g r e e o n account of the g r e a t e r vulnerability of livestock. Foremost in this category a r e t h e d e e p tissue invaders such as the screwworm (Cochliomyia hominivorax), whose attacks are a major m e n a c e to s h e e p a n d cattle ranch­ ing over wide areas, a n d the h u m a n botfly (Dermatobia hominis). T h e m o r e superficial effects of warbles a n d tissue bots (Hypod e r m a t i d a e , Oestridae), while less serious physiologically, r e d u c e the value of hides a n d pelts. Stomach bots (G aster ophilus) are also w i d e s p r e a d a n d m a r k e d l y affect the health of horses a n d cattle. Wildlife diseases caused by insects and o t h e r a r t h r o p o d s are poorly u n d e r s t o o d in Latin America. P e r h a p s the best studied a r e the so-called sylvatic forms of plasmodia ("bird malarias"), trypanosomes, a n d viruses ("jungle yellow fever"), be-

cause of t h e i r relationship to o u t b r e a k s in h u m a n s , causing such s y n d r o m e s as ma­ laria, Chagas's disease, a n d yellow fever.

References KETTLE, D. S. 1984. Medical and veterinary entomology. Wiley, Somerset, N.J. SOUTHBV, E. J. L. 1982. Helminths, arthropods and protozoa of domesticated animals. 7th ed. Lea and Febiger, Philadelphia. STEELMAN, C. D. 1976. Effects of external and internal arthropod parasites on domestic live­ stock production. Ann. Rev. Entomol. 21: 155-178. WILLIAMS, R. E. 1985. Livestock entomology. Wiley, New York.

STORED PRODUCT AND STRUCTURAL PESTS Harvesting, s t o r a g e , a n d p a c k a g i n g p r o ­ vide n o g u a r a n t e e of safety to food a n d useful items from the ravages of insects a n d mites. Many species a d a p t e d for feeding on seeds, cellulose, a n d animal tissues a n d hair are naturally attracted to items c o m p o s e d of these materials which are b r o u g h t in from the field ( B a u r 1984). T h e majority of these pests (Cotton 1960) a r e now cosmo­ politan as a result of their association with commercial p r o d u c t s that are distributed t h r o u g h o u t t h e world t h r o u g h t r a d e . T h e y survive well in w a r e h o u s e s , s t o r e r o o m s , the holds of ships, a n d in the m a r k e t p l a c e , where their p r e s e n c e a n d feeding d e g r a d e s or destroys cereals a n d grains, p a p e r , wood, fur and hides, fabrics, a n d o t h e r organic materials. S t o r e d p r o d u c t pests a r e not well investigated as a g r o u p in Latin America, with some e x c e p t i o n s (Granovsky 1976, Passoa 1983). Damage to grain in elevators a n d silos constitutes t h e largest losses to stored p r o d ­ ucts. Grain, meal, a n d flour are attacked by a variety of beetles ( H i n t o n 1945), m o t h s (Corbet a n d Tarns 1943), a n d mites (Flechtmann 1983). Beetle adults a n d larvae eat the kernels of rice, wheat, c o r n , a n d so on.

T h e s e include the rice a n d g r a n a r y weevils (Sitophilus) a n d t h e various g r a i n beetles (Tribolium a n d Tenebrio, Oryzaephilus, etc.). Fortunately, t h e d r e a d e d k h a p r a beetle (Trogodermagranarium), which p r e f e r s d r i e d vegetable p r o d u c t s b u t also attacks animal p r o d u c t s , is not now k n o w n to exist any­ w h e r e in Latin America. Its potential i n t r o ­ duction is a constant m e n a c e , however. A n u m b e r of grain beetles serve as i n t e r m e d i ­ ate hosts for h u m a n t a p e w o r m s (Hymenolepis), t h u s assuming medical i m p o r t a n c e (Cáceres a n d Guillen d e T a n t a l e á n 1972). T h e larvae of flour a n d meal m o t h s eat milled seeds a n d c o n t a m i n a t e provisions with their webbing a n d feces. T h e principal offenders in this category are the M e d i t e r r a ­ n e a n flour m o t h s (Ephestia, Anagasta), A n g o u m o i s grain m o t h (Sitotroga cerealella), a n d I n d i a n meal m o t h (Plodia interpunctella). A considerable variety of mites (Cáceres e t a l . 1989, F l e c h t m a n n 1 9 8 3 : 1 4 5 160) infest stored grains, s o m e of which actually feed on fungi g r o w i n g t h e r e a n d not on the p r o d u c e itself. Also, several types bite a n d cause dermatitis ("itch mites") in g r a n a r y workers a n d bakers. In Latin America, termites a r e the chief destroyers of finished wood p r o d u c t s . Many species are i m p o r t a n t , especially Coptotermes, which feeds n o t only o n houses a n d l u m b e r b u t o n forest trees as well. O t h e r major wood pests include powderpost beetles (especially Lyctus). Stored paper, including books, frequently is d a m ­ aged not only by these insects but by silverfish, psocids, ants, a n d bostrichid beetles. T h e leather industry is p l a g u e d by h i d e beetles (Dermestes) that riddle cowhides d u r ­ ing the t a n n i n g process a n d storage. T h e cigarette beetle (Lasioderma serricorne) a n d d r u g s t o r e beetle (Stegobium paniceum) lay waste to dried tobacco. T h e y also destroy all m a n n e r of dry animal a n d plant p r o d ­ ucts in the h o m e a n d shops (stored nuts, cereals, spices, candy, etc.). Wool g a r m e n t s a n d furs a r e subject to destruction by Webbing clothes m o t h s

STORED PRODUCT AND STRUCTURAL PESTS

99

(Tineola bisselliella) a n d case-bearing clothes m o t h s (Tinea pellionella) in Latin A m e r i c a as elsewhere.

References BAUR, F. J., ed. 1984. Insect management for food storage and processing. Amer. Assoc. Cereal Chem., St. Paul. CÁCERES, I. E., AND Z. GUILLEN DE TANTALEAN.

1972. Insectos de Lima relacionados con el cistercoide de Hymenolepis diminuta (Rudolphi, 1819), (Cestoda: Hymenolepididae). Rev. Peruana Entomol. 15: 142-147. CÁCERES, I. E., A. ELLIOT, AND I. NAKASHTMA.

1989. Ácari Prostigmata y Mesostigmata en alimentos almacenados de Lima, Huaraz e Iquitos. Rev. Peruana Entomol. 31: 13—17. CORBET, A. S., AND W. H. T. TAMS. 1943. Keys

for the identification of the Lepidoptera infecting stored food products. Zool. Soc. London Proc. B 113: 55-148. COTTON, R. T. 1960. Pests of stored grain and grain products. Burgess, Minneapolis. FLECHTMANN, C. H. W. 1983. Acaros de im­

portancia agrícola. Liv. Nobel, Sao Paulo. GRANOVSKY, T. A. 1976. Insects associated with stored grain in Paraguay, South America. Kans. Entomol. Soc. J. 49: 508. HINTON, H. E. 1945. Monograph of beetles affecting stored products. Brit. Mus. Nal. Hist., London. PASSOA, S. 1983. Lista de los insectos asociados con los granos básicos y otros cultivos se­ lectos en Honduras. Ceiba (Tegucigalpa) 25(1): 1-97.

URBAN ENTOMOLOGY U r b a n i z a t i o n is p r o c e d i n g at a r a p i d pace t h r o u g h o u t t h e world. L a r g e cities a r e b e c o m i n g e v e n larger, n e w l a n d is b e i n g t a k e n over, a n d densities within old m e t r o ­ politan c e n t e r s a r e e v e r - i n c r e a s i n g . Such g r o w t h forces c o n t a c t b e t w e e n certain k i n d s of insects that i n h a b i t h o m e s a n d buildings a n d those whose n a t u r a l habitats a r e b e i n g i n v a d e d . T h e study of this p h e ­ n o m e n o n is t h e relatively recently estab­ lished field of u r b a n e n t o m o l o g y (Ebeling 1975, F r a n k i e a n d K o e h l e r 1983). Negative effects of u r b a n insects a r e m a n y a n d d e p e n d o n t h e types of e n v i r o n ­

100

PRACTICAL ENTOMOLOGY

m e n t they occupy, including private dwell­ ings, r e s t a u r a n t s a n d o t h e r food h a n d l i n g establishments, w a r e h o u s e s , m a n u f a c t u r ­ ing plants, a n d buildings d e d i c a t e d to busi­ ness, medical care, a n d recreation. T h e principal p r o b l e m s a r e health related, n o t only from direct contact b u t t h r o u g h con­ tamination of food, b e d d i n g , a n d circulat­ ing p r o d u c t s . S o m e curious psychological s y n d r o m e s also a r e e x a c e r b a t e d , a m o n g t h e m delusory parasitosis a n d mass hyste­ rias associated with real o r i m a g i n e d micro­ scopic insects believed to infest the h u m a n skin. W o o d e n structures a n d their furnish­ ings a r e destroyed by insects, as a r e stored p r o d u c t s . Use of o u t d o o r recreational areas often exposes h u m a n s to a r t h r o p o d vectors of p a t h o g e n s . Hotels, b a t h h o u s e s , a n d like establishments foster t h e transmission of body parasites like lice a n d b e d b u g s . T h e major offenders t o h u m a n peace of m i n d a n d welfare in u r b a n areas a r e semidomesticated species, often of tropical origin, seeking t h e w a r m t h a n d h i g h h u ­ midity that prevails in o u r a b o d e s a n d w o r k i n g places (Frankie a n d Ehler 1978). T h e best e x a m p l e s of these a r e several species of cockroaches, termites, a n d silverfish that live in the walls a n d f u r n i t u r e and o t h e r w o o d e n c o m p o n e n t s of houses. Flies e n t e r t h r o u g h d o o r s a n d windows a n d both bite a n d a n n o y u s . Ants of many varieties d o likewise. Clothes m o t h s d e ­ stroy woolen fabrics, a n d grain, flour, a n d meal m o t h s a n d beetles i n v a d e t h e pantry. As virgin land is c o n v e r t e d to brick, mortar, a n d asphalt, persisting populations of native insects may b r i n g grief to the new t e n a n t s . H o u s i n g situated n e a r freshwater m a r s h e s , from which mosquitoes a n d p u n k i e s e m e r g e , may m a k e life miserable for p e o p l e in their g a r d e n s a n d even i n d o o r s . Kissing bugs (Triatominae) living in r o d e n t nests may choose h u m a n s as hosts o n their n o c t u r n a l w a n d e r i n g s . Control of domestic a n d u r b a n insect pests has special r e q u i r e m e n t s (Mallis et al. 1982, O s m u n a n d Butts 1966). P a r a m o u n t

a m o n g these is t h e n e e d to b e considerate of h u m a n sensitivities. P r o p e r t y o w n e r s and b u s i n e s s m e n c o n c e r n e d with eco­ nomic d a m a g e t o themselves a n d com­ merce m u s t deal with their fears a n d dislikes of insects a n d the d a n g e r s of using insecticides n e a r places of f r e q u e n t a n d prolonged h u m a n o c c u p a t i o n . Consider­ ation m u s t also b e given to insect species that should b e p r o t e c t e d from u r b a n sprawl a n d p r e s e r v e d in n a t u r e preserves or parks. E v e r y t h i n g n a t u r a l n e e d n o t b e destroyed in t h e n a m e of p r o g r e s s o r economic gain. A few k i n d s of desirable insects, such as butterflies, may even b e favored by u r b a n conditions a n d special f a u n a s c r e a t e d (Ruszczyk 1987).

References EBELING, W. 1975. Urban entomology. Univ. Calif. Press, Berkeley and Los Angeles. FRANKIE, G. W., AND L. E. EHLER 1978. Ecology

of insects in urban environments. Ann. Rev. Entomol. 23: 367-387. FRANKIE, G. W., AND C. S. KOEHLER, eds. 1983.

Urban entomology: Interdisciplinary perspec­ tives. Praeger, New York. MALLIS, A. 1982. Handbook of pest control: The behavior, life history, and control of household pests. 6th ed. Franzak and Foster, Cleveland. OSMUN, J. V, AND W. L. BUTTS.

1966. Pest

control. Ann. Rev. Entomol. 11: 515-548. RUSZCZYK, A. 1987. Distribution and abun­ dance of butterflies in the urbanization zones of Porto Alegre, Brazil. J. Res. Lepidop. 25: 157-178.

CONTROL OF INSECT PESTS Under n a t u r a l conditions, insect n u m b e r s are controlled by various m e a n s , princi­ pally by climatic strictures a n d by p r e d a ­ tors, parasites, parasitoids, a n d disease (Aguilar 1989, S t r o n g 1984). Insects living •n u n w a n t e d proximity to h u m a n i t y a n d competing with p e o p l e for food a n d fiber require artificial control (Martin a n d Wood­

cock 1983). Practical (applied o r economic) entomologists have devised a g r e a t m a n y strategies to deal with pest species. Today, eradication is not the goal, as in t h e past, so m u c h as r e d u c i n g d a m a g e to acceptable tolerance levels, a n a p p r o a c h r e f e r r e d to as "pest m a n a g e m e n t " (Metcalf a n d L u c k m a n 1982). M e t h o d s of pest m a n a g e m e n t a r e successful to different d e g r e e s , d e p e n d i n g o n local conditions, d a m a g e levels, a n d availability of funds. I n recent years, they have been c o m b i n e d in a p p r o p r i a t e ways to capitalize o n the best aspects of each, in the t e c h n i q u e of "integrated c o n t r o l " (Ap­ ple a n d Smith 1976, van H u i s 1981). T h i s is usually t h e most logical a n d p r o d u c t i v e a p p r o a c h , r a t h e r t h a n relying entirely o n j u s t o n e m e t h o d , primarily chemicals, to achieve quick a n d c h e a p results. T h e u s e of chemicals alone h a s n o t been totally successful because of t h e ability of insects to develop resistance to most poisons (Georghiou a n d Saito 1983) a n d t h e delete­ rious side effects that a c c o m p a n y pesticide use ( e n v i r o n m e n t a l pollution a n d d e s t r u c ­ tion of n o n t a r g e t organisms) ( G r e e n 1976). In fact, heavy a n d exclusive use of insecti­ cides may actually lead t o a d e c r e a s e in c r o p p r o d u c t i o n . A classic case of this in Latin America took place in t h e C a ñ e t e Valley in Peru (Barducci 1971). I n t h e 1920s, agricultural emphasis h e r e shifted from s u g a r c a n e to cotton. I n the following two decades, cotton pest control was accom­ plished unevenly with chemicals a n d s o m e ecological m e t h o d s , a n d yields varied. I n the 1950s, however, t r e a t m e n t s with o r ­ ganic insecticides increased greatly a n d became pervasive. Yields d r o p p e d d r a m a t i ­ cally, a n d pests increased in kinds a n d intensity of d a m a g e . Finally, i n t e g r a t e d m e t h o d s were i n t r o d u c e d , a n d after a few years, the crisis abated. A key aspect in successful i n t e g r a t e d control is vigilance a n d m o n i t o r i n g , using various kinds of t r a p p i n g a n d s a m p l i n g techniques to assess pest a n d d a m a g e levels before control m e a s u r e s a r e instigated

CONTROL OF INSECT PESTS

101

(e.g., Silveira Neto 1972). Integrated con­ trol takes advantage of the following di­ verse methods of keeping checks on injuri­ ous insects. 1. Chemical control. Insecticides (pesticides) kill insects by their chemical action, usually by interfering with some essen­ tial metabolic function, such as trans­ mission of electrical impulses across nerve synapses or blocking nutritive pathways (Corbett et al. 1984, Wilkin­ son 1976). Poisonous compounds reach their target tissues by ingestión (stom­ ach poisons), by passage through the integument and sense organs (contact poisons), or by entering the tracheal system (fumigants). Insecticides come in almost infinite va­ riety and chemical structure (Martin and Worthing 1976, Wiswesser 1976), and their mode of action, application, safety, and effectiveness comprise the complex subject of insect toxicology (Matsumura 1975). The major categories of insecti­ cides based on chemistry (Buchel 1983) are the inorganics, for example, arsenic compounds, cyanide gas, and the botani­ cals ("first-generation insecticides"), the naturally occurring types of which (rotenone, pyrethrum, nicotine) have been in use for centuries in many parts of the world. Then there are the syn­ thetic organics ("second generation"), products of modern chemistry, includ­ ing such well-known chlorinated hydro­ carbons as DDT, benzene hexachloride, and Chlordane. To these have been added recently the organophosphates (Malathion, Parathion) and the carbamates, both classes noted for their great potency. Insecticides are formulated or ap­ plied in various ways, as sprays, aero­ sols, or gases (fumigants), in pellets or granules, oils, injectates, dusts, and so on. Other chemicals are useful in combat­

102

PRACTICAL ENTOMOLOGY

ing insects by actions other than killing them. Such are repellents that prevent the pest from doing damage in the first place (Davis 1985). These are most often used against biting flies and evolved from the use of natural substances for centuries by many peoples to spare them the ravages of mosquitoes, punkies, and blackflies. Indians and rural people everywhere build smoky fires to repel insects. The Peruvian Amazon Indians rub their skin with the fruit of the Siparuna (Monimiaceae) shrub, which produces a citronella-like odor that is in­ tended to ward off mosquitoes. Very ef­ fective artificially made repellents are di­ methyl phthalate and DEET (N,Ndiethyl-3-methylbensamide). Another form of chemical control is the release of laboratory-produced pheromones to confuse the mating behav­ ior of crop pests and suppress their numbers by interfering with reproduc­ tion (Jacobson 1965). This seems to have the best potential with lepidopterous pests (Roelofs and Cardé 1977). A substance mimicking the courtship pheromone of the pink bollworm ("gossyplure") has been fairly successful in this way in cotton fields. Sex-attractant pheromones also are used in traps to catch Mediterranean ("medlure") and Mexican Fruit Flies so that their pres­ ence can be detected and monitored in infested areas. Hormone analogues, mostly growth regulators, are now known which disrupt normal growth and kill. Such sophisticated chemical at­ tacks, engineered to spurn only the of­ fensive species and operating on funda­ mental life processes so that resistance is unlikely, constitute the modern front of pest control ("third generation insec­ ticides" or semiochemicals). 2. Physical control. These methods often involve the use of special equipment, machinery, and electrical or radiationproducing devices. They are generally

costly in time and labor and rarely give general control. Some, however, are simple, such as the use of screens or barriers. A very direct approach is hand picking and destruction by gathering and disposing of the pest. This was done against larval Lepidoptera, such as sphinx caterpillars and earworms, by ancient Americans but is not practiced on a commercial scale today. However, not very long ago, bounties were of­ fered for scorpions in Durango, Mex­ ico. It is reported that from April 1785 to October 1787, prizes were paid on 506,644 scorpions in that city (Baerg 1929: 422). Pests may be caught in machines where they are killed by exposure to fumigants or excessively high or low temperatures. Most recently, electro­ magnetic radiation has been applied in different forms for control. Light draws many insects to their death in traps set at the edges of fields or around habita­ tions. Ionizing radiation is a very effec­ tive device in the war against the screwworm and various fruit flies. It is used to sterilize males in great numbers so that they may be released to flood local populations and eliminate effec­ tive reproduction ("sterile male tech­ nique"). Ultrasonic waves have been tried for insect control but are totally ineffective (Ballard and Gold 1983, Lewis et al. 1982). Other physical forces, such as lasers and radiowaves, are now undergoing trials as potential control agents. 3. Biological control. An attractive approach because it concentrates on the target organism and causes minimal harm to the environment is biological con­ trol (Cock 1985, Hoy and Hertzog 1985). Classically, this has depended on the introduction of the pest's natural enemies (Caltagirone 1957, Sweetman 1958), which include predators, para­ sites, parasitoids, and disease microor­

ganisms (Maramorosch and Sherman 1985). Many of the first three are other insects but may include insectivorous vertebrates such as birds, fish, toads, or lizards. This method is usually also selfsustaining and can maintain pest popula­ tions at low levels indefinitely as long as some hosts are left to sustain predatorparasite populations. There are many examples of biologi­ cal control in Latin America. The area serves both as the recipient of control agents and as the source of them. In the former category are five parasitic wasp species introduced into Cuba and Mex­ ico from the Middle East which finally controlled the citrus blackfly (Aleurocanthus woglumi) in the first half of this cen­ tury after many attempts (De Bach 1974: 139f, 167f.). An example of the latter case is an internal wasp parasitoid from Brazil (Tetracnemus peregrinus) which helped stop the mealybug, Pseudococcus adonidum, in parts of the United States early in this century. The Amazon fly (Metagonistylum mínense, Tachinidae) has been transported from its native South America to several areas of the Carib­ bean where it now helps to keep the sugarcane borers (Diatraea) in check (De Bach 1974: 143 f). Other forms of biological control are being devised in endlessly ingenious ways by contemporary researchers in practical entomology. Bacillus thuringiensis is a bacterium lethal to caterpillars and other larval forms. Commercial preparations of the protein crystalline inclusions in the spores may be dissemi­ nated and act like a specific insecticide (Thuricide). Genetic control (Kirschbaum 1985, Pal and Whitten 1974) takes advantage of lethal or repressive genes that entomologists artificially introduce into wild populations from laboratoryreared individuals carrying these genes. 4. Cultural control. This method of control uses ordinary farm or management prac-

CONTROL OF INSECT PESTS

103

tices to r e d u c e d a m a g e from pests as m u c h as possible. It is t h e c h e a p e s t of all control measures but must be planned far in a d v a n c e of the season of potential d a m a g e . It is also necessary to u n d e r ­ stand in detail t h e life history a n d habits of t h e insect pests involved. T h e most c o m m o n application of cultural c o n t r o l is c r o p r o t a t i o n a n d timed tilling o r soil cultivation. T h e objective is to r e m o v e t h e insect's food s o u r c e a n d modify its e n v i r o n m e n t at critical times of its life cycle. By varying the time of p l a n t i n g a n d harvesting, infestations m a y b e a v o i d e d o r m u c h r e d u c e d . T h e u s e of resistant c r o p o r a n i m a l varieties also k e e p s pest p r o b ­ lems t o a m i n i m u m (Maxwell a n d J e n ­ nings 1980). S o m e strains o r varieties of cultivars a r e m o r e o r less resistant to insect attack a n d c a n b e selected for p r o p a g a t i o n in p e s t - p r o n e areas. 5. Legal control. T h e law c a n be a p p l i e d against insect infestations a n d consider­ ably a m e l i o r a t e major p r o b l e m s . A pow­ erful w e a p o n against b o t h medical a n d a g r i c u l t u r a l pests is q u a r a n t i n e . S p r e a d of t h e o f f e n d i n g species o u t s i d e of t h e p r i m a r y a r e a is p r e v e n t e d , a n d control efforts c a n b e c o n c e n t r a t e d o n eradica­ tion. Legal m e a s u r e s a r e also i m p o r t a n t in m a k i n g s u r e that pests a r e k e p t o u t of a c o u n t r y o r r e g i o n . Historically, t h e most serious injurious insects have b e e n i m p o r t e d f r o m o t h e r places. Free of their n a t u r a l e n e m i e s in their h o m e territories, t h e i r p o p u l a t i o n s e x p l o d e in the n e w l a n d s . Laws a r e necessary t o e n f o r c e safe u s e of d a n g e r o u s insecti­ cides a n d m o v e m e n t of materials that m i g h t s p r e a d p r o b l e m species. T h e y also establish agencies to control a n d study i n j u r i o u s insects a n d r e l a t e d a r ­ t h r o p o d s , such as a g r i c u l t u r a l schools a n d e x p e r i m e n t stations, pest c o n t r o l commissions a n d b o a r d s , institutes, a n d a b a t e m e n t districts.

104

PRACTICAL ENTOMOLOGY

References AGUILAR, P. G. 1989. Las arañas como controladoras de plagas insectiles en la agricul­ tura peruana. Rev. Peruana Entomol. 3311-8. APPLE, J. L., AND R. F. SMITH. 1976. Integrated

pest management. Plenum, New York. BAERG, W. J. 1929. Some poisonous arthropods of North and Central America. 4th Int. Cong. Entomol. (Ithaca, 1928) Trans. 2: 418-438. BALLARD, J. B., AND R. E. GOLD. 1983. The

response of male German cockroaches to sonic and ultrasonic sound. Kans. Entomol. Soc.J. 56: 93-96. BARDUCCI, T. B. 1971. Ecological consequences of pesticides used to control cotton insects in the Cañete Valley, Perú. In M. T. Farvar and J. P. Milton, eds., The careless technology— ecology and international development. [Not seen.] BUCHEL, K. H. 1983. Chemistry of pesticides. Wiley, New York. CALTAGIRONE, L. 1957. Insectos entomófagos y sus huéspedes anotados para Chile. Dir. Gen. Prod. Agrar. Pesq., Santiago, Agrie. Téc. Min. Agr. 17: 16-48. COCK, M. J. W., ed. 1985. A review of biological control of pests in the Commonwealth Carib­ bean and Bermuda up to 1982. Commwealth. Inst. Biol. Contr. Tech. Comm. 9: 1-218. CORBETT, J. R., K. WRIGHT, AND A. C. BAILLIE.

1984. The biochemical mode of action of pesticides. Academic, New York. DAVIS, E. E. 1985. Insect repellents: Concepts of their mode of action relative to potential sensory mechanisms in mosquitoes (Diptera: Culicidae). J. Med. Entomol. 22: 237-243. DE BACH, P. 1974. Biological control by natural enemies. Cambridge Univ. Press, London. GEORGHIOU, G. P., AND T

SAITO. 1983. Pest

resistance to pesticides. Plenum, New York. GREEN, M. B. 1976. Pesticides—Boon or bane. Westview, Boulder, Colo.

1985- Viral insecticides for biological control. Academic, Orlando. MARTIN, H., AND D. WOODCOCK.

1983. T h e

scientific principles of crop protection. 7th ed. Arnold, London. MARTIN, H., AND C. R. WORTHING. 1976. Insec­

ticide and fungicide handbook. Blackwell, Oxford. MATSUMURA, F. 1975. Toxicology of insecticides. plenum, New York. MAXWELL, F. G., AND P. R. JENNINGS.

1980.

Breeding plants resistant to insects. Wiley, New York. METCALF, R. L., AND W. H. LUCKMAN, eds. 1982.

Introduction to insect pest management. 2d ed. Wiley, New York. PAL, R-> A N D M. J. WHITTEN. 1974. T h e use of

eenetics in insect control. Elsevier/NorthHolland, Amsterdam. ROELOFS, W. L., AND R. T. CARDÉ. 1977. Re­

sponses of Lepidoptera to synthetic sex pheromone chemicals and their analogues. Ann. Rev. Entomol. 22: 377-405. SILVEIRA NETO, S. 1972. Levantamento de in­

sectos e fluctacáo da poblacáo de pragas da ordem Lepidoptera, con o uso de armadilhas luminosas en diversas regióes do Estado de Sao Paulo. Lib. Doc, Sao Paulo. STRONG, D. R. 1984. Banana's best friend. Nat. Hist. 93(12): 50-57. SWEETMAN, H. L. 1958. The principles of bio­ logical control. Brown, Dubuque. VAN HUÍS, A. 1981. Integrated pest manage­ ment in the small farmer's maize crop in Nicaragua. Mededelingen Landbouwhogeschool 81—86, Wageningen, Netherlands. WILKINSON, C. F., ed. 1976. Insecticide biochem­ istry and physiology. Plenum, New York. WISWESSER, W. L., ed. 1976. Pesticide index. 5th ed. Entomol. Soc. Amer, College Park, Md.

VALUABLE INSECTS

HOY, M. A., AND D. C. HERTZOG, eds. 1985.

Biological control in agricultural IPM sys­ tems. Academic, Orlando. JACOBSON, M. 1965. Insect sex attractants. Wiley Interscience, New York. KIRSCHBAUM, J. B. 1985. Potential implication of genetic engineering and other biotechniques to insect control. Ann. Rev. Entomol. 30: 51-70.

T h e economics of insect life should not b e viewed only negatively. Insects a r e valu­ able, even essential, t o m a n k i n d in a variety of ways; these m a y b e categorized as (1) economic, (2) ecological, (3) scientific, a n d (4) aesthetic.

LEWIS, D. O., W. L. FAIRCHILD, AND D. J.

LEPRINCE. 1982. Evaluation of an electronic mosquito repellen Can. Entomol. 114: 699702. MARAMOROSCH, K., AND K. E. SHERMAN, eds.

Economic Value Because of t h e i r p r o d u c t s a n d services, insects have in t h e past possessed, a n d still

possess, considerable w o r t h in t h e m a r k e t ; the potential of m a n y m o r e is u n r e a l i z e d . It is n o t j u s t primitive o r extinct native cultures that profit from this r e s o u r c e ; t h e m o d e r n world does so as well. Undoubtedly, t h e most valuable a n d heavily used insect p r o d u c t s a r e h o n e y a n d wax from t h e h o n e y b e e a n d silk from t h e domestic silk m o t h . Both insects a r e t h e basis for large industries (see Stingless Bees a n d H o n e y b e e , c h a p . 12) in Latin America, a l t h o u g h t h e direct p r o d u c t i o n of raw silk h a s w a n e d from f o r m e r days (see Domestic Silk Moth, c h a p . 10). Fine silk textiles are now woven in factories, b u t they rely on raw material from t h e O r i e n t a n d elsewhere. T h e h o n e y b e e {Apis mellifera) is main­ tained by agriculturists n o t only for its p r o d u c t s b u t also for its service as a pollinator of crops. T h e m o n e t a r y value of this service in all Latin A m e r i c a n c o u n t r i e s exceeds millions of dollars annually in increased p r o d u c t i o n of fruit a n d seeds. Many insects, especially showy b u t t e r ­ flies, moths, a n d beetles, a r e collected as the r a w material t o m a k e artistic, decora­ tive, o r curiosity items for sale a r o u n d t h e world. Collectors buy m a n y of these, b u t far m o r e a r e b o u g h t by tourists a n d arti­ sans w h o may u s e only a p o r t i o n of t h e insect's body to i n c o r p o r a t e into salable pieces. In some countries, cottage i n d u s ­ tries of major economic i m p o r t a n c e have developed a r o u n d t h e insect t r a d e (see Butterflies, c h a p . 10). P h y t o p h a g o u s insects a r e used as con­ trol agents against weeds a n d o t h e r u n d e ­ sirable plants. T h e larvae of cactus m o t h s (see Cactus Moths, c h a p . 10) have b e e n e x p o r t e d to m a n y parts of the world from their native A r g e n t i n a to destroy Opuntia cactus stands that spoil vast acreages of pasture- a n d c r o p l a n d . O t h e r insects h e l p to s u b d u e explosive growths of water weeds, such as alligator weed ( B u c k i n g h a m et al. 1983) a n d the water hyacinth ( C e n t e r 1982). A lymantriid m o t h (Elnoria noyesi),

VALUABLE INSECTS

105

called malunya (or malumbia) in Peru, has b e e n c o n s i d e r e d as a possible b u t probably ineffective c o n t r o l for illegal coca cultiva­ tion ( B e r e n b a u m 1991). Insects c o n t i n u e to be a s o u r c e of food for native A m e r i c a n s in all areas (Bodenh e i m e r 1951). L a r g e m o t h a n d beetle lar­ vae a n d , to a lesser e x t e n t , ants, wasp larvae, a n d s o m e t r u e b u g s (see agave w o r m s , c h a p . 10; r h i n o c e r o s beetle, c h a p . 9; leaf c u t t e r ant, c h a p . 12) a r e t h e types most often e a t e n . T h i s was n o t e d by the early observers of life in the New World (Wallace 1853) a n d c o n t i n u e s to be d o c u ­ m e n t e d as an active c u l t u r a l practice in c o n t e m p o r a r y times ( R u d d e l 1973). Stud­ ies h a v e b e e n s a n c t i o n e d by t h e Mexican g o v e r n m e n t to d e t e r m i n e the feasibility of e x p a n d i n g local e n t o m o p h a g y to supple­ m e n t the diet of a g r o w i n g r u r a l p o p u l a ­ tion directly o r for a n i m a l feed (Elorduy d e C o n c o n i 1982).

to their use in d e t e r m i n i n g time of d e a t h as sometimes e m p l o y e d in forensic medicine (Vargas-Alvarado 1983, Keh 1985, Smith 1986).

P e r h a p s the greatest potential for profit from insects in t h e f u t u r e lies in t h e i r abili­ ties to synthesize biochemicals for medical use. S h a m a n s a n d village d o c t o r s employ insects in r e m e d i a l concoctions o r apply t h e m directly to diseased o r d a m a g e d or­ gans. T h e Aztecs u s e d t h e oil of black widow spiders (vitztle) to s t o p pain ( C u r r a n 1937). Insects may not p r o v i d e t h e raw ma­ terial for p r o d u c t i o n of quantities of these substances directly b u t g u i d e p h a r m a c o l o ­ gists to u n u s u a l c o m p o u n d s with healing qualities in a fashion paralleling t h e already p r o v e n m e t h o d with medicinal plants ("drug p r o s p e c t i n g " ) . A p o s e m a t i c species that advertise t h e p r e s e n c e of substances m a n u f a c t u r e d in t h e i r b o d i e s which a r e ca­ pable of a l t e r i n g v e r t e b r a t e metabolisms will be p r i m e c a n d i d a t e s in this process; their w a r n i n g colors may also h e l p us to find yet u n k n o w n pharmacologically po­ tent plants f r o m which they have seques­ t e r e d powerful chemical a g e n t s (Brown 1979).

Science makes f r e q u e n t use of insects as e x p e r i m e n t a l o r g a n i s m s for the study of basic biological a n d physical processes. T h e p o m a c e fly, Drosophila melanogaster, has been responsible for p r o v i d i n g insight into our basic u n d e r s t a n d i n g of genetics. Because of their small size, ease of m a i n t e n a n c e , and hardiness, insects are convenient organisms to place a b o a r d e x p e r i m e n t a l rockets to study the effects of o u t e r space o n life. Many insects, especially aquatic species, are useful as bioindicators. T h e i r p r e s e n c e or absence o r body modifications a r e charac­ teristics sensitive to changes in the atmo­ s p h e r e , to toxic substances in air or water, a n d even to radioactivity.

T h e succession of c o m m u n i t i e s of necrophilous insects in h u m a n c a d a v e r s is t h e key

106

PRACTICAL ENTOMOLOGY

Ecological Value Insects are u n s e e n a n d usually u n a p p r e c i ­ ated benefactors of m a n k i n d t h r o u g h the roles they play in ecosystems. T h e y polli­ n a t e most wild species of flowering plants, they h e l p r e d u c e t r e m e n d o u s quantities of animal a n d plant waste a n d t h e r e b y add nutrients to a n d i m p r o v e quality of soils (Seastedt 1984), they h e l p r e g u l a t e the num­ bers of o t h e r o r g a n i s m s , a n d they act as a p r i m a r y food source for all sorts of insecti­ vorous animals, most conspicuously, verte­ brates. T h e y are t h e greatest c o n v e r t e r s of plant m a t t e r to animal protein.

References B E R E NBAUM, M. 1991. Just say "Notodontid?" Amer. Entomol. 37: 196-197. BODENHEIMER, F. S. 1951. Insects as human food. Junk, T h e Hague. BROWN, JR., K.. S. 1979. Insetos aposemáticos: Indicadores naturais de plantas medicináis. Cien. Cult. 32 (suppl.): 189-200.

BUCKINGHAM, G.

R.,

D.

BOUCIAS, AND R.

F.

THERIOT. 1983. Reintroduction of the alligatorweed flea beetle (Agasicles hygrophila Selman and Vogt) into the United States from Argentina. J. Aquat. Plant Man. 21: 101-102. CENTER, T. D. 1982. T h e water hyacinth wee­ vils. Aquatics 4(2): 8, 16, 18-19. CURRAN, C. H. 1937. Insect lore of the Aztecs. Nat. Hist. 39: 196-203. ELORDUY DE CONCONI, J. R. 1982.

como fuente de proteínas en el futuro. Ed. Limusa, Mexico City. KEH, B. 1985. Scope and applications of forensic entomology. Ann. Rev. Entomol. 30: 137-154. RUDDEL, K. 1973. The human use of insects: Examples from the Yukpa. Biotropica 5: 94— 101. SEASTEDT, T. R. 1984. The role of microarthropods in decomposition and mineraliza­ tion processes. Ann. Rev. Entomol. 29: 25-46. SMITH, K. G. V. 1986. A manual of forensic entomology. Brit. Mus. Nat. Hist., London. VARGAS-ALVARADO, E. 1983. Medicina legal. 3d ed. Ed. Lehmann, San José, Costa Rica. WALLACE, A. R. 1853. On the insects used as food by the Indians of the Amazon. Entomol. Soc. London Trans, (n.s.) 2: 241-244.

Los insectos

Scientific Value

Aesthetic Value Finally, it may be a r g u e d also that insects are p a r t of the world to be valued for their own sake, a p a r t from any direct applica­ tion to which their existence may be put. T h e y should be aesthetically appreciated for their beauty, their intricacy of form, a n d t h e lessons they teach a b o u t their ways of life.

VALUABLE INSECTS

107

4

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS ■ " ' ^ . . - ■ ■ - ^

Figure 4.1 ARTHROPODS, (a) Onychophoran (Macroperipatus torquatus, Peripatidae). (b) Sea "roach" {Ligia exotica, Ligiidae). (c) Pillbug (Armadillidium vulgare, Armadillidiidae). (d) Sow bug (Porcellio laevis, Porcellionidae). (e) Sand "flea" (Orchestia platensis, Talitridae). O t h e r s i m i l a r - a p p e a r i n g lower animals a r e often c o n f u s e d with insects. A j o i n t e d body, legs, a n d a "crawly" c o u n t e n a n c e a r e all that a r e r e q u i r e d for m a n y p e o p l e to l u m p a wide variety o f terrestrial a r t h r o ­ p o d s with t h e t r u e insects. F u r t h e r m o r e , these g r o u p s , such as s p i d e r s a n d mites, are of p o p u l a r interest a n d c o n s i d e r a b l e e c o n o m i c significance. It is a p p r o p r i a t e , t h e r e f o r e , to i n c l u d e these insectlike g r o u p s (Clarke 1973, C l o u d s l e y - T h o m p son 1 9 5 8 , K a e s t n e r 1968, P a r k e r 1982) in this book. T h e fact that they m e r g e evolutionarily with t h e primitive H e x a p o d a also justifies a d d r e s s i n g t h e m . T h e y a r e dis­ cussed h e r e , p r i o r to t h e c h a p t e r s o n t h e insects themselves.

References BRUSCA, R. C , AND G. J. BRUSCA. 1990. Inverte­

brates. Sinauer, Sunderland, Mass. CLARKE, K. U. 1973. T h e biology of the Arthropoda. American Elsevier, New York. CLOUDSLEY-THOMPSON, J. L.

1958. Spiders,

scorpions, centipedes and mites. Pergamon, London. KAESTNER, A. 1968. Invertebrate zoology, ar­ thropod relatives, Chelicerata, Myriapoda. Vol. 2. Wiley Interscience, New York. PARKER, S. P., ed. 1982. Synopsis and classifica­ tion of living organisms. Vol. 2. McGraw Hill, New York.

ONYCHOPHORANS O n y c h o p h o r a . Spanish a n d Portuguese: Onicoforos. T h e s e m o d e r a t e - s i z e d ( B L 2—5 c m ) , cater­ pillarlike terrestrial a n i m a l s , n e i t h e r A n n e ­

108

lida n o r A r t h r o p o d a , form a s e p a r a t e phy­ l u m b u t c o m b i n e qualities of both g r o u p s (Marcus 1937). T h e i r annelid characteris­ tics include internally repetitious body seg­ m e n t a t i o n , an eye with a simple lens, the p r e s e n c e of n e p h r i d i a (kidneylike organs) in most body segments, a n d a soft, flexible, wormlike shape, lacking a h a r d e n e d exoskeleton. Some of their a r t h r o p o d features a r e an o p e n body cavity a n d circulatory system, modification of a pair of a p p e n d ­ ages into mandibles, claws o n t h e a p p e n d ­ ages, a b r e a t h i n g system of t r a c h e a e , a n d a n elongate dorsal h e a r t . T h e y also grow by s h e d d i n g their skins like a r t h r o p o d s . O n y c h o p h o r a n s have their o w n special structures, including a transversely wrin­ kled a n d well-pigmented i n t e g u m e n t , each fold with m a n y regularly placed papillae. T h e y also have a pair of a n n u l a t e a n t e n n a e a n d special glands in t h e m o u t h cavity used to shoot streams of slime to c a p t u r e prey a n d fend off enemies. Knowledge of t h e biology of these crea­ tures is scant. T h e y r e q u i r e m o i s t u r e and survive only in h u m i d tropical environ­ m e n t s or d a m p microhabitats in t e m p e r a t e regions. H e r e they inhabit leaf litter, rotten wood, a n d o t h e r moist retreats, such as b a n a n a stems (Young 1980) a n d cavities u n d e r bark. If agitated, they face their antagonist a n d forcefully s p u r t streams of sticky m u c u s from t h e slime glands in the m o u t h . T h e s e solidify into sticky threads that e n t a n g l e a n y t h i n g they touch, produc­ i n g a n o x i o u s mess. Silk s h o o t i n g is also used to immobilize prey.

probably consists mainly of o t h e r small invertebrates o r of partially d e c o m p o s e d leaf a n d wood tissue. O n e species is k n o w n to invade t e r m i t e galleries in r o t t i n g wood to prey o n their o w n e r s (Janvier 1975). In t h e New World tropics, t h e r e a r e eight genera of O n y c h o p h o r a , including fiftyseven species in two families (Peck 1975). Metaperipatus (Peripatopsidae) live in d a m p forests in s o u t h e r n Chile (Claude-Joseph 1928). T h e various g e n e r a of Peripatidae are m u c h m o r e w i d e s p r e a d in t h e rain forests of A m a z o n i a a n d C e n t r a l America (to s o u t h e r n Mexico) a n d t h e C a r i b b e a n . H e r e they crawl over a n d a m o n g t h e litter in search of o t h e r small invertebrates (ter­ mites, caterpillars, snails, etc.) o n which they prey. Peripatus heioisae has b e e n f o u n d in large n u m b e r s in g r o u n d - n e s t i n g termite m o u n d s in Brazil by C a r v a l h o (1942), w h o believes t h e m to be t e r m i t o p h a g o u s . Speleoperipatus speloeus is a blind a n d pale species found in caves in J a m a i c a (Peck 1975). T h e largest species, which m e a s u r e s u p to 15 centimeters in l e n g t h a n d lives in T r i n i d a d , is the collared p e r i p a t u s (Macroperipatus torquatus; fig. 4.1a) so n a m e d because of bright yellow m a r k i n g s a r o u n d t h e bases of the a n t e n n a e (Ghiselin 1985).

References CLAUDE-JOSEPH, F. 1928. Observations sur un

Péripate du Chili. Ann. Sci. Nat. Zool. Ser. lu­ l l : 285-298. GHISELIN, M. T. 1985. A movable feaster. Nat. Hist. 94(9): 5 4 - 6 1 .

JANVIER, H. 1975. Un péripate du Chili chasseur de termites. Entomologiste 31: 63-68. LEITÁO DE CARVALHO, A. 1942. Sobre "Peripatus

heioisae" do Brasil Central. Mus. Nac. Rio de Janeiro (n.s.) Zool. Bol. 2: 57-73. MARCUS, E. 1937. Sobre os Onychophoros. Insto. Biol. Sec. Agrie. Sao Paulo Arch. 8: 255-266. PECK, S. B. 1975. A review of the New World Onychophora with the description of a new cavernicolous genus and species from Ja­ maica. Psyche 82: 341-358. YOUNG, A. M. 1980. On the patchy distribution of onychophorans in two cacao plantations in northeastern Costa Rica. Brenesia 17: 143-148.

CRUSTACEANS T h e Crustacea (Bliss 1 9 8 2 - , S c h r a m 1986) are essentially m a r i n e , b u t m a n y have b e ­ c o m e a d a p t e d to fresh water, a n d some live in moist places o n land. Structurally, they are characterized primarily by t h e p r e s e n c e of b i r a m o u s a p p e n d a g e s , i n c l u d i n g m a n d i ­ bles whose g r i n d i n g surface is served by t h e i n n e r surface of t h e second primitive seg­ m e n t (gnathobasic jaws). T h e first-stage nauplius larva, a simple s w i m m i n g form with t h r e e pairs of a p p e n d a g e s a n d a single m e d i a n eye, is a u n i q u e stage f o u n d only in this g r o u p of a r t h r o p o d s . Crustaceans a r e extremely diverse a n d w i d e s p r e a d in t h e sea. Only a few types a r e habitually terrestrial a n d a p p r o p r i a t e l y in­ cluded here.

T h e s e a r e n o c t u r n a l animals. T h e i r food

CRUSTACEANS

109

References Buss, D. E., ed. 1982-. The biology of the crustácea. Vols. 1—9. Academic, New York. SCHRAM, F. R. 1986. Crustacea. Oxford, New York.

TERRESTRIAL ISOPODS Crustacea, Isopoda. Spanish: Cochinillas de la humedad, correderas (General). Portuguese: Tatuzinhos, baratinhas, bichos de conta (Brazil). Many members of this large and diverse crustacean group (Mulaik 1960, Van Name 1936) are insectlike. They live in humid locations, often near water, including in tank plants such as bromeliads. Although they usually stay near moisture, they are capable of reproducing without depositing their eggs in free water. They may be particularly abundant in leaf litter and decaying vegetation, apparently feeding on the organic debris and fungi associated with such matter. Wood lice, which are found in decomposing leaf and wood litter, are the most familiar representatives. The group is also well represented in caves (Schultz 1981). Terrestrial isopods are mostly small (BL 5—20 mm) and fairly uniform (onisciform): oval in outline and somewhat flat­ tened; body segments distinct and more or less equal, except the posterior whose lat­ eral portions are strongly curved to the rear; anterior segments may be expanded winglike to the sides. T h e thoracic region makes up most of the body length and consists of seven segments, each with a pair of simple legs. Both pairs of antennae are short, but one pair is much shorter than the other. They have stout mandibles and well-developed eyes. There are terrestrial members of only a few families of Isopoda in the Neotropics. Some of the more common are the some­ what amphibious Tylidae and Ligiidae (Ligia, fig. 4.1b; Ligidium), which live along

110

the seashore within or just above the tidal zone and inland by watercourses—"sea roaches." These have extra long antennae. The near-blind Trichoniscidae are very small (BL up to 5 mm) and inhabit dense organic litter in caves. Pillbugs (fig. 4.1c), so called because they can roll themselves into a ball, belong to the Armadillidiidae and Armadillidae. These are dull colored, often solid gray or gray-brown with lighter mottling. To resist moisture loss, the cuticle is usually tough and leathery; in some, it is quite rigid and also a good protective armor for the inter­ nal organs. In nature, the true woodlice or Porcellionidae (sowbugs) (fig. 4.Id) and Oniscidae live in accumulations of dead de­ composing plant matter. They are also abundant, sometimes very abundant, in gardens, greenhouses, and domestic situa­ tions where they are considered pests. The literature is scant on these terres­ trial arthropods; few faunal papers exist (Vandel 1972).

References S. B. 1960. Contribución al conoci­ miento de los isópodos terestres de México (Isopoda, Oniscoidea). Soc. Mex. Hist. Nat. Rev. 21(1): 79-294. SCHULTZ, G. A. 1981. Isopods (Oniscoidae) from caves in North America and northern South America. 8th Int. Cong. Speleol. Proc. 1: 551-552. VANDEL, A. 1972. Les Isopodes terrestres de la Colombie. Stud. Neotrop. Fauna 7: 147-172. VAN NAME, W. G. 1936. The American land and fresh-water isopod Crustacea. Amer. Mus. Nat. Hist. Bull. 71: 1-535. MULAIK,

rally m o r e slender and laterally com­ pressed. Both pairs of antennae are usually well developed and long. The legs are variously modified, some of the anterior often as grasping devices (raptorial) and always with three pairs specialized for walking or hopping (uropods). The exoskeleton may be thin, mineralized, or occa­ sionally heavily sclerotized. It is generally well pigmented and often pitted or other­ wise microsculptured. The dorsal plates often have lateral, winglike flanges. Like the isopods, amphipods require a moist or watery environment, not only for survival but for reproduction as well. They always return to water to deposit their eggs. There are several aquatic larval stages, the animal moving to land only with the last molts. The Talitridae are known everywhere and normally encountered burrowing in damp sand, particularly that beneath beach-stranded seaweed and other debris. The dominant Central and South Ameri­ can genus is Hyale, with many species from all shores (Barnard 1979). The widespread sand flea Orchestia platensis (fig. 4. le) is only common on Caribbean seashores, there be­ ing no members of the family on mainland South America in spite of the fact that the family is otherwise cosmopolitan. This is a neglected group in the region. Much more is to be learned regarding species present and their biology in Latin America.

Reference J. L. 1979. Litorral gammaridean Amphipoda from the Gulf of California and the Galápagos Islands. Smithsonian Contrib. Zool. 271: 1-149.

BARNARD,

TERRESTRIAL AMPHIPODA Crustacea, Amphipoda, Talytroidea, Talitridae, and other families. Beach hoppers, beach fleas, sand fleas, scuds.

ARACHNIDS

Amphipods somewhat resemble isopods in basic structure and biology but are typi-

Arachnids comprise the majority of the chelicerates and are predominantly terres­

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

Arachnida

trial (Besch 1969, Cloudsley-Thompson 1958, Savory 1977). Chelicerates are defined by the structure of the anteriormost appendages (chelicerae), which are made up of a bulky basal segment with an apical movable finger. The chelicerae are modified to form varied or­ gans, such as the venom apparatus (fangs) in spiders, piercing stylets in parasitic mites, or masticating jaws of scorpions. The several orders are quite diverse. In all, the head is undefined, being fused with the thorax (cephalothorax); in some, the abdomen is also fused into a single body complex (mites). Arachnids are an ancient stock, and they are diverse today, though poorly studied in Latin America. A great many species, espe­ cially among the spiders and mites, are yet to be discovered. Arachnids exhibit a strongly climatedependent distribution, mainly in two di­ rections. There are those in moist habitats (Uropygi, Amblypygi) and deserticolous forms (Solpugida). Some transcend these divisions and are widespread and broadly adapted (Araneae, Scorpionida); other mi­ nor groups have specialized niches, such as ectoparasitism. Several anatomical characteristics distin­ guish the arachnids. The head and thorax (leg-bearing portion) are fused into a sin­ gle unit, the cephalothorax (prosoma). The abdomen (opisthosoma) may or may not be distinct. Mouthpart appendages are composed of a single pair of pinching or piercing chelicerae, with a stationary base and movable finger. A pair of segmented sensory appendages (pedipalps) precede the four pairs of legs. Some spiders and scorpions are capable of audible stridulation (Lucas and Bücherl 1972). Arachnids are protected mainly by their secretive, commonly nocturnal habits and camouflage. Besch (1969) noted the pre­ dominance of green in the coloration of spiders and other Arachnida in South America. Many are capable of dropping

ARACHNIDS

111

a p p e n d a g e s to escape c a p t u r e ( a p p e n d o t o my) (Roth a n d Roth 1984). Q u i t e a few also p r o d u c e noxious chemicals, s o m e of which are very p o t e n t in r e p e l l i n g e n e m i e s . Most a r a c h n i d s a r e p r e d a c e o u s a n d feed by ejecting e n z y m e s o n t o t h e prey from a p r e o r a l cavity a n d s i p h o n i n g t h e resultant liquids, o r by piercing t h e prey's skin a n d sucking blood o r l y m p h . A few a r e plant feeders, using e l o n g a t e sucking m o u t h p a r t s to take sap o r o t h e r liquids from t h e host.

References BESCH, W. 1969. South American Arachnida. In E. J. Fittkau, J. lilies, H. Klinge, G. H. Schwabe, and H. Sioli, eds., Biogeography and ecology in South America. 2: 723—740. Junk, T h e Hague. CLOUDSLEY-THOMPSON,J. L. 1958. Spiders, scor­ pions, centipedes and mites. Pergamon, New York. LUCAS, S., AND W. BÜCHERL. 1972. Aparelhos

estriduladoresdoescorpiáo./f/iOjba/unwigfcMsi dorsomaculatus (Prado 1938), e da aranha caranguejeira, Theraphosa blondi (Latreille) 1804. Cien. Cult. 23: 635-637. ROTH, V. D., AND B. M. ROTH. 1984. A review of

appendotomy in spiders and other arachnids. Bull. Brit. Arachnol. Soc. 6: 137-146. SAVORY, T 1977. Arachnida. 2d ed. Academic, London.

SPIDERS Arachnida, Araneae (= Araneida). Spanish: A r a ñ a s . Portuguese: A r a n h a s . Quechua: Vilca-kuna. Tupi-Guarani: N h a n d u i . Náhuatl: T o c a m e h , sing. tócatl (Mexico). Spiders (Foelix 1982, G e r t s c h 1979, P r e ston-Mafham a n d P r e s t o n - M a f h a m 1984) c o m p r i s e a large a n d very familiar g r o u p a n d a r e fascinating a n d diverse in Latin America (Robinson 1984). T h e a b d o m e n is almost always u n s e g m e n t e d a n d nar­ rowly a t t a c h e d t o t h e c e p h a l o t h o r a x by a stalk. T h e chelicerae a r e modified into fangs c o n n e c t e d t o i n t e r n a l poison glands.

112

T h e pedipalps a r e leglike, a n d t h e r e are short terminal a b d o m i n a l a p p e n d a g e s de­ veloped in association with silk glands that o p e n between t h e m (spinnerets). T h e s e c r e a t u r e s a r e f o u n d in all natural situations, often in close association with h u m a n s (Turnbull 1973). Most a r e reclusive a n d select d a r k retreats as living q u a r t e r s . T h e y a r e often found in caves (Gertsch 1973, Brignoli 1972). O t h e r s , however, are sun loving a n d free r a n g i n g . Several kinds even inhabit t h e m a r i n e intertidal zone (Roth a n d Brown 1975). In seasonal cli­ mates, they may b e most a b u n d a n t in wet periods when m o r e prey is available than d u r i n g dry periods (Lubin 1978). A l t h o u g h spiders a r e typically solitary, a n u m b e r of species show e x t e n d e d pa­ rental care a n d even relatively p e r m a n e n t social gatherings (Burgess 1978, Buskirk 1981). In a few species, colony m e m b e r s even c o o p e r a t e in w e b building a n d nest m a i n t e n a n c e , displaying m u t u a l tolerance a n d c o m m u n i c a t i o n (Buskirk 1981, Lubin 1980). Since p r e - C o l u m b i a n times, p e o p l e in Central Mexico have b r o u g h t branches with the webs a n d spiders of a small (BL 5 m m ) social species, called el mosquero (Mallos gregalis), into their h o m e s to reduce the n u m b e r of flies that invade d u r i n g the rainy season (Simon 1909). Many individu­ als c o o p e r a t e to build a d e n s e sheet web with m a n y c h a m b e r s a n d retreats. T h e y not only exhibit g r o u p tolerance b u t prac­ tice c o m m u n a l prey c a p t u r e as well (Bur­ gess 1979, b u t see Jackson 1979 and Tietjen 1986). T h e nest is occupied and e x p a n d e d by successive g e n e r a t i o n s (Bur­ gess 1976). T h e theridiid Anelosimus eximius is t h e most w i d e s p r e a d of several c o m m u ­ nal species in t h e g e n u s in S o u t h America (Rypstra a n d T i r e y 1989). It builds giant webs (1 m by nearly 5 m o r m o r e ) contain­ ing h u n d r e d s , even t h o u s a n d s , of individu­ als w h o work, prey (Krafft a n d Pasquet 1991), a n d spin t o g e t h e r (Levi 1963). The

perception

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

and production

of

sound is d e v e l o p e d in m a n y spiders (Lejrendre 1963) b u t is only o n e of their m a n y means of c o m m u n i c a t i o n (Witt a n d Rovner 1982). T h e y also k e e p in touch by t u g g i n g and vibrating the web a n d t h r o u g h p h e r o mones a n d , m o r e rarely, vision. Many spiders a r e cryptic in f o r m , color, and behavior, a n d a considerable n u m b e r exhibit mimicry, primarily of ants (Peckham 1889). T h e y possess structural (elon­ gate body a n d legs, constricted a b d o m e n ) , behavioral (forelegs held in elevated posi­ tion like a n t e n n a e ) , a n d color modifica­ tions, all giving t h e m a close r e s e m b l a n c e to their formicine m o d e l s . In t h e N e o tropics, these mimics mostly belong to t h e families C l u b i o n i d a e , subfamily Castianeirinae (Reiskind 1969), such as Castianeira rica (Reiskind 1970, 1977), a n d Salticidae (see J u m p i n g Spiders, below). Some species even r e s e m b l e several dis­ tinct types of ants t h r o u g h variation in colors, sexual d i m o r p h i s m , a n d c h a n g e s during d e v e l o p m e n t , so-called t r a n s f o r m a ­ tional mimicry. Velvet ants (Mutillidae) a r e also mimicked. Several k i n d s of small flies are associated with spiders, as c o m m e n s a l s (Robinson a n d Robinson 1977) o r by u s i n g the web as p r o ­ tected p e r c h e s ( L a h m a n n a n d Zúñiga 1981). Symbioses involving spiders seem rare, a l t h o u g h a few u n s t u d i e d cases of in­ teractions with ants a r e k n o w n ( N o o n a n 1982). T h e r e a r e m a n y kleptoparasitic types that live in the webs of o t h e r species from which they steal u n a t t e n d e d prey (arañas ladronas, R e s t r e p o 1948). All spiders a r e obligate carnivores a n d possess v e n o m p r o d u c e d by large internal glands that e m p t y their p r o d u c t s t h r o u g h ducts o p e n i n g at t h e tips of t h e fangs. Spiders u s e poison to s u b d u e p r e y o r in defense. T h e quantity of v e n o m a n d toxic ity of most is slight, so that spider bites a r e usually of little o r n o medical i m p o r t a n c e . A few species, however, d o have the capac­ ity of h a r m i n g (Bettini a n d Brignoli 1978, Schenone 1953) o r even killing h u m a n s .

S o m e of these are f o u n d in Latin A m e r i c a , mainly in t h e g e n e r a Trechona, Phoneutria, a n d Latrodectus, whose v e n o m s a r e n e u r o toxic. Loxosceles bites, c o n t a i n i n g hemolytic toxins, while occasionally severely disfigur­ ing, a r e not lethal (Bücherl 1971). A l t h o u g h o t h e r a r t h r o p o d s spin silk, n o w h e r e is t h e process so well d e v e l o p e d a n d w e b m a k i n g so e l a b o r a t e as in t h e spiders (Savory 1952). S p i d e r silk h a s t h e highest tensile s t r e n g t h of a n y n a t u r a l fiber a n d is ideal for c o n s t r u c t i n g snares a n d traps for insects, as well as nests, retreats, a n d o t h e r s t r u c t u r e s . Webs m a y be relatively small, a m o r p h o u s , a n d loosely f o r m e d o r highly complex, symmetrical, a n d large, as in t h e o r b weavers. T h e relative a b u n d a n c e of t h e different types d e p e n d s o n t h e taxonomic composition a n d ecological characteristics of an a r e a (Lieberman-Jaffe 1981). T h e n u m b e r of species f o r m i n g t h e Neotropical spider fauna is u n k n o w n b u t m u s t b e very large. T h e r e a r e few g e n e r a l works for t h e region (Pikelin a n d Schiapelli 1963). Two major g r o u p s a r e recog­ nized, t h e L a b i d o g n a t h a (most), in which the chelicerae, o r fangs, work laterally, like pliers against each other, a n d t h e O r t h o g natha, in which t h e fangs a r e parallel a n d fold back along t h e long axis of t h e b o d y (tarantulas a n d relatives). Most spiders have eight "eyes" located on the back of the a n t e r i o r p o r t i o n of t h e cephalothorax. T h e number a n d arrange­ m e n t relative to these eyes a r e often used as identifying characteristics.

References BETTINI, S., AND S. M. BRIGNOLI. 1978. Review

of the spider families, with notes on the lesser-known poisonous forms. In S. Bettini, ed., Arthropod venoms. Springer, Berlin. Pp. 101-120. BRIGNOLI, P. M. 1972. Sur quelques araignés cavernicoles d'Argentine, Uruguay, Brésil el Venezuela récoltées par le Dr. P. Strinate (Arachnida, Araneae). Rev. Suisse Zool. 79: 361-385.

SPIDERS

113

BÜCHERL, W. 1971. Spiders. In W. Bücherl and E. E. Buckley, eds., Venomous animals and their venoms. 3. Venomous invertebrates. Academic, New York. Pp. 197-277. BURGESS, J. W. 1976. Social spiders. Nat. Hist. 234: 101-106. BURGESS, J. W. 1978. Social behavior in groupliving spider species. Zool. Soc. London Symp. 42: 6 9 - 7 8 . BURGESS, J. W. 1979. Web-signal processing for tolerance and group predation in the social spider Mallos gregalis. Simon. J. Anim. Behav. 27: 157-164. BUSKIRK, R. E. 1981. Sociality in the Arachnida. In H. R. Hermann, ed., Social insects. 2: 281 — 367. Academic, New York. FOELIX, R. F. 1982. Biology of spiders. Harvard Univ. Press, Cambridge. GERTSCH, W. J. 1973. A report on cave spiders from Mexico and Central America. Assoc. Mex. Cave Stud. Bull. 5: 141-163. GERTSCH, W. J. 1979. American spiders. 2d ed. Van Nostrand Reinhold, New York. JACKSON, R. R. 1979. Predatory behavior of the social spider Mallos gregalis: Is it cooperative? Ins. Soc. 26: 300-312.'

HAM. 1984. Spiders of the world. Facts on File New York. REISKIND, J. 1969. The spider subfamily Castianeirinae of North and Central America. Mus. Comp. Zool. (Harvard Univ.) Bull. 138163-325. REISKIND, J. 1970. Multiple mimetic forms in an ant-mimicking clubionid spider. Science 169587-588. REISKIND, J. 1977. Ant-mimicry in Panamanian clubionid and salticid spiders (Araneae: Clubionidae, Salticidae). Biotropica 9: 1—8. RESTREPO, A. 1948. La araña ladrona. Fac. Nac. Agron. (Medellin) Rev. 17: 157-168. ROBINSON, M. H. 1984. Neotropical arachnology: Historic, ecological and evolutionary aspects. 9th Cong. Latinoamer. Zool. (Are­ quipa) Inf. Final. Pp. 89-100.

KRAFFT, B., AND A. PASQUET. 1991. Synchro­

ROTH, V. D., AND W. L. BROWN. 1975. A new

PIKELIN, B. S. G., AND R. D. SCHIAPELLI. 1963

Llave para la determinación de familias de arañas argentinas. Physis 24: 43—72. PRESTON-MAFHAM, R., AND K. PRESTON-MAF-

ROBINSON,

M. H., AND B. ROBINSON.

1977.

Association between flies and spiders: Bibiocommensalism and dipsoparasitism. Psyche 84: 150-157.

nized and rhythmical activity during the prey capture in the social spider Anelosimus eximius (Araneae, Theridiidae). Ins. Soc. 38: 83-90.

genus of Mexican intertidal zone spider (Desidae) with biological and behavioral notes. Amer. Mus. Nov. 2568: 1-7.

LAHMANN, E. J., AND C. M. ZÚÑIGA. 1981. Use of

RYPSTRA, A. L., AND R. S. TIREY. 1989. Observa­

spider threads as resting places by tropical insects. J. Arachnol. 9: 339-341. LEGENDRE, R. 1963. L'audition et 1'emission de sons chez les aranéides. Ann. Biol. Ser. 4(2): 371-390. LEVI, H. W. 1963. The American spiders of the genus Anelosimus (Araneae: Theridiidae). Amer. Micro. Soc. Trans. 82: 3 0 - 4 8 .

tions on the social spider, Anelosimus domingo (Araneae, Theridiidae), in southwestern Peru. J. Arachnol. 17: 368-370. SAVORY, T H. 1952. The spider's web. Warne, London. SCHENONE, H. 1953. Mordeduras de arañas. Bol. Chileno Parasit. 8: 35-37. SIMON, E. 1909. Sur l'araignée Mosquero. Acad. Sci. (Paris) Compt. Rend. Séance 148: 736-737. TIETJEN, W. J. 1986. Social spider webs, with special reference to the web of Mallos gregalis. In W. A. Shear, ed., Spiders: Webs, behavior, and evolution. Stanford Univ. Press, Stan­ ford. Pp. 172-206. TURNBULL, A. L. 1973. Ecology of the true spi­ ders (Araneomorphae). Ann. Rev. Entomol. 18: 305-348.

LIEBERMAN-JAFFE, S. 1981. Ecology of web-

building spiders at Corcovado National Park, Costa Rica: A preliminary study. Stud. Neotrop. Fauna Environ. 16: 99-106. LUBIN, Y. D. 1978. Seasonal abundance and diversity of web-building spiders in relation to habitat structure on Barro Colorado Is­ land, Panama. J. Arach. 6: 3 1 - 5 1 . LUBIN, Y. D. 1980. Population studies of two colonial orb-weaving spiders. Zool. J. Linnaean Soc. 70: 265-287. NOONAN, G. R. 1982. Notes on interactions between the spider Eilica puno (Gnaphosidea) and the ant Camponotus inca in the Peruvian Andes. Biotropica 14: 145-148. PECKHAM, G. W. 1889. Protective resemblances in spiders. Nat. Hist. Soc. Wise. Occ. Pap. 1: 61-113.

114

W I T T , P. M., AND J. S. ROVNER, eds. 1982. Spider

communication: Mechanisms and ecological significance. Princeton Univ. Press, Princeton.

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

Tarantulas T h e r a p h o s i d a e . Spanish: T a r á n t u l a s , arañas p e l u d a s (General), matacaballos (Mexico, C e n t r a l America). Portuguese: Tarántulas, c a r a n g u e j e i r a s (Brazil). Tupi-Guarani: N h a n d u g u a c á . M y g a l o m o r p h s , bird spiders. This family is r e n o w n e d for t h e e n o r m o u s size of m a n y of its m e m b e r s . T h e largest are males of Theraphosa lablondi (fig. 4.2a), which have a leg span as great as 25 centimeters. S p e c i m e n s a r e k n o w n also with body l e n g t h s of 12 centimeters a n d with anterior f e m o r a l d i a m e t e r s u p to 8 millimeters ( G e r s c h m a n d e Pikelin a n d Schiapelli 1966). T h e b u l k i e r females a r e less expansive but m a y weigh 3 ounces (Gertsch 1979). Most species a r e smaller but are still l a r g e by s p i d e r s t a n d a r d s (leg span 7.5 to 9.5 cm); o t h e r s a r e small (less than 3 cm leg s p a n ) . Tarantulas a r e distinguished from o t h e r large hairy spiders by their large vertical fangs a n d legs t h a t h a v e only two claws, instead of t h e usual t h r e e . T h e equally sized, small eyes a r e closely g r o u p e d o n a small tubercle. T h e r e are two pairs of large spinnerets. Because of their f o r m i d a b l e size a n d hairiness, t a r a n t u l a s a r e widely r e g a r d e d with great fear a n d a r e believed to be deadly. I n spite of these attitudes, these spiders, especially t h e Mexican r e d - l e g g e d tarantula (Brachypelma smithi) from n o r t h ­ western Mexico, a r e i m p o r t e d in large numbers to t h e United States to please pet fanciers ( H e m l e y 1986). Acanthoscurria, and o t h e r s , a r e legitimately feared by Matto Grosso I n d i a n s because of their powerful bite ( B ü c h e r l 1 9 7 1 : 2 2 9 ) , a n d Hapalopus a r e k n o w n to carry p o t e n t ven­ oms (Espinoza 1966). As toxicity to h u ­ mans of some t a r a n t u l a s h a s been d o c u ­ mented, all s h o u l d b e given latitude w h e n encountered (Bettini a n d Brignoli 1978). While usually shy a n d retiring, they may become aggressive if t h r e a t e n e d . Stories of

their j u m p i n g abilities m a y b e e x a g g e r ­ ated, b u t some a r b o r e a l species a r e quite capable of leaps of a m e t e r o r m o r e over level g r o u n d (orig. obs.). Against h u m a n s a n d e n e m i e s , t a r a n t u ­ las use their bite only in defense. T h e y also discourage attack by flicking t h e hairs of the d o r s u m of t h e a b d o m e n with a h i n d leg. T h e s e a r e urticating a n d m a y lodge in the eyes o r sensitive nasal m e m b r a n e s of potential p r e d a t o r s (Cooke et al. 1973). Some Neotropical tarantulas p r o d u c e a snakelike hissing s o u n d by r u b b i n g t h e surfaces of basal segments of t h e p e d i p a l p s against opposite surfaces of t h e first legs. Adults of both sexes u s e this form of stridulation a p p a r e n t l y as a protective m e c h a n i s m ; it can b e h e a r d u p to 6 m e t e r s away (Thorns 1983). T h e s e spiders, particularly t h e females, are long lived. In n a t u r e , most probably m a t u r e in 5 to 10 years a n d m a y live several years thereafter. T h e v e n o m is normally used to s u b d u e prey. T h e i r food consists mostly of o t h e r ground-dwelling arthropods, although the largest species certainly catch a n d d e v o u r small vertebrates such as frogs, toads, liz­ ards, a n d nestling o r small birds. A f a m o u s illustration of a specimen in t h e act of feeding o n a small bird, which a p p e a r e d in M a d a m e Merian's Metamorphoses Insectorum Surinamensium (1705), is probably responsi­ ble for t h e r e p u t a t i o n of these spiders as ornithophages. T h e r e a r e b o t h terrestrial a n d a r b o r e a l tarantulas (pi. la). T h e f o r m e r s p e n d t h e day in b u r r o w s of their own construction or n a t u r a l retreats in t h e soil, e m e r g i n g at night to h u n t o r seek mates. Tree-dwelling forms h i d e a m o n g e p i p h y t e s o r in crevices or u n d e r t h e loose b a r k of d e a d trees. O t h e r s form silken nests in rolled u p leaves of terrestrial bromeliads, b a n a n a s o r Heliconia, in palm spathes, o r in t h e bristly bases of these leaves. A bit of folklore prevailing in Mexico a n d Central A m e r i c a is t h e legend of t h e

SPIDERS

115

American theraphosid. Paper delivered at the annual meeting of the Entomological Society of America, Detroit.

Jumping Spiders Salticidae. Spanish: P a p a m o s c a s (Peru).

Figure 4.2 SPIDERS, (a) Tarantula (Theraphosa lablondi, Theraphosidae). (b) Jumping spider (unidentified, Salticidae). (c) Ant-mimicking jumping spider (Aphantochilus sp., Salticidae). (d) Ant model of ant-mimicking jumping spider (Cephalotes sp., Formicidae). (e) Wolf spider (Lycosa raptoria, Lycosidae). (f) Banana spider (Phoneutria fera, Ctenidae).

matacaballo. A t o n e time, m a n y p e o p l e t h o u g h t t h a t h o o f - a n d - m o u t h disease of cattle was c a u s e d by t a r a n t u l a s . T h e spi­ d e r s w e r e p r e s u m e d t o h u n t sleeping ani­ mals a n d take a n a r r o w strip of h a i r from above t h e h o o f for its nest b u i l d i n g , using a n acid secretion t o m a k e t h e h a i r s l o u g h off (a s y m p t o m of t h e disease). T h e site of the injury suffers infection, a n d t h e h o o f may b e lost. T h e disease is actually c a u s e d by a bacillus; t a r a n t u l a s line t h e i r nests with their own silk (Gertsch 1979: 117). T h e t a x o n o m y of these spiders r e m a i n s in a n u n s e t t l e d state. A l t h o u g h their h i g h e r classification h a s b e e n o r g a n i z e d (Raven 1985), only a few of the major t h e r a p h o s i d g r o u p s have received a t t e n t i o n internally by m o d e r n w o r k e r s (Schiapelli a n d Gerschm a n d e Pikelin 1967, 1979; G e r s c h m a n d e Pikelin a n d Schiapelli 1973). A p p r o x i ­ mately f o u r h u n d r e d Latin A m e r i c a n spe­ cies a r e d e s c r i b e d , r a n g i n g t h r o u g h all climes from deserts to r a i n forest. T h e y a r e m u c h fewer a n d less c o m m o n at the h i g h e r elevations.

References

their venoms. 3. Venomous invertebrates. Academic, New York. Pp. 197-277. COOKE, J. A. L., F. H. MILLER, R. W. GROVER,

AND J. L. DUFFY. 1973. Urticaria caused by tarantula hairs. Amer. J. Trop. Med. Hyg. 22: 130-133. ESPINOZA, N. C. 1966. Acción de veneno de Hapalopus limensis. lnst. Butantan Mem. 33: 799-808. GERSCHMAN DE PIKELIN, B. S., AND R. D. SCHIA­

PELLI. 1966. Contribución al conocimiento de Theraphosa lablondi (Latreille), 1801 (Aranea: Theraphosidae). Inst. Butantan Mem. 33: 667-674. GERSCHMAN DE PIKELIN, B. S., AND R. D. SCHIA­

PELLI. 1973. La subfamilia Ischnocolinae (Araneae: Theraphosidae). Mus. Argentino Cien. Nat. Bernardino Rivadavia, Insto. Nac. Invest. Cien. Nat. Entomol. Rev. 14: 43-77. GERTSCH, W. J. 1979. American spiders. 2d ed. Van Nostrand Reinhold, New York. HEMLEY,

G.

1986. Spotlight

on

the red-

kneed tarantula trade. Traffic (U.S.A.) 6(4): 16-17. RAVEN, R.J. 1985. The spider infraorderMygalomorphae (Araneae): Cladistics and systematics. Amer. Mus. Nat. Hist. Bull. 182: 1-180. SCHIAPELLI, R. D., AND B. S. GERSCHMAN DE

PIKELIN. 1967. Estudio sistemático compara­ tivo de los géneros "Theraphosa" Walck., 1805, "Lasiodora" C. L. Koch, 1851 y "Sericopelma" Ausserer, 1975 (Aranae, Theraphosidae). Atas Simp. Biota Amazónica 5: 481-494.

BETTINI, S., AND P. M. BRIGNOLI. 1978. Review

SCHIAPELLI, R. D., AND B. S. GERSCHMAN DE

of the spider families, with notes on the lesser-known poisonous forms. In S. Bettini, ed., Arthropod venoms. Springer, Berlin. Pp. 101-120. BÜCHERL, W. 1971. Spiders. In W. Bücherl and E. E. Buckley, eds., Venomous animals and

PIKELIN. 1979. Las arañas de la subfamilia Theraphosinae (Araneae, Theraphosidae). Mus. Argentino Cien. Nat. Bernardino Rivadivia, Insto. Nac. Invest. Cien. Nat. Entomol. Rev. 5: 287-300. THOMS, E. 1983. Sound production by a South

116

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

In s p i d e r s o f this family, t h e anterior, median eyes a r e p a i r e d a n d e n o r m o u s l y enlarged a n d face f o r w a r d o n t h e steeply elevated a n t e r i o r p o r t i o n of t h e cephalothorax. T h e y afford t h e a n i m a l excellent binocular vision, e n a b l i n g it t o j u d g e dis­ tance very well. T w o pairs of additional smaller eyes a r e located b e h i n d t h e princi­ pal eyes. Good eyesight is c o r r e l a t e d with over­ sized forelegs, a n d short, s t r o n g h i n d legs, all modifications for p r e y c a p t u r e by a m ­ bush (Forster 1982). Prey is stalked by char­ acteristic j e r k i n g , o r i e n t i n g visual move­ ments of the whole body, t o within 7 to 15 centimeters distance; t h e n a t h r e a d is at­ tached to t h e s u b s t r a t e a n d t h e final dis­ tance j u m p e d . T h e s p i d e r may t h e n d r o p off the substrate o n the t h r e a d line to isolate the quarry. Extensive w e b b i n g is s p u n only for refuge a n d p r o t e c t i o n o f t h e eggs. A wide variety of insects a n d o t h e r a r a c h n i d s are taken; s o m e salticids attack a n d s u b d u e victims considerably l a r g e r t h a n themselves (Robinson a n d Valerio 1977). T h e y even pounce o n o r b spiders situated in the c e n t e r of their webs. T h e males often h a v e e x t r a l o n g chelicerae and are brightly colored, frequently in polychrome. T h e p a t t e r n s of m a n y a r e complex a n d r e s p l e n d e n t in detail, includ­ ing iridescent b l u e a n d g r e e n spots a n d scale patches. T h e entire d o r s u m of t h e abdomen is often solid, vivid r e d o r o r a n g e . Salticids a r e well k n o w n for their eye­ catching c o u r t s h i p displays, d u r i n g which the often elaborately d e c o r a t e d males vigor­ ously p o s t u r e a n d d a n c e in front of obser­ vant females (fig. 4.2b) ( C r a n e 1949; see L e h m a n 1981 for a bibliography). Many salticids a r e a n t mimics (Galiano

1967, Reiskind 1977). I n addition to hav­ ing a n antlike body, they move t h e i r slen­ d e r forelegs in front of t h e h e a d like t h e p r o b i n g a n t e n n a e of their models a n d of­ ten have e n l a r g e d p e d i p a l p s that r e s e m b l e ant mandibles. I n Peru, ants of t h e g e n u s Cephalotes (fig. 4.2d) a r e naturally well p r o t e c t e d by their heavy a r m o r a n d spined bodies. T h e y a r e closely m i m i c k e d by spiders of the g e n u s Aphantochilus (fig. 4.2c) which have a n a r r o w e d , formicoid waist, black color, a n d even a spine o n t h e a n t e r i o r p a r t of t h e c e p h a l o t h o r a x like t h e ant. T h e a n t e r i o r p a r t of t h e b o d y h a s a form that looks m u c h t h e s a m e as t h e ant's h e a d (Preston-Mafham a n d PrestonMafham 1984). T h e r e is o n e a p p a r e n t case of an a n t mimic in symbiosis with t h e leaf-nesting a n t Tapinoma melanocephalum, t h e s p i d e r using the ant's nest for s u p p o r t a n d p e r h a p s p r o t e c t i n g t h e latter from in­ vading p r e d a t o r y insects ( S h e p a r d a n d Gibson 1972). Salticidae is a large family with m a n y small g e n e r a . T h e majority of t h e N e o ­ tropical species are probably still u n k n o w n .

References CRANE, J. 1949. Comparative biology of salticid spiders at Rancho Grande, Venezuela. Pt. IV An analysis of display. Zoológica 34: 159-215. FORSTER, L. 1982. Vision and prey-catching strategies in jumping spiders. Amer. Sci. 70: 165-175. GALIANO, M. E. 1967. Salticidae (Araneae) formiciformes. VIII. Nuevas descripciones Physis 27: 27-39. PRESTON-MAFHAM,

R., AND K. PRESTON-MAF-

HAM. 1984. Spiders of the world. Facts on File, New York. REISKIND, J. 1977. Ant-mimicry in Panamanian clubionid and salticid spiders (Araneae: Clubionidae, Salticidae). Biotropica 9: 1-8. RICHMAN, D. B. 1981. A bibliography of court­ ship and agonistic display in salticid spiders. Peckhamia 2: 16-23. ROBINSON, M. H., AND C. E. VALERIO. 1977.

Attacks on large or heavily defended prey by tropical salticid spiders. Psyche 84: 1-10. SHEPARD, M„ AND F. GIBSON. 1972. Spider ant

symbiosis: Continusa spp. (Araneida: Salti-

SPIDERS

117

cidae) and Tapinoma melanocephalum (Hymenoptera: Formicidae). Can. Entomol. 104: 1951-1954.

The role of abdominal hairs. Science 182: 1153-1155. STRATTON, G. E. 1985. Behavioral studies of wolf spiders: A review of recent research. Rev. Arachnol. 6: 57-70.

Wolf Spiders Lycosidae. Spanish: A r a ñ a s lobos (General), paccha a r a ñ a s (Peru). Portuguese: A r a n h a s lobos. Wolf spiders (Stratton 1985) a r e varied in size (BL 1 0 - 2 0 m m ) , hairy, usually d a r k b r o w n , a n d fast m o v i n g . T h e y attract atten­ tion a n d a r e m u c h feared, a l t h o u g h most a r e h a r m l e s s . L a r g e species in the g e n u s Lycosa, however, may bite h u m a n s with serious c o n s e q u e n c e s . T h e v e n o m is cytotoxic a n d p r o d u c e s g r e a t local pain as well as l i n g e r i n g necrotic effects. T h e spiders are recognized by their m o d e r a t e hairiness a n d the t h r e e pairs of long, heavy, black spines a r m i n g the a n t e r i o r tibia. T h e back of t h e light b r o w n c e p h a l o t h o r a x is often m a r k e d with c o n t r a s t i n g b r o a d , longitudi­ nal bars or lines. Lycosids a r e o t h e r w i s e of m u c h i m p o r ­ tance ecologically as controllers of insect p o p u l a t i o n s , especially o n t h e g r o u n d , which is their usual h a u n t . T h e y a r e va­ g r a n t h u n t e r s with g o o d vision used in prey c a p t u r e . Very few build webs; they sit a n d wait for o t h e r s p i d e r s a n d insects that they a m b u s h , usually nocturnally. T h e y can be located at n i g h t by the b r i g h t reflectance of their eyes in a flashlight b e a m . A w i d e s p r e a d species is Lycosa raptoria (fig. 4.2e). Many have p r o n o u n c e d aggressive a n d fighting behaviors as well as highly devel­ o p e d m a t e r n a l instincts. Females carry e g g sacs a n d spiderlings for s o m e time. Spiny, k n o b b e d hairs on the back of the a b d o m e n a p p a r e n t l y p r o v i d e the stimulus a n d m e a n s of a t t a c h m e n t for this f o r m of b r o o d care (Rovner et al. 1973).

References ROVNER, J. S., G. A. HIGASHI, AND R. F. FOELIX.

1973. Maternal behavior in wolf spiders:

118

Banana Spiders C t e n i d a e , C t e n i n a e , Phoneutria. Portuguese: A r a n h a s a r m a d e i r a s (Brazil). W a n d e r i n g spiders. S p i d e r s in the g e n u s Phoeneutria a r e fairly large ( 1 0 - 1 2 cm leg s p a n , adult females BL 35—50 m m ) a n d powerfully built. T h e eight eyes are in t h r e e rows (2-4-2), the last two the largest a n d the two laterals of the second row the smallest. T h e y h a v e a short coat of grayish to pale b r o w n pelage, with larger black spines a n d chelicerae that are conspicuously clothed with long r e d hairs. T h e i n n e r surfaces of t h e t h r e e apical s e g m e n t s of the palpi a r e heavily fringed with hairs. T h e s e are essentially n o c t u r n a l , vagrant spiders, w a n d e r i n g o n t h e g r o u n d from e v e n i n g to d a w n in search of prey. T h e y do not construct webs to e n t r a p food. T h e y are also very aggressive a n d pugnacious, r e a r i n g o n the h i n d legs with t h e two pairs of forelegs raised t h r e a t e n i n g l y a n d fangs b a r e d (fig. 4.2f). T h i s exposes their u n d e r ­ sides, which a r e strongly d a r k e n e d on e i t h e r side of a c o n t r a s t i n g pale j o i n t mem­ b r a n e between the basal j o i n t of the first two pairs of legs. B a n a n a spiders r e t r e a t into d a r k places d u r i n g the day a n d frequently e n t e r dwell­ ings w h e r e they may h i d e themselves in clothes or shoes. T h e y a r e often found a m o n g b u n c h e s of b a n a n a s , which has e a r n e d t h e m their v e r n a c u l a r n a m e . T h e v e n o m of b a n a n a spiders is consid­ e r e d highly toxic, a n d a few cases of hu­ m a n d e a t h s from their bite a r e well docu­ m e n t e d (e.g., Trejos et al. 1971), although this has o c c u r r e d only in p e r s o n s who are weak or small c h i l d r e n . Bites occur fre­ quently a m o n g workers w h o h a n d l e clus­ ters of b a n a n a s . S o m e characteristic symp-

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

toms a r e local intense pain, tachycardia, salivation, d i s t u r b e d vision, sweating, priapism, a n d p r o s t r a t i o n ( S c h e n b e r g a n d Pereira Lima 1978). Two well-known species a r e P. nigriventer, from s o u t h e r n Brazil, a n d P. fera (fig. 4.2f), which inhabits A m a z o n a s . Equally large b u t less d a n g e r o u s spiders in the g e n u s Ctenus a r e easily confused with b a n a n a s p i d e r s b u t differ in the lack of d e n s e hair b r u s h o n the i n n e r palpal surfaces ( B ü c h e r l et al. 1964).

References BÜCHERL, W.,

S.

LUCAS, AND V.

DESSIMONI.

1964. Aranhas da familia Ctenidae, Sub­ familia Cteninae. I. Redescricáo dos géneros Ctenus Walckenaer 1805 e Phoneutria Perty 1833. Inst. Butantan Mem. 31: 95-102. SCHENBERG, S., AND F. A. PEREIRA LIMA.

1978.

Venoms of Ctenidae. In S. Bettini.ed., Arthro­ pod venoms. Springer, Berlin. Pp. 217—245. TREJOS, A., R. TREJOS, AND R. ZELEDÓN.

1971.

Aracnidismo por Phoneutria en Costa Rica (Araneae: Ctenidae). Rev. Biol. Trop. 19: 241-249.

Typical Orb Web Spiders Araneidae ( = A r g i o p i d a e ) . This is the most characteristic of the sev­ eral families t h a t spin o r b - s h a p e d webs (like a w a g o n w h e e l with a h u b , r a d i a t i n g struts a n d concentric circular ties). Most of the spiders in the family A r a n e i d a e a r e fairly large, often with grossly e n l a r g e d and frequently brightly colored a b d o m e n s . T h e lateral eyes a r e at a distance from the medial, the latter f o r m i n g a s q u a r e . T h e y have poor vision a n d locate prey c a u g h t in their webs by feeling t h e tension a n d vibration of the t h r e a d s (Craig 1989). T h e y wrap captives in sheets of silk d r a w n from the spinnerets by the h i n d legs. Several members of t h e family a r e favorite subjects of behavioral (Robinson a n d Olazarri 1971, Robinson a n d Robinson 1980) a n d ecological study, especially species in the genera Araneus a n d Argiope (Robinson a n d Robinson 1978), a l t h o u g h t h e r e are rela­

tively few Neotropical species in these pri­ marily Holarctic g e n e r a . A w i d e s p r e a d , c o m m o n Latin A m e r i c a n a r a n e i d is Eustala anistera (fig. 4.3a). Araneus is the larger a n d m o r e wide­ s p r e a d genus. T h e female's a b d o m e n is often i m m e n s e a n d nearly spherical a n d m a n y times is m a r k e d with c o m p l e x spot­ ted or variegate p a t t e r n s . Many species of the genus a r e n o c t u r n a l a n d t h u s a r e seldom seen a n d a p p r e c i a t e d . T h e r e a r e fewer kinds of Argiope, but they a r e m o r e conspicuous because of large size, b r i g h t colors, a n d d i u r n a l habits (Levi 1968). T h e females sit head d o w n w a r d with legs ori­ e n t e d like an "X"; they r e m a i n in the c e n t e r of their webs d u r i n g the day, often flanked above a n d below by zigzagging sheet silk structures (stabilimenta) in the web (at the tips of the legs in the silver o r b weaver). Several species are s p r e a d over m a n y parts of the globe, such as the g o l d e n o r b weaver (A. aurantia), which occurs only in the New World, from Mexico to G u a t e ­ mala, with an ovoid black a b d o m e n m a r k e d by i r r e g u l a r sublateral o r a n g e yellow streaks; the b a n d e d o r b weaver (A. trifasciata), whose similarly s h a p e d a b d o ­ m e n is white with n a r r o w black rings; a n d the silver o r b weaver (A. argentata) (fig. 4.3c), with a c o m p r e s s e d , marginally lobed a b d o m e n that is r e d , black, a n d silver spotted posteriorly a n d solid silver basally like that of the adjoining t h o r a x . T h e embryology of the last species has b e e n studied by Tse a n d Tse (1980). N a t u r a l selection has p r o d u c e d m a n y variations on the o r b web t h e m e for prey c a p t u r e . Bolas spiders (Mastophora) (fig. 4.3d), for e x a m p l e , diverge radically from the typical o r b weavers in their h u n t i n g m e t h o d , as they d o not rely passively on a web to catch prey. Instead, at night, they spin a h a n g i n g line with a sticky r o u n d globule at the tip. W h e n prey a p p r o a c h e s , the spider swings this "bola" a n d catches it on the globule ( E b e r h a r d 1980). T h e p o d a d o r a (Mastophora gasteracanthoides) or

SPIDERS

119

Figure 4.3 ORB WEB SPIDERS (ARGIOPIDAE). (a) Common orb weaver (Eustala anistera), fe­ male. (b) Golden silk spider (Nephila clavipes), female, (c) Silver orb weaver {Argiope argentata), female, (d) Bolas spider (Mastophora dizzydeani), female, (e) Spiny orb weaver (Gasteracantha cancriformis). " t r u e bolas spider," a m o d e r a t e - s i z e d (BL 11 m m ) , d a r k b r o w n , slow-moving species with l a r g e h o r n s o n the a b d o m e n , is k n o w n to w o r k e r s in v i n e y a r d s in South A m e r i c a w h o believe it bites with serious effects. It is p r o b a b l e , however, that only its f e a r s o m e a p p e a r a n c e i n c r i m i n a t e s it, a n d some o t h e r a g e n t is actually responsible for the lesions suffered by s o m e individuals w h o e n c o u n t e r it. T h e bolas s t r u c t u r e has b e e n f o u n d to c o n t a i n a volatile substance simi­ lar to t h e sex p h e r o m o n e of a favored prey, owlet m o t h s of t h e g e n u s Spodoptera. T h e male m o t h s a r e a t t r a c t e d to t h e vicinity of t h e s p i d e r w h o s e c h a n c e s of s e c u r i n g a meal a r e t h u s greatly e n h a n c e d ( E b e r h a r d 1977).

References CRAIG, C. L. 1989. Alternative foraging modes of orb web weaving spiders. Biotropica 21: 257-264. EBERHARD, W. G. 1977. Aggressive chemical mimicry by a bolas spider. Science 198: 1173-1175. EBERHARD, W. G. 1980. T h e natural history and behavior of the bolas spider Mastophora dizzydeani sp. n. (Araneidae). Psyche 87: 143-169. LEVI, H. W. 1968. T h e spider genera Gea and Argiope in America (Araneae: Araneidae). Mus. Comp. Zool. (Harvard Univ.) Bull. 136: 319-352.

120

ROBINSON, M. H., AND J. OLAZARRI. 1971. Units

of behavior and complex sequences in the predatory behavior of Argiope argentata (Fabricius): (Araneae: Araneidae). Smithsonian Contrib. Zool. 65: 1-36. ROBINSON, M. H., AND B. C. ROBINSON.

1978.

Thermoregulation in orb-web spiders: New descriptions of thermoregulatory postures and experiments on the effects of posture and coloration. Zool. J. Linnean Soc. 64: 87-102. ROBINSON, M. H., AND B. C. ROBINSON.

1980.

Comparative studies of the courtship and mating behavior of tropical araneid spiders. Pacific Ins. Monogr. 36: 1-218. TSE, M. DO C. P., AND H. D. TSE. 1980. Notas sobre o desenvolvimento da aranha Labidognatha Argiope argentata (Fabricius, 1775) (Araneae, Araneidae). Rev. Brasil. Biol. 40: 249-255. Golden Silk Spider A r a n e i d a e , Nephilinae, Nephila clavipes. Spanish: A r a ñ a d e o r o (Costa Rica). Golden-web spider, giant o r b weaver. Female Nephila clavipes a r e very l a r g e orb web spiders. It is n o t u n u s u a l to find specimens with a b o d y length of 3 to 4 centimeters a n d a leg span of 5 centimeters resting h e a d d o w n w a r d in the m i d d l e of their gigantic o r b - s h a p e d webs (fig. 4.3b). T h e web may be almost a m e t e r square. Young individuals m a k e a c o m p l e t e web; adults build only the b o t t o m p o r t i o n . T h e

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

males are m u c h smaller (BL 1 cm) a n d remain secreted in foliage at the e d g e of the web. T h e web s t r a n d s a r e exceedingly heavy and s t r o n g a n d a r e shiny gold in color. A p e r p e n d i c u l a r ladderlike s t r u c t u r e (stabilirnentum) of d e n s e , m u l t i s t r a n d e d zigzags is often c o n s t r u c t e d by a y o u n g female above a n d below h e r resting place at the hub. T h e function of this device is not known (Robinson a n d Robinson 1973). T h e webs a r e c o m m o n l y occupied by o t h e r spiders (e.g., Conopistha, Argyrodes) that are not detected o r tolerated by the web mak­ ers. T h e y a r e c o n s i d e r e d kleptoparasites, stealing prey c a u g h t in the host's web. T h e i r p r e s e n c e m a y be responsible for frequent relocation of the host to new web sites (Rypstra 1981). Robinson a n d Robin­ son (1977) o b s e r v e d milichiid flies of the genus PhyHomyza living o n the c e p h a l o t h o r ax of this Nephila in P a n a m a . T h e flies helped themselves to the juices of the spider's prey after it was w r a p p e d a n d partially digested. This is t h e only Latin A m e r i c a n species of its g e n u s , o n e c o m p r i s e d of several widely distributed species in the Old World tropical lowlands (Lubin 1983, Robinson and Robinson 1980: 34f.). T h e body is elongate, the a b d o m e n long ovoid a n d steeply inclined t o w a r d the front. T h e latter is colored olive b r o w n with a d o u b l e series of lighter, c r e a m - c o l o r e d spots a l o n g the d o r s u m . T h e legs a r e very long a n d have characteristic thick tufts of black hairs below the j o i n t s of the basal s e g m e n t s . T h e third pair of legs are s h o r t e r t h a n the others a n d lack tufts. Males a r e similar b u t much smaller, with a body length of only about 4 to 8 millimeters, a n d their legs are untufted. T h e species p r e f e r s sparse to m o d e r a t e forest vegetation in which it builds its large webs in gaps a n d c o r r i d o r s w h e r e its prey is likely to pass ( M o o r e 1977). If prey is sufficiently a b u n d a n t , different females •nay build webs c o n t i g u o u s to each o t h e r in

large aggregations. Prey usually consists of flies, bees, wasps, a n d small m o t h s a n d butterflies (Robinson a n d Mirick 1971). Females locate their r o u n d e g g sacs o n leaves at the e n d s of twigs at least 1 m e t e r a b o v e g r o u n d (Christenson a n d Wenzel 1980). Seasonal variation in e g g p r o d u c ­ tion has been studied by C h r i s t e n s o n , Wenzel, a n d L e g u m (1979). Spiderlings are c o m m u n a l a n d feed only o n immobile prey, small insects a n d their own kind (Hill a n d Christenson 1981). Recent e x p e r i m e n t s show that the spe­ cies discriminates unpalatable butterflies a n d releases t h e m u n h a r m e d from its web. Release is not accidental but results from a behavioral s e q u e n c e specifically p r o g r a m m e d to that e n d (VasconcellosNeto a n d Lewinsohn 1984). References CHRISTENSON, T

E., AND P. A. WENZEL.

1980.

Egg-laying of the golden silk spider, Nephila clavipes L. (Araneae, Araneidae): Functional analysis of the egg sac. J. Anim. Behav. 28: 1110-1118. CHRISTENSON, T E., P. A. WENZEL, AND P. LEGUM.

1979. Seasonal variation in egg hatching and certain egg parameters of the golden silk spider Nephila clavipes (Araneidae). Psyche 86: 137-147. HILL, E.

M.,

AND T

E. CHRISTENSON.

1981.

Effects of prey characteristics and web struc­ ture on feeding and predatory responses of Nephila clavipes spiderlings. Behav. Ecol. Sociobiol. 8: 1-5. LUBIN, Y. D. 1983. Nephila clavipes (Arana de oro, Golden Orb-spider). In D. H. Janzen, ed., Costa Rican Natural History. Univ. Chi­ cago Press, Chicago. Pp. 745—747. MOORE, C. W. 1977. The life cycle, habitat and variation in selected web parameters in the spider, Nephila clavipes Koch (Araneidae). Amer. Midi. Nat. 98: 95-108. ROBINSON, M. H., AND H. MIRICK. 1971.

The

predatory behavior of the golden-web spider Nephila clavipes (Araneae: Araneidae). Psyche 78: 123-139. ROBINSON, M. H., AND B. C. ROBINSON.

1973.

The stabilimenta of Nephila clavipes and the origins of stabilimentum-building in araneids. Psyche 80: 277-288. ROBINSON, M. H., AND B. C. ROBINSON.

SPIDERS

1977.

121

Associations between flies and spiders; bibiocommensalism and dipsoparasitism. Psyche 84: 150-157. ROBINSON, M. H., AND B. C. ROBINSON.

1980.

Thermoregulation in orb-web spiders: New descriptions of thermoregulatory postures and experiments on the effects of posture and coloration. Zool. J. Linnaean Soc. 64: 87-102. RYPSTRA, A. L. 1981. T h e effect of kleptoparasitism on prey consumption and web relo­ cation in a Peruvian population of the spider Nephila clavipes. Oikos 37: 179-182. VASCONCELLOS-NETO, J., AND T M. LEWINSOHN.

1984. Discrimination and release of unpalat­ able butterflies by Nephila clavipes, a Neo­ tropical orb-weaving spider. Ecol. Entomol. 9: 337-344. Spiny Orb Weavers Araneidae, Gasteracanthinae. T h e females of this category of o r b weavers (Robinson a n d R o b i n s o n 1980) are h a r d b o d i e d , a n d the a b d o m e n bears conspicu­ ous, s h a r p - t i p p e d spines, e i t h e r r a d i a t i n g from its p e r i p h e r y {Gasteracanlhd) or diverg­ ing from t h e p o s t e r i o r (Miaalhena), t h e lat­ ter giving t h e s p i d e r an a r r o w h e a d s h a p e . T h e i r a b d o m e n s a r e also colored lavishly in r e d , yellow, or white. Small d a r k d e p r e s ­ sions or pits d o t the d o r s u m as well. T h e m u c h smaller males have a cylindrical a b d o ­ m e n , lacking c o n s p i c u o u s spines (Chickering 1961). T h e a b d o m e n of males in the g e n u s Gasteracantha is squarish a n d m u c h wider t h a n long; in Micrathena, it is l o n g e r t h a n wide. T h e latter is r e p r e s e n t e d by a b o u t forty varied species t h r o u g h o u t Latin America. T h e usual Gasteracantha in the New World is t h e c o m m o n cosmopolitan G. cancriformis (fig. 4.3e; M u m a 1971), t h e g e n u s b e i n g primarily O r i e n t a l . T h e spe­ cies is d i s t r i b u t e d from Mexico to n o r t h e r n A r g e n t i n a a n d is r e c o g n i z e d by the p r e s ­ ence of six spines on the a b d o m e n . A second, poorly k n o w n species, G. tetracantha, has only four a b d o m i n a l spines a n d is restricted to P u e r t o Rico, t h e Virgin Is­ lands, a n d the B a h a m a s (Levi 1978). T h e webs of Gasteracantha a r e conspicu­

122

ous orbs found between the b r a n c h e s of s h r u b s a n d even o n buildings. T h e y are m a d e in the m o r n i n g a n d a r e usually inclined at an angle. T h e o u t e r t h r e a d s are d e c o r a t e d with flocculent tufts of silk.

References CHICKERING, A. J. 1961. The genus Micrathena (Araneae, Argiopidae) in Central America. Mus. Comp. Zool. (Harvard Univ.) Bull. 125: 392-470. LEVI, H. W. 1978. The American orb-weaver genera Colphepeira, Micrathena and Gasteracan­ tha north of Mexico (Araneae, Araneidae). Mus. Comp. Zool. (Harvard Univ.) Bull. 148: 417-442. MUMA, M. 1971. Biological and behavioral notes on Gasteracantha caucriformis (Arachnida, Araneidae). Fia. Entorno!. 54: 345-351. ROBINSON,

M.

H.,

AND B.

ROBINSON.

1980.

Comparative studies of the courtship and mating behavior of tropical araneid spiders. Pacific Ins. Monogr. 36: 1-218.

Giant Crab Spiders H e t e r o p o d i d a e (= Sparassidae) a n d Selenopidae. M e m b e r s of these two families a r e com­ monly noticed on walls inside a n d outside of buildings w h e r e their large size (leg span 7—12 cm, BL 2—3 cm) a n d swift, sideways m o v e m e n t s attract attention. M e m b e r s of both families a r e k n o w n for their ability to hide in n a r r o w cracks a n d crevices by day, e m e r g i n g at night to catch insects. They may be especially c o m m o n a r o u n d exterior house lights to which m o t h s a n d beetles are attracted ( M u m a 1953). Typically, these spiders a r e very flat, with a compressed carapace a n d a b d o m e n , the f o r m e r almost circular or slightly wider t h a n long. T h e y hold their spiny legs out flat to the sides like a crab. H e t e r o p o d i d s have four of the total of eight eyes, selenopids six, a r r a n g e d in a row along the a n t e r i o r m a r g i n of the carapace. T h e h u n t s m a n spider (sometimes also called the b a n a n a spider, b u t see Phoneutria above), Heteropoda venatoria, is the bestk n o w n species (fig. 4.4a). It is cosmopoli-

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

Figure 4.4 SPIDERS AND HARVESTMEN. (a) Huntsman spider (Heteropoda venatoria, Heteropodidae). (b) Black widow (Latrodectus mactans, Theridiidae), female, (c) South American violin spider (Loxosceles laeta, Loxoscelidae). (d) Harvestman (Prionostemma sp., Gagrellidae). (e) Harvestman (Gonyleptus janthinus, Gonyleptidae). tan in distribution, probably i n t r o d u c e d from Asia. It is very c o m m o n in h u m a n habitations t h r o u g h o u t the w a r m lowlands of Latin A m e r i c a . H e r e it is valued as a predator of cockroaches, a n d it is k n o w n at times also to kill a n d eat scorpions a n d even small bats. S o m e details of its life history have b e e n elucidated (Ross et al. 1982). It is capable of p r o d u c i n g a faintly audible buzz or h u m by m e a n s of leg oscillations while c o u p l e d to the substra­ tum by tarsal adhesive hairs; the s o u n d apparently plays a role in c o u r t s h i p (Rovner 1980).

References MUMA, M. H. 1953. A study of the spider family Selenopidae in North America, Central Amer­ ica, and the West Indies. Amer. Mus. Nov. 1619: 1-55. Ross, J., D. B. RICHMAN, F. MANSOUR, A. TRAMBARULO, AND W. H. WHITCOMB. 1982.

The life cycle of Heteropoda venatoria (Lin­ naeus) (Araneae: Heteropodidae). Psyche 89: 297-305. ROVNER, J. S. 1980. Vibration in Heteropoda venatoria (Sparassidae): A third method of sound production in spiders. J. Arachnol. 8: 193-200.

Widow Spiders Theridiidae, Latrodectus. Spanish: Viudas negras (General); a r a ñ a s naranjas (Venezuela); cul r o u g e , 24-horas (West

Indies); lucachas (Peru); guiños, pallus (Chile); h u y u r o s micos (Bolivia); rastrojeras, a r a ñ a s del lino ( A r g e n t i n a ) ; a r a ñ a s capulinas, p o - k o - m o o (Mexico); a r a ñ a s bravas ( s o u t h e r n South America). Náhuatl: T z i n t l a t l a u h q u e h , sing, tzintlatlauhqui (var. chintatlahuc). Female "black widow" spiders (Latrodectus mactans) a r e medium-sized (BL 8—15 m m ) a n d j e t black, with a large, n a k e d , globose a b d o m e n having a characteristic r e d d i s h hourglass m a r k i n g on the u n d e r s i d e (fig. 4.4b). In o t h e r species of the g e n u s , the b a c k g r o u n d color may vary from white to r e d d i s h - b r o w n , with beautiful r e d a n d yel­ low lines or spots a d o r n i n g the d o r s u m . T h e males are four to five times smaller but with legs almost as long as the females'. T h e eight eyes a r e in two rows. All are widely feared for their v e n o m o u s qualities. I n d e e d , they b e a r a highly p o t e n t n e u r o t o x i c v e n o m (Bettini a n d Maroli 1978). S y m p t o m s of t h e bite begin with a s h a r p local pain that gradually moves from the w o u n d a r e a to o t h e r p a r t s of t h e body, c o n c e n t r a t i n g finally in t h e a b d o m e n or legs. O t h e r effects are n a u s e a , dizziness, fainting, a n d shock, occasionally with a fatal o u t c o m e . In spite of the potential serious­ ness of envenomization by these spiders, they are reluctant to bite or to inject m u c h

SPIDERS

123

v e n o m so s h o u l d n o t b e c o n s i d e r e d really dangerous. Because of their medical i m p o r t a n c e , they h a v e b e e n investigated m o r e t h a n o t h e r s p i d e r s , particularly in A r g e n t i n a , w h e r e t h e several species a r e n o w fairly well studied embryologically (González 1981, 1984) a n d ecologically (Schnack et al. 1983, Estévez et al. 1984). T h e y a r e con­ trolled naturally by p r e d a c e o u s m u d a n d spider wasps (Sphecidae, Pompilidae) a n d parasitoids a m o n g t h e chalcidoid wasps (Desantisca, E u r y t o m i d a e ) a n d c h l o r o p i d flies (Pseudogaurax) (Pérez Rivera 1980, Sabrosky 1966). Black widows a r e shy a n d largely n o c t u r ­ nal. D u r i n g t h e day, they rest in their finely t h r e a d e d , a m o r p h o u s webs, which they construct in p r o t e c t e d , cool, dry, d a r k r e ­ treats. T h e s e a r e often soil fissures o r spaces a m o n g d e b r i s , in w o o d piles, refuse piles, o r u n d e r h o u s e s ( A n d e r s o n 1972). T h e t a x o n o m y of these spiders is diffi­ cult a n d still unsettled o w i n g to their great variability a n d o v e r l a p in s t r u c t u r a l fea­ tures. Levi (1959) o n c e recognized only t h r e e basic species, L. mactans, L. geomét­ ricas, a n d L. curacaviensis, b u t h e c o n c e d e s that this is a n oversimplification a n d t h a t t h e r e a r e probably several Latin A m e r i c a n species (Levi 1983). Various o t h e r species have b e e n d e s c r i b e d (Carcavallo 1959) a n d reclassifications p r o p o s e d (see review of Bettini a n d Maroli 1 9 7 8 : 149f.), b u t t h e g e n u s Latrodectus is in n e e d of a c o m p l e t e revision u s i n g m o d e r n analytic t e c h n i q u e s .

References ANDERSON, M. P. 1972. Notes on the brown widow spider, Latrodectus geometricus (Araneae: Theridiidae) in Brazil. Great Lakes Entomol. 5: 115-118. BETTINI, S., AND M. MAROLI. 1978. Venoms of

Theridiidae, genus Latrodectus. In S. Bettini, ed., Arthropod venoms. Springer, Berlin. Pp. 149-212. CARCAVALLO, R. U. 1959. Una nueva Latrodectus y consideraciones sobre las especies del género en la República Argentina (Arach. Theridiidae). Neotropica 5: 8 5 - 9 4 .

124

ESTF.VEZ, A. L., A. GONZÁLEZ, ANDJ. A. SCHNACK.

1984. Estadísticos vitales en especies Argenti­ nas del género Latrodectus Walckenaer (Araneae, Theridiidae). II. Latrodectus antheratus (Babcock), Latrodectus corallinus Abalos y Latrodectus diaguita Carcavallo. Physis, Sec. C, 42(102): 29-37. GONZÁLEZ, A. 1981. Desarrollo postembrionario de Latrodectus mirabilis, Latrodectus corallinus y Latrodectus antheratus (Araneae, Theridiidae). Physis, Sec. C, 39(97): 8 3 - 9 1 . GONZÁLEZ, A. 1984. Desarrollo postembrionario y evolución de los órganos mecanorreceptores de Latrodectus diaguita Carcavallo, y estudio de la tricobotriotaxia de Latrodectus quartus Aba­ los (Araneae, Theridiidae). Physis, Sec. C, 42(102): 1-5. LEVI, H. W. 1959. T h e spider genus Latrodectus (Araneae, Theridiidae). Amen Micro. Soc. Trans. 78: 7 - 4 3 . LEVI, H. W. 1983. On the value of genitalic struc­ tures and coloration in separating species of widow spiders (Latrodectus sp.) (Arachnida: Araneae: Theridiidae). Naturwiss. Ver. Ham­ burg Verh. (n.f.) 26: 195-200. PÉREZ RIVERA, R. A. 1980. Distribución geo­

gráfica, potencial reproductivo y enemigos naturales de la viuda negra en Puerto Rico. Carib.J. Sci. 15: 79-82. SABROSKY, C. W. 1966. Three new Brazilian species of Pseudogaurax with a synopsis of the genus in the Western Hemisphere (Díptera: Chloropidae). Dept. Zool., Sec. Agrie, Sao Paulo, Pap. Avul. 19: 117-127. SCHNACK,

j . A.,

A.

GONZÁLEZ,

AND A.

L.

ESTÉVEZ. 1983. Estadísticos vitales en especies Argentinas del género Latrodectus Walckenaer (Araneae, Theridiidae). 1. Latrodectus mirabilis Holmberg. Neotropica 29(82): 141-152.

Violin Spiders Loxoscelidae, Loxosceles. Spanish: d e las rincones (Chile).

Arañas

Loxosceles a r e shy, s e d e n t a r y spiders that occupy a wide variety of d a r k , secretive habitats in n a t u r a l a n d d o m e s t i c situations, usually rock crevices o r hollows u n d e r rocks, u n d e r debris a n d loose bark, or at cave e n t r a n c e s . T h e y a r e c o m m o n in cor­ n e r s a n d niches in a d o b e brick houses and o t h e r domestic s t r u c t u r e s . T h e i r irregular webs a r e large, with thick, very sticky

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

threads. T h e y usually r e m a i n o n their webs, which they c o n t i n u e to e n l a r g e as long as they live. Violin s p i d e r s carry a v e n o m capable of severely injuring h u m a n s ( S c h e n o n e a n d Suárez 1978). T h e v e n o m ' s tissue-destroy­ ing capability h a s b e e n well established. Clinical signs from bites r a n g e from mild necrosis to systemic reactions b u t rarely death. A l t h o u g h t h e v e n o m o u s n e s s of only a few species is r e c o r d e d , it a p p e a r s that all species of t h e g e n u s a r e toxic. Shy and retiring, they never c o m e forth to bite intentionally. Cases of e n v e n o m i z a t i o n , "loxoscelism" (Biicherl 1961), a r e nor­ mally only c a u s e d by specimens that have accidentally crawled into beds o r o n t o clothing a n d that have bitten in defense when c o m p r e s s e d . T h e s e spiders a r e recognized by their small to m e d i u m size (BL 5—20 m m ) , light to m e d i u m - b r o w n color, a n d long, thin legs. T h e back of t h e c a r a p a c e often carries a d a r k o u t l i n e in t h e s h a p e of a violin. They a r e u n i q u e in having t h e six, equalsized eyes f o r m i n g a transverse row, in three diads. T h e legs a n d b o d y a r e thickly clothed with a b u n d a n t , fine, basally feath­ ered hairs lying between long, erect, toothed hairs. A l t h o u g h spiders of t h e g e n u s Loxosceles are known f r o m Africa, they a r e most diverse t h r o u g h o u t t h e Americas. H e r e seventy-four species a r e f o u n d (58 in Mex­ ico a n d C e n t r a l A m e r i c a , 6 in t h e West Indies, 30 in S o u t h America). Loxosceles laeta (fig. 4.4c) is a large species. It h a s gained notoriety because of its t e n d e n c y to live in u r b a n settings, because it h a s been i n t r o d u c e d into n e w areas of t h e world by c o m m e r c e , a n d because of its reputation for b e i n g especially toxic. As a result, its biology h a s b e e n studied in some detail (e.g., Galiano 1967, S c h e n o n e et al. 1970, Lowrie 1980). C o m p l e t e bibli­ ographies o n violin spider biology a n d taxonomy a r e available (Gertsch 1967, Gertsch a n d E n n i k 1983).

References BÜCHERL, W. 1961. Aranhas do género Loxo­ sceles e loxoscelismo na América. Cien. Cult. 13: 213-224. GALIANO, M. E. 1967. Ciclo biológico y desarollo de Loxosceles laeta (Nicolet, 1849) (Araneae, Scytodidae). Acta Zool. Lilloana 23: 431-464. GERTSCH, W.J. 1967. T h e spider genus Loxosceles in South America (Araneae, Scytodidae). Amer. Mus. Nat. Hist. Bull. 136: 1 17-174. GERTSCH, W. J., AND F. ENNIK. 1983. T h e spider

genus Loxosceles in North America, Central America, and the West Indies (Araneae, Loxoscelidae). Amer. Mus. Nat. Hist. Bull. 175: 264-360. LOWRIE, D. C. 1980. Starvation longevity in Loxosceles laeta (Nicolet) (Araneae). Entomol. News 91: 130-132. SCHENONE, H., A. ROJAS, AND H. REYES. 1970.

Prevalence of Loxosceles laeta in houses in central Chile. Amer. J. Trop. Med. Hyg. 19: 564-567. SCHENONE, H., AND G. SUÁREZ. 1978. Venoms of

Scytodidae, genus Loxosceles. In S. Bettini, ed., Arthropod venoms. Springer, Berlin. Pp. 247-275.

HARVESTMEN Opiliones ( = Phalangida). Spanish: Macacos (Mexico). T h e r e is a n i n o r d i n a t e diversity of O p i ­ liones in t h e Neotropical Region (Kaestner 1937), w h e r e t h e majority of t h e world's a p p r o x i m a t e l y 5,000 species a r e f o u n d (Roewer 1923). T h e r e a r e two d o m i n a n t g r o u p s , t h e Cyphopalpitores a n d Laniatores (Martens 1986). T h e former, typified by t h e G a g rellidae, mostly exhibit a small (BL 4—6 mm), simple, oval b o d y s u s p e n d e d in t h e center of t h e immensely long, s l e n d e r (al­ most filamentous) legs, t h e second pair of which a r e usually t h e longest (fig. 4.4d). T h e pedipalps a r e s l e n d e r with weak claws apically, a n d t h e coxae of t h e legs a r e sepa­ rated ventrally by a breast plate. T h e Lania­ tores, best k n o w n in t h e Neotropics by t h e family Gonyleptidae, have relatively s h o r t e r legs of varied length a n d stoutness, t h e h i n d pair often longest a n d heaviest, with very

HARVESTMEN

125

large coxal s e g m e n t s a n d e l a b o r a t e spines or o t h e r excrescences (fig. 4.4e). T h e pedipalps a r e stout, with g r a s p i n g claws at the tip. T h e h a r d body is also often s p i n e d , a n d t h e coxae of t h e legs t o u c h along t h e midline ventrally. B o t h g r o u p s typically pos­ sess eight legs a n d a p r o s o m a divided d o r sally into t h r e e p a r t s by two transverse grooves. T h e a b d o m e n is distinctly seg­ m e n t e d a n d c o n t i n u o u s with t h e p r o s o m a . T h e t h r e e - s e g m e n t e d chelicerae have long blades, a n d t h e p e d i p a l p s a r e leglike b u t always m u c h s h o r t e r t h a n t h e legs. Most species in this o r d e r live in h u m i d retreats, b e n e a t h rocks, in tree b a r k crev­ ices, a n d in niches o n t h e forest floor. O t h e r s r o a m freely o n t h e g r o u n d o r o n tree t r u n k s o r o t h e r vegetation. All p r e f e r s h a d e a n d moist c o n d i t i o n s . T h e i r food consists of o t h e r small i n v e r t e b r a t e s (mites, springtails, even snails). S o m e m a y take only plant d e t r i t u s . H a r v e s t m e n p r o t e c t themselves in a vari­ ety of ways ( C o k e n d o l p h e r 1987). T h e y a r e secretive, often n o c t u r n a l , cryptically col­ o r e d a n d f o r m e d , a n d practice a p p e n d o t o m y (voluntary d r o p p i n g of legs). W h e n d i s t u r b e d , they can eject defensive qui­ ñ o n e s a n d p h e n o l s (Roach et al. 1980) from paired r e p u g n a t o r i a l g l a n d s o n t h e a n t e r i o r e d g e of t h e p r o s o m a . Vonones sayi from Pan­ a m a initially discharges a clear fluid contain­ ing q u i ñ o n e s from its g l a n d s a n d t h e m o u t h a n d t h e n d i p s t h e tips of its forelegs into t h e m i x t u r e a n d b r u s h e s t h e m against t h e of­ f e n d i n g a g e n t (Eisner et al. 1971). Generally, t h e sexes m e e t fortuitously, eggs a r e laid o n moist substrata, a n d n o p a r e n t a l care is exhibited. B u t t h e males of at least o n e species, Zygopachylus albomarginis construct a n d g u a r d a nest of b a r k detri­ tus o n fallen trees which females visit to c o p u l a t e a n d lay t h e i r eggs ( R o d r i g u e z a n d G u e r r e r o 1976). T h e o r d e r is little s t u d i e d in Latin A m e r ­ ica. Soares a n d Soares (1948, 1949, 1955) p r o v i d e a catalog of most of t h e g e n e r a . Ringuelet's (1959) extensive review of t h e

126

A r g e n t i n i a n fauna is of g e n e r a l utility. Many taxa a r e easily r e c o g n i z e d by their beautiful coloration a n d s t r a n g e body forms.

References COKENDOLPHER, J. C. 1987. Observations on the

defensive behaviors of a Neotropical Gonyleptidae (Arachnida, Opiliones). Rev. Arachnol. 7: 59-63. EISNER, T.,

A.

F. KLUGE, J.

E. CARREL, AND

J. MEINWALD. 1971. Defense of phalangid: Liquid repellent administered by leg dab­ bing. Science 173: 650-652. KAESTNER, A. 1937. Ordnung der Arachnida: Opiliones Sunderval = Weberknechte. In W. Kiihkenthal and T. Krumbach, ed., Handbuch der Zoologie. W. Gruyter & Co., Berlin. Band 3, Halfte 2, Lief. 9, (2) Teil. Pp. 385-496. MARTENS, J. 1986. Die Grossgliederung der

Opiliones und die Evolution der Ordnung (Arachnida). Actas 10th Cong. Int. Arach. (Jaca, Spain) 1:289-310. RINGUELET, R. A. 1959. Los arácnidos Ar­

gentinos del orden Opiliones. Mus. Arg. Cien. Nat. "Bernardino Rivadavia," Inst. Nac. Cien. Nat. Zool. Rev. 5: 127-439, pi. 1-20. ROACH, B., T. EISNER, AND J. MEINWALD.

1980.

Defensive substances of opilionids. J. Chem. Ecol. 6: 511-516. RODRÍGUEZ T., C. A., AND S. GUERRERO B., S.

1976. La historia natural y el comporta­ miento de Zygopachylus albomarginis (Cham­ berlain) (Arachnida, Opiliones: Gonyleptidae). Biotropica 8: 242-247. ROEWER, C. F. 1923. Die Weberknechte der

Erde, systematische Bearbeitung der bisher bekannten opiliones. G. Fischer, Jena. SOARES, B. A. M., AND H. E. M. SOARES. 1948.

Monografía dos géneros de Opilióes Neotrópicos. Arq. Zool. Sao Paulo 5: 553-635. SOARES, B. A. M., AND H. E. M. SOARES. 1949.

Familia Gonyleptidae, continuacáo. Arq. Zool. Sao Paulo 7: 151-239. SOARES, B. A. M., AND H. E. M. SOARES. 1955.

Monografía dos géneros de Opilióes Neotrópicos. Arq. Zool. Sao Paulo 8: 225-302.

MITES AND TICKS Acari ( = Acariña, A c a r i d a ) . T h e mites a r e t h e most diverse a n d speciesrich g r o u p of a r a c h n i d s a n d also t h e most

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

difficult to c h a r a c t e r i z e (Krantz 1978, H u g h e s 1959, F l e c h t m a n n 1975). Most a r e very small, s o m e m i n u t e (BL 1 to 2 m m ) . T h e body is fused into o n e piece, with n o separation b e t w e e n t h e p r o s o m a a n d unsegmented o p i s t h o s o m a ( a b d o m e n ) , a n d usu­ ally has an ovoid s h a p e , s o m e w h a t flattened in most ticks, e l o n g a t e o r q u a d r a t e in some mites. T h e m o u t h p a r t s e x t e n d from a par­ tial, headlike s t r u c t u r e anteriorly, f o r m e d from fusion of basal p e d i p a l p a l a n d m o u t h part s e g m e n t s . T h e first active stage of d e ­ velopment is a "larva" that h a s six legs. Once considered a natural group, the mites a n d ticks a r e now t h o u g h t to have multiple origins, a n d at least t h r e e (possi­ bly u n r e l a t e d ) major g r o u p s a r e presently recognized by specialists: Opilioacariformes (Notostigmata), Parasitiformes, and Acariformes. T h e first is a primitive assemblage, c o m p o s e d of comparatively large (1 m m o r m o r e ) , elongate, longlegged, leathery mites, s o m e w h a t resem­ bling h a r v e s t m e n ( H o f f m a n n a n d Vaz­ quez 1986). T h e second a r e medium-sized to large, well sclerotized, a n d with a tracheal system, o p e n i n g t h r o u g h p a i r e d ventrolateral spiracles (stigmata). Many of these a r e parasitic, i n c l u d i n g t h e atypical ticks. T h e A c a r i f o r m e s a r e mostly small and have a poorly f o r m e d tracheal system, and t h e b o d y is often divided into two regions by a transverse furrow. Within each category, major s u b g r o u p i n g s a r e recognized a c c o r d i n g to t h e p r e s e n c e o r absence a n d position of t h e stigmata: hid­ den (Cryptostigmata), a n t e r i o r (Prostigmata), b e t w e e n t h e second a n d f o u r t h coxae (Mesostigmata), n e a r t h e posterior coxae (Metastigmata), dorsally (Notostig­ mata), o r absent (Astigmata). All these g r o u p s a r e well r e p r e s e n t e d in Latin America (Schuster 1969). T h e fauna is immense, a n d t h o u g h m a n y t h o u s a n d s of species a r e n o w k n o w n , these a r e surely only a fraction of those that must exist. Anything a p p r o a c h i n g even a g e n e r a l r e ­ view is well b e y o n d t h e scope of this book.

Only a very few of t h e b e t t e r - k n o w n types of special i m p o r t a n c e ecologically o r eco­ nomically in t h e A m e r i c a n tropics can be treated h e r e . All biotic types a r e r e p r e s e n t e d a m o n g mites. T h e r e a r e free-living a n d parasitic forms on animals a n d plants. Many p a r a ­ sitic forms a r e vectors of diseases (Oldlield 1970). Free-living forms dwell in t h e soil a n d on vegetation w h e r e they feed o n sap a n d tissue fluids, sometimes causing consid­ erable d a m a g e to c r o p s ( J e p p s o n et al. 1975). Some cause galls (Eriophyidae), a n d a few infest flour a n d o t h e r stored g r a i n products. T h e r e a r e m a n y parasites of vertebrates, including h u m a n s . T h e y may be of consid­ erable medical a n d veterinary i m p o r t a n c e (Baker et al. 1956, F l e c h t m a n n 1973). Most are external biters (Dermanyssus), b u t a few b u r r o w into t h e skin (scabies mite) o r lodge d e e p in body cavities (nasal chiggers) o r skin p o r e s (hair follicle mite). T h e y a r e often highly allergenic. Pneumonyssus (Halarachnidae) a n d m e m b e r s of o t h e r families infest t h e lungs a n d respiratory tracts of snakes a n d birds. Many a r e specific ecto­ parasites of bats (e.g., Chirodiscidae, Chirorhynchobiidae, Spelaeorhynchidae, Spinturnicidae) a n d o t h e r characteristic Neotropical m a m m a l s , such as Archemyobia latipilis in o p o s s u m s (Fain et al. 1981). Pollen- a n d nectar-feeding species of Rhinoseius a n d Proctolaelaps (Ascidae) a r e phoretic o n h u m m i n g b i r d s , living in t h e nares a n d using t h e m for t r a n s p o r t a t i o n from flower to flower (Colwell 1979). T h e ticks a r e highly modified bloodsuckers o n all v e r t e b r a t e o r d e r s . O t h e r mites live in bird a n d m a m m a l nests, scavenging o n t h e host's food o r organic detritus t h e r e i n . Such is Hypoaspis dasypus, k n o w n only from a r m a d i l l o b u r ­ rows (Menzies a n d S t r a n d t m a n n 1952). Many a r e associates of o t h e r a r t h r o p o d s (Bischoff d e Alzuet 1978, M a u r i a n d Bischoff d e Alzuet 1972) in various ways, as parasites, p r e d a t o r s , a n d c o n s u m e r s of

MITES AND TICKS

127

e x u d a t e s from t h e host. O t h e r s practice phoresy, using t h e host for t r a n s p o r t only. S o m e of t h e b e t t e r - k n o w n e x a m p l e s of t h e latter a r e Macrocheles, which a r e often very n u m e r o u s on t h e bodies o f d u n g beetles (Evans a n d H y a t t 1963). Mites of diverse g r o u p s l o d g e in t h e t y m p a n i c cavities of m o t h s (Treat 1975). Several kinds of mites a r e also associated with stingless bees ( F l e c h t m a n n a n d d e C a m a r g o 1974). T h e varroa m i t e (Varroa jacobsonii), a n o t o r i o u s apicultural pest in t h e O l d W o r l d a n d n o w established in all major b e e k e e p i n g areas in South A m e r i c a , is associated with t h e honey­ bee. Arrehenurus is a w i d e s p r e a d aquatic ge­ nus, ectoparasitic o n m o s q u i t o larvae a n d pupae. E n t i r e families, for e x a m p l e , A n a l g i d a e , Cheyletidae, D e r m o g l y p h i d a e , a r e a d a p t e d to life o n t h e feathers of b i r d s ( G a u d a n d Atyeo 1979). Ophiomegistus ( A n t e n n o p h o r i dae) a r e f o u n d o n snakes a n d lizards. Iguanacarus (Trombiculidae) lives in t h e nasal fossae of t h e m a r i n e i g u a n a of t h e Galápagos Islands ( V e r c a m m e n - G r a n d j e a n 1965). S o m e major g r o u p s a r e aquatic, such as the primarily m a r i n e s u p e r f a m i l i e s Halacaroidea a n d freshwater H y d r a c h n o i d e a (Cook 1980). T h e latter c a n b e very a b u n ­ d a n t in tropical p o n d s a n d lakes w h e r e they form a p a r t of t h e p l a n k t o n c o m m u ­ nity (Gliwicz a n d Biesiadka 1975).

References BAKER, E. W., T. M. EVANS, D. J. GOULD, W. B. HULL, AND H. L. KEEGAN. 1956. A manual of

parasitic mites of medical or economic impor­ tance. Nat. Pest Contr. Assoc. Tech. Publ. BISCHOFF DE ALZUET, A. 1978. Ácaros asociados

a artrópodos de interés sanitario. Neotropica 24:145-149. COLWELL, R. K. 1979. T h e geographic ecology of hummingbird flower mites in relation to their host plants and carriers. Rec. Adv. Acarol. 2: 461-468. COOK, D. R. 1980. Studies on Neotropical water mites. Amer. Entomol. Inst. Mem. 31: 1-645. EVANS, G. O., AND K. H. HVATT. 1963. Mites of

the genus Macrocheles Latr. (Mesostigmata)

128

associated with coprid beetles in the collection of the British Museum (Natural History). Brit. Mus. Nat. Hist. Bull. 9: 327-401.

Eriophyids Eriophyidae. Spanish: Eriófidos (General), ácaros d e agallas. Gall mites.

u n d e r t h e bracts, w h e r e their feeding causes scarring o n t h e nuts, leading t o r e d u c e d copra yields.

FAIN, A., E. MÉNDEZ, AND F. S. LUKOSCHUS.

1981. Archemyobia (Nearchemyobia) latipilis sp. n. (Acari: Prostigma: Myobiiidae) parasitic on marsupials in Panama and Brazil. Rev. Biol. Trop. 29: 7 7 - 8 1 . FLECHTMANN, C. H. W. 1973. Ácaros de im­

portancia médico-veterinaria. Liv. Nobel, Sao Paulo. FLECHTMANN, C. H. W. 1975. Elementos de

Acarologia. Liv. Nobel, Sao Paulo. FLECHTMANN, C. H. W., AND C. A. DE CAMARGO.

1974. Acari associated with stingless bees (Meliponidae, Hymenoptera) from Brazil. 4th Int. Cong. Acarol. Proc. Pp. 315-319. GAUD, J., AND W. T ATVEO. 1979. Co-évolution

des acariens sarcoptiformes plumicoles et de leurs hótes. Acarologia 21: 291-306. GLIWICZ, Z. M., AND E. BIESIADKA. 1975. Pelagic

water mites (Hydracarina) and their effect on the plankton community in a Neotropical man-made lake. Arch. Hydrobiol. 76: 65-88. HOFFMANN, A., AND M. VÁZQUEZ.

1986. Los

primitivos ácaros Opilioacáridos en México. Fol. Entomol. Méx. 67: 53-60. HUGHES, T E. 1959. Mites or the acari. Athlone, Univ. London, London. JEPPSON, L. R., H. H. KEIFER, AND E. W. BAKER.

1975. Mites injurious to economic plants. Univ. California, Berkeley. KRANTZ, G. W. 1978. A manual of acarology. 2d ed. Ore. St. Univ. Bookstores, Corvallis. MAURI, R., AND A. BISCHOFF DE ALZUET. 1972.

Ácaros asociados a artrópodos. Soc. Argen­ tina Entomol. Rev. 34: 151-159. MENZIES, G C , AND R. W. STRANDTMANN. 1952.

A new species of mite taken from nest of armadillo. Entomol. Soc. Wash. Proc. 54: 265-269. OLDFIELD, G N. 1970. Mite transmission of plant viruses. Ann. Rev. Entomol. 15: 343—380. SCHUSTER, R. 1969. Die terrestrische Milbenfauna Südamerikas in zoogeographischer Sicht. In E. J. Fittkau, J. lilies, H. Klinge, G H. Schwabe, and H. Sioli, eds., Biogeography and ecology in South America. 2: 7 4 1 763. Junk, T h e Hague. TREAT, A. E. 1975. Mites of moths and butter­ flies. Cornell Univ., Ithaca.

T h e s e mites a r e m a r k e d by structural sim­ plification: they a r e wormlike a n d so small as to b e barely visible to t h e u n a i d e d eye (BL 0.2 m m ) ; only t h e a n t e r i o r two pairs of legs a r e d e v e l o p e d ; a tracheal system is absent, a n d r e s p i r a t i o n is by c u t a n e o u s diffusion. T h e cuticle bears conspicuous fine rings over t h e whole body. All species of e r i o p h y i d s a r e obligate plant feeders a n d constitute t h e m a i n fam­ ily of mites injurious t o cultivated plants. Physical d a m a g e is caused by mechanical feeding a n d inoculation of h a r m f u l vi­ ruses. Host r e s p o n s e s t o feeding a r e leaf curling a n d adventitious tissue formation, including twigs, blisters, a n d galls. A serious a n d w i d e s p r e a d pest species is the citrus b u d mite (Eriophyes [= Acería] sheldoni) (fig. 4.5a). It is k n o w n in all citrusgrowing areas of C e n t r a l a n d S o u t h A m e r ­ ica. T h e species infests t h e b u d , d e v e l o p i n g blossoms, a n d area b e n e a t h t h e b u t t o n of citrus fruits as well as leaf axil b u d s . T h e growing tissue m a y be completely d e ­ stroyed o r a l t e r e d t o form s t u n t e d o r misshapen s t r u c t u r e s , especially in t h e fruit, which m a y a s s u m e g r o t e s q u e shapes, becoming unfit for m a r k e t . A n o t h e r very injurious species is t h e coconut mite (Eriophyes guerreronis) (Moore and A l e x a n d e r 1987). Populations d e v e l o p

Reference MOORE, D., AND L. ALEXANDER. 1987. Aspects

of migration and colonization of the coconut palm by the coconut mite, Eriophyes guer­ reronis (Keifer) (Acari: Eriophyidae). Bull. Entomol. Res. 77: 641-650.

House Dust Mites Pyroglyphidae,

Dermatophagoides.

H o u s e dust, consisting of h u m a n a n d p e t skin particles, food fragments, hair, cotton, paper, wool a n d synthetic fibers, soil, a n d like material, provides food a n d shelter for a c o m m u n i t y of exceedingly m i n u t e o r g a n ­ isms d o m i n a t e d by a r t h r o p o d s , including m a n y kinds of mites, a n d fungi (Van Bronswijk 1979, 1981). A m o n g t h e latter are most c o m m o n l y t h e h o u s e d u s t mites of the g e n u s Dermatophagoides, which is r e p r e ­ sented in Latin America by both t h e wide­ s p r e a d species D.farinae (fig. 4.5b) a n d t h e localized D. pteronyssinus. D. neotropicalis is a species sometimes also c o n s i d e r e d a p a r t of this complex (Fain a n d Van Bronswijk 1973). As m a n y as 3,000 such mites m a y inhabit a g r a m o f d u s t (Arlian et al. 1979). T h e y a r e m o r e c o m m o n in t h e d u s t o n mattresses a n d b e d r o o m floors t h a n else­ w h e r e in t h e h o u s e . Dust h a s long b e e n k n o w n to cause allergies a n d a s t h m a in sensitive p e r s o n s .

VERCAMMEN-GRANDJEAN, P. H. 1965. Iguana­

carus, a new subgenus of chigger mite irom nasal fossae of the marine iguana in the Gala­ pagos Islands, with a revision of the genus Vatacarus Southcott (Acariña, Trombiculidae). Acarologia 7 (suppl.): 266-274.

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

Figure 4.5 MITES, (a) Citrus bud mite (Eriophyes sheldoni, Eriophyidae). (b) American house dust mite (Dermatophagoides farinae, Pyroglyphidae). (c) Spider mite (Tetranychus telarius, Tetranychidae). (d) Mold mite (Tyrophagus putrescentiae, Acaridae). (e) Soil mite (Oribatula minuta, Oribatulidae). (f) Follicle mite (Demodex folliculorum, Demodicidae).

MITES AND TICKS

129

N u m e r o u s studies have r e p o r t e d a correla­ tion between s y m p t o m s a n d t h e presence of h o u s e d u s t mites, a l t h o u g h a direct relationship is n o t yet clearly established (Leeks 1973). W h a t little h a s b e e n d o n e on these mites a n d their effect on health in Latin A m e r i c a h a s c e n t e r e d in Colombia (Mulla a n d Sánchez M e d i n a 1980, Charlet et al. 1979) a n d Chile (Casanueva a n d Artigas 1985).

References ARLIAN, L. G., 1. L. BERNSTEIN, C. L.JOHNSTON,

AND J. S. GALLAGHER. 1979. Ecology of house dust mites and dust allergy. Rec. Adv. Acarol. 2: 185-195. CASANUEVA, M. E., AND J. N. ARTIGAS.

1985.

Distribución geográfica y estacional de los ácaros del polvo de habitación en Chile. Gayana. Zool. 49: 3 - 7 5 . CHARLET, L. D., M. S. MULLA, M. SANCHEZMEDINA, AND M. A. REYES. 1979. Species

composition and population trends of mites in various climatic zones of Colombia. J. Asthma Res. 16: 131-148. FAIN, A., AND J. E. M. H. VAN BRONSWIJK. 1973.

On a new species of Dermalophagoides (D. neolropicalis) from house dust, producing both normal and heteromorphic males (Sarcoptiformes: Pyroglyphidae). Acarologia 15: 181-187. LECKS, H. 1. 1973. T h e mite and house dust allergy: A review of current knowledge and its clinical significance. Clin. Pediat. 12: 514— 517. MULLA, M. S., AND M. SÁNCHEZ-MEDINA,

ed.

1980. Domestic acari of Colombia, bionomics, ecology and distribution of allergenic mites, their role in allergic disease. COLCIENC1AS, Bogotá. VAN BRONSWIJK, J. E. M. H. 1979. House dust as an ecosystem. Rev. Adv. Acarol. 2: 167-172. VAN BRONSWIJK, J. E. M. H. 1981. House dust biology for allergists, acarologists, and mycologists. NIB Publ., Zeist, The Netherlands.

Spider Mites Acarida, T e t r a n y c h i d a e . Spanish: A r a ñ i t a s , a r a ñ a s rojas (General). Portuguese: Ácaros d e teia, ácaros rajados, ácaros v e r m e l h o s (Brazil). T h e s e mites often live a m i d masses of fine silk w e b b i n g o n t h e u n d e r s i d e s of leaves

130

which they spin from glands located in t h e p r o s o m a , a habit e a r n i n g t h e m their ver­ nacular n a m e (Baker 1979, Helle a n d Sabelis 1985). T h e feeding of large n u m ­ bers of these mites o n commercial plants often causes severe d a m a g e . Leaves de­ velop pale blotches a n d may eventually totally d r y u p . Plants lose their vigor a n d die (Huffaker et al. 1969). Consequently, many species a r e recognized pests in Latin America (Pritchard a n d B a k e r 1955). Sec­ o n d to t h e eriophyids, they a r e probably the most agriculturally i m p o r t a n t plantfeeding g r o u p of mites. A c o m p l e x of species, t h e two-spotted spider mite g r o u p (Tetranychus bimaculatus, T. telarius [fig. 4.5c], a n d T. cinnabarinus) is especially trouble­ some to cotton, beans, a n d vegetable crops in general, as well as to flowers, especially in g r e e n h o u s e s (Vereau et al. 1978). S p i d e r mites a r e g r e e n , yellow, o r a n g e , or r e d a n d small (BL always less than 1 m m ) , often with d a r k blotches o n either side of t h e d o r s u m .

sclerotized, d a r k - p i g m e n t e d , h a r d - b o d i e d mites, most with a body length of less t h a n 1 millimeter. Some have e n l a r g e d m o u t h parts, a n d they often have h i n g e d , wing­ like structures e x t e n d i n g from t h e sides of the body u n d e r which they can tuck their legs to form a c o m p a c t ball for protection. T h e leg segments a r e sometimes swollen, giving t h e legs a n o d u l a r a p p e a r a n c e .

T h e principal o f f e n d i n g species in Latin America b e l o n g i n g to this category a r e Tyrophagus putrescentiae (Acaridae) (fig. 4.5d), Carpoglyphus lactis ( C a r p o g l y p h i d a e ) , and Pyemotes ventricosus (Pyemotidae).

(Acari) of Antarctica with special reference to the Antarctic Peninsula and South Shetland Islands. Pacific Ins. 7: 453-468. WILLIAMS, E. C. 1941. An ecological study of the floor fauna of the Panamanian rain forest. Chicago Acad. Sci. Bull. 6: 63-124.

Reference

Follicle Mites

FLECHTMANN, C. H. W. 1983. Ácaros de im­

Demodicidae, Demodex. Portuguese: Cravos (Brazil).

portancia agrícola. Liv. Nobel, Sao Paulo.

References BAKER, E. W. 1979. Spider mites revisited—a review. Rec. Adv. Acarol. 2: 387-394. HELLE, W.,

ucts, feeding directly on t h e substance o r secondarily o n fungi g r o w i n g on it. A few are parasitoids of insects living on t h e substratum. T h e y a r e all very small mites (BL mostly 1 m m o r less), pale a n d with long body setae. T h e y may d e v e l o p e n o r m o u s popula­ tions a n d be very destructive both in indus­ trial (warehouses, g r a n a r i e s , holds of ships) a n d h o u s e h o l d (larders, c u p b o a r d s , medicine chests) e n v i r o n m e n t s . T h e y may literally replace t h e s u b s t r a t u m with t h e mass of their collective bodies. Some a r e known to bite h u m a n s w o r k i n g a r o u n d stored p r o d u c t s , causing m i n o r rashes ("baker's itch").

AND M.

W. SABELIS, eds.

1985.

Spider mites: Their biology, natural enemies and control. Vol. 1, Pt. A, 405; Pt. B, 458. Elsevier, Amsterdam.

Soil Mites Oribatulatidae ( = Cryptostigmata). Moss mites, beetle mites.

Acariformes, various families, including Acaridae and Carpoglyphidae.

Soil mites (Balogh 1988) form a very large and c o m p l e x g r o u p , c o m p r i s e d of m a n y families a n d a b o u t 4 5 0 species as presently known in S o u t h A m e r i c a , far short of t h e probable actual n u m b e r . Balogh (1972) lists about 2 7 0 Neotropical g e n e r a , includ­ ing cosmopolitan a n d pan-tropical taxa. These mites a r e very a d a p t a b l e to environ­ mental stresses a n d r e p r e s e n t o n e of t h e few types of terrestrial a r t h r o p o d s surviv­ ing on t h e h a r s h fringes of Antarctica a n d the most southerly Atlantic islands (Wallwork 1965). By their very n u m b e r s , they play an i m p o r t a n t role in t h e d e c o m p o s i ­ tion of o r g a n i c substances in t h e soil (Wil­ liams 1941; see special habitats, c h a p . 2).

Q u i t e a n u m b e r of mite species a r e as­ sociated with stored food a n d fiber prod-

Soil mites, such as t h e c o m m o n Oribatula minuta (fig. 4.5e), a r e generally very well

HUFFAKER, C. B., J. A. MCMURTRY, AND M. VAN

DE VRIE. 1969. T h e ecology of tetranychid mites and their natural control. Ann. Rev. Entomol. 14: 125-174. PRITCHARD, A. E., AND E. W. BAKER. 1955. A

revision of the spider mite family Tetrany­ chidae. Pac. Coast Entomol. Soc. Mem. 2: 1472. VEREAU, W. V, M. CUEVA, AND D. OJEDA. 1978.

Biología de la "arañita roja del algodonero" Tetranychus cinnabarinus (Boisduval) (Acariña, Tetranychidae). Rev. Peruana Entomol. 21: 50-54.

Stored Product Mites

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

References BALOGH, J. 1972. The oribatid genera of the world. Akadémiai Kiadó, Budapest. BALOGH, J. 1988. Oribatid mites of the Neo­ tropical Region. 1. The soil mites of the world. Vol. 2. WALLWORK, J. A. 1965. The

Cryptostigmata

T h e s e mites (Desch a n d N u t t i n g 1971) a r e all ectoparasites of m a m m a l s a n d r e m a r k ­ ably modified for residence in pits in t h e skin; rarely, they secondarily invade t h e lymphatic a n d circulatory system. T h e y seem to be rigidly host specific (Nut­ ting 1974), certain species b e i n g k n o w n from several Latin A m e r i c a n animals, in­ c l u d i n g t h e domestic h o r s e (Demodex equi), d o g (D. canis), s h e e p (D. ovis), pig (D. phylloides), goat (D. caprae), cat (D. cati), a n d cow (D. bovis). M o r e species a r e b e i n g discov­ e r e d in wild m a m m a l s as well, for e x a m p l e , guinea pigs, mice, m o n k e y s (Lebel a n d Nut­ ting 1973), a n d bats (Desch a n d N u t t i n g 1972). T h e entire life cycle is s p e n t on t h e host, all stages being f o u n d in skin pustules a n d hair follicles on t h e body, especially a b o u t t h e face ( Q u i n t e r o 1978). T r a n s f e r to a new host normally requires close contact, possibly in infancy o r in t h e nest. H u m a n s also h a r b o r a particular species,

MITES AND TICKS

131

D. folliculorum (fig. 4.5f). T h e s e a r e m i n u t e (BL 0.1—0.4 m m ) , with a n e l o n g a t e , almost w o r m l i k e body. T h e o p i s t h o s o m a is trans­ versely striated. T h e legs a r e s h o r t a n d stubby w i t h o u t setae a n d a r e located close t o g e t h e r anteriorly (Desch a n d N u t t i n g 1977). T h e mite inhabits t h e hair follicles a n d sebaceous g l a n d s , particularly a r o u n d the nose a n d eyelids b u t s o m e t i m e s else­ w h e r e , such as t h e scalp. Its p r e s e n c e is usually i n n o c u o u s b u t h a s b e e n k n o w n to i n d u c e acnelike c o n d i t i o n s . It is s u r p r i s ­ ingly c o m m o n ; p r o b a b l y the majority of the human population anywhere unknowingly serves as host t o this species. I n animals, severe s y m p t o m s c a n d e ­ velop w h e n t h e mites i n t e r f e r e with secre­ tion a n d w h e n they i n v a d e t h e skin a n d b l o o d s t r e a m ( N u t t i n g 1975).

References DESCH,

D.

E.,

AND W. B.

NUTTING.

1971.

Demodicids (Trombidiformes: Demodicidae) of medical and veterinary importance. 3d Int. Cong. Acarol. Proc. Pp. 499-505. DESCH, D. E., AND W. B. NUTTING. 1972. Para­

sitic mites of Surinam. VII: Demodex longissiw n. sp. from Carollia perspicillata and D. molossi n. sp. from Molossvs molossus (Demo­ dicidae: Trombidiformes); Meibomian com­ plex inhabitants of Neotropical bats (Chiroptera). Acarologia 14: 35—53. DESCH, C. E., AND W. B. NUTTING. 1977. Mor­

phology and functional anatomy of Demodex folliculorum (Simon) of man. Acarologia 19: 422-462. LEBEL,

R.

R.,

AND W.

B.

NUTTING.

1973.

Demodectic mites of subhuman primates. I: Demodex saimirí sp. n. (Acari: Demodicidae) from the squirrel monkey, Saimiri sciureus. J. Parasit. 59: 719-922. NUTTING, W. B. 1974. Synhospitaly and speciation in the Decodicidae [sic] (Trombidi­ formes). 4th Int. Cong. Acarol. Proc. Pp. 2 6 7 272. NUTTING, W. B. 1976. Pathogenesis associated with hair follicle mites (Acari: Demodicidae). Acarologia 17: 493-506. QUINTERO, M. T. 1978. Frecuencia de ácaros en especies de animales domésticos. Vet. Mex. 9: 111-114.

Scabies Mite Sarcoptidae, Sarcoptes scabiei. Spanish: Acaro d e la sarna. Portuguese: A c a r i a n o d a sarna (Brazil). T h e r e a r e several strains o r subspecies of this mite (Mellanby 1943, 1985), a d a p t e d to different m a m m a l hosts (horses, dogs, pigs, s h e e p , cattle) i n c l u d i n g h u m a n s , in which it can cause a g e n e r a l skin affliction called scabies o r m a n g e (Arlian 1989, Gor­ d o n a n d U n s w o r t h 1947). T h e species s p e n d s its e n t i r e life cycle o n t h e host. Females a r e t h e infective stage. T h e y lay their eggs in t u n n e l s m a d e by b u r r o w i n g t h r o u g h t h e s u b c u t a n e o u s layers of t h e skin. T h e adults live a m o n t h o r m o r e . T h e b u r r o w i n g a n d f e e d i n g (on blood) of t h e mites of all stages cause e x t r e m e itching. T h e sinuous t u n n e l s a r e n e a r t h e skin's surface a n d c a n b e seen as delicate gray lines, o n h u m a n s usually b e t w e e n t h e

fingers a n d toes, b e h i n d t h e k n e e , a n d o n the genitalia. Pimples o r vesicles m a y form on the affected skin which when scratched, become infected a n d cause sores a n d scabs. Animals may d e v e l o p large areas of leath­ ery e n c r u s t m e n t s a n d lose most of their body hair ( m a n g e , sarna pira). T h e s e a r e very small mites (females 0.4 m m in d i a m e t e r , males 0.2 m m ) . T h e y a r e r o t u n d with very short, stubby legs. T h e third pair in t h e male a n d t h e t h i r d a n d fourth pairs in t h e female (fig. 4.6a) a r e tipped with a very long bristle; t h e o t h e r legs a r e t i p p e d with small, stalked suckers. T h e i n t e g u m e n t is coarsely striated a n d bears spinelike projections of two sizes o n the posterior half of the d o r s u m . T h e species is cosmopolitan in distribu­ tion. Infestations s p r e a d t h r o u g h close con­ tact b e t w e e n infected hosts. Sarcoptic mange in livestock causes r e d u c t i o n in meat, milk, a n d wool p r o d u c t i o n a n d even the d e a t h of severely afflicted animals.

References ARLIAN, L. G. 1989. Biology, host relations, and epidemiology of Sarcoptes scabiei. Ann. Rev. Entomol.34: 139-161. GORDON, R. M., AND K. UNSWORTH. 1947. A

review of scabies since 1939. Carib. Med. J. 9: 56-71. MELLANBY, K. 1943. Scabies. Oxford Univ. Press, London. MELLANBY, K. 1985. Biology of the parasite [scabies mite]. In M. Orkin and H. 1. Maibach, eds., Cutaneous infestations and insect bites. Marcel Dekker, New York. Pp. 9-18.

Chiggers

Figure 4.6 MITES AND TICKS, (a) Scabies mite (Sarcoptes scabiei, Sarcoptidae), female. (b) Common, or "sweet potato" chigger (Eutrombicula batatas, Trombiculidae). (c) cayenne tick (Amblyomma cajennense, Ixodidae), male, (d) Tropical horse tick (Dermacentor nitens, Ixodidae), male, (e) Fowl tick (Argas miniatus, Argasidae).

132

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

Trombiculidae, Eutrombicula. Spanish: Coloradillos, bichos colorados (General); patatas ( S u r i n a m ) ; callocallo, isangos (Peru); celembas (Ecuador). Náhuatl: T l a l z a h u a t l (Mexico). Portuguese: Bichos colorados. Tupi-Guaraní: Micuims (Brazil). French: Bétes r o u g e s (Cayenne). Harvest mites, red bugs, r e d mites. Worldwide, various g e n e r a of t h e family Trombiculidae a r e t h e i n f a m o u s biting

mites k n o w n as chiggers (Sasa 1961, W h a r ton a n d Fuller 1952). A m o n g t h e eightyseven g e n e r a in America ( B r e n n a n a n d Goff 1977, 1978), t h e m a i n o f f e n d e r is Eutrombicula, whose larvae attach to h u ­ m a n s , causing severe, itching rashes (trombidiosis) a n d o t h e r allergenic reactions. Parascoschoengastia (Euschoengastia) nunezi has b e e n responsible for d e r m a t o s i s in Mexico ( A n d r a d e 1947). In the n o r m a l life cycle, t h e larvae feed o n m a m m a l s , birds, o r reptiles (Wrenn a n d Loomis 1984). After e n g o r g i n g , they d r o p off p e r m a n e n t l y to c o n t i n u e d e v e l o p m e n t as free-living, p r e d a c e o u s terrestrial mites ( n y m p h s a n d adults). T h e adults a r e fairly large (BL 2 - 3 m m ) , frequently b r i g h t r e d or o r a n g e , a n d covered with a velvety pelage of feathery hairs. T h e y a r e usually found j u s t b e n e a t h t h e surface of loose soil, in cracks, b u r r o w s , leaf litter, h u m u s , or decaying wood w h e r e they feed o n t h e eggs a n d early stages of o t h e r m i n u t e a r t h r o p o d s , such as springtails. Structurally, t h e six-legged larval chig­ gers a r e recognizable u n d e r high magnifi­ cation by a pair of u n i q u e k n o b b e d o r p l u m o s e hairs arising from a r e c t a n g u l a r plate o n t h e a n t e r i o r - d o r s a l surface of the body (idiosoma; Goff et al. 1982, W h a r t o n et al. 1951). T h e m o u t h p a r t s consist of a pair of heavy piercing chelicerae a n d a pair of palpi, which t o g e t h e r give t h e a p p e a r ­ ance of a h e a d . C o n t r a r y to p o p u l a r belief, the chigger does not b u r r o w into t h e skin o r imbed even this false h e a d . Only the tips of the medial m o u t h p a r t e l e m e n t s a r e in­ serted. T h e salivary secretions contain pow­ erful histolytic enzymes that b r e a k d o w n d e e p epithelial cells. T h e tissue h a r d e n s a r o u n d t h e chelicerae, f o r m i n g a feeding canal (stylostome), t h r o u g h which the host's dissolved cells a n d l y m p h a r e s i p h o n e d (Allred 1954). After the chigger d e p a r t s , it is the persistence of this foreign object that p r o d u c e s the long-lasting itching a n d irrita­ tion so characteristic of its bite. There

a r e nearly

eighty

species of

MITES AND TICKS

133

Eutrombicula in Latin A m e r i c a (Loomis a n d W r e n n 1984, H o f f m a n n 1970). Only a few are r e g u l a r h u m a n pests, a n d these n o r ­ mally attack lizards, snakes, g r o u n d dwelling birds, o r m a m m a l s . Species of o t h e r g e n e r a with r e g u l a r m a m m a l hosts seem rarely to cause a n allergic reaction w h e n they d o attach t o m a n . N o n e t r a n s m i t p a t h o g e n i c o r g a n i s m s . However, Leptotrombidium, k n o w n vectors of rickettsioses else­ w h e r e , a n d Pseudoschoengastia c a n cause d e r m a t o s i s a n d a r e potentially trouble­ some. T h e most c o m m o n chiggers attacking h u m a n s t h r o u g h o u t A m e r i c a b e l o n g to t h e Eutrombicula alfreddugesi g r o u p , c o m p r i s e d of several poorly d i s t i n g u i s h e d species (Wil­ liams 1946). T h e i r larvae a r e a b u n d a n t in transition areas b e t w e e n forest a n d grass­ land, a l o n g s w a m p m a r g i n s , a n d from sea level to nearly 3 0 0 0 m e t e r s elevation. T h e y normally parasitize almost a n y terrestrial v e r t e b r a t e a n d readily t r a n s f e r t o m a n . A n o t h e r w i d e s p r e a d species of major medical i m p o r t a n c e t h r o u g h o u t Latin A m e r i c a is t h e so-called sweet p o t a t o chigger, Eutrombicula batatas (fig. 4.6b) ( M i c h e n e r 1946). T h i s is a n a n i m a l of sunlit places, t h e u n f e d larvae e m e r g i n g o n t o t h e surface of t h e soil s o m e t i m e s in large n u m b e r s , especially in grassy areas. It m a y b e c o m e very a b u n d a n t a r o u n d h o m e s a n d villages w h e r e d o m e s t i c ani­ mals, particularly chickens (Canfalonieri a n d d e C a r v a l h o 1973), a r e n u m e r o u s . T h e n o r m a l hosts a r e principally birds a n d small terrestrial m a m m a l s .

BRENNAN, J. M., AND M. L. GOFF, 1978. T h r e e

new monotypic genera of chiggers (Acari: Trombiculidae) from South America. J. Med. Entomol. 14: 541-544. CANFALONIERI, U. E. C., AND L. P. DE CARVALHO.

1973. Ocorréncia de Trombicula (Eutrombicula) batatas (L.) em Callus gallus domesticus L. no estado do Rio de Janeiro (Acariña, Trombicu­ lidae). Rev. Brasil. Biol. 33: 7-10. GOFF, M. L., R. B. LOOMIS, W. C. WELBOURN,

AND W. J. WRENN. 1982. A glossary oí chigger terminology (Acari: Trombiculidae). J. Med. Entomol. 19: 221-238. HOFFMANN, A. M. 1970. Estudio monográfico de los trombicúlidos de México (Acariña: Trombiculidae). Primera parte. Ese. Nac. Cien. Biol. México An. 18: 191-263. LOOMIS, R. B., AND W. J. WRENN. 1984. System-

atics of the pesi chigger genus Eutrombicula (Acari: Trombiculidae). Acarology 6(1): 1 5 2 159. MICHENER, C. D. 1946. Observations on the

habits and life history of a chigger mite, Eutrombicula batatas (Acariña: Trombiculi­ dae). Entomol. Soc. Amer. Ann. 39: 101-118. SASA, M. 1961. Biology of chiggers. Ann. Rev. Entomol. 6: 221-244. WHARTON, G. W., AND H. S. FULLER. 1952. A

manual of the chiggers. Entomol. Soc. Wash. Mem. 4: 1-185. WHARTON, G. W., D. W. JENKINS, J. M. BRENNAN, H. S. FULLER, G. M. KOHLS, AND C. B. PHILLIP.

1951. T h e terminology and classification of trombiculid mites (Acariña: Trombiculidae). J. Parasitol. 37: 13-31. WILLIAMS, R. W. 1946. A contribution to our

knowledge of the bionomics of the common North American chigger, Eutrombicula al­ freddugesi (Oudemans), with a description oí a rapid collecting method. Amer. J. Trop. Med. 26: 243-250. WRENN, W. J., AND R. B. LOOMIS. 1984. Host

selectivity in the genus Eutrombicula (Acari: Trombiculidae). Acarology 6(1): 160—165.

References

Ticks

ALLRED,

I x o d o i d e a . Spanish: G a r r a p a t a s (General); c o n c h u d a s , plateadas, pinolillos (Mexico). Portuguese: C a r r a p a t o s , c a r r a p a t o s pólvoras, c a r r a p a t o s fogo, c a r r a p a t i n h o s (Brazil-larvae). Náhuatl: M a z a a t e m i m e h , sing, mazaatémitl (Mexico).

D. M. 1954. Observations on the

stylostome (feeding tube) of some Utah chig­ gers. Utah Acad. Sci. Arts & Letters Proc. 3 1 : 61-63. ANDRADE, R. M. 1947. Trombididiasis por Neo-

schóngastia nuñezi Hoffmann, Med.'(Mexico) 77: 219-240.

1944. Gaceta

BRENNAN, J. M., AND M. L. GOFF. 1977. Keys to

the genera of chiggers of the Western Hemi­ sphere (Acariña: Trombiculidae). J. Parasitol. 63: 554-566.

134

Ticks c o m p r i s e a n isolated a n d very special­ ized g r o u p of mites d i s t i n g u i s h e d by well-

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

developed, multifaceted spiracles situated lateral to t h e t h i r d coxae o r b e h i n d t h e fourth coxae ( O b e n c h a i n a n d G a l u n 1982, Oliver 1989). T h e i n t e g u m e n t is leathery to hard, a n d they a r e all considerably larger than most mites (BL 2—5 m m ) . T h e i r m o u t h p a r t s a r e u n i q u e : t h e digits of t h e chelicerae have lateral teeth, a n d between the p e d i p a l p s is a m e d i a n holdfast o r g a n with r e c u r v e d t e e t h (called t h e h y p o s t o m e ) . T h e first tarsus h a s a dorsal heat-sensitive organ in a large pit (Haller's o r g a n ) to assist in finding v e r t e b r a t e hosts o n which they feed. Ticks a n c h o r o n t o t h e host's skin by the h y p o s t o m e a n d d o n o t d r o p off until they finish feeding. If forcibly r e m o v e d , t h e hypostome ( e r r o n e o u s l y t h o u g h t to b e t h e head by most people) m a y r e m a i n in t h e wound a n d fester. After molting, ticks wait o n t h e tips of twigs o r leaves, forelegs o u t s t r e t c h e d a n d ready to snag a n y animal b r u s h i n g past. On t h e potential host's a p p r o a c h , t h e legs wave frantically in r e s p o n s e to various stimuli: primarily to c a r b o n dioxide ex­ haled d u r i n g r e s p i r a t i o n b u t also to vibra­ tion, body heat, o r t h e host's s h a d o w pass­ ing over t h e m . Two m a i n families of ticks a r e recog­ nized, t h e " h a r d ticks" (Ixodidae) a n d "soft ticks" (Argasidae). H a r d ticks a r e flat­ tened, a n d all stages b e a r a thick, t o u g h plate (scutum) dorsally. T h i s plate covers only about half t h e a n t e r i o r e n d of t h e body in larvae, n y m p h s , a n d females; t h e entire body in males. It may o r may not be o r n a m e n t e d with silver. Soft ticks a r e baglike, w i t h o u t plates in postlarval stages. T h e m o u t h p a r t s a r e h i d d e n by t h e over­ hanging body ( c a m e r o s t o m a ) . T h e y a r e primarily nest i n h a b i t a n t s a n d parasites of birds a n d small m a m m a l s (mainly bats) in semitropical a n d tropical areas. T h e y a r e very resistant t o water loss a n d a r e charac­ teristic of d e s e r t faunas. Ticks a r e ectoparasitic in all stages, feed>ng primarily o n t h e blood of m a m m a l s , birds, reptiles, a n d a m p h i b i a n s . Usually,

individuals attach to different host species in their various stages (larva, n y m p h , adult) a n d feed only o n c e o n each. S o m e , however, r e m a i n o n a single host t h r o u g h ­ out their lifetime. T h e body, especially that of t h e female, is elastic a n d capable o f e n o r m o u s distension w h e n t h e s t o m a c h fills with blood o r eggs. Specimens m a y reach t h e size of a g r a p e after 5 to 6 days of feeding. Adult males of some species a p ­ parently d o not feed at all. T h e r e a r e m a n y m o r e ixodid t h a n argasid species (114 a n d 5 8 , respectively) in the Neotropics (Keirans pers. c o m m . ) . T h e g e n e r a a r e Ixodes, Dermacentor (including Anocenter; Yunker et al. 1986), Haemaphysalis, Boophilus, Amblyomma, Rhipicephalus, a n d Aponomma. Several h a r d tick species a r e of special i m p o r t a n c e . T h e s o u t h e r n cattle tick (Boo­ philus microplus) is prevalent over most of Latin America. It may b e very a b u n d a n t in some areas (Rawlins 1979) a n d very trou­ blesome to cattle, t r a n s m i t t i n g babesiosis. T h e tropical h o r s e tick (Dermacentor nitens, fig. 4.6d) is distributed from Mexico t o A r g e n t i n a a n d o n t h e G r e a t e r Antilles. It infests t h e ears of its hosts (horses mainly but also cattle, deer, a n d goats) w h e r e it u n d e r g o e s its c o m p l e t e d e v e l o p m e n t . Amblyomma is t h e largest g e n u s , with fifty Neotropical species ( J o n e s et al. 1972). Many exhibit beautiful colors a n d o r n a m e n t a t i o n . T h e y a r e often large, flat, a n d almost circular in outline. Hosts a r e varied, usually m a m m a l s , b u t also reptiles, including turtles (Ernst a n d E r n s t 1977), a n d often birds, in t h e larval a n d n y m p h a l stages. T h e cayenne tick (Amblyomma cajennense, fig. 4.6c), k n o w n as mostacilla o r carrapato estrela, is a general nuisance in all p a r t s of t h e Neotropics w h e r e it m e n a c e s b o t h m a n a n d livestock ( H o f f m a n n 1962). T h e lar­ vae may swarm in t h o u s a n d s in grass a n d low h e r b a g e . Very little h a s b e e n published o n its n a t u r a l biology ( D r u m m o n d a n d W h e t s t o n e 1975), a l t h o u g h m e t h o d s for

MITES AND TICKS

135

artificial c u l t u r e have b e e n w o r k e d o u t (Travassos a n d Vallejo-Freire 1944). T h e tropical b o n t tick (A. variegatum) has b e e n i n t r o d u c e d from Africa to t h e Carib­ b e a n , w h e r e it is associated with s t r e p t o thricosis, a bacterial skin disease of livestock (Garris 1987), a n d h e a r t w a t e r (caused by the rickettsia Cowdria ruminantium). Ixodes pararicinus (once confused with the E u r o p e a n castor b e a n tick, / . ricinus) is k n o w n widely as a p a r a s i t e of cattle in t h e s o u t h e r n half of S o u t h A m e r i c a (Keirans et al. 1985). Soft tick g e n e r a in Latin A m e r i c a a r e Otobius, Antricola, Argas, Nothoaspis, a n d Ornithodoros ( J o n e s a n d Clifford 1972). Notable r e g i o n a l taxa a r e as follows. Argas a r e small a n d f o u n d o n o d d hosts. T h e twenty Latin A m e r i c a n species a r e mostly associated with b i r d s (owls, fowl, etc.) ( H o o g s t r a a l et al. 1979). A. persicus is t h e c o s m o p o l i t a n fowl tick ("adobe tick," "tam­ pan") a n d o n e of t h e m o s t i m p o r t a n t poultry parasites. A. miniatus (fig. 4.6e), however, may m o r e often be t h e o f f e n d e r in Latin A m e r i c a generally, A. moreli in Peru (Keirans et al. 1979). A. transversus attaches to t h e neck a n d t h r o a t skin of giant G a l á p a g o s tortoises ( H o o g s t r a a l et al. 1973). All a r e restricted to dry niches in d e s e r t to s a v a n n a life zones. Ornithodorus talaje a n d O. rudis a r e com­ m o n N e o t r o p i c a l species. T h e y feed o n wild r o d e n t s a n d most d o m e s t i c animals a n d m a n a n d a r e vectors o f t h e r e l a p s i n g fever s p i r o c h e t e Borrelia in G u a t e m a l a , P a n a m a , a n d C o l o m b i a . O. darwini a n d O. galapagensis h a v e b e e n d e s c r i b e d from t h e G a l á p a g o s i g u a n a s (Kohls et al. 1969, K e i r a n s et al. 1980). Antricola a r e f o u n d o n bats o r in their g u a n o . M e m b e r s of b o t h tick families a r e p u r ­ veyors of n u m e r o u s serious diseases of man and domestic animals (Arthur 1961, H o o g s t r a a l 1981). Particularly serious to h u m a n h e a l t h in Latin A m e r i c a a r e spot­ ted fevers c a u s e d by rickettsial o r g a n i s m s ( H o o g s t r a a l 1967) a n d r e l a p s i n g fevers

136

b r o u g h t on by Borellia. Rocky M o u n t a i n spotted fever is e n d e m i c in m a n y parts of Latin America (Mexican spotted fever, fiebre m a n c h a d a , Tobia fever, Sao Paulo fever) w h e r e it may be t r a n s m i t t e d by almost any tick that lives o n t h e m a m ­ malian hosts for t h e p a t h o g e n s . Extensive use of o n e major vector, Amblyomma cajennense, has b e e n applied to a t t e m p t s to develop a vaccine for this virulent disease (Travassos a n d Vallejo-Freire 1944). Tickb o r n e viral diseases seem not to be a regional p r o b l e m , b u t in general, these a r e not well studied in Latin America. A m o n g animals, domestic a n d wild, ticks transmit Texas cattle fever or red water fever (tristeza) caused by t h e proto­ zoan Babesia bigemina, in various parts of the A m e r i c a n tropics. T h e vector is usually Boophilus microplus. T h e disease destroys the red blood cells a n d has a very high mortality rate. S o m e n a t u r a l control of these vectors by fire ants in Mexico has b e e n discovered (Butler et al. 1979). Ticks also cause local t r a u m a a n d inflam­ mation at t h e site of a t t a c h m e n t a n d some­ times elicit a paralysis by t h e injection of s o m e neurologically active substance in their saliva. Fortunately, tick paralysis ( M u r n a g h a n a n d O ' R o u r k e 1978) is usu­ ally temporary, d i s a p p e a r i n g w h e n t h e tick is r e m o v e d , b u t recovery may be delayed a n d can cause d e a t h in y o u n g o r sensitive p e o p l e , especially w h e n t h e bite focus is on the neck at t h e base of t h e skull. T h e nonexistent tick "Ixodes maloni" was r e c o r d e d from a G u i a n a n tepui by Sir A r t h u r C o n a n Doyle in his fiction classic, The Lost World (Hoogstraal 1972).

References ARTHUR, D. R. 1961. Ticks and disease. Row, Peterson, Evanston, 111. BUTLER, J. F., M. L. CAMINO, AND T. M. PÉREZ.

1979. Boophilus microplus and the fire ant Solenopsis geminata. Rec. Adv. Acarol. 1: 469472. DRUMMOND, R. O., AND T. M. WHETSTONE. 1975.

Oviposition of the cayenne tick, Amblyomma

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

cajennense (F), in the laboratory. Entomol. Soc. Amer. Ann. 68: 214-216. ERNST,

C.

H.,

AND

E.

M.

ERNST.

1977.

Ectoparasites associated with Neotropical tur­ tles of the genus Callopsis (Testudines, Emydidae, Batagurinae). Biotropica 9: 139-142. GARRÍS, G. 1. 1987. Amblyomma variegatum (Acari: Ixodidae): Population dynamics and hosts used during an eradication program in Puerto Rico. J. Med. Entomol. 24: 82-86. HOFFMANN, A. 1962. Monografía de los lxo-

doidea de México. 1. Rev. Soc. Mex. Hist. Nat. 23: 191-307. HOOGSTRAAL, H. 1967. Ticks in relation to human diseases caused by Rickettsia species. Ann. Rev. Entomol. 12: 377-420. HOOGSTRAAL, H. 1972. Ixodes maloni Doyle, 1912 (nomen nudum) (Ixodoidea: Ixodidae) parasitizing humans in Brazil. Entomol. Soc. Amer. Bull. 18: 141. HOOGSTRAAL, H. 1981. Changing patterns of tickborne diseases in modern society. Ann. Rev. Entomol. 26: 75-99. HOOGSTRAAL,

H.,

C. M. CLIFFORD, AND J.

E.

KEIRANS. 1973. Argas (Microargas) transversus (Ixodoidea: Argasidae) of Galápagos giant tor­ toises: Description of the female and nymph. Entomol. Soc. Amer. Ann. 66: 727-732. HOOGSTRAAL, H., C. M. CLIFFORD, J. E. KEIRANS, AND H. Y. WASSEF. 1979. Recent

developments in biomedical knowledge of Argas ticks (Ixodoidea: Argasidae). Rec. Adv. Acarol. 2: 269-278. JONES, E. K., AND C. M. CLIFFORD. 1972.

The

systematics of the subfamily Ornithodorinae (Acariña: Argasidae). V.A. revised key to larval Argasidae of the Western Hemisphere and description of seven new species of Ornithodoros. Entomol. Soc. Amer. Ann. 65: 730-740. JONES, E. K., C. M. CLIFFORD, J. E. KEIRANS, AND

G. M. KOHLS. 1972. The ticks of Venezuela (Acariña: Ixodoidea) with a key to the species of Amblyomma in the Western Hemisphere. Brigham Young Univ. Biol. Ser. 17(4): 1-40. KEIRANS, J. E., C. M. CLIFFORD, A. A. GuGLIELMONE, AND A . J . MANGOLD. 1 9 8 5 . Ixodl'S

(Ixodes) pararicinus, n. sp. (Acari: Ixodoidea: Ixodidae), a South American cattle tick long confused with Ixodes ricinus. J. Med. Entomol. 22: 401-407. KEIRANS,

J.

E.,

C.

M.

CLIFFORD,

AND

H.

HOOGSTRAAL. 1980. Identity of the nymphs and adults of the Galápagos iguanid lizard parasites, Ornithodoros (Alectorobius) darwini and O. (A.) galapagensis (Ixodoidea: Arga­ sidae). J. Med. Entomol. 17: 427-438.

KEIRANS, J.

E.,

H.

HOOGSTRAAL,

AND C.

M.

CLIFFORD. 1979. Observations on the subgenus Argas (Ixodoidea: Argasidae: Argas). 16. Argas (A.) moreli, new species, and keys to Neotropical species of the subgenus. J. Med. Entomol. 15: 246-252. KOHLS,

G.

M.,

HOOGSTRAAL.

C.

M.

CLIFFORD,

1969. Two new

AND

H.

species of

Ornithodoros from the Galápagos Islands (Acariña: Argasidae). J. Med. Entomol. 6: 75-78. MURNAGHAN, M. F , AND F. J. O'ROURKE.

1978.

Tick paralysis, In S. Bettini, ed.. Arthropod venoms. Springer, Berlin. Pp. 419-464. OBENCHAIN, F D., AND R. GALUN, eds.

1982.

The physiology of ticks. Pergamon, New York. OLIVER, JR., J. H. 1989. Biology and systematics of ticks (Acari: Ixodida). Ann. Rev. Ecol. Syst. 20: 397-430. RAWLINS, S. C. 1979. Seasonal variation in the population density of larvae of Boophilus microplus (Canestrini) (Acari: Ixodoidea) in Jamaican pastures. Bull. Entomol. Res. 69: 87-91. TRAVASSOS, J., AND A. VALLEJO-FREIRE.

1944.

Criacáo artificial de Amblyomma cajennense para o preparo de vacina contra a febre maculosa. lnst. Butantan Mem. 18: 1-91. YUNKER, C. E., J. E. KEIRANS, C. M. CLIFFORD,

AND E. R. EASTON. 1986. Dermacentor ticks (Acari: Ixodoidea: Ixodidae) of the New World: A scanning electron microscope atlas. Entomol. Soc. Wash. Proc. 88: 609-627.

WHIP SCORPIONS Uropygi ( = Pedipalpida, in part). Spanish: Vinagrosos, vinegrones (General), vinagrillos (Mexico). V i n e g a r r o o n s . A whiplike flagellum e x t e n d i n g from t h e tip of t h e a b d o m e n gives this g r o u p its c o m m o n n a m e . O t h e r distinctive c h a r a c t e r ­ istics include a l o n g e r t h a n wide p r o s o m a , spiderlike chelicerae, a n d massive pedipalps e q u i p p e d with short b u t s t r o n g spines (Weygoldt 1972). T h e first legs a r e feelerlike a n d slightly l o n g e r t h a n t h e oth­ ers, which a r e relatively short a n d held close to the body. A pair of defensive glands that o p e n n e a r t h e a n u s secrete formic a n d acetic acid, a m o n g o t h e r c o m -

WHIP SCORPIONS

137

Figure 4.7 WHIP SCORPIONS AND PSEUDOSCORPION. (a) Vinegarroon (Mastigoproctus giganteus, Elyphonidae). (b) Tailless whip scorpion (Heterophrynus longicornus, Phrynidae). (c) Pseudoscorpion (Chelifer cancroides, Cheliferidae). p o u n d s (e.g., caprylic acid, a wetting agent). T h e s e c o m p o u n d s m a y be forcibly expelled from t h e a n u s as a m e a n s of chemical p r o t e c t i o n (Eisner et al. 1961). T h e s e animals a r e n o c t u r n a l p r e d a t o r s that live o n t h e g r o u n d , for e x a m p l e , u n d e r litter a n d stones a n d in s t u m p s (Kaestner 1968: 1 1 7 - 1 1 9 ) . Mastigoproctus giganteus (fig. 4.7a) actively excavates t u n ­ nels in t h e g r o u n d w h e r e it r e m a i n s d u r i n g the d a y t i m e . T h o s e most often e n c o u n ­ tered a r e t h e vinagrillos (so-called because of t h e i r v i n e g a r o d o r w h e n d e f e n d i n g themselves with t h e i r chemical sprays) of the g e n u s Mastigoproctus. S p e c i m e n s visit the vicinity of artificial lights at night, particularly in d e s e r t a r e a s , to feast o n insects a t t r a c t e d to t h e illumination. T h e y a r e large (BL to 7 c m ) , d a r k b r o w n to black, a n d heavily sclerotized. I n spite of their f e a r s o m e scorpionlike a p p e a r a n c e , they d o n o t bite o r sting. W h i p scorpions p r e f e r h u m i d condi­ tions a n d a r e mostly f o u n d in tropical climes. T h o s e that inhabit arid regions a p p e a r only after rains. T h e r e a r e r e p r e s e n t a t i v e s of two fami­ lies in Latin A m e r i c a (Rowland a n d C o o k e 1973). Most a r e of t h e family E l y p h o n i d a e , t h e largest g e n u s b e i n g Mastigoproctus with twelve species. T h e o n e species of Thelyphronellus is restricted to G u y a n a a n d n o r t h e a s t ­

138

e r n Brazil, as is Arnauromastigon, t h e single g e n u s a n d species of t h e o t h e r family, Hypoctonidae.

References EISNER,

T.,

J.

MEINWALD,

A.

MONRO, AND

R. GHENT. 1961. Defense mechanisms of ar­ thropods. I. The composition and function of the spray of the whip scorpion, Mastigoproctus giganteus (Lucas) (Arachnida, Pedipalpida). J. Ins. Physiol. 6: 272-298. KAESTNER, A. 1968. Invertebrate zoology, ar­ thropod relatives, Chelicerata, Myriapoda. Vol. 2. Wiley Interscience, New York. ROWLAND, J. M., AND J. A. L. COOKE.

1973.

Systematics of the arachnid order Uropygida ( = Thelyphonida). J. Arachnol. 1: 5 5 - 7 1 . WEYGOLDT,

P.

1972.

Geisselskorpione

und

Geisselspinnen (Uropygi und Amblypygi). Zeit. Kolner Zoo. 15(3): 95-107.

TAILLESS WHIP SCORPIONS Amblypygi ( = Pedipalpida, in part). Spanish: Frinos ( P a n a m a ) . W h i p spiders. T h e s e a r a c h n i d s (Weygoldt 1972) some­ what r e s e m b l e t h e w h i p scorpions but differ in lacking a t e r m i n a l a p p e n d a g e or tail a n d having spinous p e d i p a l p s . They a r e greatly flattened a n d have tremen­ dously elongated forelegs; t h e pedipalps a r e n o t as massive as t h e f o r m e r g r o u p but

TERRESTRIAL ARTHROPODS AND PRIMITfVE INSECTS

possess a f o r m i d a b l e a r r a y of long, very s h a r p - t i p p e d spines. T h e walking legs a r e long a n d s l e n d e r a n d splayed o u t to t h e side. T h e y lack spray glands. Amblypygids (Kaestner 1968) live in tropical to subtropical areas (Beck 1968), always u n d e r fairly h u m i d conditions, u n ­ der stones, u n d e r b a r k a n d in hollow logs, in litter, a n d so o n . T h e y s h u n light a n d a r e well-known cave inhabitants. T h e y occa­ sionally r u n o n t h e t r u n k s of large trees o r even t h e walls of h o m e s at night, w h e r e they search in d a r k n e s s with their sensitive, a n t e n n i f o r m (whiplike) forelegs a n d s n a r e with their spiny p e d i p a l p s . Little m o r e is known of their habits in n a t u r e ; s o m e laboratory studies have b e e n c o m p l e t e d (Weygoldt 1977). T h e r e a r e two s u b o r d e r s , Pulvillata a n d Apulvillata ( Q u i n t e r o 1986), each contain­ ing several families a n d m a n y g e n e r a , found from t h e s o u t h e r n United States to the n o r t h e r n half of S o u t h America. A m o n g t h e former, t h e r e a r e n i n e g e n e r a with varied habits. Paracharon a n d Tricharinus a r e small, blind, a n d live in t h e nests of ants a n d t e r m i t e s ( Q u i n t e r o 1986). O t h e r dominant g e n e r a a r e Chirinus in South America a n d Charinides in t h e C a r i b b e a n (Quintero 1983). T h e New World species of Apulvillata a r e placed in five g e n e r a : Phrynus, t h e most w i d e s p r e a d , Paraphrynus (Mullinex 1975), Heterophrynus (fig. 4.7b), Acanthophrynus, a n d Trichodamon (MelloLeitáo 1935). T h e last is t h e only New World m e m b e r of its otherwise O l d World family, D a m o n i d a e , a n d is f o u n d in Brazil (Quintero 1976). Phrynus a r e sometimes m u c h feared by locals for t h e i r large size (BL to 2.8 cm, leg span of 20 c m ) , ugliness, a n d wrongly suspected v e n o m o u s n e s s . T h e y a r e truly harmless c r e a t u r e s . T h e y d e f e n d t h e m ­ selves only by p i n c h i n g a n d carry n o sting or poisonous bite. I n this g e n u s , t h e pedipalpal spines form a kind of basket for catching prey w h e n i n t e r p o s e d .

References BECK, L. 1968. Aus den Regenwaldern

am

Amazonas 11. Natur Museum 98(2): 71-80. KAESTNER, A. 1968. Invertebrate zoology, ar­ thropod relatives, Chelicerata, Myriapoda. Vol. 2. Wiley Interscience, New York. MELLO-LEITÁO, C. 1935. Sobre o género Trichodamon M.-L. fnst. Butantan Mem. 10: 297-302. MULLINEX, B. L. 1975. Revision oí Paraphrynus Moreno (Amblypygida: Phrynidae) for North America and the Antilles. Calif. Acad. Sci. Occ Pap. 116: 1-80. QUINTERO, JR., D. 1976. Trichodamon and Da­

monidae, new family status (Amblypygi: Arachnida). Brit. Arachnol. Soc Bull. 3: 222-227. QUINTERO, JR., D. 1980. Systematics and evolu­ tion oí Acanthophrynus Kraepelin (Amblypygi, Phrynidae). 8th Int. Cong. Arachnol. (Vi­ enna) Proc. Pp. 341-347. QUINTERO,

JR., D.

1983.

Revision

of

the

amblypygid spiders of Cuba and their rela­ tionships with the Caribbean and continental American amblypygid fauna. Stud. Fauna Curacao Carib. Is. 196: 1—54. QUINTERO, JR., D. 1986. Revision de la clasifi­ cación de Amblypygidos pulvinados: Crea­ ción de subórdenes, una nueva familia y un nuevo género con tres nuevas especies (Arach­ nida: Amblypygi). 9th Int. Cong. Arachnol. (Panama City) Proc. Pp. 203-212. WEYGOLDT,

P.

1972.

Geisselskorpione

und

Geisselspinnen (Uropygi und Amblypygi). Zeit. KólnerZoo. 15(3): 95-107. WEYGOLDT, P 1977. Coexistence of two species of whip spiders (Genus Heterophrynus) in the Neotropical rain forest (Arachnida, Amblypy­ gi). Oecologia 27: 363-370.

FALSE SCORPIONS Pseudoscorpionida. Spanish: Q u e r n i t o s (General). Book scorpions. False scorpions all possess scorpionlike p e d ­ ipalps a n d general s h a p e b u t differ from those a r a c h n i d s most conspicuously by their m u c h smaller size (BL of most 2—4 m m ) a n d t h e lack of a s e g m e n t e d tail. Also missing a r e t h e scorpion's characteristic ventral pectines. T h e fingers of t h e pedi­ palps usually contain v e n o m glands o p e n -

FALSE SCORPfONS

139

ing at t h e s h a r p tips. T h e y also have silk glands, located in t h e p r o s o m a , with ducts o p e n i n g at t h e tip of t h e chelicera's mov­ able article. Silk is used to m a k e cocoonlike nests a b o u t twice t h e animal's size in which to molt, pass t h e winter, o r oviposit. T h e r e a r e t h r e e s u b o r d e r s , all with m e m ­ bers in t h e Neotropics. T h e region's f a u n a is poorly k n o w n ; Beier (1932) listed a b o u t 180 species, to which a u t h o r i t i e s have since a d d e d a p p r o x i m a t e l y 4 0 0 species; s o m e 100 to 2 0 0 m o r e m a y r e m a i n u n d i s c o v e r e d (Mahnert pers. comm.). T h e Amazonian f a u n a is especially diverse ( M a h n e r t 1979, M a h n e r t et al. 1986). Phoresy is practiced by m a n y species in this o r d e r , usually only by females, a n d is a p p a r e n t l y i m p o r t a n t to t h e m as a m e a n s of dispersal ( M u c h m o r e 1971). T h e bestk n o w n hosts a r e l o n g - h o r n e d beetles, espe­ cially t h e h a r l e q u i n beetle, u n d e r whose elytra several species travel. T h e best k n o w n of t h e s e is Cordylochernes scorpioides, which c o n s t r u c t s a "safety h a r ­ ness" of silk o n t h e beetle's a b d o m e n to which they cling w h e n t h e host flies (Zeh a n d Zeh 1991). A few p s e u d o s c o r p i o n s have b e c o m e associated with civilization. Cheiridiurn museorum, Chelifer cancroides (fig. 4.7c), a n d Withius piger (in Chile) have s p r e a d recently from t h e O l d World to South A m e r i c a in g r a i n s h i p m e n t s in which they h a v e s e c r e t e d themselves to feed o n s t o r e d p r o d u c t pests. P s e u d o s c o r p i o n s a r e typically cryptic, living in soil a n d litter, u n d e r bark, b e n e a t h rocks, a n d in similar r e t r e a t s . Many a r e cavernicolous ( M u c h m o r e 1972). T h e r e are several m a r i t i m e species, living u n d e r stones a n d s h o r e d e b r i s such as driftwood a n d s t r a n d e d seaweed a n d in rock crevices, w h e r e they s e a r c h for p r e y (Lee 1979). C o u r t s h i p a n d m a t i n g a r e c o m p l e x be­ haviors m u c h like t h o s e of scorpions. T h e process b e g i n s with a f o r m of d a n c e in which t h e m a l e stops close to t h e female, shakes his a b d o m e n , a n d waves his pincers. H e t h e n d r o p s a t h r e a d from his v e n t r a l

140

genital o p e n i n g . T h i s h a r d e n s into a pedes­ tal ( s p e r m a t o p h o r e ) on which h e leaves seminal fluid. T h e female straddles the pillar a n d draws t h e fluid into h e r sexual o p e n i n g , d u r i n g which process t h e male may advance a n d shake h e r by t h e legs to e n s u r e that t h e fluid becomes d e t a c h e d within h e r body.

References BEIER, M. 1932. Pseudoscorpionida. Das Tierreich 57: 1-258, 58: 1-294. LEE, V. F. 1979. The maritime pseudoscorpions of Baja California, México (Arachnida: Pseu­ doscorpionida). Calif. Acad. Sci. Occ. Pap. 131: 1-38. MAHNERT, V. 1979. Pseudoskorpione (Arach­ nida) aus dem Amazonas-Gebiet (Brasilien). Rev. Suisse Zool. 86: 719-810. MAHNERT, V., J. ADIS, AND P. F. BÜHRNHEIM.

1986. Key to the families of Amazonian Pseudoscorpiones (Arachnida). Amazoniana 10: 21-40. MUCHMORE, W. B. 1971. Phoresy by North and Central American pseudoscorpions. Roches­ ter Acad. Sci. Proc. 12: 79-97.

head, which is fused with t h e t h o r a x , a n d basal a b d o m i n a l s e g m e n t s . T h e latter is followed by a long six-segmented tail, t h e last article of which (the telson) is b u l b o u s and s h a r p t i p p e d , constituting t h e sting. T h e latter c o n t a i n s p a i r e d poison glands with ducts o p e n i n g in a slit a short distance from t h e tip. Scorpion sexual behavior is peculiar. T h e male first faces t h e female a n d grasps her p e d i p a l p s o r chelicerae with his own like a p p e n d a g e s . T h e p a r t n e r s advance and retreat in a kind of d a n c e ( " p r o m e ­ nade a d e u x " ) for a time until a suitable place is f o u n d for m a t i n g . T h e m a l e expels a s p e r m p a c k e t to t h e g r o u n d which is picked u p by t h e female's genitalia after more " d a n c i n g . " T h e n t h e female frees herself from t h e male a n d goes to a b u r r o w or o t h e r shelter; t h e m a l e goes away. It is not t r u e that t h e female kills t h e male. T h e latter m a y even live to ultimately m a t e with several o t h e r females.

pseudoscorpions, mainly cavernicolous, from Mexico. Amer. Micros. Soc. Trans. 91: 2 6 1 276. ZEH, D. W., ANDJ. A. ZEH. 1991. Novel use of silk by the harlequin beetle-riding pseudoscorpion, Cordylochernes scorpioides (Pseudoscor­ pionida, Chernetidae). ]. Arach. 19: 153-154.

Females give birth to active larval scorpi­ ons. T h e s e m i g r a t e to their m o t h e r ' s back where they r e m a i n until their first molt, when they leave a n d begin i n d e p e n d e n t lives. I n most species, after o n e to t h r e e years a n d several m o r e molts, they attain adulthood. A r a r e few of these a r a c h n i d s lead a semisocial existence (Polis a n d Lourenco 1986).

SCORPIONS

Scorpions feed entirely o n o t h e r a r t h r o ­ pods (including o t h e r scorpions), mainly spiders a n d insects that they e n c o u n t e r

MUCHMORE, W. B. 1972. New diplosphyronid

a n d overcome d u r i n g their nightly r a m blings. Certain d e s e r t species may go with­ out water for several m o n t h s b u t will d r i n k it readily if available; yet scorpions from h u m i d forest e n v i r o n m e n t s m a y d i e in a few days if forced to go without water. Scorpion v e n o m p r o d u c e s n e u r o t o x i c circulatory a n d muscular effects in h u m a n s . Several species a r e of major medical i m p o r ­ tance (Keegan 1980); those in Latin A m e r ­ ica b e l o n g to t h e g e n e r a Tityus a n d Centruroides, both in t h e family B u t h i d a e . Tityus (Diniz 1978, B ü c h e r l 1978) a r e f o u n d in varied habitats. Many a r e h u m i d forest inhabitants, living u n d e r t h e bark of d e a d trees a n d a m o n g g r o u n d litter. T h e y a r e an entirely Neotropical g e n u s of wide distribu­ tion continentally a n d a r e p r e s e n t o n sev­ eral of t h e Antilles. T h e g e n u s is also t h e region's largest, with over 130 species ( L o u r e n c o 1978). T h e deadliest species is T serrulatus (fig. 4.8a), which causes t h e d e m i s e of m a n y p e o p l e , especially very y o u n g children, in s o u t h e a s t e r n Brazil. Its v e n o m h a s b e e n characterized chemically a n d pharmacologically (Possani et al. 1977). In t h e d r i e r p a r t s of Mexico a n d C e n t r a l America, species of "bark scorpions" (Cen­ truroides) a r e t h e most d a n g e r o u s (Bücherl 1971, S t a h n k e 1978); t h e g e n u s is also found in t h e West Indies a n d in p a r t s of South America (Sissom a n d L o u r e n c o 1987). A m o n g t h e over fifty species (Santiago-Blay p e r s . c o m m . , S t a n h k e a n d Calos 1977) is the well-known D u r a n g o scor-

Scorpionida. Spanish: Escorpiones (General), alacranes. Portuguese: Escorpióes, lacraus (Brazil). Náhuatl: C o l o m e h , sing, cótotl (Mexico). Scorpions (Polis 1990, Williams 1987) are characterized by their eight legs, a pair of large p i n c h e r s o r p e d i p a l p s (like lobster claws) at t h e front, a n d a m u c h smaller but heavy pair of jaws (chelicerae) a r o u n d the m o u t h . O n t h e u n d e r s i d e , a pair of comblike "pectines" a r e c o n s p i c u o u s struc­ tures. T h e m a i n body is divided into a large front t r u n k s e g m e n t , c o m p r i s i n g the

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

Rgure 4.8 SCORPIONS AND SUN SPIDER, (a) Forest scorpion (Tityus serrulatus, Buthidae). scorpion (Centruroides suffusus, Buthidae). (c) Sun spider (Eremobates sp., (b) Durango __. Erematobatidae)

SCORPIONS

141

pión, C. suffusus (fig. 4.8b) (Atúnez 1950), which was responsible for 1,608 d e a t h s be­ tween 1890 a n d 1926 in t h e city of Dur a n g o , which h a d a p o p u l a t i o n of a b o u t 40,000 in this p e r i o d . Actually, t h e d e a t h rate from scorpions is h i g h e r in o t h e r areas in Mexico, o v e r 110 p e r h u n d r e d t h o u s a n d in some years in s o u t h e r n states such as Colima (Mazzotti a n d Bravo-Becherelle 1963). C. limpidus is a n even m o r e p o t e n t spe­ cies a n d is c o n s i d e r e d t h e most d a n g e r o u s scorpion in Mexico, responsible for m o r e t h a n 50 p e r c e n t of t h e 100,000 stings p e r year in t h e c o u n t r y ( a c c o r d i n g to Possani et al. [1980], w h o h a v e also described t h e qualities of its v e n o m ) . F r o m a public health s t a n d p o i n t , scorpi­ ons a r e t h e most i m p o r t a n t v e n o m o u s ani­ mals of Mexico. D u r i n g t h e p e r i o d 1 9 4 0 1949 a n d 1 9 5 7 - 1 9 5 8 , a total of 20,352 p e r s o n s w e r e killed by scorpion stings (Mazzotti a n d B r a v o - B e c h e r e l l e 1963). Small a r b o r e a l scorpions a r e also k n o w n in S o u t h A m e r i c a . O t h e r s , so-called field scorpions, p r e f e r t h e soil of d a m p places a n d a r e f o u n d u n d e r stones, especially a l o n g rivers. D e s e r t o r s e m i d e s e r t types, like Centruroides, a r e often b u r r o w e r s in places p r o t e c t e d from t h e heat. Many live in old buildings, u n d e r houses, a n d in g a r a g e s a n d often e n t e r o c c u p i e d premises w h e r e they c o m e into contact with h u ­ m a n s . T r u e cave-dwelling scorpions a r e comparatively r a r e (Francke 1 9 8 1 , Lou­ r e n c o & F r a n c k e 1985, Mitchell 1968). Most s c o r p i o n s a r e c o n t i n e n t a l , a l t h o u g h a few exist o n oceanic o r offshore c o n t i n e n ­ tal islands, such as Coco, t h e Antilles, a n d in t h e Gulf of California. Overall, in Latin A m e r i c a , t h e r e a r e s o m e 48 g e n e r a a n d m o r e t h a n 4 0 0 species allocated to 7 families ( L o u r e n c o pers. c o m m . ) . T h e largest g e n e r a a r e Tityus, d o m i n a n t in S o u t h A m e r i c a ( L o u r e n c o 1978), a n d Centruroides, which occurs mostly in C e n t r a l A m e r i c a a n d Mexico ( S t a h n k e a n d Calos 1977). N o g e n e r a l

142

taxonomic t r e a t m e n t exists for t h e e n t i r e area, a l t h o u g h a t t e m p t s have b e e n m a d e to review t h e South A m e r i c a n f a u n a a n d general distributional p a t t e r n s ( A r m a s 1982, L o u r e n c o 1986, Mello-Leitáo 1945). A key to t h e g e n e r a of B u t h i d a e is available (Vachon 1977).

References ATÚNEZ, F. 1950. Los alacranes en el folklore de Durango. Priv. publ., Aguascalientes, Mexico. BÜCHERL, W. 1971. Classification, biology and venom extraction of scorpions. In W. Bücherl and E. E. Buckley, eds., Venomous animals and their venoms. III. Venomous inverte­ brates. Academic, New York. Pp. 317—347. BÜCHERL, W. 1978. Venoms of Tityinae. A. Systematics, distribution, biology, venomous apparatus, etc., of Tityinae; venom collection, toxicity, human accidents and treatment of stings. In S. Bettini, ed., Arthropod venoms. Springer, Berlin. Pp. 371—379. DE ARMAS, L. F. 1982. Algunos aspectos zoogeográficos de la escorpionfauna antillana. Poeyana238: 1-17. DE MELLO-LEITÁO, C. 1945. Escorpióes Sul-

Americanos. Mus. Nac. (Rio de Janeiro) Arq. 40: 7-468. DINIZ, C. R. 1978. Venoms of Tityinae. B. Chemical and pharmacologic aspects oí Tityine venoms. In S. Bettini, ed., Arthropod venoms. Springer, Berlin. Pp. 379-394. FRANCKE, O. F. 1981. A new genus of troglobitic scorpion from Mexico (Chactidae, Megacorminae). Amer. Mus. Nat. Hist. Bull. 170: 23-28. KEEGAN, H. L. 1980. Scorpions of medical importance. Univ. Mississippi, Jackson. LouRENgo, W. R. 1978. Sur les difficultés rencontrées dans la revision du genre Tityus (Scorpiones, Buthidae). Zool. Soc. London Symp. 42: 502. LOURENCO, W. R. 1986. Les modeles de distri­ bution géographique de quelques groupes de scorpions néotropicaux. Soc. Biogeogr. Comp. Rend. 62: 6 1 - 8 3 .

MITCHELL, W. R. 1968. Typhlochactas, a new

genus of eyeless cave scorpions from Mexico (Scorpionidae, Chactidae). Ann. Speleol. 23: 753-777. POLIS, G. A. 1990. T h e biology of scorpions. Stanford Univ. Press, Stanford. POLIS, G.

A.,

AND W.

R.

LOURENCO.

1986.

Sociality among scorpions. 10th Cong. Int. Arachnol. (Jaca, Spain) Actas 1: 111-115. POSSANI, L. D., A. C. ALAGÓN, P. L. FLETCHER,

JR., AND B. W. ERICKSON. 1977. Purification

and properties of mammalian toxins from the venom of the Brazilian scorpion Tityus serrulatus Lutz and Mello. Arch. Biochem. Biophys. 180: 394-403. POSSANI, L. D., P. L. FLETCHER, J R , A. B. C. ALAGÓN, A. C. ALAGÓN, ANDJ. Z.JULIÁ. 1980.

Purification and characterization of a mam­ malian toxin from venom of the Mexican scorpion, Centruroides limpidus tecomanus Hoff­ mann. Toxicon 18: 175—183. SISSOM, W. D., AND W. R. LOURENCO. 1987.

The

genus Centruroides in South America (Scorpi­ ones, Buthidae). J. Arachnol. 15: 11-28. STAHNKE, H. L. 1978. T h e genus Centruroides (Buthidae) and its venom. In B. Bettini, ed., Arthropod venoms. Springer. Berlin. Pp. 277-307. STAHNKE, H. L., AND M. CALOS. 1977. A key to

the species of the genus Centruroides Marx (Scorpionida: Buthidae). Entomol. News 88: 111-120. VACHON, M. 1977. Contribution á l'étude des scorpions Buthidae du nouveau monde. I. Complement á la connaissance de Microtityus rickyi Kj.-W. 1956 de l'ile de la Trinité. II. Description d'une nouvelle espéce et d'un nouveau genre Mexicains: Darchenia bernadettae. III. Cié de determination des genres de Buthidae du nouveau monde. Acta Biol. Venezuelica 9: 283-302. WILLIAMS, S. C. 1980. Scorpions of Baja Califor­ nia, Mexico and adjacent islands. Calif. Acad. Sci. Occ. Pap. 135: 1-12. WILLIAMS, S. C. 1987. Scorpion bionomics. Ann. Rev. Entomol. 32: 275-295.

LouRENgo, W. R., AND O. F. FRANCKE. 1985.

Revision des connaissances sur les scorpions cavernicoles (troglobies) (Arachnida, Scorpi­ ons). Mem. Biospél. 12: 3—7. MAZZOTTI, L., AND M. A. BRAVO-BECHERELLE.

1963. Scorpionism in the Mexican republic. In H. L. Keegan and W. V. Macfarlane, eds., Venomous and poisonous animals and nox­ ious plants of the Pacific Region. Macmillan, New York. Pp. 119-131.

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

stricted, f o r m i n g a headlike a n t e r i o r por­ tion. T h e p e d i p a l p s a r e similar to t h e walking legs b u t a r e elevated w h e n in use a n d p r o v i d e d with adhesive o r g a n s at t h e tip. Typically, their soft bodies a r e covered with short, silky pelage. Most a r e m e d i u m sized (BL 2 - 4 cm). Sun spiders p r e f e r w a r m , arid habitats a n d a r e most a b u n d a n t in t h e Chilean, Peruvian, a n d Mexican deserts. S o m e oc­ c u r at high elevations in t h e A n d e s . At night, they r u n rapidly over t h e substra­ t u m a n d occasionally a r e seen n e a r houses, w h e r e they c o m e to catch insects attracted to lights. T h e y s p e n d t h e daylight h o u r s a n d winter m o n t h s in g r o u n d b u r r o w s of their o w n m a k i n g ; some b u r r o w in pithy o r rotten wood. A l t h o u g h large specimens are aggressive a n d f o r m i d a b l e in a p p e a r ­ ance, they a r e without v e n o m - d e l i v e r i n g capabilities a n d a r e i n n o c u o u s to h u m a n s . T h e y feed o n t h e o t h e r g r o u n d - d w e l l i n g a r t h r o p o d s that they e n c o u n t e r o n t h e i r nightly w a n d e r i n g s .

SUN SPIDERS Solpugida ( = Solifuga). W i n d scorpions. Members of this o r d e r a r e distinguished from o t h e r a r a c h n i d g r o u p s by their h u g e , forward-projecting, scissorlike chelicerae. The p r o s o m a b e h i n d t h e chelicerae is con­

Latin A m e r i c a n species a r e almost all f o u n d in t h e two families A m m o t r e c h i d a e (widespread; 61 species in several g e n e r a ) a n d E r e m o b a t i d a e (far n o r t h e r n Mexico a n d Baja California; 21 species in two g e n e r a ) ( M u m a 1970, 1976). Eremobates (fig. 4.8c) is a major g e n u s . Two species, Amacata penai ( M u m a 1971) a n d Syndaesia mastix (Maury 1980), from t h e A t a c a m a Desert a n d western A r g e n t i n a , respec­ tively, r e p r e s e n t t h e family Daesiidae ( = Amacataidae).

References MAURY, E. A. 1980. Presencia de la family Daesiidae en América del Sur con la de­ scripción de un nuevo género (Solifugae). J. Arachnol. 8: 59-67. MUMA, M. H. 1970. A synoptic review of North American, Central American, and Wesl In­ dian Solpugida (Arthropoda: Arachnida). Arths. Fla. Neighborhood Land Areas 5: 1 — 62. MUMA, M. H. 1971. T h e solpugids (Arachnida, Solpugida) of Chile, with descriptions of a

SUN SPIDERS

143

new family, new genera, and new species. Amer. Mus. Nov. 2476: 1-23. MUMA, M. H. 1976. A review oí solpugid families with an annotated list of Western Hemisphere solpugids. West. New Mex. Univ, Off. Res. Pub. 2(1): 1-33.

MYRIAPODS M y r i a p o d s (Camatini 1980) a r e all terres­ trial b u t take to moist to h u m i d e n v i r o n ­ m e n t s . T h e y a r e e l o n g a t e (often wormlike), with m a n y similar b o d y s e g m e n t s from which arise o n e pair of a p p e n d a g e s each (these t r u e s e g m e n t s have fused in the milli­ pedes). T h e m o u t h p a r t s a r e m a n d i b u l a t e a n d t h e legs several j o i n t e d , without basal lobes or subdivisions. T h e r e is r e a s o n to believe that m y r i a p o d s a n d insects s h a r e a c o m m o n m u l t i l e g g e d ancestor. Reference CAMATINI, M., ed. 1980. Myriapod biology. Academic, New York.

MILLIPEDES D i p l o p o d a . Spanish: Milpiés. Portuguese: C ó n g o l o s , piolhos d e cobra (Brazil). Quechua: Pachac c h a q u i . Like c e n t i p e d e s , which they superficially r e s e m b l e , millipedes gain their n a m e from an a b u n d a n c e of legs. Two pairs arise from most a p p a r e n t s e g m e n t s , a condition cre­ ated by e m b r y o n i c fusion of a l t e r n a t e b o d y somites, each c a r r y i n g o n e pair of legs, a pair of tracheal o p e n i n g s , a n d a ventral n e r v e c o r d g a n g l i o n , to f o r m d o u b l e somites (diplosomites). T h e r e s u l t i n g secon­ dary segment (diplosegment), therefore, has two ganglia a n d four legs a n d spiracles, all of which a r e located in the m e t a z o n i t e , r e p r e s e n t i n g the p o s t e r i o r of t h e fused somites. T h e p r o z o n i t e , r e p r e s e n t i n g the a n t e r i o r of t h e fused somites, lacks a p p e n d ­ ages a n d can t h u s b e telescoped into the

144

p r e c e d i n g metazonite, resulting in a m o r e c o m p a c t body form. T h e four anter i o r m o s t segments have o n e leg or n o legs, a secondary loss associated with the ability to curl the h e a d backward into a protective posture. T h e head of millipedes b e a r s o n e pair of short a n t e n n a e , o n e pair of internal mandibles, used to masticate decaying wood a n d o t h e r vegetation, a n d a flat­ t e n e d , platelike s t r u c t u r e , the gnathochilarium, which forms the a n t e r i o r floor of the m o u t h . T h e latter consists of several small sclerites whose s h a p e a n d arrange­ m e n t differ a m o n g o r d e r s a n d a r e , there­ fore, taxonomically useful. T h e subclass D i p l o p o d a is divided into t h r e e s u p e r o r d e r s , each r e p r e s e n t e d in the Neotropical region, but t h e vast majority belong to t h e s u p e r o r d e r H e l m i n t h o m o r p h a , c o n t a i n i n g calcified forms that are generally e l o n g a t e with cylindrical or flat­ tened bodies. T h e r e p r o d u c t i v e tracts o p e n on s e g m e n t 3, but in males, o n e or both pairs of legs on s e g m e n t 7 a n d occa­ sionally the a n t e r i o r pair on s e g m e n t 8 are modified into copulatory s t r u c t u r e s called g o n o p o d s . T h e g o n o p o d s transfer sperm packets to the female, a n d their configura­ tions are of p r i m a r y significance at the generic a n d specific levels. Pairs of milli­ pedes are often e n c o u n t e r e d e n r a p t u r e d , stretched o u t or coiled t o g e t h e r venter to venter in a many-legged copulatory em­ brace. After copulation, males of some large "flat-backed" species r i d e on the backs of females for several days (Heisler 1983). N i n e of the eleven h e l m i n t h o m o r p h or­ d e r s occur in the Neotropics, b u t the Polydesmida, Spirostreptida, a n d Spirobolida a r e d o m i n a n t . T h e Polydesmida are generally characterized by lateral expan­ sions of the d o r s u m called "paranota,' which i m p a r t a flattened a p p e a r a n c e to the animals, h e n c e the n a m e "flat-backed" milli­ pedes. Seventeen polydesmid families oc­ c u r in t h e Neotropics (Hoffman 1979),

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

Figure 4.9 MILLIPEDES AND CENTIPEDES, (a) Spirobolid millipede (Orthoporus sp., Spirostreptidae). (b) Polydesmid millipede (Barydesmus sp., Platyrhacidae). (c) Giant centipede (Scolopendra gigantea, Scolopendridae). (d) House centipede (Scutigera coleoptrata, Scutigeridae). many of which a r e c o m p r i s e d of m i n u t e , cryptic forms. T h e d o m i n a n t families with large-bodied f o r m s a r e Platyrhacidae a n d Chelodesmidae. C o m m o n platyrhacid g e n e r a a r e Nyssodesmus (Costa Rica) a n d Barydesmus (Costa Rica to Peru a n d western Brazil; fig. 4.9b), whose adults r a n g e from 70 to 100 millime­ ters in l e n g t h . Only slightly smaller are Psammodesmus ( P a n a m a to Peru) a n d the diverse assemblage of forms usually as­ signed to Amplinus, Pycnotropis, a n d Polylepiscus (Mexico to Peru). In t h e Chelo­ desmidae, Chondrodesmus, with a r o u n d forty species, is a b u n d a n t from Mexico to Brazil. T h e S p i r o s t r e p t i d a a n d Spirobolida contain phenotypically similar cylindrical forms of variable l e n g t h a n d d i a m e t e r ; they are distinguished by details of the h e a d , exoskeleton, a n d g o n o p o d s . Nearly thirty Neotropical s p i r o s t r e p t i d g e n e r a , in t h r e e families, are c u r r e n t l y recognized (Hoff­ man 1979, K r a b b e 1982), b u t the most widespread is Orthoporus (fig. 4.9a), with around forty species that r a n g e from the southwestern United States t h r o u g h o u t most of South A m e r i c a to Brazil. Vilcastreptus hoguei is a large, d a r k - b o d i e d form with pinkish legs, seen by t h o u s a n d s of tourists *ach year in the I n c a n r u i n s of Machu Kcchu (Hoffman 1988). Of the five Neotropical spirobolid fami*Wi the Rhinocricidae is d o m i n a n t . T h e s e millipedes vary greatly in size a n d are

readily recognized as b e l o n g i n g to the family, but a satisfactory generic a r r a n g e ­ m e n t has not b e e n attained. T h e Latin A m e r i c a n millipede f a u n a is diverse but poorly k n o w n . H o f f m a n (1969) r e p o r t e d some 4 7 0 species from Brazil, a n d Loomis (1968) r e p o r t e d a r o u n d 750 from Mexico a n d Central America. T h e s e are the only counts available. T h e total fauna doubtlessly consists of several t h o u ­ sand species, probably less t h a n 20 p e r c e n t of which have been discovered, m u c h less n a m e d a n d described. A l t h o u g h deserticolous forms exist, these are c r e a t u r e s mostly of d a n k , h u m i d habitats. T h e y live in r o t t e n logs, a m o n g leaf litter, in the soil, u n d e r stones a n d loose tree bark, in caves, a n d are a major c o m p o n e n t of the forest g r o u n d f a u n a ; a few a r e symbiotic with a r m y ants (Loomis 1959). Millipedes a r e harmless c r e a t u r e s , p r o ­ tected primarily by their h a r d exoskele­ ton. However, most p r o d u c e d r o p l e t s of caustic or noxious secretions from a series of lateral p o r e s on most s e g m e n t s (Eisner et al. 1978). T h e liquid usually oozes out, b u t some large millipedes, such as the Peruvian platyrhacid Barydesmus (orig. obs.), the Costa Rican Nyssodesmus python (Heisler 1983), a n d Rhinocricus lethifer, a J a m a i c a n rhinocricid (Loomis 1936), can forcibly squirt the fluid J to 4 d e c i m e t e r s or m o r e .

MILLIPEDES

145

T h e active chemicals in these secretions are diverse. Most o r d e r s h a v e single-cham­ b e r e d glands that p r o d u c e b e n z o q u i n o n e s , cresols, o r a l d e h y d e s that can blister o r tan h u m a n skin (Burtt 1947). Polydesmids give off h y d r o g e n cyanide from t h e o u t e r p a r t of t w o - c h a m b e r e d g l a n d s , from n o n t o x i c m a n d e l o n i t r i l e p r e c u r s o r s in t h e i n n e r c o m p a r t m e n t ( W o o d r i n g a n d B l u m 1963). Diplopods also a d o p t a defensive p o s t u r e by coiling with t h e v u l n e r a b l e h e a d tucked in t h e center. Two N e o t r o p i c a l polydesmoid families can roll u p into balls o r s p h e r e s , an a d a p t i v e f e a t u r e that h a s evolved i n d e p e n d e n t l y in o t h e r polydesmoid families a n d a n o t h e r subclass in o t h e r p a r t s of t h e world. M e m b e r s of t h e class h a r b o r a diverse microflora in t h e g u t which functions to release n u t r i e n t s a n d r e d u c e toxins from ingested plant m a t t e r (Sakwa 1974). Discussions of t h e a n a t o m y a n d g e n e r a l biology of millipedes a r e available in several g e n e r a l textbooks o n a r t h r o p o d s (Kaestner 1968: 3 8 9 - 4 2 9 ) . Refer to Loomis (1968) a n d H o f f m a n (1979) for bibliographies a n d taxonomy. I m p o r t a n t regional t a x o n o m i c works have b e e n written by Loomis (1936, 1964, 1968) a n d J e e k e l (1963).

References BURTT, E. 1947. Exúdate from millipedes with particular reference to its injurious effects. Trop. Dis. Bull. 44: 7 - 1 2 . EISNER, T., D. ALSOP, K. HICKS, AND J. MEIN-

WALD. 1978. Defensive secretions of millipeds. In S. Bettini, ed., Arthropod venoms. Springer, Berlin. Pp. 41—72. HEISLF.R, I. L. 1983. Nyssodesmus python (milpiés, large forest-floor millipede). In D. H. Janzen, ed., Costa Rican natural history. Univ. of Chicago Press, Chicago. Pp. 747-749. HOFFMAN, R. L. 1969. T h e origin and affinities of the southern Appalachian diplopod fauna. In P. C. Holt, ed., T h e distributional history of the biota of the southern Appala­ chians. Virginia Polytech. lnst., Blacksburg. Pp. 221-246. HOFFMAN, R. L. 1979. Classification

of the

Diplopoda. Mus. Hist. Nat., Geneva. HOFFMAN, R. L. 1988. A new genus and species

146

of spirostreptoid millipedes from the eastern Peruvian Andes. Myriapodologica 2: 29-36. JEEKEL, C. A. W. 1963. Diplopoda of Guiana ( 1 5). Nat. Stud. Suriname Nederlandse Antillen 4(27): 1-157. KAESTNER, A. 1968. Invertebrate zoology. Vol. 2. Wiley Interscience, New York. KRABBE, E. 1982. Systematik der Spirostreptidae (Diplopoda: Spirostreptomorpha). Natur. Ver. Hamburg Abh. (n.f.) 24: 1-476. LOOMIS, H. F. 1936. T h e millipedes of Hispaniola, with descriptions of a new family, new genera, and new species. Mus. Comp. Zool. (Harvard Univ.) Bull. 80: 1-191. LOOMIS,

H. F. 1959. New

myrmecophilous

millipeds from Barro Colorado Island, Canal Zone and Mexico. Kans. Entomol. Soc. J. 32: 1-7. LOOMIS, H. F. 1964. T h e millipedes of Panama (Diplopoda). Fieldiana: Zoology 47: 1-136. LOOMIS, H. F. 1968. A checklist of the milli­ pedes of Mexico and Central America. U.S. Nati. Mus. Bull. 266: 1-137. SAKWA, W. N. 1974. A consideration of the chemi­ cal basis of food preference in millipedes. Zool. Soc. London Symp. 32: 329-346. WOODRING, J. P., AND M. S. BLUM. 1963. Anat­

omy and physiology of repugnatorial glands of Pachydesmus. Entomol. Soc. Amer. Ann. 56: 448-453.

CENTIPEDES C h i l o p o d a . Spanish: Ciempiés, e s c o l o p e n d r a s (General); alacranes (Puerto Rico). Portuguese: Centopéias, lacraias (Brazil). Náhuatl: Petlazolcoameh, sing, petlazolcoatl (Mexico). S c o l o p e n d e r s . C e n t i p e d e s (Kaestner 1968: 3 5 6 - 3 8 8 , Lewis 1981) a r e slender, very elongate, and m a n y s e g m e n t e d a r t h r o p o d s , resembling millipedes only in a very g e n e r a l way. They are distinguished by bodies with only one pair of legs p e r s e g m e n t . T h e latter are clearly distinct, n o t fused into pairs. T h e n u m b e r of s e g m e n t s varies from 15 to 181; n o species has an even n u m b e r of pairs of legs a n d never, t h e r e f o r e , has o n e h u n d r e d legs, as t h e g r o u p ' s c o m m o n n a m e suggests. C e n t i p e d e s possess a single pair of long,

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

very flexible a n t e n n a e a n d m a n d i b u l a t e , forward-projecting m o u t h p a r t s . T h e a n t e riormost legs a r e modified into fourjointed, sharply p o i n t e d fangs (forcipules) connected to i n t e r n a l v e n o m glands. T h e s e u b i q u i t o u s a r t h r o p o d s exhibit a large size r a n g e . T h e majority a r e rela­ tively small (BL 1-5 cm). S o m e very large chilopods b e l o n g to t h e S c o l o p e n d r o m o r pha (Bücherl 1974). Scolopendra gigantea (fig. 4.9c) of t h e West Indies is gigantic, attaining a b o d y length of 27 centimeters (Bücherl 1971). Because of their g r e a t size, such c e n t i p e d e s a r e greatly feared. T h e y sometimes bite h u m a n s , causing pain a n d often a local inflammation b u t usually nothing m o r e serious, n o t w i t h s t a n d i n g hor­ ror tales in t h e literature (Minelli 1978). Scolopenders (Scolopendra spp.) a r e most often t h e o f f e n d i n g species. It is probably a myth that they leave a w o u n d with each leg if they crawl on b a r e skin, a l t h o u g h t h e legs a r e t i p p e d with s h a r p claws that m a y cause a prickling sensation. T h e r e is n o evidence that t h e legs contain v e n o m glands, a l t h o u g h s o m e c e n t i p e d e s p r o d u c e noxious chemicals from o t h e r parts of t h e body. T h e chemical composition of centi­ pede v e n o m is still virtually u n k n o w n . Bücherl (1946, 1971), e x p e r i m e n t i n g with five of the largest a n d most c o m m o n Brazil­ ian species o n laboratory animals, con­ cluded that t h e v e n o m from actual bites, while capable of giving intense p a i n , was too weak to seriously h a r m h u m a n s . In fact, it most probably can be h a n d l e d with impunity. S o m e even have been g a t h e r e d for use as h u m a n food. Von H u m b o l d t a n d Bonpland (1852: 1:157) saw C h a y m a s In­ dian children d r a g 4 5 - c e n t i m e t e r - l o n g cen­ tipedes o u t of g r o u n d b u r r o w s a n d d e v o u r them directly.

m o n in dwellings. C e n t i p e d e s e m e r g e at night to prey o n o t h e r small surfacedwelling invertebrates, especially e a r t h ­ w o r m s a n d insects. S o m e Scolopendra take small vertebrates (mice, lizards, toads) in captivity, a n d this may be their chief food in n a t u r e also, which may explain t h e generally large size of t h e m e m b e r s of this g e n u s . T h e y a r e quick a n d easily s u b d u e their prey with a v e n o m o u s bite from t h e a n t e r i o r fangs. T h e t a x o n o m y of c e n t i p e d e s is still u n ­ settled. Classifications of h i g h e r g r o u p s vary greatly, a n d many species r e m a i n u n described. Four o r d e r s a r e usually recog­ nized: the many-legged, wormlike, soildwelling G e o p h i l o m o r p h a ; t h e shortbodied L i t h o b i o m o r p h a , with 15 pairs of legs; t h e S c u t i g e r o m o r p h a , which has 15 pairs of very long, spiderlike legs; a n d t h e large, flattened S c o l o p e n d r o m o r p h a , with 21 o r 23 pairs of legs. T h e last is t h e most familiar g r o u p , with m a n y m e m b e r s belong­ ing to t h e g e n u s Scolopendra. T h e ten Neotropical species r a n g e over t h e n o r t h ­ e r n half of South America a n d t h e Carib­ bean islands of Jamaica a n d T r i n i d a d . Centi­ pedes a r e c o m m o n l y called alacranes in P u e r t o Rico, a n a m e that should be r e ­ served for Scorpionida (Santiago-Blay 1985).

References BÜCHERL, W. 1946. Acáo do veneno dos escolopendroinorfos do Brazil sobre alguns ani­ máis de laboratorio. Inst. Butantan Mem. 19: 181-197. BÜCHERL, W. 1971. Venomous chilopods or centipedes. In W. Bücherl and E. E. Buckley, eds., Venomous animals and their venoms. 3: Venemous invertebrates. Academic, New York. Pp. 169-191. BÜCHERL, W. 1974. Die Scolopendromorpha

Centipedes a r e most at h o m e in w a r m , humid retreats w h e r e they h i d e by day under stones o r logs on t h e g r o u n d , be­ neath loose b a r k , in r o t t i n g wood, a n d in caves a n d similar niches. T h e h o u s e centi­ pede (Scutigera coleoptrata; fig. 4.9d) is com­

der neotropischen region. Zool. Soc. London Symp. 32: 99-133. KAESTNER, A. 1968. Invertebrate zoology. Vol. 2. Wiley Interscience, New York. LEWIS, J. G. E. 1981. The biology of centipedes. Cambridge Univ. Press, Cambridge. MINELLI, A. 1978. Secretions of centipedes. In

CENTIPEDES

147

S. Bettini, ed., Arthropod venoms. Springer, Berlin. Pp. 73-85. SANTIAGO-BLAY, J. 1985. Aclaraciones en torno a los significantes zoológicos de la voz "alacrán" en Puerto Rico. Ciencia 12(2): 43-45. VON H ü M B O L D T , A . , AND A . BONPLAND. 1 8 5 2

[1814-1825]. Personal narrative of travels to the equinoctial regions of America, during the year 1799-1804. 3 vols. Henry Bohn, London. Translated by Thomasina Ross.

HEXAPODS Possession of t h r e e pairs of legs, h e n c e the n a m e H e x a p o d a , is a c o n s t a n t f e a t u r e of the Insecta b u t also of t h r e e additional primitive o r d e r s , a m o n g t h e m the Collembola. Because of peculiarities in the anat­ omy of this assemblage they a r e s e p a r a t e d from t h e t r u e insects as t h e subclass Parainsecta. Larval mites a n d ticks also have six legs b u t a r e m e m b e r s of the very distinct s u b p h y l u m Chelicerata. (See Evolu­ tion a n d Classification, c h a p . 1.)

SPRINGTAILS Collembola. Spanish a n d Portuguese: Colémbolos (General). T h e s e m i n u t e to small (BL 0.3—10 m m , average 2 m m ) , soft-bodied insect relatives take t h e i r c o m m o n n a m e in English from a forked a p p e n d a g e located o n t h e u n d e r ­ side of t h e a b d o m e n which is used as a springlike device to p r o p e l t h e animal u p w a r d . T h i s f o r m of locomotion is used when t h e insect is d i s t u r b e d a n d can send it m a n y times t h e length of its body into the air or to the side. Collembola also possess a t u b u l a r s t r u c t u r e (collophore) on the m i d v e n t r a l p a r t of t h e first a b d o m i n a l s e g m e n t . T h i s is t i p p e d with a bilobed, eversible sac t h o u g h t to have a function in o s m o r e g u l a t i o n . Six n o r m a l walking legs a r e p r e s e n t . T h e h e a d b e a r s four-jointed a n t e n n a e a n d poorly f o r m e d eyes. Springtails a r e often

148

in parallel with

insects because of their six legs a n d general form, but they differ in several f u n d a m e n ­ tal ways, including the p r e s e n c e of muscles in all a n t e n n a l segments (basal only in true insects), water-repellent substances in the cuticle (not found in insects), a n d complete cleavage in the e m b r y o (polar in insects). T h e elongate chewing o r sucking m o u t h parts a r e also uniquely w i t h d r a w n into a d e e p p o u c h in the h e a d . A l t h o u g h seldom seen a n d rarely a p p r e ­ ciated, these a r e extremely n u m e r o u s in­ habitants of m a n y different ecological re­ gimes (Christiansen 1964). T h e y prefer d a m p microhabitats w h e r e they are pro­ tected from wetting by a h y d r o p h o b i c cuticle (Ghiradella a n d Radigan 1974) and a r e highly characteristic soil a n d litter ani­ mals. T h e collembolan fauna a n d popula­ tion densities of t h e m i d l a t i t u d e tropical habitats, however, a r e m u c h less diverse t h a n that of t e m p e r a t e regions, probably because of the comparatively p o o r litter layer. T h e y are found often a m o n g decompos­ ing plant matter, on which they feed di­ rectly, or on associated fungi a n d algae. A few occur along the seashore, even within the intertidal zone, a n d a r e really marine organisms. O t h e r s live on the surface of fresh water a n d even o n snow. A few are symbiotic in a n t a n d t e r m i t e nests. Some eke o u t a b a r e existence o n Antarctic shores ( R a p o p o r t 1971). A considerable cave f a u n a also exists (Palacios 1983). T h e r e are also economically important forms that attack m u s h r o o m s , onions, to­ matoes, s u g a r c a n e , a n d alfalfa. Studies on the t a x o n o m y a n d biology of the Latin A m e r i c a n collembolan fauna are only beginning. It is estimated that the ap­ proximately 800 k n o w n species (Mari Mutt a n d Bellinger 1989) may r e p r e s e n t only 25 p e r c e n t of the total n u m b e r actually living in the region. T h e Latin A m e r i c a n fauna has representatives of all the world's twelve c o m m o n families (Palacios 1990), plus a r a r e new family (Coenaletidae) that has re-

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

Figure 4.10 SPRINGTAILS AND THYSANURANS. (a) Springtail (Temerías surinamensis, Sminthuridae). (b) Springtail (Ctenocyrtinus prodigus, Entomobryidae). (c) Silverfish (Ctenolepisma longicaudata, Lepismatidae). (d) Rock hopper (Neomachillelus scandens, Meinertellidae).

cently b e e n f o u n d o n the island of G u a d e ­ loupe (Bellinger 1985). R e p r e s e n t a t i v e spe­ cies of the large Latin A m e r i c a n f a u n a a r e Temeritas surinamensis ( S m i n t h u r i d a e ; fig. 4.10a) a n d Ctenocyrtinus prodigus ( E n t o m o ­ bryidae; fig. 4.10b).

References BELLINGER, P. F. 1985. A new family of Collem­ bola (Arthropoda, Tracheata). Carib. J. Sci. 21:117-123. CHRISTIANSEN, K. 1964. Bionomics of Collem­ bola. Ann. Rev. Entomol. 9: 147-178. GHIRADELLA, H.,

AND W. RADICAN. 1974.

Col­

lembolan cuticle: Wax layer and antiwetting properties. J. Ins. Physiol. 20: 310-306. MARI MUTT, J. A., AND P. F. BELLINGER.

1989.

Catalog of Neotropical Collembola. Flora and Fauna, Gainesville. PALACIOS, J. G. 1983. Collemboles cavernicoles du Mexique. Pedobiologia 25: 349-355. PALACIOS, j . G. 1990. Diagnosis y clave para determinar las familias de los Collembola de la región Neotropical. Man. Guias Est. Microartr. (UNAM, Mexico) 1: 1-15. RAPOPORT, E. H. 1971. The geographical distri­ bution of Neotropical and Antarctic Collem­ bola. Pacific Ins. Monogr. 25: 99-1 18.

THYSANURANS Thysanura, sensu lat. T h y s a n u r a n s a r e c o n s i d e r e d to c o m p r i s e the most primitive o r d e r of t r u e insects because of their c o m p l e t e winglessness, weak sclerotization, a n d p r e s e n c e of vesti­ gia], j o i n t e d a p p e n d a g e s o n the u n d e r s i d e

of the a b d o m e n . I m m a t u r e s grow g r a d u ­ ally into adults without a p p r e c i a b l e c h a n g e in form (i.e., n o m e t a m o r p h o s i s ) . T h e y also b e a r t h r e e long, many-jointed tails e x t e n d i n g from the tip of the a b d o m e n , a n d the equal thoracic s e g m e n t s have r a t h e r large lateral lobelike e x p a n s i o n s . T h e y are generally small (most B L 1 0 - 1 5 mm). T h e body is covered with scales b u t may be b a r e except for n u m e r o u s fine bristles. T h e r e are two s u b o r d e r s , the silverfish (Zygentoma) a n d the bristletails (Microcoryphia) (Remington 1954). T h e s e catego­ ries have been variously recognized a n d n a m e d by taxonomists, a n d s o m e confu­ sion r e m a i n s o n the best way to classify t h e group. M e m b e r s of both g r o u p s a r e widely distributed a n d mostly secretive, o c c u r r i n g in all sorts of general habitats such as in rotten wood, in rock crevices, a n d in h u ­ m u s a n d g r o u n d litter. S o m e h a v e highly specialized habits, living in bird's nests, associating with social insects, or living in caves (Wygodzinsky 1967). Many have b e e n t r a n s p o r t e d t h r o u g h o u t the world by h u m a n traffic. Free-living t h y s a n u r a n s , es­ pecially those r e q u i r i n g a w a r m e n v i r o n ­ m e n t , often b e c o m e a d a p t e d to domiciles.

References REMINGTON, C. L. 1954. The suprageneric classi­ fication of the order Thysanura (Insecta). Entomol. Soc. Amer. Ann. 47: 277-286.

THYSANURANS

149

WYGODZINSKY, P. 1967. On the geographical

distribution of the South American Microcoryphia and Thysanura (Insecta). In D. Deboutteville and E. Rapoport, eds., Biologie de L'Amérique Australe. 3: 505— 524. Ed. Cent. Nat. Recher. Sci., Paris.

Silverfish Z y g e n t o m a ( = L e p i s m a t o i d e a ; formerly T h y s a n u r a , in part). Spanish: Pececitos d e plata, pescaditos p l a t e a d o s (General). Portuguese: Tracas d o s livros (Brazil). Silverfish (so called because of their shiny, slick a p p e a r a n c e ) a r e familiar cosmopoli­ tan h o u s e h o l d pests. However, t h e classic domestic species, Lepisma saccharina, h a s b e e n f o u n d in Latin A m e r i c a only sporadi­ cally in t h e cooler h i g h l a n d s of Brazil, Bolivia, a n d A r g e n t i n a . T h e y a p p a r e n t l y d o n o t tolerate h u m i d , tropical conditions. Lepisma wasmanni is a soil-dwelling resi­ d e n t of t h e lomas of coastal Peru b u t seems to h a v e b e e n i n t r o d u c e d from t h e Mediterranean region. T h e long-tailed h o u s e silverfish (Ctenolepisma longicaudata, fig. 4.10c) is t h e species most c o m m o n l y e n c o u n t e r e d i n d o o r s in Latin A m e r i c a (Wygodzinsky 1967). It has a thick b o d y covering of slick, slate-colored scales. It usually occurs in d a m p situations, but its precise r a n g e in t h e region is u n k n o w n . Stylifera gigantea is also c o m m o n but seldom occurs i n d o o r s . T h e r e a r e m a n y o t h e r species in various g e n e r a , but k n o w l e d g e of t h e i r distribution a n d habits is r u d i m e n t a r y (Wygodzinsky 1967). M e m b e r s of this s u b o r d e r a r e character­ ized by their cylindrical o r s o m e w h a t flat­ t e n e d s h a p e , t h o r a x that is n o t a r c h e d , m o u t h p a r t s that a r e directed forward, a n d small eyes that a r e set widely a p a r t . T h e lateral lobes of the t h o r a x a r e only slightly e x p a n d e d . T h e g r o u p as a whole r e q u i r e s w a r m e r e n v i r o n m e n t s t h a n t h e bristletails, which p r e f e r cooler, usually m o u n t a i n o u s climates. T h r e e of t h e world's four families a r e

150

f o u n d in Latin America (Paclt 1963, 1967; Wygodzinsky 1972). T h e Lepismatidae is the largest a n d most diverse a n d contains several domestic types as well as many native species. Several of t h e latter a r e associated symbiotically with social insects, as a r e the Nicoletiidae. M e m b e r s of the latter family a r e s u b t e r r a n e a n , often inhab­ iting caves (Mexico a n d Cuba) a n d living in ant a n d termite nests. Cave forms lack eyes and integumentary pigment. T h e Maindroniidae, with its single bizarre genus, Maindronia, has only b e e n e n c o u n t e r e d u n d e r d r y i n g seaweed along arid Peruvian a n d Chilean coasts. T h e s e silverfish have a very elongate, p i g m e n t e d , unsealed body.

details a r e m u c h less seen, completely wild insects, living secretly in all sorts of d r y o r moist hideaways, u n d e r stones a n d bark, in leaf litter, a n d a m o n g rocks at t h e seashore. T h e y a r e terrestrial o r littoral, n o c t u r n a l or crepuscular, a n d a r e highly active, r u n ­ ning swiftly a n d , if t h r e a t e n e d , j u m p i n g violently to escape c a p t u r e . T h e largest a n d most w i d e s p r e a d Neo­ tropical g e n u s is Neomachillelus (fig. 4.10d), which inhabits spaces u n d e r tree b a r k a n d the soil surface. It is f o u n d over t h e e n t i r e

area except for Patagonia (Wygodzinsky 1959). T h e s o u t h e r n t e m p e r a t e regions are inhabited by Machiloides a n d Machilmus. T h e latter t e n d s t o w a r d arid zones. T h e g e n u s Meinertellus is r e p r e s e n t e d by several species in n o r t h e a s t e r n S o u t h America.

Reference WYGODZINSKY, P. 1959. Thysanura and Ma­

chilida of the Lesser Antilles and northern South America. Stud. Fauna Curacao Carib. Isl. 36: 28-49.

References PACLT, J. 1963. Thysanura Fam. Nicoletiidae. Genera Insectorum 216: 1—58. PACLT, J. 1967. Thysanura Fam. Lepidotrichidae, Maindroniidae, Lepismatidae. Gen­ era Insectorum 218: 1-86. WYGODZINSKY, P. 1959. Thysanura and Ma-

chilida of the Lesser Antilles and northern South America. Stud. Fauna Curacao Carib. Is. 36: 28-49. WYGODZINSKY, P. 1967. On the geographical

distribution of the South American Microcoryphia and Thysanura (Insecta). In D. Deboutteville and E. Rapoport, eds., Biologie de L'Amérique Australe. 3: 505524. Ed. Cent. Nat. Recher. Sci., Paris. WYGODZINSKY, P. 1972. A review of the silver­ fish (Lepismatidae, Thysanura) of the United States and the Caribbean area. Amer. Mus. Nov. 2481: 1-26.

Bristletails Microcoryphia ( = A r c h a e o g n a t h a , Machiloidea, Machilida). Rock j u m p e r s . While silverfish tend to be cylindrical or slightly flattened, bristletails a r e com­ pressed laterally. T h e h e a d is r o t a t e d down­ ward so that t h e m o u t h p a r t s project ventrally, a n d t h e t h o r a x is strongly arched. T h e eyes a r e well developed a n d contigu­ ous, with m a n y facets. T h e lobular sides of the t h o r a x a r e large a n d a p p r e s s e d to the sides. Also, in contrast to silverfish, bns-

TERRESTRIAL ARTHROPODS AND PRIMITIVE INSECTS

THYSANURANS

151

Ó

ORTHOPTEROIDS AND OTHER ORDERS

Most of this c h a p t e r treats those o r d e r s closely associated taxonomically with t h e O r t h o p t e r a . For c o n v e n i e n c e , several addi­ tional o r d e r s , a l t h o u g h not related, a r e included also.

ORTHOPTEROIDS Orthopterodea. T h i s is an assemblage of primitive insects that have close affinities with t h e o r d e r O r t h o p t e r a . T h e classification of t h e major g r o u p s of o r t h o p t e r o i d s is very u n s e t t l e d . Kevan (1977) p r o v i d e s a n exhaustive a n d a u t h o r i t a t i v e s c h e m e , a l t h o u g h tentative. T h e t e r m " o r t h o p t e r o i d s " traditionally re­ fers to several o r d e r s with similar g e n e r a l ­ ized b o d y f o r m , m o d e r a t e to large size, ovoid h e a d with m a n d i b u l a t e m o u t h p a r t s , usually n a r r o w , long, s o m e w h a t t h i c k e n e d fore wings, e n l a r g e d p r o t h o r a x (often p r o ­ l o n g e d b a c k w a r d ) , a n d l e a t h e r y (fore) or m e m b r a n o u s (hind) wings (when p r e s e n t ) with a c o m p l e x , reticulate vein p a t t e r n . Internally, the p r e s e n c e of very n u m e r o u s Malpighian tubules is also characteristic. With the major e x c e p t i o n of m a n t i d s a n d s o m e species a n d g r o u p s (e.g., Decticinae) t h a t occasionally take p r e y o r feed o n insect carcasses, o r t h o p t e r o i d s a r e basi­ cally all vegetarians. N y m p h s of s o m e g r o u p s often visit flowers; a l t h o u g h they feed o n petals a n d a n t h e r s , they m a y benefit t h e plant by effecting pollination (Schuster 1974). O r t h o p t e r o i d s e m p l o y a wide variety of

152

protective devices, often highly specialized a n d used in all combinations a n d multi­ ples. Relatively few a r e involved in mim­ icry, b u t a great m a n y a r e cryptically col­ o r e d , some very realistically, to mimic leaves, sticks, stones, a n d o t h e r inert ob­ jects. T h i s is often c o m b i n e d with startling displays, involving eyespots or aposematic color fields, coupled with r e p u g n a n t secre­ tions a n d o m i n o u s s o u n d s . Most have spiny legs a n d powerful jaws, both of which are e m p l o y e d in active defense. N o c t u r n a l or very secretive habits afford safety to o t h e r s , a n d most can either j u m p or fly to escape persistent attack (Robinson 1969). S o u n d p r o d u c t i o n is particularly well d e v e l o p e d in the o r t h o p t e r o i d s , primarily by males for the attraction of females. Both sexes a r e well e q u i p p e d with h e a r i n g or­ g a n s ( t y m p a n a ) located in various parts of the body. Males may sing alone o r exhibit " c h o r u s i n g , " that is, calling by two or more conspecific males, involving interactions with each other. C h o r u s i n g takes many forms, d e p e n d i n g on the precise timing of the multiple s o u n d s (Greenfield a n d Shaw f983). In some primitive p h a n e r o p t e r i n e katydids, the female has a series of minute knoblike stridulatory o r g a n s o n the wing base which a r e used to answer calling males.

References GREENFIELD, M.

D.,

AND K.

C.

SHAW.

1983.

Adaptive significance of chorusing with spe­ cial reference to the Orthoptera. In D. T.

Gwynne and G. K. Morris, eds., Orthopteran mating systems. Westview, Boulder, Colo. Pp. 1-27. KEVAN, D. K. M C E . 1977. Suprafamilial classifi­ cation of "orthopteroid" and related insects; a draft scheme for discussion and consid­ eration. Lyman Entomol. Mus. Res. Lab. (McGill Univ.), Mem. 4, Spec. Publ. 12: Appendix, 1-26. ROBINSON, M. H. 1969. T h e defensive behav­ iours of some orthopteroid insects from Pan­ ama. Royal Entomol. Soc. London Trans. 121: 281-303. SCHUSTER, J. C. 1974. Saltatorial Orthoptera as common visitors to tropical flowers. Biotropica6: 138-140.

Long-Horned Orthopteroids Grylloptera ( = Ensifera) These a r e mainly the katydids a n d crickets with long a n t e n n a e of well over thirty segments. T h e a u d i t o r y o r g a n s a r e located on the fore tibiae a n d the stridulatory organs o n the basal p o r t i o n s of the fore wings.

KATYDIDS Tetdgoniidae. Spanish: E s p e r a n z a s (General), p u l p o n e s (Costa Rica), langostas v e r d e s ( A r g e n t i n a ) , saltamontes n o c t u r n o s ( P a n a m a ) , grillos voladores (Peru). Portuguese: Esperanzas. Katydids a r e also called l o n g - h o r n e d grass­ hoppers a n d i n d e e d r e s e m b l e those o r t h o p terans except for their very long, manysegmented, whiplike a n t e n n a e . T h e y may be fully winged a n d capable of flight, or they may h a v e very short wings, the h i n d wing completely lost a n d the fore wings stubby. W h e n wings a r e p r e s e n t , a n area at the base of t h e thickened fore wing is often modified in t h e males into a stridulatory o r s o u n d - p r o d u c i n g o r g a n . O n e wing has a roughened r i d g e (file), which is r u b b e d against an o p p o s i n g s h a r p e d g e (scraper) on the other. T h e action sets b o t h wings into rapid vibration a n d p r o d u c e s the fa­

miliar chirping, buzzing, lisping, clicking, o r s n a p p i n g s o u n d s of these noisy crea­ tures. Small t y m p a n a a r e located in slots o n either side of the base of the front tibiae a n d in pockets on the sides of t h e t h o r a x n e a r the h i n d e d g e of the thoracic shield. T h e female ovipositor may be strongly compressed for incising leaves a n d wood or valvelike for inserting eggs in t h e g r o u n d . C o m p r e s s e d forms a r e short a n d strongly curved or long a n d sword like. T h i s is a large family in Latin A m e r i c a , with some 1,350 k n o w n species; this n u m ­ ber probably will increase by 30 p e r c e n t or m o r e ultimately (Nickle pers. c o m m . ) . T h e subfamilies may be recognized by use of Rentz's key (1979), which recognizes b r o a d e r categories t h a n Kevan's (1977) classification. Several new g e n e r a a n d spe­ cies a m o n g the shield-backed katydids from Chile a n d A r g e n t i n a have b e e n a d d e d to the f a u n a by Rentz a n d G u r n e y (1985). T h e y show affinities with o t h e r s of the subfamily from Australia a n d western N o r t h America (ibid., 70). T h e c o m m o n English n a m e of these insects derives from the song of the N o r t h A m e r i c a n species, Pterophylla camellifolia, which s o u n d s like the plaintive p h r a s e , "kate-she-did," "kate-did-she-did," or "katy-did." A folktale exists c o n c e r n i n g the accountability of a fictitious lady in the d e a t h of a lover w h o s p u r n e d her. Local n a m e s in Spanish a n d P o r t u g u e s e m e a n " h o p e , " in r e f e r e n c e to the g r e e n color of so m a n y species in the family, the symbolic color of this e m o t i o n . Katydids are an e x t r e m e l y i m p o r t a n t link in v e r t e b r a t e food chains, as they are utilized by m a n y birds, bats, m o n k e y s , lizards, a n d snakes. Consequently, they have evolved a rich a r r a y of a n t i p r e d a t o r defenses (Belwood 1990). Forest katydids in P a n a m a a r e major p r e y for foliagegleaning bats. To some d e g r e e , the bats have a d a p t e d to r e d u c e d acoustic p r o d u c ­ tion by these katydids, w h o use a n alterna­ tive form of c o m m u n i c a t i o n ("substrate

KATYDIDS

153

transmitted tremulation"), rather than the usual singing, to attract mates (Belwood a n d Morris 1987). Like o t h e r o r t h o p t e r o i d s , katydids pos­ sess a rich r e p e r t o i r e of d e f e n s e tactics. A device not exhibited by o t h e r s but used by the katydid Ancistrocercus in Costa Rica a n d Eremopedes colonialis in Mexico (Rentz 1972: 54) is association with wasps (Downh o p e r a n d Wilson 1973).

References BELWOOD, J. J. 1990. Anti-predator defences and ecology of Neotropical forest katydids, especially the Pseudophyllinae. In W. J. Bailey and D. C. F. Rentz, The Tettigoniidae: Biol­ ogy, systematics and evolution. Springer, Ber­ lin. Pp. 8-26. BELWOOD, J.J., AND G. K. MORRIS. 1987. Bat

pre-

dation and its influence on calling behavior in Neotropical katydids. Science 238: 6 4 - 6 7 . DOWNHOPER, J.

F,

AND D.

E. WILSON.

D.

C.

F

1979.

Comments on

RENTZ, D. C. F, AND A. B. GURNEY. 1985.

The

shield-backed katydids of South America (Orthoptera: Tettigoniidae, Tettigoniinae) and a new tribe of Conocephalinae with genera in Chile and Australia. Entomol. Scandinavica 16: 69-119.

Broad-winged Katydids Tettigoniidae, P s e u d o p h y l l i n a e . Not all pseudophyllines have b r o a d wings as their c o m m o n n a m e implies, n o r a r e they leaflike as indicated by their scientific n a m e . P e r h a p s a majority of the a p p r o x i m a t e l y 600 described Neotropical species a r e small to m e d i u m , spindle-shaped insects t h a t sing little a n d d o not look like leaves. T h e adults of m e m b e r s of some tribes, especially t h e fol­ lowing, d o have very b r o a d fore wings and closely resemble leaves.

1973.

Wasps as a defense mechanism of katydids. Amer. Midi. Natur. 89: 451-455. KEVAN, D. K. M C E . 1977. Suprafamilial classifi­ cation of "orthopteroid" and related insects; a draft scheme for discussion and consider­ ation. Lyman Entomol. Mus. Res. Lab. (McGill Univ.), Mem. 4, Spec. Publ. 12: Appen­ dix, 1-26. RENTZ, D. C. F. 1972. Taxonomic and faunistic comments on decticine katydids with the description of several new species (Orthoptera: Tettigoniidae: Decticinae). Acad. Nat. Sci. Philadelphia Proc. 124: 41-77. RENTZ,

description of two new subfamilies. Austr. J. Zool 27: 991-1013.

the

classification of the orthopteran family Tetti­ goniidae, with a key to subfamilies and

Leaf Katydids Tettigoniidae, Pseudophyllinae, primarily the tribe Pterochrozini. Spanish: Esperanzas hojas (General). Portuguese: Bichos folhas (Brazil). Many katydids in this g r o u p of j u s t u n d e r o n e h u n d r e d species a p p e a r incredibly like the leaves a m o n g which they live. T h e y are g r e e n to b r o w n in general color with an outline s h a p e , even with d r i p tip, and m a r k i n g s imitating in every detail the color a n d s t r u c t u r e of leaves, c o m p l e t e to midribs, side veins, a n d even t r a n s p a r e n t

and discolored areas r e s e m b l i n g insect feeding holes, leaf m i n e r t u n n e l s , chewed edges, a n d m o l d spots (pi. 3g)! This r e s e m b l a n c e is f u r t h e r e n h a n c e d with behavior, for those species that dwell in living trees often walk with a slow u n d u l a t i n g gait that makes t h e m look like a leaf gently swaying in the breeze. Inhabit­ ants of litter a n d d e a d vegetation t e n d to sit very still, a n g l e d on their sides a n d fitting perfectly with t h e b r o w n , d r i e d leaves o n t h e forest floor. T h e latter t e n d also to be very large, a p p r o a c h i n g the sizes of the p a r t s of plants they resemble. Celidophylla albimacula (BWL 8 cm) is a rare, gigantic species from C e n t r a l A m e r ­ ica ( H o g u e 1979) with yellow-green wings which looks like a large w i t h e r e d leaf. T h e principal g e n e r a a r e Pterochroza, Mimetica, Tanusia, Cycloptera, a n d Typophyllum (Vignon 1930). A Brazilian species made famous by H e n r y Walter Bates in the related tribe Pterophyllini, a m o n g which are also m a n y leaf-shape forms, is the tananá {Thliboscelus hypericifolius = Chlorocoelus tanana; fig. 5.1a). The males produce a very loud and not un­ musical noise by rubbing together the over­ lapping edges of their wing-cases. The notes are certainly the loudest and most extraordi­ nary that I ever heard produced by an orthopterous insect. The natives call it the Tananá, in allusion to its music, which is a sharp, resonant stridulation resembling the syllables ta-na-ná, ta-na-ná, succeeding each other with little intermission. It seems to be rare in the neighborhood. When the natives capture one, they keep it in a wicker-work cage for the sake of hearing it sing. (Bates 1892: 129)

p a t t e r n is especially well d e v e l o p e d in gen­ era like Cycloptera (fig. 5.1c). W h e n folded, the lower p o r t i o n of t h e wings a r e often rolled u n d e r a n d h i d e the insect's a b d o ­ m e n . In repose, the tips of the fore wings project beyond or are equal to t h e l e n g t h of the h i n d wings. T h e n y m p h s are also leaf- or twiglike, often with very c o m p r e s s e d bodies a n d flattened leg segments a n d erect crests on the back of the p r o t h o r a x . T h e y rest on the u p p e r surfaces of leaves with their long legs s p r e a d spiderlike a n d bodies pressed close to the surface. Because they a r e a r b o r e a l a n d n o c t u r n a l a n d so difficult to see even d u r i n g t h e light of day, these katydids a r e seldom collected, a n d little is k n o w n of their biology. T h o s e r a r e adults that c o m e to artificial lights constitute the majority of specimens in m u s e u m collections. M e m b e r s of the related tribe Cocconotini, primarily the g e n u s Cocconotus (fig. 5 . I d ) , are also large (BWL 8—9 cm) a n d have very long a n t e n n a e , two to t h r e e times the body length. T h e y habitually rest in rolled leaves of b a n a n a s , heliconias, gingers, cannas, a n d the like, h e a d out­ ward, only with the tips of the a n t e n n a e e x p o s e d , testing the outside world while the rest of the insect r e m a i n s h i d d e n . T h e y are mostly d a r k gray-brown a n d elongate, their wings rolled a r o u n d the body. Some m e m b e r s of t h e subfamily are wingless. A giant species f o u n d in t h e canopy of the Peruvian rain forest is Panoploscelus (tribe Eucocconotini). It is well protected by its great size a n d s t r e n g t h a n d sharply spined legs.

References

Figure 5.1 BROAD-WINGED KATYDIDS (TETTIGONIIDAE). (a) Tananá {Thliboscelus hypericifolius). (b) Eye-winged Katydid (Pterochroza ocellata). (c) Leaf katydid {Cycloptera speculata). (d) Long-winged leaf katydid (Cocconotus sp.).

154

ORTHOPTEROIDS AND O T H E R ORDERS

Pterochroza ocellata (pi. lb) is an A m a z o n i a n species with g r e e n i s h or b r o w n i s h - r e d , brown-black-splotched fore wings that cam­ ouflage it well on d e a d leaves. But if this first line of d e f e n s e fails, it fans o u t its h i n d w «ngs to display large t h r e a t e n i n g eyespots at their o u t e r c o r n e r s (fig. 5.1b). T h e leaf

BATES, H. W. 1892. The naturalist on the River Amazons. John Murray, London. BEIER, M. 1960. Orthoptera, Tettigoniidae (Pseu­ dophyllinae II). Das Tierreich 74: 1—396. BEIER, M. 1962. Orthoptera, Tettigoniidae (Pseu­ dophyllinae I). Das Tierreich 73: 1—468. HOGUE, C. L. 1979. A third specimen of Celido­ phylla albimacula (Orthoptera: Tettigoniidae)

KATYDIDS

155

and remark on the emergence of Díptera from insect carrion. Entomol. News 90: 151. VIGNON,

M.

P.

1930.

Recherches

sur

les

sauterelles-feuilles de l'Amérique tropicale. Premiere partie. Revision du groupe des "Pterochrozae" (Phasgonuridae, Pseudophyllinae). Mus. Nati. Hist. Natur. (Paris), 6 Ser., Arch. 5: 57-214.

Narrow-winged Katydids Tettigoniidae, P h a n e r o p t e r i n a e . Many of these katydids a r e leaf mimics but not q u i t e so r e m a r k a b l y as those b e l o n g i n g to the b r o a d - w i n g e d g r o u p (fig. 5.2d). T h e fore wings a r e generally l o n g e r a n d n a r r o w e r a n d m o r e plainly m a r k e d , g r e e n to b r o w n . In a few species, the h i n d wings a r e brightly colored, such as Vellea, which has a b r o a d scarlet zone basally. W h e n t h e wings a r e folded, t h e tips of the h i n d pair project b e y o n d t h e apices of the fore pair. Females have a globose h e a d a n d short, u p t u r n e d ovipositor. T h e o u t e r p a r t of the fore tibia in cross section is s q u a r e a n d with a flat o r slightly concave, dorsal surface. T h e thoracic auditory pock­ ets a r e large a n d e x p o s e d . Most n a r r o w - w i n g e d katydids are me­ dium-sized to small (BWL 2 5 - 5 0 m m ) , but Steirodon (formerly Peucestes) is gigantic (BWL 10—12 cm) a n d possesses a coarsely s e r r a t e c o r o n a a r o u n d the p e r i p h e r y of the p r o t h o r a c i c shield (fig. 5.2a) (Nickle 1985). N y m p h s of the g e n u s a r e strongly compressed and have a conspicuous dark ocellate spot laterally, o n t h e wing p a d .

T h e g r o u p is p h y t o p h a g o u s a n d arbo­ real. S o m e habitually rest o n lichencovered tree t r u n k s , b l e n d i n g in perfectly o n account of their own mottled white, g r e e n , a n d black body m a r k i n g s . T h e re­ semblance is increased by lobular excres­ cences o n the legs a n d body exactly like the folióse form of the plant. A n excellent e x a m p l e is Dysonia fuscifrons (fig. 5.2b) from Mexican cloud forests w h e r e the perpetually h u m i d a t m o s p h e r e s u p p o r t s a lush growth of epiphytes ( D a m p f 1939). M e m b e r s of the tribe Pleminiini (e.g., Championica) a r e large, with mottled green a n d b r o w n m a r k i n g s that camouflage t h e m on moss- a n d lichen-covered tree b a r k w h e r e they habitually rest. In Brazil, Scaphura a n d Aganacris (fig. 5.2c) mimic t a r a n t u l a hawk wasps (Pepsis) a n d a r e not only colored the same, with d e e p blue body a n d rust o r d a r k wings, but have a waspish a t t i t u d e as well, walking jerkily a n d rapidly a n d waving their o r a n g e - t i p p e d a n t e n n a e . At the a p p r o a c h of d a n g e r , they also flutter their wings and t u r n u p the a b d o m e n in mockery of an a t t e m p t to sting. T h e g r o u p is u n i q u e in possessing the only katydids whose females answer the calling songs of the males with a signal, t h o u g h weak, of their own (Nickle and Carlysle 1975). T h e interval between the male a n d female s o u n d s is i m p o r t a n t in d e t e r m i n i n g the correct association and d r a w i n g the male to the female. T h e chro-

Figure 5.2 NARROW-WINGED KATYDIDS (TETTIGONIIDAE). (a) Giant narrow-winged leaf katy­ did (Steirodon sp.). (b) Lichen-mimicking katydid (Dysonia fuscifrons). (c) Tarantula hawk-mimicking katydid (Aganacris sp.). (d) Common leaf-mimicking katydid (undetermined).

156

ORTHOPTEROIDS AND OTHER ORDERS

niosomes of a n u m b e r of Neotropical spe­ cies of this subfamily have b e e n described by F e r r e i r a ( 1 9 7 7 ) . T h e subfamily contains a p p r o x i m a t e l y 600 n a m e d species in Latin America.

References DAMPF, A. 1939. Un caso de fitomimetismo en un ortóptero Mexicano. Ese. Nac. Cien. Biol. Anal. 1: 525-533. FERREIRA, A. 1977. Cytology of Neotropical Phaneropteridae (Orthoptera-Tettigoniidae). Genética 47: 8 1 - 8 6 . NICKLE, D. A. 1985. A new steirodont katydid from Colombia (Orthoptera: Tettigoniidae). Entomol. News 96: 11-15. NICKLE, D.

A.,

AND T.

C.

CARLYSLE.

1975.

Morphology and function of female sound producing structures in ensiferan Orthoptera with special emphasis on the Phaneropte­ rinae. Int. J. Ins. Morph. Embryol. 4: 159168.

Cone-Headed and Meadow Katydids Tettigoniidae, C o n o c e p h a l i n a e , Copiphorini and Conocephalini. Many of these closely related g r o u p s of 140 to 150 species of katydids h a v e power­ fully d e v e l o p e d j a w s a d a p t e d for seed eating or, in s o m e cases, p r e d a t i o n . T h e i r name, however, derives from the strongly projecting a n t e r i o r p o r t i o n of t h e h e a d ; this may be only a simple convexity in meadow katydids (Conocephalini) or an elongate c o n e with a tubercle at its base

(this often n o t c h e d below) in t h e coneh e a d e d katydids (Copiphorini). T h e m e a d o w katydids, such as Conocephalus (fig. 5.3a), a r e relatively small (BWL seldom over 25 m m ) . T h e y f r e q u e n t m a r s h e s or v e r d a n t areas, usually living close to the g r o u n d a n d in a g g r e g a t i o n s . T h e i r stridulatory activity is mainly d i u r n a l or crepuscular. C o n e - h e a d e d katydids a r e larger (BWL often 5—6 cm) a n d occupy diverse habitats, b u t m a n y a r e associated with grasses whose seeds they eat. T h e i r two-part songs (ticks, giving way to buzzes) are h e a r d from d u s k to d a w n a n d a r e often of the c h o r u s i n g type. Males stridulate for several h o u r s each e v e n i n g from e x p o s e d perches, a p p a r e n t l y in sexual competition. Some n o n s i n g i n g males also r e m a i n close to the singer, taking a d v a n t a g e of his call to steal confused females that c o m e n e a r (Greenfield 1983). O n e South A m e r i c a n c o n e - h e a d , Copiphora (fig. 5.3b), is fairly large (BWL 4—6 cm) a n d has bright blue, r e d , a n d yellow colors o n the a b d o m e n . T h e s e a r e b r o u g h t into view w h e n the insect is d i s t u r b e d , usually by tipping t h e h e a d d o w n against the s u b s t r a t u m a n d elevating t h e r e a r p a r t of the body u p w a r d ; the wings also may be o p e n e d or raised to b e t t e r e x p o s e these colors, w a r n i n g would-be p r e d a t o r s of p r e ­ sumably noxious body fluids. Neoconocephalus (fig. 5.3d) is a n o t h e r c o m m o n Neotropical c o n e - h e a d , whose large size (BWL 5 - 6 cm) a n d characteristic

Rgure 5.3 KATYDIDS (TETTIGONIIDAE). (a) Meadow katydid (Conocephalus sp.). (b) Multicol­ ored katydid (Copiphora sp.). (c) Spike-headed katydid (Panacanthus sp.), nymph, (d) Cone-headed katydid (Neoconocephalus sp.).

KATYDIDS

157

loud, p e n e t r a t i n g calls h a v e m a d e t h e m attractive subjects for collectors a n d field studies of b e h a v i o r (Greenfield 1990). Many of t h e species a r e barely distinguish­ able morphologically b u t can be s e p a r a t e d readily by t h e distinctive s o u n d p a t t e r n s of the males (Walker a n d Greenfield 1983). T h e e n o r m o u s h e a d of the n y m p h of the s p i k e - h e a d e d katydid (Panacanthus; fig. 5.3c) has b u l b o u s eyes a n d is grotesquely a d o r n e d with a p e r i p h e r a l crown of large teeth a n d a multispiked h o r n o n the fore­ h e a d . A S o u t h A m e r i c a n g e n u s , Coniungoptera, has b e e n recently a d d e d to the c o n e - h e a d g r o u p (Rentz a n d G u r n e y 1985). It is associated with Nothophagus in t h e s o u t h e r n beech forests a n d is most closely related to an Australian g e n u s , a n o t h e r e x a m p l e of an a m p h i n o t i c distribution.

References GREENFIELD, M. C. 1983. Unsynchronized cho­ rusing in the cone-headed katydid Neoconocephalus a/finis (Beauvois). J. Anim. Behav. 31: 102-112. GREENFIELD, M. C. 1990. Evolution of acoustic communication in the genus Neoconocephalus: Discontinuous songs, synchrony, and inter­ specific interactions. In W. J. Bailey and D. C. F. Rentz, The Tettigoniidae: Biology, systematics and evolution. Springer, Berlin. Pp. 71-97. RENTZ, D. C ,

AND A. B. GURNEY. 1985.

The

shield-backed katydids of South America (Orthoptera: Tettigoniidae, Tettigoniinae) and a new tribe of Conocephalinae with genera in Ghile and Australia. Entomol. Scandinavica 16: 69-119. WALKER, T

J.,

AND M.

D. GREENFIELD.

1983.

Songs and systematics of Caribbean Neoconocephalus (Orthoptera: Tettigoniidae). Amer. Entomol. Soc. Trans. 109: 357-389.

CRICKETS Gryllidae. Spanish: Grillos (General). Portuguese: Grilos (Brazil). Although many orthopteroids produce highly a u d i b l e a n d c o m p l e x s o u n d s , it is the crickets that a r e best k n o w n for their

158

musical talents. Males can m a k e a variety of notes, most characteristically, a series of short, pulsed chirps o r c o n t i n u o u s soft trilling, m o r e melodious a n d less rasping t h a n the calls of katydids. T h e s o u n d is p r o d u c e d by t h e vibration of m e m b r a n o u s areas of the fore wings. T h e s e a r e set into motion w h e n the scraper n e a r t h e base of o n e wing is d r a w n across the o p p o s i n g file of the other. T h i s acoustic behavior plays a major role in pair formation. Generally, the male gives a species-specific calling s o n g to which females r e s p o n d by a p p r o a c h i n g him. Males of g r o u n d - d w e l l i n g types call with the wings raised at a b o u t a 4 0 - d e g r e e angle to t h e long body axis; those inhabit­ ing vegetation hold their fore wings u p at a right angle to the body axis. T h e s e posi­ tions a r e related to s o u n d - p r o d u c i n g effi­ ciency in their habitats (Forrest 1982). T h e Central A m e r i c a n long-legged Amphiacusta maya, f o u n d d u r i n g the day in hollow trees a n d u n d e r o v e r h a n g i n g banks, mates in g r o u p s consisting of b o t h sexes a n d subadult n y m p h s . Male c o u r t s h i p chirp­ ing a p p e a r s to be m o r e of a w a r n i n g to o t h e r males than a signal to females, to r e d u c e fighting that r e d u c e s successful mat­ ing frequency (Boake 1984). T h e habits of related species, such as A. annulipes (fig. 5.4a), a r e mostly known. T h e r e is evidence that crickets also r e s p o n d with evasive be­ havior to the ultrasonic e m a n a t i o n s of bats, o n e of their chief p r e d a t o r s (Doherty and H o y 1985). Crickets may be flattened or cylindrical in cross section. Most a r e small to m e d i u m sized (BL 5 - 2 5 m m ) b u t often have very m u c h e n l a r g e d , muscled h i n d f e m o r a , mak­ ing t h e m good j u m p e r s . T h e n u m b e r of tarsal s e g m e n t s in the legs is r e d u c e d to three, not five like most o t h e r o r t h o p t e r o i d s . T h e female's ovipositor is t u b u l a r a n d the cerci frequently large a n d conspicuous. T h e r e a r e t h r e e major cricket types: (1) "bush crickets," which are small to m e d i u m sized, brownish, a n d a r b o r e a l (several sub-

ORTHOPTEROIDS AND OTHER ORDERS

Figure 5.4 CRICKETS, (a) Long-legged cricket (Amphiacusta annulipes, Gryllidae). (b) Bush cricket (Eneoptera surinamensis, Gryllidae). (c) Tree cricket (Oecanthus sp., Gryllidae), male. (d) Ground cricket (Gryllus sp., Gryllidae). (e) Mole cricket (Scapteriscus sp., Gryllotalpidae). families; a w i d e s p r e a d species is Eneoptera surinamensis; fig. 5.4b); (2) "tree crickets" (Oecanthinae, especially Oecanthus, fig. 5.4c, a n d Neoxabea), which a r e also f o u n d in vegetation b u t a r e delicate, pale g r e e n , a n d translucent, with s l e n d e r bodies a n d legs and almost horizontal h e a d s , males having broad, oval v/ings a n d thoracic glands that produce a m a t i n g p h e r o m o n e (Walker 1967); (3) " g r o u n d a n d field crickets" (Gryllinae a n d o t h e r subfamilies), which are small to large a n d robust a n d generally live on the g r o u n d . A m o n g t h e last g r o u p a r e some species of native Gryllus, many g o i n g u n d e r the aggregate n a m e G. assimilis (fig. 5.4d) (Aguilar a n d Saenz 1970). T h e related European h o u s e cricket (Acheta domesticus) has been i n t r o d u c e d into Latin A m e r i c a n houses from E u r o p e b u t is not generally common. A l r e a d y w i d e s p r e a d a n d increas­ ing its r a n g e rapidly is t h e I n d i a n h o u s e cricket (Grylloides supplicans). Probably because of their familiarity and the pleasant s o u n d s emitted by the domestic species, n u m e r o u s superstitions and folk beliefs have built u p a r o u n d crickets. T h e y a r e almost universally con­ sidered a sign of good luck, a l t h o u g h sometimes a h a r b i n g e r of d e a t h . In Caraguatatuba, Brazil, a black cricket in a r o o m •s a signal of sickness, a gray o n e a sign of money, a n d g r e e n , of h o p e . I n Rio G r a n d e do Sul, Brazil, killing a cricket is t h o u g h t to bring rain ( L e n k o a n d P a p a v e r o 1979).

In contrast to these mostly positive ways of viewing crickets, t h e leanings of these insects toward p o p u l a t i o n explosions a n d their taste for c r o p s stands t h e m in p o o r stead with the farmer. N o c t u r n a l a n d at­ tracted by light a n d food, p l a g u e s of mil­ lions may invade fields a n d nearby u r b a n areas, c o n s u m i n g leaves a n d covering everything, even piling u p in drifts m a n y centimeters d e e p in p r o t e c t e d places. T h e habits of native crickets a r e n o t well k n o w n a m o n g the poorly studied Latin A m e r i c a n fauna. Crickets a r e generally h e r b i v o r o u s on grasses, h e r b s , a n d o t h e r plants b u t take insect p r e y o p p o r t u n i s t i ­ cally. Young tree crickets may be entirely p r e d a c e o u s on scale insects, a p h i d s , a n d o t h e r small insects a n d t h u s may b e of considerable benefit as n a t u r a l control of these pests. T h e family is large, a l t h o u g h only s o m e 500 species a r e now listed for Latin A m e r ­ ica ( C h o p a r d 1 9 6 7 - 6 8 ) ; many species a r e awaiting description. As a g r o u p , they are widely distributed a n d a r e found in almost all life zones.

References AGUILAR, P. G., AND D. SAENZ. 1970.

Algunas

variaciones morfológicas en el grillo común de la costa central. Rev. Peruana Entomol. 13: 76-86. BOAKE, C. R. B. 1984. Natural history and acoustic behavior of a gregarious cricket. Behaviour 89: 241-250. CHOPARD, L. 1967-68. Gryllides. In M. Beier,

CRICKETS

159

ed., Orthopterorum Gravenhage.

catalogus. W. Junk,

DOHERTY, J., AND R. HOY. 1985. T h e acoustic

behavior of crickets: Some views of genetic coupling, song recognition, and predator de­ tection. Quart. Rev. Biol. 60: 457-472. FORREST, T. G. 1982. Acoustic communication and baffling behaviors of crickets. Fla. Entomol. 65: 33-44. LENKO, K., AND N. PAPAVERO. 1979. lnsetos no

folclore. Sec. Cult. Cien. Tech, Sao Paulo. WALKER T J . 1967. Revision of the Oecanthinae (Gryllidae: Orthoptera) of America south of the United States. Entomol. Soc. Amer. Ann. 60:784-796.

Mole Crickets O r t h o p t e r a , Gryllotalpidae, Scapteriscus a n d Neocurtilla. Spanish: Grillotopos (General), c h a n g a s ( P u e r t o Rico), p e r r i t o s d e m o n t e , playacuros (Peru). Portuguese: Grilos t o u p e i r o s , frades, macacos, c a c h o r r i n h o s , d a g u a s , p a q u i n h a s (Brazil). G r o u n d p u p p i e s . Mole crickets (fig. 5.4e), as their n a m e implies, live in s u b t e r r a n e a n galleries. T h e y a r e aptly specialized for d i g g i n g by the modified forelegs, which a r e powerful excavating tools. T h e s e g m e n t s a r e massive a n d powerfully muscled a n d b e a r heavy spines o n t h e u n d e r m a r g i n to form cut­ ting chisels a n d s c r a p e r s . O t h e r identifying features of these insects a r e their cricket­ like f o r m , fairly l a r g e size ( B L 3—4 c m ) , m e d i u m b r o w n color, a n d abbreviated fore wings b e y o n d t h e tips of which t h e twisted h i n d wings project for s o m e distance. Also, the e n t i r e surface of t h e b o d y is covered with a short, sparse, yellowish velvet. Female m o l e crickets deposit their eggs in a cluster in t h e e n l a r g e d c h a m b e r at t h e e n d of a side gallery. T h e n y m p h s r e m a i n u n d e r g r o u n d , f e e d i n g directly o n h e r b a ­ ceous plant roots o r o n stems of plants that they pull d o w n into t h e i r u n d e r g r o u n d passages. T h e adults feed in t h e s a m e way but m a y also leave t h e b u r r o w at night to disperse a n d find m a t e s . T h e y a r e fre­

160

quently attracted to artificial lights. T h e i r food most often consists of seedlings or small crops growing in t h e friable soil they prefer for their diggings. S o m e species take considerable animal m a t t e r as food as well a n d should b e c o n s i d e r e d omnivores (Castner a n d Fowler 1984a). A l t h o u g h superficially similar, m a n y dis­ tinct species in various g e n e r a (especially Scapteriscus) occur in Latin A m e r i c a with varied life-styles (Fowler a n d d e Vascon­ celos 1989). T h e best-known mole cricket, because of its taste for commercial plant­ ings, is t h e changa of P u e r t o Rico (Barrett 1902, T h o m a s 1928). Formerly t h o u g h t to be o n e species, Scapteriscus "vicinus," (actu­ ally didaciylus b u t misidentified by all early a u t h o r s ) , it m a y b e c o m p r i s e d of two o r m o r e as indicated by t h e existence of populations with d i v e r g e n t m a t i n g songs; o n e was n a m e d S. imitatus (Nickle a n d Castner 1984). T h e s e a n d S. abbreviatus have been i n t r o d u c e d to P u e r t o Rico from South America (Castner a n d Fowler 19846) a n d h a v e b e c o m e pests of t u r f a n d agricul­ t u r e . Scapteriscus oxydaclylus is t h e largest South A m e r i c a n mole cricket a n d is a pest of rice a n d other crops cultivated along the banks of t h e A m a z o n River w h e n the waters a r e receding. S o m e n a t u r a l control is w r o u g h t by t h e cricket's parasitoid ene­ mies in t h e sphecid wasp g e n u s Larra (Castner 1984, Castner a n d Fowler 19846). Male mole crickets u s e songs to call mates (Forrest 1983). T h e s o u n d is a bro­ ken trill at a pulse rate of a b o u t 57 to 68 p e r second a n d of low frequency. It is emitted by t h e insect u n d e r g r o u n d , either from closed b u r r o w s o r t h r o u g h special funnel-shaped o p e n i n g s to a u g m e n t the s o u n d ("acoustic horns") (Bennet-Clark 1970). Females occasionally also make s o u n d s of short d u r a t i o n a n d for u n k n o w n p u r p o s e s (Ulagaraj 1976). Stridulation is accomplished by friction between scraper a n d file elements on t h e fore wings, much like t r u e crickets (Bennet-Clark 1970).

ORTHOPTEROIDS AND OTHER ORDERS

GRASSHOPPERS

References BARRETT, O. W. 1902. T h e changa, or mole cricket {Scapteriscus didactylus Latr). Puerto Rico Agrie. Exper. Sta. Bull. 2: 1-19. BENNET-CLARK, H. C. 1970. T h e mechanism

and efficiency of sound production in mole crickets. J. Exper. Biol. 52: 619-652. CASTNER, J- L. 1984. Suitability of Scapteriscus spp. mole crickets [Ort.: Gryllotalpidae] as hosts of Larra bicolor [Hym.: Sphecidae]. Entomophaga 29: 323-329. CASTNER, J- L., AND H. G. FOWLER 1984a. Gut

content analysis of Puerto Rican mole crickets (Orthoptera: Gryllotalpidae: Scapteriscus). Fla. Entomol. 6 7 : 4 7 9 - 4 8 1 . CASTNER, J- L., AND H.

G.

FOWLER.

19846.

Distribution of mole crickets (Orthoptera: Gryllotalpidae: Scapteriscus) and the mole cricket parasitoid Larra bicolor (Hymenoptera: Sphecidae) in Puerto Rico. Fla. Ento­ mol. 67: 481-484. FORREST, T. G. 1983. Phonotaxis and calling in Puerto Rican mole crickets. (Orthoptera: Gryllotalpidae). Entomol. Soc. Amer. Ann. 76: 797-799. FOWLER, H. G., AND H. L. DE VASCONCELOS.

1989. Preliminary data on life cycles of some mole crickets (Orthoptera, Gryllotalpidae) of the Amazon Basin. Rev. Brasil. Entomol. 33: 134-141.

Acrididae. Spanish: Saltamontes, saltarines, saltones (General); t u c u r a s (Argentina). Náhuatl: C h a p u l t i n s , sing. chapolin (Mexico). Quechua: Chili c u t u . Portuguese: Gafanhotos. G r a s s h o p p e r s a r e so familiar that they h a r d l y n e e d description. T h e characteris­ tics of e n l a r g e d h i n d f e m u r (containing t h e j u m p i n g muscles) c o m b i n e d with short, stout a n t e n n a e always distinguish t h e m from similar o r t h o p t e r o i d s . T h e dorsal p a r t of t h e p r o t h o r a x f o r m s a b r o a d collar, folding over t h e s e g m e n t o n t h e sides, a n d is often crested middorsally. B o t h sexes also possess a n auditory o r g a n , visible as a circular m e m b r a n e , o n either side of t h e basal a b d o m i n a l segment. Most h a v e fully developed wings, t h e fore wing b e i n g elon­ gate a n d parallel sided, t h e h i n d wing b r o a d , fanlike, a n d reticulate veined. Many types have r e d u c e d o r virtually n o wings a n d a r e flightless, particularly s o m e high A n d e a n (Melanoplus; R o b e r t s 1973) a n d rain forest (Rowell 1978) types.

NICKLE, D. A., AND J. L. CASTNER 1984. Intro­

duced species of mole crickets in the United States, Puerto Rico, and the Virgin Islands (Orthoptera: Gryllotalpidae). Entomol. Soc. Amer. Ann. 77: 450-465. THOMAS, W. A. 1928. T h e Puerto Rican mole cricket. U.S. Dept. Agr., Farm. Bull. 1561: 1-8. ULAGARAJ, S. M. 1976. Sound production in mole crickets (Orthoptera: Gryllotalpidae: Scapteriscus). Entomol. Soc. Amer. Ann. 69: 299-306.

Short-Horned Orthopteroids O r t h o p t e r a ( = Caelifera) Grasshoppers m a k e u p t h e majority of this group, which is characterized by short antennae with less t h a n thirty s e g m e n t s . Auditory o r g a n s , w h e n present, a r e lo­ cated o n t h e t e r g u m of t h e first a b d o m i n a l segment. Stridulatory s t r u c t u r e s a r e o n t h e distal p o r t i o n s of t h e wings.

T h e female uses h e r short ovipositor to excavate holes in t h e soil into which s h e deposits u p to o n e h u n d r e d e l o n g a t e eggs in a mass c e m e n t e d t o g e t h e r with a viscid secretion. Both n y m p h s a n d adults feed o n vegetation a n d w h e n a b u n d a n t , a r e devas­ tating c r o p pests. C e r t a i n species, espe­ cially in t h e g e n u s Schistocerca, form e n o r ­ m o u s m i g r a t i n g swarms that can destroy fields over wide areas. T h e s e a r e t h e lo­ custs, most f a m o u s in t h e O l d World b u t r e p r e s e n t e d by a species in Latin A m e r i c a n o less imposing o r devastating. A l t h o u g h g r a s s h o p p e r s have b e e n used as sustenance by p e o p l e s in Africa, t h e Middle East, a n d o t h e r parts of t h e world, both in history a n d today, they seem to b e of relatively m i n o r i m p o r t a n c e as a food in tropical America. Exceptions in t h e past w e r e t h e natives of several West I n d i a n islands w h o , as M a r t y r ( B o d e n h e i m e r

GRASSHOPPERS

161

1951: 301) r e c o r d s , a t e g r a s s h o p p e r s a n d stored t h e m for t r a d e . P a d r e Florian P a u c h e , w h o lived with t h e Mocovies Indi­ ans of A r g e n t i n a in t h e e i g h t e e n t h century, described their practice o f catching, cook­ ing, a n d e a t i n g t h e locusts (Schistocerca) that periodically p l a g u e d their land (Lieb e r m a n n 1948). G r a s s h o p p e r s were i m p o r t a n t in t h e diet of t h e Aztecs; o n e species was called acáhapali, arrow, in N á h u a t l , because of its s h a p e a n d because it m a d e a distinctive buzz in flight ( C u r r a n 1937). A c c o r d i n g to Kevan (1977), t h e w o r d chapulli a p p a r e n t l y r e f e r r e d to m a n y edible species, particu­ larly Sphenarium, a n d f o r m e d t h e basis for m a n y p l a c e - n a m e s in c e n t r a l Mexico d u r ­ ing t h e p e r i o d of t h e Aztec civilization. A hill in Mexico City, f a m o u s to t h e Aztecs a n d w h e r e Maximilian's castle is located, was n a m e d chapultepec, c o m b i n i n g t h e word for g r a s s h o p p e r with hill (tepeque). G r a s s h o p p e r s also a p p e a r as design motifs in t h e a r t of early M i d d l e A m e r i c a n cul­ t u r e s ( B o n i n g 1971). G r a s s h o p p e r s live in all habitats a n d generally h a v e highly specialized life forms to a d a p t to prevailing e n v i r o n m e n t a l condi­ tions. T h e y s e e m to p r e f e r o p e n habitats a n d a r e most n u m e r o u s a n d diverse in grasslands. T h e r e a r e indications that u n d e r s t o r y species d e c r e a s e as o n e e n t e r s tropical forests (Brodey 1975), b u t distinct c a n o p y (Roberts 1 9 7 3 , D e s c a m p s 1976) a n d u n d e r s t o r y (Rowell 1978) assemblages a r e evident. T h e r e a r e even successional stages o f these categories following clear­ ing of forest ( A m e d e g n a t o a n d Descamps 1980). Many g r a s s h o p p e r s can m a k e s o u n d s by frictional contact between various p a r t s of the body, usually o n e of t h e h i n d leg s e g m e n t s against t h e fore wings. In t h e winged l u b b e r s , r a p i d closing of t h e h i n d wings b r i n g s s e r r a t e veinlets o n this wing in contact with s c r a p e r veins o n t h e fore wing, p r o d u c i n g a s n a p p i n g o r rattling s o u n d . A c r a c k i n g is often h e a r d from

162

b a n d wings in flight. It is t h o u g h t to be caused by t h e partial folding a n d rapid e x p a n s i o n of t h e h i n d wing (Uvarov 1966: 176f.). In addition to t h e locusts (Schistocerca), which mass in flight, t h e n y m p h s of some lubbers also a r e m i g r a t o r y o n foot. Young lubbers also form d e n s e g r o u p s o n t h e tops of plants. T h e y a r e m o r e brightly colored than t h e adults a n d a r e certainly distaste­ ful, a fact advertised m o r e effectively en masse t h a n individually. Lowland a n d lower m o n t a n e forest g r a s s h o p p e r s have been f o u n d to have n a r r o w feeding prefer­ ences (Rowell et al. 1984). Probably, many such feeding specialists a r e sequestering toxic chemicals from their hosts (Rowell 1978). G r a s s h o p p e r s a r e essentially land in­ sects, b u t a few have semiaquatic habits in South America. T h e best-studied examples a r e Marellia remipes, Paulinia acuminata (fig. 5.5b), a n d Cornops aquaticum (Pauliniidae), which live on b r o a d , floating leaves of water lilies a n d o t h e r aquatic plants. T h e y frequently fall into t h e water after a hasty j u m p . T h e y show clear m o r p h o l o g i c a l and behaviorial a d a p t a t i o n s to life in water including p a d d l e - s h a p e d h i n d legs in some species (Bentos a n d Lorier 1991). Females often insert their eggs into t h e stems of water plants (Cornops) o r place t h e m on the u n d e r s i d e s of floating leaves (Marellia and Paulinia) (Carbonell 1959, 1964). T h e so-called b a n d - w i n g g r a s s h o p p e r s sport brightly colored r e d , blue, purple, g r e e n , yellow, o r o r a n g e h i n d wings. To this g r o u p b e l o n g t h e c o m m o n , wide­ s p r e a d , arid-land species, Trimerotropis pallidipennis (fig. 5.5a), recognized by its trans­ lucent yellow h i n d wings. T h e fore wings are usually cryptically colored to m a t c h the gravel o r sandy soil o n which t h e species habitually rests. S o m e grass-loving types (Achurum; fig. 5.5d) a r e very slender a n d elongate, better to h i d e a m o n g t h e blades on which they rest.

ORTHOPTEROIDS AND OTHER ORDERS

With their h i n d legs held o u t at right

Figure 5.5 GRASSHOPPERS, (a) Band-wing grasshopper (Trimerotropis pallidipennis, Acrididae). (b) Aquatic grasshopper (Paulinia acuminata, Pauliniidae). (c) American locust (Schistocerca piceifrons, Acrididae). (d) Grass-mimicking grasshopper (Achurum sumichrasti, Acrididae). (e) Eumastacid grasshopper (Eumastax sp., Eumastacidae).

angles to t h e body, Eumastax (Eumasta­ cidae; fig. 5.5e) bask o n leaves in t h e w a r m pools of sunlight that flood into t h e forest u n d e r g r o w t h below holes in t h e c a n o p y (pi. lc). T h e s u n b e a m s excite t h e irides­ cence in t h e g r e e n s , blues, a n d yellows o n the bodies of these forms, m a k i n g t h e m glow like jewels. Uvarov (1977: 42If.) p r e s e n t s a faunistic s u m m a r y of Latin A m e r i c a n g r a s s h o p p e r s ; A m e d e g n a t o (1974) reviews t h e g e n e r a . T h e g e o g r a p h i c a n d evolutionary history of the regional forms h a s b e e n traced by various a u t h o r s (Carbonell 1977, A m e d e ­ gnato a n d D e s c a m p s 1979).

References AMEDEGNATO, C. 1974. Les genres d'Acridiens néotropicaux, leur classification par families, sous-families el tribus. Acrida 3: 193-203. AMEDEGNATO,

C,

AND M. DESCAMPS.

1979.

History and phylogeny of the Neotropical acridid fauna, Metaleptea. Soc. Panamer. Acridiologia2(l): 1-10. AMEDEGNATO,

C,

AND M.

DESCAMPS.

1980.

Evolution des populations d'Orthopteres d'Amazonie du nord-ouest dans les cultures traditionnelles et les formations secondaires d'origine anthropique. Acrida 9: 1-33. BENTOS, A., AND E. LORIER 1991. Acridomorfos

acuáticos (Orthoptera, Acridoidea). I. Adapta­ ciones morfológicas. Rev. Brasil. Entomol. 35: 631-653. BODENHEIMER, F. S. 1951. Insects as human food. Junk, T h e Hague. BONING, K. 1971. Mesoamerikanische Heu-

schreckendarstellungen. Anz. Schádlingskunde Pffanzenschutz 44: 185-189. BRODEY, K. 1975. A study of grasshopper species composition in primary and secon­ dary growth in Costa Rica. Entomol. News 86: 207-211. CARBONELL, C. S. 1959. The external anatomy of the South American semiaquatic grasshop­ per Marellia remipes Uvarov (Acridoidea, Pau­ liniidae). Smithsonian Misc. Coll. 137: 61-97. CARBONELL, C. S. 1964. Habitat, etología y ontongenia de Paulinia acuminata (Dg.), (Ac­ ridoidea, Pauliniidae) en el Uruguay. Soc. Uruguayo Entomol. Rev. 6: 39-48. CARBONELL, C. S. 1977. Origin, evolution, and distribution of the Neotropical acridomorph fauna (Orthoptera): A preliminary hypothe­ sis. Soc. Entomol. Argentina Rev. 36: 153-175. CURRAN, C. H. 1937. Insect lore of the Aztecs. Nat. Hist. 39: 196-203. DESCAMPS, M. 1976. La faune dendrophile néotropicale. I. Revue des Proctolabinae (Orth. Acrididae). Acrida 5: 63-167. KEVAN, D. K. McE. 1977. The American Pyrgomorphidae (Orthoptera). Soc. Entomol. Ar­ gentina Rev. 36: 3-28. LIEBERMANN, J. 1948. Curiosidades históricas acerca de la langosta. Aim. Min. Agrie. Ganad. Buenos Aires 23: 434-438. ROBERTS, H. R. 1973. Arboreal Orthoptera in the rain forests of Costa Rica collected with insecticide: A report on the grasshoppers (Acrididae), including new species. Acad. Nat. Sci. Philadelphia Proc. 125: 49-66. ROWELL, C. H. F. 1978. Food plant specificity in Neotropical rain-forest acridids. Entomol. Exper. Appl. 24: 651-662. ROWELL, C. H. F , M. ROWELL-RAHIER, H. E. BAKER, G. COOPER-DRIVER, AND L. D. GÓMEZ.

f984. The palatability of ferns and the ecol-

GRASSHOPPERS

163

ogy of two tropical foresl Biotropica 15:207-216. UVAROV, B. 1966,

grasshoppers.

1977. Grasshoppers

and

locusts: A handbook of general acridology. 2 vols. Cambridge Univ. Press, Cambridge.

Locusts Acrididae, C y r t a c a n t h a c r i d i n a e , Schistocerca. Spanish: Langostas, langostas voladoras (General). Portuguese: G a f a n h o t o s d e p r a g a . A l t h o u g h n o t all easily distinguished, twenty to thirty k i n d s of Schistocerca inhabit tropical A m e r i c a . T h e y a r e all large (BL 4 6 cm), s l e n d e r g r a s s h o p p e r s with e x p a n ­ sive wings that e x t e n d well beyond t h e tip of t h e a b d o m e n w h e n folded. Females a r e m u c h larger t h a n t h e males. T h e fore wings of both sexes a r e varied in coloration b u t a r e generally dull, yellowish with ir­ r e g u l a r brownish spots; t h e h i n d wings a r e m o r e o r less pelucid. T h e s e forms a r e variable in behavior as well as in coloration. C e r t a i n of t h e m p r o d u c e large s w a r m s a n d m i g r a t e periodi­ cally, like t h e i n f a m o u s O l d World migra­ tory locust (S. gregaria) t o which they a r e closely related. S o m e of this variation may be caused by d e v e l o p m e n t a l influences, especially c r o w d i n g , l e a d i n g to so-called phases. T h e r e is c o n s i d e r a b l e confusion in the literature r e g a r d i n g t h e exact species status, interrelationships, a n d genetic sig­ nificance of all these f o r m s a n d phases. I n o n e t a x o n o m i c study based o n e x t e r n a l morphology, t h e m i g r a t o r y types have b e e n relegated to a single subspecies (S. paranensis) within a w i d e s p r e a d species, 5. americana, w h e r e v e r they o c c u r in C e n t r a l o r S o u t h A m e r i c a (Dirsh 1974), while oth­ ers classify t h e locusts differently (Harvey 1981). Hybridization e x p e r i m e n t s , how­ ever, show t h e p i c t u r e to be still m o r e c o m p l e x : a t least 5. piceifrons (fig. 5.5c), 5. americana, a n d S. cancellata should be con­ s i d e r e d s e p a r a t e species (Harvey 1979, J a g o et al. 1982), t h e first being t h e t r u e s w a r m i n g pest (Harvey 1983).

164

W h a t e v e r its correct n a m e , this locust is well k n o w n in Latin A m e r i c a for its ravages to crops a n d r a n g e l a n d from p r e h i s t o r y to m o d e r n times. Descriptions of the invasions rival t h e stories of t h e O l d World species r e c o u n t e d in t h e Bible a n d o t h e r historical writings. While crossing t h e d r y p a m p a s of A r g e n t i n a in March 1835, C h a r l e s Darwin (1962 [1845]: 3 3 0 - 3 3 1 ) wrote, Shortly before arriving at the village and river of Luxan, we observed, to the south, a ragged cloud of a dark reddish brown color. At first we thought it was caused by some great fire on the neighboring plains, but we soon found that it was a swarm of lo­ custs. . . . The sound of their wings was as the sound of chariots of many horses run­ ning to battle. . . . The sky, seen through the advanced guard, appeared like a mezzotinto engraving; but the main body was impervi­ ous to sight. . . . When they alighted, they were more numerous than the leaves in the field, and the surface became reddish in­ stead of green. T h e s e events a r e k n o w n in many parts of the region, most c o m m o n l y a n d regularly in n o r t h e r n A r g e n t i n a a n d s o u t h e r n Brazil but also to a limited extent in coastal Chile a n d t h e A n d e a n m o u n t a i n valleys of Peru, Ecuador, a n d Colombia as well as in scat­ tered localities in C e n t r a l A m e r i c a and Mexico. T h e a n n u a l p a t t e r n s of b r e e d i n g a r e strongly influenced by t h e w e a t h e r a n d its seasonal variations. B r e e d i n g is confined to t h e period of s u m m e r rains. O u t b r e a k s occur in t h e driest p a r t s of t h e species' r a n g e , triggered by t h e occasional abun­ d a n t rainfalls that foster t h e insect's high r e p r o d u c t i v e capacity ( H u n t e r a n d Cosenzo 1990, Waloff a n d Pedgley 1986). In most s o u t h e r n areas, t h e r e a r e two a n n u a l generations, b e g i n n i n g with mating a n d oviposition in N o v e m b e r a n d Decem­ ber. T h e n y m p h s h a t c h a n d a r e active and g r o w i n g from D e c e m b e r to April. T h e y become adults t h r o u g h April a n d May,

ORTHOPTEROIDS AND OTHER ORDERS

when t h e m i g r a t i o n s to alternative b r e e d ­ ing g r o u n d s take place. H e r e a second m a t ' n g a n d e g g laying ensues, a n d t h e young of this g e n e r a t i o n p r o d u c e a whole new b r o o d o f adults by S e p t e m b e r to November which migrates back to t h e areas of original habitat ( D a g u e r r e 1953). O t h e r Schistocerca species a n d subspecies overlap t h e r a n g e of t h e t r u e locusts in diverse p a r t s of Latin America, all of a totally s e d e n t a r y n a t u r e . T h e s e include endemic species o n m a n y isolated islands in the West I n d i e s , B e r m u d a , a n d t h e Galápagos a n d Revillagigedo archipelagos.

South American locusl, Schistocerca cancellata (Orthoptera: Acrididae) in Argentina. Bull. Entomol. Res. 80: 295-300. JAGO, N. D., A. ANTONIOU, AND J. P. GRUNSHAW.

1982. Further laboratory evidence for the separate species status of the South American locusl {Schistocerca cancellata Serville) and the Central American locust (Schistocerca piceifrons piceifrons) (Acrididae, Cyrtacanthacridinae). J. Nat. Hist. 16: 763-768. WALOFF, Z., AND D. E. PEDGLEY. 1986. Compara­

tive biogeography and biology of the South American locust, Schistocerca cancellata (Ser­ ville), and the South African desert locust, S. gregaria flaviventris (Burmeister) (Orthoptera: Acrididae): A review. Bull. Entomol. Res. 76: 1-20.

References

Lubber Grasshoppers

DAGUERRE, J. B. 1953. Vida de la langosta voladora. Reun. Com. Interamer. Perm. Antiacrid (Puerto Alegre), 1952: 55-79. DARWIN, C. 1962 [1845]. T h e voyage of the Beagle. Edited by L. Engel. Doubleday-Amer. Mus. Nat. Hist, New York. DlRSH, V. M. 1974. Genus Schistocerca. Junk (Ser. Entomol. 10), T h e Hague. HARVEY, A. W. 1979. Hybridization studies in the Schistocerca americana complex. I. The specific status of the Central American Lo­ cust. Biol. J. Linnean Soc. 12: 349-355.

Romaleidae.

HARVEY, A. W. 1981. A reclassification of the

Schistocerca americana complex. Acrida 10: 61-77. HARVEY, A. W. 1983. Schistocerca piceifrons (Walker), the swarming locust of tropical America: A review. Bull. Entomol. Res. 73: 171-184. HUNTER, D. M., AND E. L. COSENZO. 1990.

The

origin of plagues and recent outbreaks ol the

T h i s g r a s s h o p p e r family is f o u n d only in the New World a n d consists c u r r e n t l y of forty-eight g e n e r a a n d at least d o u b l e that n u m b e r of species, distributed over all t h e Neotropical Region b u t n o t o n t h e Antilles except C u b a (Rehn a n d G r a n t 1959). T h e g r o u p is of very diverse m o r p h o l o g y but is generally m a d e u p of quite large, heavybodied forms, including t h e biggest of t h e world's g r a s s h o p p e r s , Tropidacris (fig. 5.6a), with body lengths of 8 to 9 c e n t i m e t e r s . T h e y a r e fully winged to a p t e r o u s , a n d many h a v e partially d e v e l o p e d wings n e v e r used for flight. T h e i n t e g u m e n t is often g r a n u l a t e o r with p r o n o u n c e d , t u b e r c u l a t e points. T h e thoracic disk usually h a s a

Rflure 5.6 LUBBER GRASSHOPPERS (ROMALEIDAE). (a) Giant grasshopper (Tropidacris cristata). (b) Lubber grasshopper (Taenopoda varipennis). (c) Rainbow-winged grasshopper (77tenacris gloriosa), (d) Independence grasshopper (Chromacris speciosa).

GRASSHOPPERS

165

definite crest b u t may be wide o r simple, like that of o t h e r acridids. T h e v e r n a c u l a r n a m e of these g r a s s h o p ­ p e r s refers to t h e terrestrial habits of t h e majority of species, which is dictated by their flightlessness ("landlubbers"). Unable to fly, they rely o n o t h e r m o d e s of selfp r o t e c t i o n , often releasing r e p u g n a n t se­ cretions from g l a n d s o r t h e m o u t h a n d emitting t h r e a t e n i n g s o u n d s . Release of n o x i o u s liquids o r foams from t h e mesothoracic spiracle can b e a c c o m p a n i e d by a hissing a n d buzzing of t h e wings in an i m p o s i n g threat display. T h e s e a r e abilities of t h e c o m m o n g e n u s Taenopoda, for e x a m p l e , which is r e p r e ­ s e n t e d by several species, i n c l u d i n g t h e well-known T. eques, in t h e d r i e r parts of n o r t h e r n Mexico, d o w n t h r o u g h C e n t r a l A m e r i c a to P a n a m a , w h e r e T. varipennis (fig. 5.6b) takes over. T h e y a r e usually fond of mimosas a n d acacias, t h e i r p r i m a r y food plants. Taenopoda a r e m o d e r a t e l y large (BL 6—7 cm) a n d h a v e only slightly d i m i n u t i v e wings. T h e fore wings a r e basically g r e e n , with fine, yellow reticulation o r d a r k blotches. T h e n y m p h s a r e black with r e d gilding a n d a r e g r e g a r i o u s . T h e y a p p a r ­ ently rely o n unpalatability for protection, b e i n g so c o n s p i c u o u s in color a n d v u l n e r a ­ ble on e x p o s e d , h e r b a c e o u s vegetation. A n o t h e r n o r t h e r n species is Brachystola magna, which p r e f e r s grasslands o n the Mexican p l a t e a u a n d is confined to a life on t h e g r o u n d because of its almost com­ plete winglessness a n d inability to climb. T h e fore wing is only a small disk, t h e h i n d wing n o t m u c h m o r e t h a n a vestigial fan. L u b b e r s of t h e g e n u s Chromacris a r e usually a glossy g r e e n with yellow m a r k ­ ings a n d striking r e d o r yellow wings. T h e h i n d wings a r e various s h a d e s of r e d , o r a n g e , o r yellow with c o n t r a s t i n g black b a n d s . T h e y occur in t h e h u m i d p o r t i o n s of t h e A m e r i c a n tropics from Mexico to A r g e n t i n a . All show a p r e f e r e n c e for solanaceous a n d c o m p o s i t e plants a n d have g r e g a r i o u s j u v e n i l e s like o t h e r r o m a -

166

leids. C. speciosa (fig. 5.6d) is the most wider a n g i n g a n d variable species. It is green a n d yellow, t h e colors of t h e Brazilian flag, a n d has b e e n d u b b e d t h e " i n d e p e n d e n c e g r a s s h o p p e r " (gafanhoto da independencia) in the republic ( O h a u s 1990: 2 3 3 , as C. miles; Roberts a n d Carbonell 1982).

References OHAUS, F. 1990. Bericht über eine entomologische Reise nach Centralbrasilien. Stettiner Entomol. Zeit. 61: 164-273. REHN, J. A. G., AND H. J. GRANT, JR. 1959.

A

reviewof theRomaleinae(Orthoptera; Acrididae) found in America north of Mexico. Acad. Nat. Sci. Philadelphia Proc. I l l : 109-271. ROBERTS, H. R., AND C. S. CARBONELL. 1982. A

revision of the grasshopper genera Chromacris and Xestotrachelus (Orthoptera, Romaleidae, Romaleinae). Calif. Acad. Sci., Ser. 4, Proc. 43: 43-58. Giant Grasshoppers R o m a l e i d a e , Tropidacris, Titanacris. Birdwing g r a s s h o p p e r s , rainbow grasshoppers. Tropidacris a n d Titanacris a r e unusually large, fully winged g r a s s h o p p e r s . T h e first g e n u s contains only t h r e e species, which a r e 10 to 13 c e n t i m e t e r s from h e a d to wing tips a n d have a w i n g s p a n of 25 centimeters o r m o r e . A w i d e s p r e a d b u t not c o m m o n species is Tr. cristata. T h e second g e n u s , in which t h e r e a r e seven species (e.g., T. velazquezii, T. gloriosa; D e s c a m p s a n d Car­ bonell 1985), is only slightly smaller, 10 to 11 centimeters in length a n d with a wingspan of u p to 23 centimeters. Aside from their great size, these grass­ h o p p e r s a r e spectacular in flight o r when crepitating from p e r c h e s , w h e n they dis­ play their brilliantly colored h i n d wings. In Tropidacris, these wings a r e generally red with a solid black m a r g i n a l z o n e giving way to a finely c h e c k e r e d gray o r bluish pattern toward t h e wing base. T h e fore wings are gray-green, splotched with gray, a n d the posterior p a r t of t h e p r o n o t u m is flat and yellow a n d g r e e n speckled. T h e h i n d wing

ORTHOPTEROIDS AND O T H E R ORDERS

0f

Titanacris gloriosa (fig. 5.6c) is a veritable rainbow, b r i g h t blue basally, g r a d i n g to crimson anteriorly, a n d t h e n g r e e n over the apical p o r t i o n . T h e fore wings a r e uniformly leaf g r e e n , as is t h e p r o n o t u m ; the latter h a s a s e r r a t e d crest r u n n i n g down t h e e n t i r e m i d d l e to distinguish it from Tropidacris, whose crest r u n s over t h e anterior half of t h e p r o n o t u m only. In spite of their conspicuousness, n o t much has b e e n r e c o r d e d r e g a r d i n g t h e life habits of these e n o r m o u s o r t h o p t e r a n s . They a r e occasionally f o u n d o n s h r u b b y vegetation b u t m o r e normally inhabit t h e crowns of trees a n d a r e particularly active on hot days, s t r i d u l a t i n g a n d readily taking flight. Tr. cristata has b e e n e n c o u n t e r e d o n plants of t h e g e n u s Quassia (Simaroubaceae) which contain repellent chemicals that the insect may sequester (Rowell 1983). T h e i m m a t u r e s a r e g r e g a r i o u s . Young (pers. comm.) r e p o r t s t h e species c o m m o n on Erythrina (Papilionaceae) in Costa Rica.

References DESCAMPS, M.,

AND C.

S. CARBONELL.

1985.

Revision of the Neotropical arboreal genus Titanacris (Orthoptera, Acridoidea, Romalei­ dae). Soc. Entomol. France Ann. (n.s.) 21: 259-285. ROWELL, H. F. 1983. Tropidacris cristata (saltamonte o Chapulín gigante, giant red-winged grasshopper). In D. H. Janzen, ed., Costa Rican natural history. Univ. Chicago Press, Chicago. Pp. 772-773.

flage, especially a m o n g grasses, which a r e a d o m i n a n t habitat. S o m e information re­ g a r d i n g internal a n a t o m y a n d cytology has b e e n p r o v i d e d by Ferreira (1978). T h e r e a r e 153 n a m e d species in seven­ teen g e n e r a , b u t m a n y m o r e probably exist. S o m e k n o w n species may be f o u n d to be c o m p o s e d of multiple species w h e n details of their anatomy, such as in t h e genitalia, a r e studied (Descamps 1973). Apioscelis (fig. 5.7a) is a typical species. T h e s e o r t h o p t e r a n s (Carbonell 1977, Mello-Leitáo 1939) a r e sticklike in s h a p e , fairly large (adult BL 5—10 cm), a n d easily mistaken for walkingsticks ( P h a s m a t o d e a ) (Liana 1972). T h e slightly dilated h i n d femur, elongate t h o r a x , a n d vertical, c o n e s h a p e d g r a s s h o p p e r - t y p e h e a d distinguish t h e m , however. Also, t h e a n t e n n a e a r e short a n d have few s e g m e n t s like t h e g r a s s h o p p e r s , which a r e their closer rela­ tives b u t from which they differ by having a vertical h e a d a n d an elongate p r o t h o r a x that lacks a dorsal shield. T h e p o r t i o n of the head a n t e r i o r to t h e eyes may be extremely p r o l o n g e d in s o m e g e n e r a . Wings a r e completely missing in almost all a n d at most a r e m i n u t e vestiges, a n d they lack tympanic a n d stridulatory o r g a n s . T h e y a r e mostly dull colored, solid olive to b r o w n , a l t h o u g h s o m e have black, r e d , o r yellow m a r k i n g s .

References Jumping Sticks Proscopiidae. Spanish: Palitos vivientes d e antenas cortas (General). Portuguese: Chicos m a g r o s , g a f a n h o t o s d e marmeleiro, g a f a n h o t o s d e j u r e m a , María moles (Brazil). J u m p i n g sticks a r e f o u n d only o n t h e South A m e r i c a n c o n t i n e n t , as far n o r t h as Panama, a n d o n t h e C a r i b b e a n island of Bonaire. T h e i r biology is incompletely u n ­ known. Specimens a r e usually seen resting on vegetation, their cryptic stick form a n d colors giving t h e m a m e a s u r e of c a m o u ­

CARBONELL, C. S. 1977. Superfam. Proscopoidea, Fam. Proscopiidae. In M. Beier, ed., Orthopterum Catalogus. Junk, The Hague. Pp. 1-29. DESCAMPS, M. 1973. Notes préliminaires sur les genitalia de Proscopoidea (Orthoptera Acridomorpha). Acrida 2: 77—95. FERREIRA, A. 1978. Contribuicáo ao estudo da evolucáo dos Proscopiidae (Orthoptera-Proscopoidea). Studia Entorno!. 20: 221-233. LIANA, A. 1972. Etudes sur les Proscopiidae (Orthoptera). Polska Akad. Nauk, lnst. Zool., Ann. Zool. 29: 381-459. MELLO-LEITÁO, C. 1939. Estudio monográfico de los Proscópidos. Mus. La Plata Rev., Nov. Ser., Zool. 8(1): 279-449.

GRASSHOPPERS

167

t h r o u g h an o p e n i n g o n e i t h e r side b e h i n d the head ( M o r e n o 1940). T h i s is a r a t h e r robust stick insect, fairly c o m m o n in parts of s o u t h e r n A r g e n t i n a a n d Chile w h e r e it is known by several v e r n a c u l a r s in Q u e ­ chua (chinchemoyo, "stinking chest") a n d Mapuche (tabolongo, chirindango, fitquilén) (Schneider 1934).

Figure 5.7 JUMPING STICK AND WALKINGSTICKS. (a) Jumping stick (Apioscelis sp., Proscopiidae). (b) Giant walkingstick (Phibalosoma phyllinum, Phibalosomatidae). (c) Chinchemoyo (Paradoxomorpha crassa, Anisomorphidae). (d) Winged walkingstick (Pseudophasma ?, Pseudophasmatidae).

WALKING STICKS Phasmatodea (= Phasmatoptera, P h a s m i d a , P h a s m o d e a ) . Spanish: Bichos palitos, zacatones (General); c h i n c h e m o y o s , palotes (Chile); c a m p a m o c h a s (Mexico); palitos vivientes (Peru). Portuguese: Bichos p a u . Phasmids. A greatly a t t e n u a t e d , sticklike body a n d legs a r e t h e h a l l m a r k s of these c o m m o n but s e l d o m seen insects. T h e i r resem­ blance to twigs a n d leaves is so perfect that they a r e usually discovered in their n a t u r a l h a u n t s only by accident. H u m a n eyes can play time a n d again over a s p e c i m e n in its resting place a n d n e v e r see it. S o m e in central S o u t h A m e r i c a a r e a m o n g t h e larg­ est insects as m e a s u r e d by l e n g t h . Bactridium grande (female B L 26 cm) is t h e biggest; Phibalosoma phyllinum (fig. 5.7b) a n d Otocrania aurita b o t h a r e over 20 centi­ m e t e r s long. T h e r e a r e b o t h w i n g e d (fig. 5.7d) a n d wingless stick insects, a t e n d e n c y toward the latter m o r e in females t h a n in males. W h e n p r e s e n t , t h e h i n d wings only a r e well f o r m e d , a n d t h e n they a r e large, fanlike, a n d efficient flight o r g a n s . T h e fore wings are m u c h r e d u c e d , often only scalelike cups o r e l o n g a t e leathery plates. I n g e n e r a l , t h e g r o u p ( B e d f o r d 1978) shows a n u m b e r of interesting biological as

168

well as structural features, b u t little specific information is available on t h e Neotropical species (Willig et al. 1986, Zapata and Torres 1970). All feed o n plants a n d a r e a p p a r e n t l y fairly host specific. P a r t h e n o ­ genesis occurs in a n u m b e r of E u r o p e a n forms a n d presumably also occurs in re­ gional mantids. An u n e x p l a i n e d character­ istic of some female walkingsticks is a bright r e d color at t h e base of t h e fore femur. Individuals can r e g e n e r a t e lost limbs, a n d those of s o m e species can c h a n g e color to m a t c h their b a c k g r o u n d . Defensive behaviors, c o u p l e d with their cryptic forms a n d colors, a r e well devel­ o p e d a n d parallel to some e x t e n t those of mantids. T h e y include (1) r o c k i n g motions in which t h e body is swayed from side to side by flexing of t h e legs at t h e u p p e r joints, an action t h o u g h t to e n h a n c e resem­ blance to plant parts m o v i n g in t h e breeze; (2) active escape by d r o p p i n g o r flight; (3) flashing of wings to startle o r display aposematic colors; (4) catalepsy, o r freez­ ing o r feigning d e a t h ; (5) s o u n d produc­ tion by wing rattling; (6) fighting with the legs; a n d (7) release of r e p u g n a n t o r caus­ tic chemicals. Paradoxomorpha (fig. 5.7c) can fire an aerosol spray that can blister hu­ m a n skin. T h e substance has b e e n ana­ lyzed as c o n t a i n i n g ethyl e t h e r o r orthoformic acid a n d is p r o d u c e d by a pair of i m m e n s e glands in t h e t h o r a x that exit

ORTHOPTEROIDS AND OTHER ORDERS

A single species m a y employ a variety of these tactics. Pterinoxylus spinulosus from Panama e n h a n c e s its resting p o s t u r e resem­ blance to sticks by closely folding t h e m i d and h i n d legs, which t h e n look like small shoots from t h e m a i n stick (Robinson 1968). phasmid eggs a r e curiously s h a p e d , look­ ing like seeds with a h a r d shell a n d either smooth a n d shiny a n d u n i c o l o r o u s o r heav­ ily sculptured a n d p a t t e r n e d . All have an operculum. T h e y a r e placed carefully o n the host plant o r d i s p e r s e d by flicking movements o f t h e a b d o m e n o r scattered indiscriminately. Stick insects a r e f o u n d in all habitats b u t are most a b u n d a n t a n d diverse in h u m i d forests. Two species, Bostra scabrinota a n d Libethra minúscula, a r e typical of t h e lomas or seasonal h e r b l a n d s of t h e coastal Peru­ vian desert. D e v e l o p m e n t a l stages exhibit conspicuous c h a n g e s of color c o r r e s p o n d ­ ing to t h e flourishing a n d w a n i n g of this vegetation that they mimic. T h e y pass t h e long dry season as eggs o n t h e soil surface (Aguilar 1970). Four c e r a t o p o g o n i d gnats in t h e subgenus Microhelea of Forcipomyia attach to stick insects a n d suck their blood. With t h e exception of o n e I n d o n e s i a n e x a m p l e , these p h a s m i d parasites, called "stick ticks," are known only from tropical A m e r i c a (Wirth 1971). T h e o r d e r h a s b e e n newly reclassified into six families (Bradley a n d Galil 1977) and its world distribution reviewed ( G u n ther 1953). A high p e r c e n t a g e , p e r h a p s 30 percent of m o r e t h a n 2,500 world species, live in Latin A m e r i c a , i n c l u d i n g t h e West Indies (Moxey 1972).

References AGUILAR, P. G. 1970. Los "palitos vivientes de Lima." 1. Phasmatidae de las lomas. Rev. Peruana Entomol. 13: 1-8. BEDFORD, G. O. 1978. Biology and ecology of the Phasmatodea. Ann. Rev. Entomol. 23: 125-149. BRADLEY, J. C., AND B. S. GALIL. 1977.

The

taxonomic arrangement of the Phasmatodea with keys to the subfamilies and tribes. Entomol. Soc. Wash. Proc. 79: 176-208. GUNTHER, K. 1953. Uber die taxonomische Gliederung und die geographische Verbreitung der lnsektenordnung de Phasmatodea. Zeit. Entomol. 3: 541-563. MORENO, A. 1940. Glándulas odoríferas en Paradoxomorpha. Mus. La Plata Not. (Zool. 45) 5: 319-323. MOXEY, C. F. 1972. T h e stick-insects (Phasma­ todea) of the West Indies: Their systematics and biology. Ph.D. diss., Harvard Univ. [Not seen.] ROBINSON, M. H. 1968. T h e defensive behavior of Pterinoxylus spinulosus Redtenbacher, a winged stick insect from Panama. Psyche 75: 195-207. SCHNEIDER, C. O. 1934. Las emanaciones del

chinchemoyo Paradoxomorpha crassa (Blanch, Kirby). Rev. Chilena Hist. Nat. 38: 44-46. Wi LLIG, M.

R.,

R.

W. GARRISON, AND A. J.

BAUMAN. 1986. Population dynamics and natural history of a Neotropical walkingstick, Lamponium portoricensis Rehn (Phasmatodea: Phasmatidae). Texas J. Sci. 38: 121-137. WIRTH, W. W. 1971. A review of the "stick-ticks," Neotropical biting midges of the Forcipomyia subgenus Microhelea parasitic on walkingsticks (Díptera: Ceratopogonidae). Entomol. News 82: 229-245. ZAPATA, S., AND E. TORRES. 1970.

Biología y

morfología de Bacteria granulicollis (Blanchard) (Phasmida). Univ. Chile, Cen. Esl. Entomol. Publ. 10: 23-42.

COCKROACHES Blattodea ( = Blattaria). Spanish: Cucarachas. Portuguese: B a r a t a s . Quechua: Utiuti. Cockroaches a r e r e g a r d e d with disgust by nearly e v e r y o n e . A l t h o u g h widely b e ­ lieved to carry disease, their i m p o r t a n c e as mechanical vectors is probably o v e r r a t e d

COCKROACHES

169

(Roth and Willis 1957). Some evidence to the contrary does exist (Gazivoda and Fish 1985). It is a shame that a few unsavory types have given a bad name to an otherwise diverse and biologically fascinat­ ing group of insects, one begging investi­ gation in Latin America where a large and distinctive fauna exists which is practically unknown. Cockroaches are among the most primi­ tive and ancient of winged insects. Their origin dates back 250 million years to the Carboniferous period. Fossil records show that they were very abundant at that time and very little different in structure from their modern descendants. Nocturnal cockroaches are mostly oval, much flattened and dull-colored, with pli­ able wings, all adaptations for life in dark, confined spaces. Others are teardrop shaped, convex, with brightly hued, hard, beetlelike wings; these features relating them to an exposed, diurnal existence on vegetation, tree bark, and tree trunks. All cockroaches have a well-developed, thin layer of grease or wax on the outside of the cuticle which protects them from desiccation and gives them their slick feel. The head is horizontal but bent backward so that the mouthparts project toward the rear, being situated almost between the bases of the front legs. T h e head is often entirely concealed from above by a large, widely expanded, and flattened disklike shield (the pronotum), out from under which project the very long, whiplike anten­ nae. Wings are usually present and large with a highly complex, reticulate venation. They are often abbreviated in the female and in many species may be absent alto­ gether from both sexes. The fore wing is narrow and elongate and more rigid in texture than the hind wing, which is broad, fanlike, and membranous. The legs are well spined. A pair of elongate, segmented cerci project from the apex of the abdo­ men. These appendages are richly en­ dowed with external sense organs adapted

170

for perception of vibrations, sound, and air movements. Female cockroaches give birth in differ­ ent ways. They normally encapsulate their eggs in groups of few to many within a hard, darkly pigmented case (ootheca), which is deposited in the proper environ­ ment. Some retain the case in the birth canal where it often remains partly ex­ truded until the young hatch. Others keep it entirely within the abdomen, and the nymphs hatch inside the mother, spending up to several days in a kind of uterus before being born alive. Cockroaches actively secrete a variety of exocrine substances, such as sex lures, aggregation stimulants, and defensive com­ pounds of all kinds (Roth and Alsop 1978). The glands that produce these chemicals are located either in the head (mandibular glands) or on the abdomen, usually on the back of the more posterior segments. A great deal is known about the anat­ omy, physiology, and natural history of the few common domestic and semidomestic forms (Cornwell 1968, Guthrie and Tindall 1968), but information is scarce con­ cerning these aspects in the majority of wild species. One study (Schal and Bell 1986) indicates a vertical segregation of species in the layered vegetation of forests. Many genera are represented in bromeliad "terraria" (Rocha and Lopes 1976); Epilampra (fig. 5.8a) has been observed to swim well (Crowell 1946), even having tubular spiracles at the rear of the abdo­ men, possibly functioning as elementary snorkels. Several species mimic other in­ sects. For example, lycid beetles of the genus Calopteron are mimicked by the cock­ roaches Holocampsa and Paratropes, lampyrid beetles Cratomorphus and Aspisoma by Achroblatta luteola (figs. 5.8b, 5.8c), and the giant fungus beetle Erotylus by Plectoptera. Most of these are diurnal and commonly seen running on forest understory vegeta­ tion; the firefly mimics are nocturnally active like their models, the similarity be-

ORTHOPTEROIDS AND OTHER ORDERS

Figure 5.8 COCKROACHES, (a) Aquatic cockroach (Epilampra sp., Epilampridae). (b) Fireflymimicking cockroach (Achroblatta luteola, Allicolidae). (c) Firefly model of firefly-mimicking cockroach [Aspisoma sp., Lampyridae). (d) Myrmecophilous cockroach (Myrmecoblatta sp., Atticolidae). (e) Leaf cockroach (Pseudomops sp., Blatellidae).

ine effective, however, during the daytime when the insects are at rest and visible to predators. The lycid mimic Paratropes may be responsible for pollination of plants in the canopy of Central American rain for­ ests (Perry 1978). The fat body of cockroaches is packed with intracellular bacteria (bacteroids) that contribute to the synthesis of amino acids used in metabolism. The alimentary canal of some wood-feeding types also contain symbiotic protozoa, which, like those of certain termites, assist in the digestion of cellulose (Roth and Willis 1960). Ecological niches occupied by tropical American cockroaches are extremely var­ ied. Many of these insects are associated casually with vegetation, usually seen sit­ ting on the upper surfaces of leaves (Plectoptera), or are disposed to feed on the fruit, leaves, bark, roots, and other parts of living plants of particular species; others consume the wood of rotting logs. Pseudomops (fig. 5.8e) are common, small, brightly colored forms (head orange and wings and pronotum shining dark purple, the latter margined with yellow) that perch on the leaves of low plants in forests and clear­ ings, hopping from place to place in the daytime. Blaberus depends on the excre­ ment or dead bodies of other creatures and lives in animal burrows, hollow trees, or caves. A number of species live symbiotically with ants (Bolivar 1905), such as the minute Attiphila, found in the fungus gar­

dens of leaf cutter ants (Atta), and Myrmeco­ blatta (fig. 5.8d), a genus associated with carpenter ants (Deyrup and Fisk 1984). Cockroaches have been found in all climes, from hot deserts and cold mountaintops to warm, humid lowland rain forests. Everywhere they are important reducers of leaf litter and wood. In the inundation forests of Amazonia, Epilampra feeds on dead leaves and insect carcasses. In any one area, as much as 5.6 percent of the yearly leaf fall and other organic detritus may be turned over by members of this genus (Irmler and Furch 1979). There is some tendency for adult cock­ roaches in lowland forests to disperse their activity over a long time, thereby possibly reducing interspecific competi­ tion in nonseasonal locations; their occur­ rence does not follow this pattern where well-defined wet and dry seasons are present (Wolda and Fisk 1981). A number of cockroaches are capable of sound production. They accomplish this by rubbing the abdomen against the wings (Blaberus), by stridulation of roughened areas on the thorax (Panchlora), or by other devices (Roth and Hartman 1967). Of the 3,500 known species in the world, probably over 30 percent are found in Latin America (Princis 1962-1971). No general guide to their classification is avail­ able, although there are some good local treatments with broad applicability (Fisk and Wolda 1979, Bruijning 1959).

COCKROACHES

171

References BOLIVAR, I. 1905. Les blattes myrmécophiles. Schweizerischen Entomol. Ges. Mitt. 11: 134-141. BRUIJNING, C. F. A. 1959. T h e Blattidae of

Surinam. Stud. Fauna Suriname Guianas 2(4): 1-103. CORNWELL, P. B. 1968. T h e cockroach. Vol. 1.

Hutchinson, London. CROWELL, H. H. 1946. Notes on an amphibious cockroach from the Republic of Panama. Entomol. News 57: 171-172. DEYRUP, M., AND F. FISK. 1984. A myrmeco-

philous cockroach new to the United States (Blattaria: Polyphagidae). Entomol. News 95: 183-185. FISK, F. W., AND H. WOLDA. 1979. Keys to the

cockroaches of central Panama. Stud. Neotrop. Fauna Environ. 14: 177-201. GAZIVODA, P., AND D. FISH.

1985. Scanning

electron microscope demonstration of bacte­ ria on tarsi of Blatella germánica. New York Entomol. Soc. J. 93: 1064-1067. GUTHRIE, D. M., AND A. R. TINDALL. 1968. T h e

biology of the cockroach. Arnold, London. IRMLER, U , AND K. FURCH. 1979. Production,

energy, and nutrient turnover of the cock­ roach Epilampra irmleri Roch e Silva and Aguilar in a Central-Amazonia inundation forest. Amazoniana 6: 497-520. PERRY, D. R. 1978. Paratropes bilunata (Orthoptera; Blattidae): An outcrossing pollinator in a Neotropical wet forest canopy. Entomol. Soc. Wash. Proc. 80: 656-657. PRINCIS, K. 1962-1971. Blatariae. In M. Beier, ed., Orthopterorum Catalogus. Junk, T h e Hague. Pts. 3, 4, 6, 7, 8, 11, 13, 14. ROCHA E SILVA ALBUQUERQUE, I., AND S. M. RODRIGUES LOPES. 1976. Blattaria de bro-

mélia (Dictyoptera). Rev. Brasil. Biol. 36: 873-901. ROTH, L. M., AND D. W. ALSOP. 1978. Toxins of

Blattaria. In S. Bettini, ed., Arthropod ven­ oms. Springer, Berlin. Pp. 465-487. ROTH, L. M., AND H. B. HARTMAN. 1967. Sound

production and its evolutionary significance in the Blattaria. Entomol. Soc. Amer. Ann. 60: 740-752. ROTH, L. M., AND E. R. WILLIS.

1957. T h e

medical and veterinary importance of cock­ roaches. Smithsonian Misc. Coll. 134(10): 1-147. ROTH, L. M., AND E. R. WILLIS.

1960. T h e

biotic associations of cockroaches. Smithso­ nian Misc. Coll. 141: 1-470. SCHAL,

C,

AND W. J.

BELL.

1986.

Vertical

community structure and resource utilization

172

in Neotropical forest Entomol. 11: 411-423.

cockroaches.

Ecol.

WOLDA, H., AND F. W. FISK. 1981. Seasonality of

tropical insects. II. Blattaria in Panama, f Anim. Ecol. 50: 827-838. Domestic Cockroaches Less t h a n thirty-five species of cockroaches live in intimate closeness with h u m a n s . T h e y frequent h o m e s , r e s t a u r a n t s , hotels, a n d all o u r dwellings, w h e r e they scavenge food leavings a n d whatever edible organic m a t t e r they c a n find (Cornwell 1968). All are O l d World in origin, probably carried to Latin America in t h e last four centuries a b o a r d ships. S o m e less a d a p t a b l e species have spotty distributions, reflecting to some d e g r e e their points of arrival, while o t h e r s have s p r e a d virtually everywhere. Probably t h e most w i d e s p r e a d a n d fre­ quently seen is t h e A m e r i c a n cockroach, or encarachan (Periplaneta americana; fig. 5.9a), which infests warm dwellings t h r o u g h o u t the world (Bell a n d Adiyodi 1981). It prefers a heated, moist e n v i r o n m e n t a n d is c o m m o n o u t d o o r s in tropical p o r t i o n s of A m e r i c a as well as i n d o o r s . It is often found in sewers. A medium-sized cock­ roach (BWL 2 8 - 4 4 m m ) , its body is shin­ ing r e d d i s h - b r o w n , with a paler yellow area a r o u n d t h e e d g e of t h e h e a d shield. T h e fully d e v e l o p e d wings e x t e n d well beyond t h e a b d o m e n in t h e male b u t only j u s t overlap t h e a b d o m e n in t h e female. Females p r o d u c e u p to ninety free oothecae with about twelve eggs each. A related a n d similarly ubiquitous spe­ cies is t h e Australian cockroach (Periplaneta australasiae; fig. 5.9b), which is a general pest in cooler areas w h e r e v e r moist artifi­ cial e n v i r o n m e n t s prevail. A b o u t t h e same size (BWL 3 0 - 3 5 m m ) a n d facies as P. americana, it differs primarily in having well-defined, bright yellow "shoulders" (elongate m a r k s at t h e o u t e r bases of the fore wings). Its biology is also similar to that of t h e A m e r i c a n cockroach, but it seems n o t t o b e a d e n i z e n o f sewers. T h e

ORTHOPTEROIDS AND OTHER ORDERS

Figure 5.9 COCKROACHES, (a) American cockroach (Periplaneta americana, Blattidae). (b) Aus­ tralian cockroach (Periplaneta australasiae, Blattidae). (c) Brown-banded cockroach (Supella longipalpa, Blatellidae). (d) Harlequin cockroach (Neostylopyga rhombifolia, Blattidae). (e) Madeira cock­ roach' (Leucophaea maderae, Oxyhaloidae). (f) Surinam cockroach (Pycnoscelus surinamensis, Pycnoscelididae). original h o m e of b o t h of these Periplaneta species was probably tropical Africa. T h e b r o w n - b a n d e d cockroach (Supella longipalpa, s o m e t i m e s called S. supellectilium; fig- 5.9c) is small ( B W L 1 0 - 1 4 m m ) , with c o m p l e t e wings in t h e male a n d short­ ened wings in t h e female. It is buff in general color b u t with two suffuse, trans­ verse black o r b r o w n b a n d s . It is highly domiciliary, t a k i n g u p residence in furni­ ture a n d f o r m i n g colonies in d r a w e r s , behind pictures o n t h e wall, o n book­ shelves, a n d in like places. H e r e t h e adults and n y m p h s h i d e by day, e m e r g i n g at night to feed o n a n y available fodder, including g l u e , sizing o n books, wallpaper, food scraps, a n d even plain paper. T h e h a r l e q u i n cockroach (Neostylopyga rhombifolia; fig. 5.9d) is medium-sized (BL 2 0 - 2 5 m m ) a n d completely without wings in both sexes, a l t h o u g h t h e fore wings a r e represented by small stubs below t h e o u t e r corners of t h e h e a d shield. It h a s a striking color p a t t e r n of d e e p yellow m a r b l i n g o n a shining, brownish-black b a c k g r o u n d . Its biology is largely u n s t u d i e d , a l t h o u g h it is common b o t h i n d o o r s a n d o u t . T h e M a d e i r a cockroach (Leucophaea maderae; fig. 5.9e), k n o w n as barata cascuda in Brazil, h a s b e c o m e widely established around t h e C a r r i b e a n , w h e r e it often is a serious pest in h o m e s a n d w a r e h o u s e s , a n d in many parts of S o u t h a n d C e n t r a l A m e r ­ ica. I n tropical e n v i r o n m e n t s , it lives out­

d o o r s a n d is c o m m o n in s u g a r c a n e a n d b a n a n a plantations. It is a fairly large cockroach ( B W L 4 - 5 c m ) , with a m p l e wings in b o t h sexes. It is pale b r o w n to tawny olive in g e n e r a l color, t h e fore wings m a r k e d with a d a r k spatter p a t t e r n over most of their surface; t h e basal a n d a n t e ­ rior marginal areas a r e contrastingly plain, except for a d a r k linear a r c r u n n i n g obliquely across t h e wing base. Adults a r e slow moving b u t capable of d e f e n d i n g themselves well with a n offensive odor. T h e y also stridulate. Females b e a r twentyfive to thirty live y o u n g at a time. A l t h o u g h first found in South A m e r i c a , the S u r i n a m cockroach (Pycnoscelus surinam­ ensis; fig. 5.9f), k n o w n as barata de pau podre in Brazil, is most likely of O r i e n t a l origin. Because it is p a r t h e n o g e n e t i c , even u n ­ related females c a n start thriving p o p u l a ­ tions wherever they h a p p e n t o b e c a r r i e d , a n d t h e species h a s b e c o m e established widely in Latin America. Away from civili­ zation, it occurs u n d e r stones a n d loose litter a n d can b u r r o w superficially into t h e soil. It is medium-sized ( B W L 1 8 - 2 4 m m ) , shining b r o w n , with d a r k s h o u l d e r streaks a n d a contrasting black h e a d shield. Wings a r e complete in b o t h sexes. It often h a s a pale b a n d along t h e a n t e r i o r m a r g i n of t h e h e a d shield, t h e posterior m a r g i n of which is strongly sinuate. A n o t h e r domiciliary cockroach from Af­ rica is t h e lobster cockroach (also called t h e

COCKROACHES

173

Figure 5.10 COCKROACHES, (a) Cinereous cockroach (Nauphaeta cinérea, Oxyhaloidae). (b) Oriental cockroach (Blatta orientalis, Blattidae). (c) Oriental cockroach egg case, (d) German cockroach egg case, (e) German Cockroach (Blatella germánica, Blatellidae). (f) Death's-head cock­ roach (Blaberus giganteus, Blaberidae). (g) Cuban cockroach (Panchlora nivea, Panchloridae). c i n e r e o u s cockroach, Nauphaeta cinérea; fig. 5.10a), which derives its n a m e f r o m a lobster-shaped design o n t h e h e a d shield. It is large ( B W L 2 5 - 2 9 m m ) , with welld e v e l o p e d , ash-colored wings that a r e j u s t short of c o v e r i n g t h e a b d o m e n in both sexes. Males stridulate w h e n c o u r t i n g fe­ males. T h e females f o r m o o t h e c a e with twenty-six to forty eggs that h a t c h as she e x t r u d e s t h e capsule f r o m h e r b r o o d sac. T h e original h o m e of t h e O r i e n t a l cock­ roach (Blatta orientalis; fig. 5.10b) is N o r t h Africa. Its b o d y is s h i n i n g black, except for t h e r e d d i s h - b r o w n s h o r t wings of t h e male; these o r g a n s a r e m e r e stubs in t h e female. It is a m e d i u m - s i z e d species ( B W L 20—24 m m ) t h a t p r o d u c e s u p to e i g h t e e n large o o t h e c a e (5 by 10 m m ) (fig. 5.10c). T h e latter c o n t a i n r a t h e r oversized eggs that d e v e l o p free of t h e p a r e n t . T h e species is of local o c c u r r e n c e in dwellings b u t is often f o u n d o u t d o o r s w h e r e t h e climate is m o d e r a t e . A n o t h e r very w i d e s p r e a d v a g r a n t cock­ r o a c h is t h e G e r m a n c o c k r o a c h {Blattella germánica; fig. 5.10e). It is small ( B W L 10— 15 m m ) a n d pale buff in color, with distinct, parallel, wide b a n d s o n t h e h e a d shield. Wings a r e nearly completely f o r m e d , only the tip of t h e a b d o m e n b e i n g left e x p o s e d in b o t h sexes. T h e species p r e f e r s a w a r m , moist a m b i e n c e a n d is t h e r e f o r e c o m m o n in kitchens, l a r d e r s , a n d r e s t a u r a n t s , rarely in b e d r o o m s . I t h a s t h e capacity to build u p

174

e n o r m o u s p o p u l a t i o n s w h e n food is ade­ q u a t e . T h e o o t h e c a e a r e d r o p p e d free by the female a n d contain o n t h e average thirty-seven eggs (fig. 5.10d). T h e species is considered an i n d o o r pest, b u t in warm e n v i r o n m e n t s , heavy infestations m a y over­ flow o u t d o o r s .

References BELL, W. J., AND K. G. ADIYODI, eds. 1981. The

American cockroach. Chapman Hall, London. CORNWELL, P. B. 1968. T h e cockroach. Vol. 1.

Hutchinson, London.

Giant Cockroaches Blaberidae, Blaberinae, Blaberini, Blaberus. Spanish: Cascudas (General); cucarachas m a n d i n g a s , cucarachas m a m a , cargatables (Peru). Cockroaches of t h e Divine Face, d e a t h ' s - h e a d cockroaches. A p a r t from being very big (the largest species, B. colosseus, reaches a B W L to 8 cm), t h e giant cockroaches a r e recognized by their fully d e v e l o p e d , shiny, light brown fore wings, these usually m a r k e d with a large squarish spot in t h e m i d d l e a n d a diffuse spot sometimes o n t h e o u t e r third. T h e r e can be also an elongate thin dark bar in t h e s h o u l d e r area of t h e wing, paralleling t h e front m a r g i n . I n addition, they have a sharply defined, square to trapezoidal black patch resting against the h i n d m a r g i n in t h e c e n t e r of t h e oval head

ORTHOPTEROIDS AND OTHER ORDERS

shield in which some see t h e i m a g e of a h u m a n face, o r d e a t h ' s - h e a d . This g e n u s contains twelve very similar but separate species, readily distinguished only by t h e s t r u c t u r e of t h e genitalia (Roth 1969)- All n o r m a l l y inhabit caves, rock crevices, hollow trees, a n d cavities u n d e r loose tree b a r k . Blaberus parabolicus (tranga bakkas) lives u n d e r houses in S u r i n a m a n d becomes a pest i n d o o r s w h e n attracted to light at night (Bruijning 1959). A d u l t s a n d the yellow-spotted, trilobitelike n y m p h s are often f o u n d t o g e t h e r a n d exhibit subsocial b e h a v i o r (Gautier 1978, Schal 1983). Males a s s u m e a d o m i n a n t r a n k in dense g r o u p s , their status distinguishable by their erect p o s t u r e a n d aggressiveness. An a g g r e g a t i o n p h e r o m o n e , c o n t a i n i n g undecane a n d t e t r a d e c a n e a n d o t h e r com­ pounds of u n k n o w n function (Brossut et al. 1973), is secreted from m a n d i b u l a r glands in all stages a n d by b o t h sexes. Females also p r o d u c e a volatile sex p h e r o ­ mone that acts as a p r i m a r y releaser of complex a n d l e n g t h y m a l e c o u r t s h i p activi­ ties. T h e r e p e r t o i r e includes intense in­ terínale fighting over calling females (Wendelken 1977). N y m p h s a r e a d e p t at b u r r o w i n g in t h e soil or r o t t i n g wood to escape p r e d a t i o n . Adults lead m o r e e x p o s e d lives a n d a r e likelier to avoid h a r m by flying, releasing offensive o d o r s , o r kicking with their s h a r p spined legs (Crawford a n d CloudsleyT h o m p s o n 1971). T h e y a r e often attracted to artificial light d u r i n g t h e i r n o c t u r n a l wanderings. T h e food of giant cockroaches consists of organic d e t r i t u s that accumulates in their d a n k niches, including bat g u a n o , rotting wood a n d fruits, seeds a n d o t h e r decomposing vegetation, a n d d e a d insects or animals at times. T h e two c o m m o n species, Blaberus craniifer a n d B. giganteus (fig. 5.10f) (Schal 1983), a r e easy to m a i n t a i n in c u l t u r e a n d are used in m a n y laboratories for physio­

logical research (Lefeuvre 1960). Adults can live for as long as twenty m o n t h s (Piquett a n d Fales 1953). Females inter­ nally incubate their oothecae; t h e latter contain a b o u t thirty to forty eggs. O t h e r very large Neotropical cock­ roaches a r e t h e four species of S o u t h A m e r i c a n Megaloblatta (Blatellidae, Nyctiborinae). T h e y often a r e over 8 c e n t i m e ­ ters in length. A C o l o m b i a n specimen, whose overall length (head to wing tips) was m e a s u r e d at 10 c e n t i m e t e r s ( G u r n e y 1959), is t h e largest cockroach r e c o r d e d .

References BROSSUT, R., P. DUBOIS, AND J. RIGAUD.

1973.

Le grégarisme chez Blaberus craniifer: lsolemenl et identification de la phéromone. J. Ins. Physiol. 20: 529-543. BRUIJNING, C. F. A. 1959. The Blattidae of

Surinam. Stud. Fauna Suriname 2(4): 1-103. CRAWFORD,

C.

S.,

AND J.

L.

Guianas

CLOUDSLEY-

THOMPSON. 1971. Concealment behavior of nymphs oí Blaberus giganteus L. (Dictyoptera: Blattaria) in relation to their ecology. Rev. Biol. Trop. 18: 5 3 - 6 1 . GAUTIER, J. Y. 1978. Le comportement social de Blaberus colosseus en milieu naturel; plasticité du systéme social. Ins. Soc. 25: 289-301. GURNEY, A. B. 1959. T h e largest cockroach (Orthoptera, Blattoidea). Entomol. Soc. Wash. Proc. 61: 133-134. LEFEUVRE, J. C. 1960. A propos de Blabera craniifer Burmeister 1838 (Insecte dictyoptére). Soc. Scien. Bretagne Bull. 35: 145-161. PIQUETT, P. G., AND J.

H. FALES. 1953.

Life

history of Blaberus giganteus. J. Econ. Ento­ mol. 46: 1089-1090. ROTH, L. M. 1969. T h e male genitalia of Blattaria. I. Blaberus spp. (Blaberidae: Blaberi­ nae). Psyche 76: 217-250. SCHAL, C. 1983. Blaberus giganteus (cucaracha, giant cockroach, giant drummer, cockroach of the Divine Face) and Xestoblatta hamata (cucaracha). In D. H. Janzen, ed., Costa Rican natural history. Univ. Chicago Press, Chicago. Pp. 693-696. WENDELKEN, P. W. 1977. T h e evolution of

courtship phenomena in Blaberus and related genera with reference to sexual selection. Diss. Abstr. B37(8): 3816.

COCKROACHES

175

MANTIDS

Green Cockroaches P a n c h l o r i d a e , Panchlorini, Panchlora. T h e best k n o w n of the forty or so species in this g e n u s ( G u r n e y a n d Roth 1972) a r e pale, t r a n s l u c e n t g r e e n , o r s o m e s h a d e of gray or c r e a m , the fore wings also with some d a r k mottling. T h e y a r e m e d i u m sized (BWL 25—30 m m ) a n d elongateellipsoidal in outline s h a p e . Both sexes have c o m p l e t e wings a n d a s m o o t h , u n ­ m a r k e d h e a d shield. Occasional pink indi­ viduals t u r n u p , the color a n a p p a r e n t s y m p t o m of viral infection (Roth a n d Willis 1960). Curiously, the virus, Serratia marcescens, is also a h u m a n p a t h o g e n causing septicemia with a h i g h mortality (Appel pers. c o m m . ) . T h e s e c o n s p i c u o u s cockroaches a r e com­ m o n t h r o u g h o u t Latin A m e r i c a . S o m e spe­ cies, such as Panchlora nivea (formerly cubensis) (fig. 5.10g), a r e t r a n s p o r t e d widely by c o m m e r c e , especially in s h i p m e n t s of tropi­ cal fruits. In n a t u r e , they live in the r o t t i n g t r u n k s of palms w h e r e they feed o n the d e c o m p o s i n g b r o w n fibers of the t r u n k , t h r o u g h which they m a k e t u n n e l s . Females m a k e a thin-walled, pale ootheca that may be e x t r u d e d d u r i n g forma­ tion a n d t h e n r e t r a c t e d into a b r o a d sac w h e r e the eggs d e v e l o p . W h e n the e m ­ bryos m a t u r e , the o o t h e c a is r e e x t r u d e d . T h e n y m p h s free themselves from their d e v e l o p i n g m e m b r a n e s as t h e ootheca is forced o u t a n d d r o p to t h e s u b s t r a t u m (Roth a n d Willis 1958). References GURNEY, A.

B.,

AND L.

M.

ROTH.

1972.

A

generic review of the cockroaches of the subfamily Panchlorinae (Dictyoptera, Blattaria, Blaberidae). Entomol. Soc. Amer. Ann. 65: 521-532. ROTH, L.

M.,

AND E.

R.

WILLIS. 1958.

The

biology of Panchlora nivea, with observations on the eggs of other Blattaria. Amer. Entomol. Soc. Trans. 83: 195-208. ROTH, L.

M.,

AND E.

R.

WILLIS.

1960.

The

biotic associations of cockroaches. Smithso­ nian Misc. Coll. 141: 1-470.

176

M a n t o d e a (= Mantoidea, M a n t e o d e a ) . Spanish: Mantidos, a d i v i n a d o r e s (General); tata dios, m a m b o r e t á s , comepiojos (Argentina); Santa Teresas (Peru). Portuguese: Louvas a D e u s (Brazil). Preying m a n t i d s , p r a y i n g m a n t i d s . Mantises. A l t h o u g h m a n y o t h e r o r t h o p t e r o i d s take insects as food, they are basically vegetar­ ian. T h e m a n t i d s , however, a r e all rapa­ cious carnivores. T h e principal feature identifying t h e m is their m e a n s of catching p r e y — t h e highly specialized, raptorial forelegs. T h e f e m u r is large, c a r r y i n g pow­ erful a n d quick-acting muscles for closing the spined, g r a s p i n g tibia against it, to form a vise from which escapes a r e few. Mantids wait in a m b u s h for passing insect prey, often with their f o r e p a r t s erect and foreleg segments closed a n d held in a p r a y i n g position, for which they a r e called "praying mantids." T h i s a n d o t h e r h u m a n o i d behavior, such as the way they follow m o t i o n with their mobile, t r i a n g u l a r h e a d s a n d the large, " p u p i l e d " eyes, has g e n e r a t e d much folklore. Mantid m e a n s "soothsayer" or "diviner," a n d they are believed by some p e o p l e to possess occult p o w e r s or to be worthy of being r e g a r d e d as sacred. In Amazonia, the sex of an u n b o r n child can be l e a r n e d from a m a n t i d placed nearby the e x p e c t a n t m o t h e r (pde mesa). If it just moves its forelegs, the infant will be fe­ male; if the insect j u m p s o n t o s o m e o n e , a male can be e x p e c t e d ( L e n k o a n d Papavero 1979: 1 1 - 1 2 ) Mantids a r e mostly fairly large (BWL 3 15 cm), with an elongate p r o t h o r a x and otherwise slender body a n d walking legs. Adults usually b e a r full wings, although the females' wings a r e often abbreviated. T h e y a r e cryptically colored in leafy greens a n d b r o w n s . T h e d e a d leaf m a n t i d (Acan­ thops falcataria; fig. 5.11a) is a c o m m o n

ORTHOPTEROIDS AND OTHERORDERS

Figure 5.11 MANTIDS (MANTIOAE). (a) Dead leaf mantid (Acanthops falcataria). (b) Bark mantid (Liturgusa sp.). (c) Leaf mantid (Choeradodis rhombicollis). (d) Common mantid (Stagmotoptera sp.). (e) Horned mantid (Vafes sp.). example of t h e latter. It has b r o a d , b r o w n wings with t h e exact crinkly t e x t u r e a n d twisted s h a p e of a d e a d leaf. O t h e r s are marked with lichenose p a t t e r n s a n d fre­ quent tree t r u n k s in moist forests, well camouflaged against the e n c r u s t e d bark (e.g., Liturgusa; fig. 5.1 lb). T h e leaf m a n t i d (Choeradodis rhombicollis; fig. 5.1 Ic) is u n i q u e in form (BWL 6 5 - 7 0 rum), the p r o t h o r a c i c d o r s u m being greatly e x p a n d e d laterally to form a flat, r h o m b o i d plate covering the h e a d completely. T h e wings are wide a n d the whole d e p r e s s e d . With its g r e e n color a n d strongly veined wings, it looks incredibly like a living leaf. Female m a n t i d s flood their eggs at the time they a r e laid with a whitish, frothy secretion from glands off the oviduct. They attach t h e mass to t r e e t r u n k s , limbs, rocks, a n d o t h e r rigid substrata, often in very e x p o s e d situations. T h e secretion dries to form a h a r d , protective encase­ ment for the eggs. Each species' case has its own characteristic s h a p e . Methods of d e f e n s e e m p l o y e d by m a n ­ tids have b e e n well studied ( C r a n e 1952, Robinson 1969). T h e r e a r e four general strategies: (1) r e s e m b l a n c e to i n a n i m a t e objects, including cryptic structural a n d color mimicry of leaves, sticks, bark, a n d so on, combined with stillness or swaying; (2) active flight, i n c l u d i n g d o d g i n g , j u m p i n g and d r o p p i n g , t h r e a t a n d flying; (3) star­ tling displays, consisting primarily of fac•ng the e n e m y with wings raised a n d fore­ legs splayed a p a r t ; a n d (4) active attack by

striking with t h e forelegs. S o m e m a n t i d s have an imperfect eyespot in t h e m i d d l e of the h i n d wing. W h e n the insect is m o ­ lested, the wings are elevated a n d these spots threateningly displayed. Many also e x p o s e a d a r k m a r k i n g o n t h e i n n e r sur­ face of the fore f e m u r w h e n the legs a r e s p r e a d in a t h r e a t p o s t u r e . Few adult m a n t i d s are involved in t r u e mimicry com­ plexes. In Belize, Mantoida maya n y m p h s , in their earliest stages, are very small a n d have the s h a p e a n d attitude of Camponotus ants (Jackson a n d D r u m m o n d 1974). T h e Neotropical Region is rich in its variety of m a n t i d s . T h e latest reviews (Beier 1 9 3 3 - 1 9 3 5 , Giglio-Tos 1927) r e c o r d approximately 300 species in some 74 g e n e r a . Many species certainly await discov­ ery, a n d the g r o u p n e e d s a c o m p r e h e n s i v e m o n o g r a p h . Some local reviews (e.g., Beebe et al. 1952) are helpful for the m o r e c o m m o n types. D o m i n a n t g e n e r a are Stagmomantis a n d Stagmotoptera (fig. 5.1 Id), which a r e large a n d usually g r e e n ; Litur­ gusa, which are very c o m m o n , small (BWL 2 - 3 cm), flattened forms that r u n actively o n tree t r u n k s ; slender, sticklike Angela a n d Vates (fig. 5.1 le), which have a s h a r p spine o n the forehead a n d leaflike flanges o n the mid- a n d h i n d leg s e g m e n t s .

References BEEBE, W., J. CRANE, AND S. HUGHES-SCHRADER.

1952. An annotated list of the mantids (Orthoptera, Mantoidea) of Trinidad, B.W.I. Zoológica 37: 245-258, PI. I - V I I I .

MANTIDS

177

BEIER, M. 1933-1935. Mantodea. General lnsectorum 196: 1-37: 197: 1-10; 198: 1-9; 200: 1 - 3 2 ; 2 0 1 : 1-10; 203: 1-146. CRANE, J. 1952. A comparative study of innate defensive behavior in Trinidad mantids (Orthoptera, Mantoidea). Zoológica 37: 259-293. GIGLIO-TOS, E. 1927. Orthoptera, Mantidae. Das Tierreich 50: 1-707. JACKSON, J. F., AND B. A. DRUMMOND III. 1974.

A Batesian ant-mimicry complex from the Mountain Pine Ridge of British Honduras, with an example of transformational mim­ icry. Amer. Midi. Nat. 9 1 : 248-251. LENKO, K., AND N. PAPAVERO. 1979. Insetos no

folclore. Conselho Estad. Artes Cien. Hum., Sao Paulo. ROBINSON, M. H. 1969. T h e defensive behav­ iour of some orthopteroid insects from Pan­ ama. Royal Entomol. Soc. London Trans. 121:281-303.

EARWIGS D e r m a p t e r a . Spanish: Tijeretas. Portuguese: T e s o u r a s . Earwigs a r e most easily recognized by t h e pincers o r forcepslike cerci b o r n e o n t h e e n d of t h e a b d o m e n . T h e i r tips a r e strongly i n c u r v e d , a n d they often have i n t e r n a l teeth. T h e cerci, normally larger in males t h a n females, a r e s h o r t to almost as long as t h e body. T h e y a r e used to c a p t u r e a n d m a n i p u l a t e p r e y as well as for defense. Dermaptera

a r e otherwise

monoton­

ously similar, somber-colored insects, most small t o medium-sized (BL 1—2.5 cm), with chewing m o u t h p a r t s , short t h r e a d l i k e or beadlike a n t e n n a e , a n d unspecialized, simi­ lar legs. T h e fore wings a r e thickened and leathery a n d very short, m e e t i n g in a straight line d o w n the back, almost like the elytra of some beetles. T h e h i n d , o r flight wings are m e m b r a n o u s a n d r o u g h l y circu­ lar a n d fold fanlike along m a n y radially a r r a n g e d creases. A p p r e c i a b l e anatomical variation in their body s t r u c t u r e occurs primarily in t h e length a n d s h a p e of the cerci, for e x a m p l e , in Metresura ruficeps, they are considerably l o n g e r t h a n the body (fig. 5.12c). S o m e species have reddish (Carcinophora americana; fig. 5.12a) o r yel­ low (Doru lineare; fig. 5.12b) areas o n the fore wings. Scientifically, this is o n e of t h e most neglected o r d e r s of insects, even though they are c o m m o n p l a c e . Earwigs are, on the whole, n o t p o p u l a r owing to their appear­ ance a n d their u n w e l c o m e presence in g a r d e n s a n d h o m e s . While they d o cause some economic d a m a g e to stored food, roots, a n d shoots of t e n d e r y o u n g vegeta­ bles, some a r e p r e d a c e o u s a n d help to control populations of o t h e r m o r e serious pests. Urban earwigs are all adventives, thought to have m a d e their way from E u r o p e to Latin America with t h e earliest colonists,

Figure 5.12 EARWIGS, (a) Giant earwig {Carcinophora americana, Anisolabiidae). (b) Lined ear­ wig {Doru lineare, Forficulidae). (c) Earwig {Metresura ruficeps, Anisolabiidae). (d) Shore earwig {Labidura riparia, Labiduridae). (e) Flat earwig {Sparatta pelvimetra, Sparattidae).

178

ORTHOPTEROIDS AND OTHER ORDERS

possibly in the ballast of ships o r o n n u r s e r y stock. O n e of t h e most w i d e s p r e a d is t h e ring-legged earwig {Euborellia annulipes), so called because of t h e faint d a r k b a n d a r o u n d t h e f e m o r a a n d tarsi of each leg. Also characteristic a r e t h e d a r k a n t e n n a e with t h e t h i r d to fifth subapical segments contrastingly pale. It is medium-sized (BL 1.5 cm) a n d wingless. T h e forceps of both sexes are very short a n d stout. T h e s h o r e (or striped) earwig {Labidura riparia; fig. 5.12d) is a n o t h e r nearly ubiqui­ tous species. It is m u c h larger t h a n t h e other adventive species (BL 2.5 cm) a n d much lighter colored, generally t a n a n d with d a r k , longitudinal stripes on t h e wings a n d d o r s u m of the a b d o m e n , these markings c o n t r a s t i n g sharply with t h e lighter adjacent areas. T h e forceps of males a r e almost as long as t h e a b d o m e n , slender, a n d smoothly incurved. T h e spe­ cies prefers d a m p habitats n e a r water, including t h e sea. It is a g e n e r a l p r e d a t o r and scavenger a n d t h u s is to b e c o n s i d e r e d a beneficial insect. It is also a good flier a n d often attracted to lights (Gross a n d Spink 1971). T h e m a r i t i m e earwig {Anisolabis marí­ tima) also is a cosmopolitan species b u t is not so w i d e s p r e a d as t h e f o r e g o i n g as it occurs only n e a r t h e seashore. It is fairly large (BL 2 cm), wingless, a n d with a shiny black to d a r k b r o w n body a n d pale yellow­ ish legs. T h e forceps a r e short, stout, a n d strongly i n c u r v e d in the male. It is s t r a n g e that t h e E u r o p e a n earwig (Forfícula auricularia), so c o m m o n almost everwhere else, has n o t s u c c e e d e d in invad­ ing Latin A m e r i c a . It is k n o w n t h e r e from only a few, f a r n o t h e r n localities. In t h e Neotropics, a m u c h m o r e varied and less-known native f a u n a lives in all climes a n d situations (Brindle 1968). T h e 300 o r so r e m a i n i n g species, in fifty-eight genera, mostly belong to t h e family LabiMae (Reichardt 1 9 6 8 - 1 9 7 1 ; S t e i n m a n n 1973, 1975). T h e biggest g e n u s is Marava.

T h e Pygidicranidae is a family of primitive earwigs, containing several S o u t h A m e r i ­ can species, whose closest relatives a r e in s o u t h e r n Africa (Brindle 1984). T h e native earwigs dwell in all sorts of d a m p , secluded habitats, u n d e r stones, in rotten wood, in a b a n d o n e d t e r m i t e nests, in cracks in rocks, a n d t h e like. Sparatta (fig. 5.12e) is particularly flattened as a n adaptation for living u n d e r bark. Earwigs a r e generally m o r e conspicuous in d a m p forests, b u t t h e r e are d e s e r t a n d m o u n t a i n dwellers as well. In contrast to their dullcolored semidomestic c o u n t e r p a r t s , m a n y are m a r k e d with b r i g h t p a t t e r n s , often spotted red o r yellow. Typically, earwigs a r e active at night, w h e n they forage for food. A l t h o u g h most are o m n i v o r o u s , a n u m b e r a p p e a r to b e at least partly carnivorous, feeding primarily on o t h e r insects. T h e y use their forceps to seize a n d hold t h e victim a n d curve t h e a b d o m e n forward to access t h e mandibles.

References BRINDLE, A. 1968. The Dermaptera of Surinam and other Guyanas. Stud. Fauna Suriname Guyanas 36: 1—60. BRINDLE, A. 1984. The Esphalmeninae (Dermap­ tera: Pygidicranidae): A groupof Andean and southern African earwigs. Syst. Entomol. 9: 281-292. GROSS, H. R., JR., AND W. T. SPINK. 1971. Flight

habits of the striped earwig Labidura riparia. Entomol. Soc. Amer. Ann. 64: 746-748. REICHARDT, H. 1968-1971. Catalogue of New World Dermaptera (Insecta). Dept. Zool. Sec. Agrie. (Sao Paulo), Pap. Avul. Zool. 21: 1 8 3 193 (Pi. I. Introduction and Pygidicranoidea); 22: 35-46 (Pt. II. Labioidea, Carcinophoridae); 23: 83-109 (Pt. III. Labioidea, Labiidae); 24: 161-184 (Pt. IV: Forficuloidea); 24: 221-257 (Pt. V: Additions, corrections, bibli­ ography and index). STEINMANN, H. 1973. A zoogeographical check­ list of world Dermaptera. Fol. Entomol. Hungarica 26: 145-154. STEINMANN, H. 1975. Suprageneric classifica­ tion of Dermaptera. Acad. Sci. Hungaricae Acta Zool. 21: 195-220.

EARWIGS

179

TERMITES Isoptera. Spanish: Comejenes, hormigas blancas, palomillas de San Juan, polillas de la madera (General). Portuguese: cupins (sing, cupim), formigas brancas, formigas de asa, tucurus, aleluias do cupim (alates) (Brazil). Termites are the analogues in tropical soils of earthworms in temperate regions. All Neotropical termites, except Kalotermitidae, live in the soil or maintain a close connection between nest and soil (pi. If). Their physical burrowing to construct nests and digestion of plant structural material (cellulose) add significantly to soil fertility and earn termites a place in nature as very beneficial insects, notwithstanding the few species that damage man's struc­ tures (Snyder 1924). Their role as soil animals has been studied more in other regions (Lee and Wood 1971) but is cer­ tainly similar in Latin America. Recently, it has been realized that ter­ mites also have the potential to alter the environment in other ways, principally by releasing large amounts of methane, car­ bon dioxide, and hydrogen gases into the atmosphere as by-products of cellulose digestion. T h e greatest emissions come from natural tropical wet savanna and areas of human disturbance, such as cleared, burned, and cultivated lands, where abundant wood resources are avail­ able (Zimmerman et al. 1982). Termites live in nests (termitaria) of their own construction. T h e form and location of these structures vary among the different kinds of termites. They may be wholly subterranean with mounds cover­ ing or linked to subsoil chambers by tun­ nels or arboreal masses off the ground but with runways communicating with the soil surface. Mounds may be large, rising 3 to 4 meters aboveground and forming con­ spicuous edifices in the landscape, particu­ larly noticeable in open, flat country (Lacher et al. 1986). Such are the nests of

180

Cornitermes cumulans, common in pastures, cultivated land, and savannas in southern Brazil (Redford 1984). Arboreal nests may also be large, obvious, ovoid structures, but these are lodged on branches of trees and shrubs. The materials used for construction de­ pend on the termites' feeding habits and availability. They usually consist of clay soil, excreta, and plant fragments mixed with saliva. The nest generally has an inner laby­ rinth of chambers, including special cen­ tral rooms for the royal pair, where eggs are laid and the brood is raised (and in some cases, where food is stored and fungus cultivated). This is surrounded by a protective wall, itself sometimes perforated with galleries that lead to the exterior. There may also be long tunnels running to the surface, along the ground, or on the trunks and main limbs of trees. There is no clear correlation between nest architecture and termite systematics (Noirot 1977). It is difficult to generalize about the biology of the fauna because so very little is known about only a few species (Matthews 1977, Araújo 1970). Amitermes constructs nests with a very tall portion aboveground, some with umbrellalike lateral projections that function as rain-shedding devices. Syntermes (fig. 5.13a) is restricted to South America and is conspicuous because of the large size of individuals of most species (BL up to 17 mm or more) and the enormous volcano-shaped nests of some. Workers cut fragments of leaves and grass stalks and transport them to undergound galleries where they become stores for later consumption. Neocapritermes soldiers (fig. 5.13b) carry large, asymmetrical man­ dibles that may be snapped crosswise explo­ sively, emitting an audible click and driving the sharp angulate tips into the skin of any animal holding it. Nests of many species provide an invit­ ing abode for an assemblage of other higher animals, including nesting birds.

ORTHOPTEROIDS AND OTHER ORDERS

Figure 5.13 TERMITES, (a) Common tropical termite (Syntermes dims, Termitidae), reproductive male, (b) Crooked jaw termite (Neocapritermes braziliensis, Termitidae), head of worker, (c) Common tropical termite (Syntermes sp., Termitidae), physogastric queen from macropterous female. (d) Nasute termite (Nasutitermes costalis, Termitidae) worker, (e) Nasute termite soldier, (f) Nasute termite nest.

Many reptiles, such as legless lizards, feed on termites and occupy termitaria almost continuously. In Amazonia, in the forest away from normal sunlit riverbanks, fe­ males of the crocodilian Paleosuchus trigonatus find ground nests a convenient incubator for their eggs (Magnusson et al. 1985). According to locals, the Amazonian tortoise (Geochelone) also employs nests in this manner (orig. obs.). Many mammals specialize on termites as food, particularly the so-called anteaters. They eat more termites than ants, tearing open the nests with their powerful front claws. There are some true commensal termitophiles also among the insects and arthro­ pods. These are other termites, silverfish, scale insects (Termitococcus, Margarodidae), and even tiger beetles that live in intimate association with termite colonies. Many beetles, particularly some rove beetles, are highly modified for life among termites (Kistner 1969). Abondoned nests of one kind of termite can be taken over by termites of a different species. As is true of termites in other parts of the world, there are those habitually found in man-made structures that can cause considerable damage when they feed on important wooden members. Some of these, such as Cryptotermes brevis (Wolcott 1957) and Coptotermes havilandi, are intro­ duced from other places in the world, but some indigenous species are major pests as

well, for example, Incisitermes snyderi and Coptotermes niger. Termites are social insects and exhibit complex group behavior similar to that of ants and social bees and wasps. They are fundamentally different, however, in hav­ ing gradual metamorphosis and thus not having to provide for helpless larvae and pupae. Individuals comprising a colony are really members of a large family of sibling progeny started by a single paired winged reproductive male (fig. 5.13a) and female. The female queen is accompanied throughout her life by the attendant male and may live many years, growing in size to tremendous proportions (fig. 5.13c). T h e offspring consist of morphologically and functionally different castes that may be produced from newly hatched immatures, depending on the requirements of the colony. Control of this development de­ pends on pheromones given off by mem­ bers of the reproductive castes. Sterile castes are the workers and sol­ diers, wingless individuals in which the growth of the reproductive organs is sup­ pressed. The former are the most numer­ ous and generalized in form. They are responsible for all foraging activity and care for the eggs, larvae, and queen. Soldiers have similar bodies but highly modified mouthparts and an enlarged, strongly sclerotized head. There are two well-defined types, those with large promi-

TERMITES

181

n e n t m a n d i b l e s a n d t h e nasutes with a snoutlike p r o l o n g a t i o n o n t h e front of t h e h e a d a n d vestigial m a n d i b l e s . Both actively d e f e n d t h e nest against attackers, by biting a n d p i n c h i n g o r by t h e emission of viscid, noxious secretions (Deligne et al. 1 9 8 1 ; Prestwich 1983a, 19836). T h e h e a d of t h e soldiers is so modified that they c a n n o t feed themselves. D e p e n d i n g o n t h e species, t h e colony's p r i m a r y food is d e a d wood a n d o t h e r vegetative p a r t s of plants (usually d r y ) , roots, h u m u s , d u n g , a n d fungi, a l t h o u g h the last a r e n o t c u l t u r e d in Neotropical forms as in s o m e in Africa. T o a large extent, termites a r e d e p e n d e n t o n u n i q u e , symbiotic flagellate p r o t o z o a a n d bacteria for digestion of cellulose a n d o t h e r com­ plex polysaccharides in their diet. Mem­ bers of t h e family T e r m i t i d a e lack t h e p r o t o z o a , a l t h o u g h bacteria a r e p r e s e n t which a s s u m e t h e s a m e digestive function.

ARAÚJO, R. L. 1977. Catálogo dos Isoptera do novo mundo. Acad. Brasileira Cien., Rio de Janeiro. COWAN, F. 1865. Curious facts in the history of insects. Lippincott, Philadelphia.

I n times past in Brazil, large, h a r d termitaria have b e e n hollowed o u t a n d used for ovens (Sou thy, in C o w a n 1 8 6 5 : 134). T h e s a m e nests w e r e pulverized a n d used as a k i n d of c e m e n t t o m a k e "con­ c r e t e " floors for t h e early settlers of this land. T h e r e is j u s t o n e review of Neotropical termites (Araiijo 1970). I n t h e Neotropical region, t h e r e a r e sixty-two g e n e r a , contain­ ing 4 0 8 species (Araiijo 1970, 1972, 1977). Most b e l o n g t o t h e family T e r m i t i d a e . G e n e r a l i n f o r m a t i o n o n these insects is available in K r i s h n a a n d W e e s n e r (1969— 70). Very useful bibliographies of t e r m i t e literature to 1978 have b e e n compiled by S n y d e r (1956, 1 9 6 1 , 1968) a n d Ernst a n d Araiijo (1986).

MAGNUSSON, W. E., A. P. LIMA, AND R. M. SAM-

References ARAÚJO, R. L. 1970. Termites of the Neotropical Region. In K. Krishna and F. M. Weesner, Biology of termites. 2: 527-576. Academic, New York. ARAÚJO, R. L. 1972. Súmula faunistica dos Isoptera americanos. Cien. Cult. 24(3): 253-256.

182

DELIGNE, J., A. QUENNEDEY, AND M. S. BLUM.

1981. T h e enemies and defense mechanisms of termites. In H. R. Hermann, Social insects. 2: 1-76. Academic, New York. ERNST, E., AND R. L. ARAÚJO. 1986. A bibliogra­

phy of termite literature 1966-1978. Wiley & Sons, Somerset, N.J. KISTNER, D. H. 1969. T h e biology of termitophiles. In K. Krishna and F. M. Weesner, Biology of termites. 1: 525-557. Academic, New York. KRISHNA, K., AND F. M. WEESNER.

1969-70.

Biology of termites. 2 vols. Academic, New York. LACHER, JR., T. E., I. EGLER, C. J. R. ALHO, AND

M. A. MARES. 1986. Termite community com­ position and mound characteristics in two grassland formations in central Brazil. Biotropica 18: 356-359. LEE, K. E., AND T. G. WOOD. 1971. Termites and

soils. Academic, London. PAIO. 1985. Sources of heat for nests of Paleosuchus trigonatus and a review of crocodilian nest temperatures. J. Herpet. 19: 199-207. MATTHEWS, A. G. A. 1977. Studies on termites from the Mato Grosso State, Brazil. Acad. Brasileira Cien., Rio de Janeiro. NOIROT, C. 1977. Nest construction and phylogeny in termites. 8th Int. Union Study Soc. Ins. (Netherlands 1977) Proc. Pp. 177-180. PRESTWICH, G. D. 1983a. Chemical systematics of termite exocrine secretions. Ann. Rev. Ecol. Syst. 14: 287-311. PRESTWICH, G. D. 19836. T h e chemical defenses of Termites. Sci. Amer. 249: 78-87. REDFORD,

K. H.

1984. T h e termitaria of

Cornitermes cumulans (Isoptera, Termitidae) and their role in determining a potential keystone species. Biotropica 16: 112—119. SNYDER, T. E. 1924. Damage by termites in the Canal Zone and Panama and how to prevent it. U.S. Dept. Agrie. Dept. Bull. 1232: 1-25. SNYDER, T. E. 1956. Annotated, subject-heading bibliography of termites 1350 B.C. to A.D. 1954. Smithsonian Misc. Coll. 130: 1-305. SNYDER, T. E. 1961. Supplement to the anno­ tated, subject-heading bibliography of ter­ mites 1955 B.C. to A.D. 1960. Smithsonian Misc. Coll. 143: 1-137. SNYDER, T. E. 1968. Second supplement to the annotated subject-heading bibliography of

ORTHOPTEROIDS AND OTHER ORDERS

termites 1961 — 1965. Smithsonian Misc. Coll. 152(3): 1-188. WOLCOTT, G. N. 1957. Inherent natural resis­ tance of woods to the attack of the West Indian dry-wood termite, Cryptotermes brevis Walker. J. Agrie. Univ. Puerto Rico 41: 2 5 9 31L ZIMMERMAN, P. R., J. P. GREENBERG, S. D. WANDIGA, AND P. J. CRUTZEN. 1982. Termites:

A potentially large source of atmospheric methane, carbon dioxide, and molecular hy­ drogen. Science 218: 563-565.

Nasute Termites Termitidae, N a s u t i t e r m i t i n a e , Nasutitermes. Spanish: C o m e j e n e s c o m ú n (General). T h e large ( 1 - 2 m d i a m e t e r ) , r o u n d o r ovoid, d a r k b r o w n nests of these ubiqui­ tous tropical termites p u n c t u a t e t h e moist lowland l a n d s c a p e t h r o u g h o u t Latin A m e r ­ ica (fig- 5.13f, pi. 4a). T h e y a r e c o n s t r u c t e d of a paste of chewed wood a n d fecal cement p u t into place by myriad workers. T h e paste h a r d e n s into a c a r t o n substance that is soft a n d p a p e r y o n t h e o u t s i d e a n d increasingly h a r d e r t o w a r d t h e inside. T h e outer e n v e l o p e is c o n t i n u o u s , f o r m i n g a protective shell f o r t h e l a b y r i n t h i n e inte­ rior. Nests u n d e r g o continual e x p a n s i o n during their existence ( J o n e s 1979). Runways, c o v e r e d with this s a m e c a r t o n substance, lead from t h e m a i n nest to remote f o r a g i n g sites. T h e s e t u n n e l s e n d abruptly o n t h e surfaces of d e a d wood which t h e w o r k e r s p e n e t r a t e to feed. T h e presence of t h e s e r u n w a y s is what distin­ guishes t e r m i t e nests from t h e o t h e r arbo­ real p a p e r o r c a r t o n a n t a n d wasp nests with which they a r e often confused. Not all species build such elevated s t r u c t u r e s ; some, such as Nasutitermes fulviceps, live o n the soil surface a n d a r e partly s u b t e r r a ­ nean (Talice e t al. 1969). T h e t u n n e l s m a y also lead to fence posts, t e l e p h o n e poles, a n d t h e f o u n d a ­ tions of h o u s e s , whose substance is d e ­ voured with e q u a l gusto. Consequently, these termites a r e c o n s i d e r e d pests a n d often r e q u i r e control. T h e evidence for a

beneficial role for Nasutitermes as n i t r o g e n fixers is inconclusive (Bradley et al. 1983). T h e soldiers a r e best k n o w n a n d a r e usually seen w h e n they a p p e a r at t h e surface of nests that a r e b e i n g d a m a g e d . T h e y a r e small (BL 3 - 4 m m ) , with b r o w n ­ ish, p i g m e n t e d bodies a n d d a r k b r o w n h e a d s b e a r i n g a conspicuous, e l o n g a t e beak o n t h e front (fig. 5.13e). T h e y also possess an effective chemical defense. W h e n a b r e a k occurs in t h e nest surface, they immediately s w a r m t o t h e site a n d r e m a i n t h e r e until t h e i n t r u d e r leaves o r until they a r e ravished. As a m e a n s of repelling attackers, they ooze o r squirt a n irritating, thick, e n t a n g l i n g substance (na­ sute glue) from their long p o i n t e d snouts. T h i s sticky, smelly secretion is p r o d u c e d by large glands in t h e h e a d a n d contains volatile t e r p e n o i d s (Prestwich 1982). It is used mostly against o t h e r insect invaders of t h e colony b u t is effective also in ward­ ing off t h e attacks of t e r m i t o p h a g o u s verte­ brates, including a n t e a t e r s (Tamanduá, Myrmecophaga). T h e s e chemicals a r e topical irritants that affect t h e skin a n d m u c o u s m e m b r a n e s of t h e nostrils a n d m o u t h (Lubin a n d M o n t g o m e r y 1981). I n c u r s i o n s by animals a n d t h e formation of nest holes by trogons, parakeets, a n d o t h e r birds often d o considerable d a m a g e to nests a n d may force a b a n d o n m e n t by their occu­ p a n t s . Ants of t h e g e n u s Azteca a r e some­ times f o u n d living within Nasutitermes nests a n d m a y exclude t h e o w n e r s from their rightful h o m e . T h e workers (fig. 5.13d) a r e a b o u t t h e same size as t h e soldiers b u t have a r o u n d h e a d , a r e pale, a n d generally r e m a i n d e e p within t h e nest, c a r i n g for y o u n g a n d t h e q u e e n , even d u r i n g times of o u t w a r d threat. A l t h o u g h workers have n o special w e a p o n s , they can bite effectively a n d sometimes j o i n with t h e soldiers in aggres­ sive e n c o u n t e r s , especially against o t h e r termites ( T h o r n e 1982). Q u e e n s m a y attain g r e a t size ( B L 3—6 cm) by t h e e n l a r g e m e n t of t h e a b d o m e n

TERMITES

183

with fat a n d eggs. Normally, t h e r e is o n e p e r colony, b u t Nasutitermes corniger is k n o w n to be facultatively polygynous (mul­ tiple q u e e n s ) ( T h o r n e 1982). M e m b e r s vary greatly in n u m b e r , usu­ ally from five t h o u s a n d to six t h o u s a n d , d e p e n d i n g o n a g e , species, a n d health of the colony. B u t m u c h m o r e n u m e r o u s p o p u l a t i o n s a r e possible. S o m e Nasutitermes corniger nests may c o n t a i n 800,000 to a million individuals ( T h o r n e a n d Noirot 1982, Laffitte a n d A b e r d e S z t e r m a n 1976). R e p r o d u c t i o n is seasonal. S w a r m i n g of the winged male a n d female r e p r o d u c t i v e stages usually occurs after t h e first showers of t h e incipient rainy season a n d m a y c o n t i n u e for m a n y m o n t h s into t h e wet p a r t of t h e year. Many of t h e sixty-seven Neotropical spe­ cies of Nasutitermes place their nests in trees, o r at least off t h e g r o u n d in s h r u b b y vegetation. T h e walls of t h e r a r e terrestrial nests a r e s h o w n to contain n u t r i e n t s in excess of t h e s u r r o u n d i n g soil, t h u s concen­ trating substrate richness for plant g r o w t h a n d c o n t r i b u t i n g to patchy vegetation pat­ t e r n s in t h e A m a z o n Basin (Salick et al.

LUBIN, Y. D. 1983. Nasutitermes (comején, hormiga blanca, nasute termite). In D. H. Janzen, ed., Costa Rican natural history. Univ. Chicago Press, Chicago. Pp. 743-744. LUBIN, Y. D., AND G. G. MONTGOMERY.

1981.

Defenses of Nasutitermes termites (Isoptera Termitidae) against Tamanduá anteaters (Edentata, Myrmecophagidae). Biotropica 13: 66-76. MCMAHAN, E. A. 1970. Radiation and the

termites at El Verde. In H. T Odum, ed., A tropical rain forest: A study of irradiation and ecology at El Verde, Puerto Rico. U.S. AEC, Washington, D.C. Pp. E-105-122. PRESTWICH, G. D. 1982. From tetracycles to marcocycles: Chemical diversity in the de­ fense secretions of nasute termites. Tetrahe­ dron 38: 1911-1919. SALICK, J., R. HERRERA, AND C. F.JORDAN. 1983.

Termitaria: Nutrient patchiness in nutrientdeficient rain forests. Biotropica 15: 1—7. TALICE, R. V, S. L. MOSERA, R. CAPRIO, AND

A. M.S. DE SPRECHMANN. 1969. Estructura de

los termiteros. Univ. Uruguay, Dept. Biol. Gen. Exper. Publ. 2: 1-20. THORNE, B. L. 1982. Termite-termite interac­ tions: Workers as an agonistic caste. Psyche 89: 133-150. THORNE, B. L. 1982. Polygyny in termites: Multiple primary queens in colonies of Nasutitermes corniger (Motschuls) (Isoptera: Termitidae). Ins. Soc. 29: 102-117.

Figure 5.14 INSECTS OF VARIOUS ORDERS, (a) Web spinner (Clothoda urichi, Clothodidae). (b) Barklouse (Poecilopsocus iridescens, Psocidae). (c) Booklouse (Liposcelis bostrychophila, Lipuscelidae). (d) Black hunter (Leptothrips mali, Phlaeothripidae). (e) Greenhouse thrip {Heliothrips haemorrhoidalis, Thripidae). T h e silk f o r m i n g these labyrinths issues from glands in t h e basal s e g m e n t of t h e forelegs. T h i s s e g m e n t is inflated in nymphs a n d adults a n d clearly distin­ guishes these insects from all o t h e r s . O t h e r identifying features a r e t h e usually com­ plex, asymmetrical genitalia of t h e male, and, in alate species (many lack wings altogether o r have only small alar buds), wings that h a v e p i g m e n t e d b a n d s along the veins a l t e r n a t i n g with clear stripes. Web s p i n n e r s a r e m a n d i b u l a t e with fili­ form a n t e n n a e with n u m e r o u s segments.

ganic m a t t e r of plant origin that they find in their i m m e d i a t e habitat. A l t h o u g h Latin A m e r i c a is a major cen­ ter of evolution of t h e order, t h e embiid f a u n a has b e e n studied only to a limited d e g r e e (Ross 1943, 1944, 1984). Several h u n d r e d may actually exist, b u t only a b o u t 150 species in five families a r e presently described. T h e family C l o t h o d i d a e is con­ fined to South A m e r i c a (including Trini­ d a d a n d P a n a m a ) . Chelicera (Anisembiidae) is t h e d o m i n a n t g e n u s , with m a n y species in semiarid e n v i r o n m e n t s . Web s p i n n e r s a r e found in widely varied habitats, from h u m i d forests to deserts. O n e species is k n o w n from t h e Galápagos Islands, a n o t h e r from t h e f o g - d a m p e n e d lomas of coastal Peru (Ross 1966). A r o u n d h u m a n habitations, t h e most c o m m o n is Oligotoma saundersii, a "weed species," s p r e a d by m a n from India. Its males a r e attracted to lights.

References ARAÚJO, R. L. 1970. Termites of the Neotropical Region. In K. Krishna and F. M. Weesner, Biology of termites. 2: 527—576. Academic, New York.

WEB SPINNERS Embiidina ( = E m b i o p t e r a , Embiodea). Embiids.

A web s p i n n e r ' s b o d y is elongate a n d very supple. Flexible wings a n d s h o r t legs allow it to m o v e with g r e a t ease, even backward as easily as f o r w a r d , t h r o u g h its galleries. T h e s e insects a r e well p r o t e c t e d by their ability to r e t r e a t deeply within their silken tent, which n o t only forms a physical b a r r i e r to e n t r y by such p r i m a r y enemies as ants b u t hides t h e m from t h e eyes of larger p r e d a t o r s .

BRADLEY, R. S., L. A. DE OLIVEIRA, AND A. GOMEZ

While t h e majority of embiids (Ross 1970) are secretive a n d u n k n o w n except to the specialist, some species a r e very conspicu­ ous because of t h e extensive webs they construct o n tree t r u n k s a n d limbs. At times, almost t h e entire boles of large trees may be covered with these filmy mats that show little organized s t r u c t u r e save branch­ ing galleries. It is within these passages that the web s p i n n e r s live, a n d they may be seen t h r o u g h t h e walls as they move back a n d forth.

Embiids a r e g r e g a r i o u s . Typical colonies consist of a single female living in t h e midst of its brood (Edgerly 1988). A l t h o u g h they •hare in c o m m o n a c o m p l e x of galleries, they should b e c o n s i d e r e d subsocial, for they l a c k c a s t e S ; division of labor, o r o t h e r characteristics of t h e t r u e insect societies. At least o n e T r i n i d a d i a n web spinner, Gmhoda urichi (fig. 5.14a), is facultatively | » m m u n a l (Edgerly 1987). T h e food of embiids consists of bark, ~d leaves, moss, lichens, a n d o t h e r or­

1983, G o o d l a n d 1965). G e n e r a l i n f o r m a ­ tion o n t h e g e n u s is s p a r s e (Lubin 1983, Araujo 1970).

BANDEIRA. 1983. Nitrogen fixation in Nasuti­ termes in central Amazonia. In P. Jaisson, Social insects in the tropics. 1st Int. Symp. (Cocoyoc 1980), Int. Union Stud. Soc. Ins. and Soc. Mexicana Entomol. Proc. 2: 235-244. GOODLAND, R. J. A. 1965. On termitaria in a

savanna ecosystem. Can. J. Zool. 46: 641—650. JONES, R. J. 1979. Expansion of the nest of Nasutitermes costalis. Ins. Soc. 26: 322-342. LAFFITTE, S. AND A. ABER DE SZTERMAN.

1976.

Comportamiento interespecifico en Nasuti­ termes fulviceps (Silvestri, 1901) con otras ter­ mites. Rev. Biol. Uruguay 4: 59—65.

184

THORNE, B. L., AND C. NOIROT. 1982. Ergatoid

reproduction in Nasutitermes corniger (Motschulsky): Isoptera, Termitidae. J. Ins. Morph. Embryol. 11: 213-226.

ORTHOPTEROIDS AND OTHER ORDERS

References EDGERLY, J. S. 1987. Colony composition and some costs and benefits of facultatively com­ munal behavior in a Trinidadian webspinner Clothoda urichi (Embiidina: Clothodidae). En­ tomol. Soc. Amer. Ann. 80: 29-34. EDGERLY, J. S. 1988. Maternal behavior of a web­ spinner (Order Embiidina): Mother-nymph associations. Ecol. Entomol. 13: 263-272. Ross, E. S. 1943. Métodos de recoleción, crianza y estudio de los Embiópteros (Ins. Embioptera). Rev. Entomol. 14: 441-446. Ross, E. S. 1944. A revision of the Embioptera

WEB SPINNERS

185

of the New World. U.S. Nati. Mus. Proc. 94: 401-504. Ross, E. S. 1966. A new species of Embioptera from the Galápagos Islands. Calif. Acad. Sci. Proc. (Ser. 4) 34: 499-504. Ross, E. S. 1970. Biosystematics of the Embiop­ tera. Ann. Rev. Entomol. 15: 157-171. Ross, E. S. 1984. A classification of the Embiidina of Mexico with descriptions of new taxa. Calif. Acad. Sci. Occ. Pap. 140: 1-50.

HEMIPTEROIDS PSOCIDS Psocoptera ( = C o r r o d e n t i a ) . Barklice. Psocids (New 1987) a r e free-living insects that feed o n microflora a n d o r g a n i c debris on surfaces of vegetation o r on o t h e r sur­ faces. T h e r a n g e of food includes fungi ( h y p h a e a n d spores), yeasts, lichens, o r f r a g m e n t s of a n i m a l o r vegetable matter. Most a r e a r b o r e a l ( T h o r n t o n 1985) a n d a r e f o u n d on t h e b a r k o r leaves of trees, b u t m a n y also occur in g r o u n d litter. T h e r e t e n d s to b e a h i g h e r p r o p o r t i o n of leaf f r e q u e n t e r s in t h e c a n o p y t h a n n e a r t h e g r o u n d in Neotropical forests that have been s a m p l e d for psocids ( B r o a d h e a d a n d Evans 1979, B r o a d h e a d a n d Wolda 1985, W o l d a a n d B r o a d h e a d 1985). S o m e d w e l l i n the nests of m a m m a l s o r birds, b u t n o n e a r e parasitic like t h e i r close relatives, t h e biting a n d sucking lice. M e m b e r s of t h e family Trogiidae m a k e s o u n d s by d r u m m i n g t h e a b d o m e n against t h e substrate. Psocids a r e frequently g r e g a r i o u s as n y m p h s o r adults or b o t h a n d may even g r o u p t o g e t h e r u n d e r a c o m m u n a l web. Several c o s m o p o l i t a n types a r e com­ monly f o u n d i n d o o r s u n d e r h u m i d condi­ tions a n d a r e c o n s i d e r e d h o u s e h o l d pests. T h e s e a r e sometimes called "booklice" (fig. 5.14c) because of their habit of feeding on paper, sizing, a n d glue in book b i n d i n g s ( B r o a d h e a d 1946). T h e y a r e m i n u t e insects (BL 1 m m o r less) b u t a r e usually noticeable against a white p a p e r b a c k g r o u n d . All a r e

186

similar wingless (or n e a r wingless) forms brownish in color, a n d with s l e n d e r legs T h e r e a r e several species whose p r o p e r n a m e s a r e confused in t h e literature. T h e widely used n a m e Liposcelis divinatoria has b e e n declared invalid by psocid taxonomists; L. bastrychophila is t h e species to which it formerly r e f e r r e d ( L i e n h a r d 1990). T h e s e insects a r e a p p a r e n t l y widespread in Latin America, b u t t h e real e x t e n t of their o c c u r r e n c e h a s n o t been d o c u m e n t e d be­ cause of a lack of collecting a n d t h e uncer­ tainty of their identification. Psocids a r e related to lice (Lyal 1985) a n d a r e louselike in g e n e r a l appearance, b u t adults usually have wings that a r e held rooflike over t h e a b d o m e n w h e n a t rest. T h e wings have few veins, a n d t h e fore pair a r e m u c h larger t h a n t h e h i n d pair. Polymorphism is c o m m o n in some fami­ lies, t h e usual alternate form involving the r e d u c t i o n o r loss of wings. A u n i q u e devel­ o p m e n t is t h e bulging clypeal region on the front of t h e r o u n d h e a d , which is unusually movable at t h e neck for an insect. T h e p r o t h o r a x is r e d u c e d . T h e legs are slender a n d simple with a reduced n u m b e r of tarsal s e g m e n t s . A l t h o u g h some c o m m o n pest species a r e m i n u t e , most wild psocids a r e small (BL 1—2 m m ) a n d drably colored, gray or b r o w n , frail insects. In h u m i d tropical lowland forests, t h e r e a r e some much l a r g e r a n d quite colorful forms. Poecilopsocus iridescens, Psocidae (fig. 5.14b), of A m a z o n i a n Peru is a p p r o x i m a t e l y 12 milli­ m e t e r s long, with d a r k blue, white, a n d red wing m a r k i n g s a n d long a n t e n n a e . Possi­ bly, they a r e mimics of mirid o r reduviid bugs (Mockford pers. c o m m . ) . T h i s is a m u c h larger o r d e r in Latin America t h a n indicated by published lists (e.g., Smithers 1967). At present, there are at least 780 species described in 9 6 genera a n d several h u n d r e d m o r e that a r e certain to b e f o u n d (Mockford pers. comm.). Some speciose, typical regional g e n e r a a r e Thrysophorus, Ceratipsocus, a n d Graphocaecilius-

ORTHOPTEROIDS AND O T H E R ORDERS

A great m a n y species certainly a r e undis­ covered in t h e tropical p o r t i o n s of t h e region- As with o t h e r insect g r o u p s , t h e r e is a progressive increase in psocid diversity from t e m p e r a t e to tropical forests (Broad­ head 1983). S o m e show a m p h i n o t i c G o n d wanaland distributions with close relatives ¡ n Australia (e.g., Drymopsocus, Elipsocidae), b u t t h e s o u t h e r n t e m p e r a t e g r o u p s are mostly e n d e m i c a n d distinct from t h e rest of A m e r i c a to t h e n o r t h (New 1987).

References BROADHEAD, E. 1946. The book louse and other library pests. Brit. Book News 68: 7 7 - 8 1 . BROADHEAD, E. 1983. T h e assessment of faunal diversity and guild size in tropical forests with particular reference to the Psocoptera. In S. L. Sutton, T. C. Whitmore, and A. C. Chadwick, eds. Tropical rain forest: Ecology and management. Blackwell, Oxford. Pp. 107-119. BROADHEAD, E., AND H. A. EVANS. 1979.

The

diversity and ecology of Psocoptera in tropi­ cal forests. 4th Int. Symp. Trop. Ecol. [Pan­ ama] Acta 1: 185-196. BROADHEAD, E.,

AND H.

WOLDA.

1985.

The

diversity of Psocoptera in two tropical forests in Panama. J. Anim. Ecol. 54: 739-754. LIENHARD, C. 1990. Revision of the western Palaearctic species of Liposcelis Motschulsky (Psocoptera: Liposcelidae). Zool. Lb. Sys. 117: 117-174. LYAL, C. H. C. 1985. Phylogeny and classifica­ tion of the Psocodea, with particular refer­ ence to the lice (Psocodea: Phthiraptera). Syst. Entomol. 10: 145-165. NEW, T. R. 1987. Biology of the Psocoptera. Oriental Ins. 21: 1-109. SMITHERS, C. N. 1967. A catalogue of the Pso­ coptera of the world. Austr. Zool. 14: 1-145. THORNTON, I. W. B. 1985. T h e geographical

and ecological distribution of arboreal Psocop­ tera. Ann. Rev. Entomol. 30: 175-196. WOLDA, H.,

AND E. BROADHEAD.

1985.

Sea­

sonally of Psocoptera in two tropical forests in Panama. J. Anim. Ecol. 54: 519-530.

THRIPS "Thysanoptera. These a r e all very small insects (BL of most ~2 m m ) , a l t h o u g h "giant" forms a r e

found in t h e tropical forests of Latin A m e r ­ ica. T h e largest is t h e Peruvian Dasythrips regalis, which reaches a body l e n g t h of 12 millimeters. T h e o r d e r is only recently becoming well k n o w n generally ( A n a n t h a k r i s h n a n 1984, Lewis 1973). T h r i p s a r e characterized structurally mostly by their u n i q u e wings; b o t h pairs a r e very slender a n d elongate, w i t h o u t well-defined o r extensive venation a n d with very long hair fringes. Many species a r e wingless, however, a n d o t h e r features, such as t h e asymmetric m o u t h p a r t s located on a conical beak on t h e u n d e r s i d e of t h e h e a d , must be called on to define t h e m . Only t h e m a n d i b l e of t h e left side is d e v e l o p e d a n d is used to p u n c h holes in the e p i d e r m i s of plants to release t h e s a p , which is then sucked u p . T h e y also have a protrusible, saclike p a d at t h e a p e x of each leg. Most thrips live on plants, from which they take their liquid n o u r i s h m e n t . T h e b a n a n a flower thrips (Frankliniella párvula; H a r r i s o n 1963) a n d o t h e r s a r e often com­ m o n on flowers, w h e r e their feeding m a y result in injured fruit; s o m e h i d e in curled leaves (called queima in Brazil) or galls, which a r e caused by their feeding. T h e vegetarians may be very n u m e r o u s a n d cause extensive e c o n o m i c d a m a g e to com­ mercially valuable plants directly o r by introducing pathogenic microorganisms. A large n u m b e r of species a r e associated with t h e coconut palm (Sakimura 1986). A very different g r o u p lives on d e a d twigs a n d a m o n g leaf litter a n d soil w h e r e they a r e p r e d a c e o u s o n o t h e r m i n u t e in­ sects a n d mites o r feed on t h e fungi (hyphae a n d spores) associated with t h e early stages of decay ( M o u n d 1977). S o m e of t h e p r e d a t o r y types a r e c o n s i d e r e d b e n e ­ ficial when they attack pests. A n e x a m p l e is the black h u n t e r (Leptothrips mali; fig. 5.14d), which takes all sorts of injurious insects, including a p h i d s , scale insects, mites, a n d o t h e r t h r i p s . S o m e species feed on termites. T h r i p s also a r e pollinators, for

THRIPS

187

e x a m p l e , the b a n a n a flower thrips, which f r e q u e n t s the flowers of cacao in T r i n i d a d (Billes 1941). D e v e l o p m e n t in s o m e m e m b e r s of the o r d e r exhibits a parallel with that of t h e h o l o m e t a b o l o u s insects, the last n y m p h a l instar being quiescent a n d r e s e m b l i n g a pupa. T h e h i g h e r classification of t h e o r d e r has recently b e e n clarified ( M o u n d et al. 1980). Most of t h e eight families h a v e Latin A m e r i ­ can r e p r e s e n t a t i v e s . However, primarily only the species with pest status a r e k n o w n . T h e majority b e l o n g to t h e two ubiquitous families T h r i p i d a e a n d P h l a e o t h r i p i d a e . T h e s e include s o m e cosmotropical species such as t h e g r e e n h o u s e t h r i p s (Heliothrips haemorrhoidalis; fig. 5.14e), citrus thrips (Scirtothrips), tobacco a n d cotton t h r i p s (Frankliniella a n d Thrips), gladiolus t h r i p s (Taeniothrips simplex), a n d b a n a n a t h r i p s (Chaetanaphothrips). Native species a r e very poorly k n o w n in the r e g i o n . A bizarre Mexican a n d J a m a i ­ can t h r i p is Arachisothrips, in which the leading e d g e of the fore wing is ballooned into a hollow, p e a n u t - s h a p e d o u t g r o w t h with a reticulate surface ( S t a n n a r d 1952). It lives in rain forest g r o u n d cover, b u t the adaptiveness of this s t r a n g e structural fea­ t u r e is u n k n o w n . O n e Brazilian species, with its n e a r e s t relative in S i n g a p o r e , com­ prises t h e a b e r r a n t family U z e l o t h r i p i d a e . Franklinothrips vespiformis a n d relatives mimic various g e n e r a of ants in Mexico ( J o h a n s e n 1983).

References ANATHAKRISHNAN, T. N. 1984. Bioecology of

thrips. Indira, Oak Park, Mich. BILLES, D. J. 1941. Pollination of Theobroma cacao L. in Trinidad, B.W.I. Trop. Agrie. (Trinidad) 18: 151-156. HARRISON, J. O. 1963. Notes on the biology of the banana flower thrips Frankliniella párvula, in the Dominican Republic. Entomol. Soc. Amer. Ann. 56: 664-666. JOHANSEN, R. M. 1983. Nuevos estudios acerca del mimetismo en el género Franklinothrips

188

Back (Insecta: Thysanoptera), en Mexico. Inst. Biol. Univ. Nac. Aut. México Ann. (Ser Zool. 1)53: 133-156. LEWIS, T. 1973. Thrips, their biology, ecology and economic importance. Academic, Lon­ don. MOUND, L. A. 1977. Species diversity and the systematics of some New World leaf litter Thysanoptera (Phlaeothripinae; Glytothripini). Syst. Entomol. 2: 225-244. MOUND, L. A., B. S. HEMING, AND J. M. PALMER.

1980. Phylogenetic relationships between the families of recent Thysanoptera (Insecta). Zool. J. Linnean Soc. 69: 111-141. SAKIMURA, K. 1986. Thrips in and around the coconut plantations in Jamaica, with a few taxonomic notes (Thysanoptera). Fla. Ento­ mol. 69: 348-363. STANNARD, L. J. 1952. Peanut-winged thrips. Entomol. Soc. Amer. Ann. 45: 327-330.

NERVE-WINGED INSECTS Neuroptera. T h e most characteristic feature of adult n e u r o p t e r a n s is their well-developed wing venation, with highly c o m p l e x vein-branch­ ing p a t t e r n s , e n d twigging of the main veins, diverse polygonal cells in the middle areas, a n d f r e q u e n t stair-step pathways of m a n y vein b r a n c h e s . A few small groups a r e atypical, however, in having much simplified venation or wings very much r e d u c e d in size (brachypterous). Four equal size a n d s h a p e wings a r e the rule; they a r e held roofwise over the body when not in use for flight. T h e wing m e m b r a n e is usually clear but may sometimes be b r o w n p i g m e n t e d or rarely may have bright color spots a n d fields; it is com­ pletely whitish o p a q u e in the dusty wings (Coniopterygidae). T h e m o u t h p a r t s are m a n d i b u l a t e , a n d the h e a d possesses long, m a n y - s e g m e n t e d , filiform a n t e n n a e and well-formed c o m p o u n d eyes. A few fami­ lies have well-developed ovipositors. Most n e u r o p t e r a n adults a r e rather drably colored in cryptic g r e e n s , browns, a n d grays. Several u n r e l a t e d types, how­ ever, form an aposematic mimetic complex

ORTHOPTEROIDS AND OTHER ORDERS

in s o u t h e r n S o u t h America, all having a similar pair of b r i g h t r e d stripes o n the nrothorax n e a r the o p e n i n g s of scent glands that p r o d u c e a noxious, skunklike (contains skatol) odor. S o m e also have conspicuous p a t t e r n s o n the wings. A m o n g these are a n t lions ( M y r m e l e o n t i d a e ) , such Glenurus, a n d Maracandula, a s Dimares, some chrysopids, a n d mantispids. T h e mantispid g e n u s Climaciella mimics large social wasps, w h e r e a s t h e g e n u s Anchieta resembles stingless bees. T h e largest m a n ­ tispid in t h e world, Drepanicus gayi from Chile, looks like a g r e e n katydid. A n t lions are cryptically p a t t e r n e d to match their daytime resting sites (Stange 1970). T h e g e n e r a l f o r m of t h e larvae varies greatly, b u t m a n y r e s e m b l e g r o u n d beetle larvae. S o m e h a v e peculiarly modified elon­ gate, c u r v e d jaws, f o r m e d like tongs to grasp insect prey. Each has a n internal canal t h r o u g h which t h e juices of the food are s i p h o n e d . Many larvae are hairy o r

spiny. T h e larvae of s o m e C h r y s o p i d a e b e a r hooked bristles o n their backs to which they fix m i n u t e bits a n d pieces of debris to give t h e m a kind of camouflage (a habit that parallels the d e c o r a t o r crabs in coral reefs a n d t h e N o r t h A m e r i c a n Chrysopa slossonae, which attaches bits of the wax secretions of its woolly a p h i d prey to its body to mask it from recognition by the ants that p r o t e c t this a p h i d (Eisner et al.

1978). Such "trash-carrying lacewings" be­ long to the g e n e r a Leucochrysa a n d Ceraeochrysa. Larval habitats vary considerably a n d include vegetation, sandy soil, bark crevices, a n d cavities u n d e r objects o n the g r o u n d . A few ant lion larvae (Glenurus) are also trash carriers. H a r d l y a n y t h i n g has b e e n published o n the biology of the i m m a t u r e s of the Neotropical fauna except for a few benefi­ cial types such as t h e C h r y s o p i d a e ( N u n e z 1989a, 19896) a n d H e m e r o b i i d a e , the lar­ vae of which are voracious p r e d a t o r s of h o m o p t e r o u s pests (aphids a n d scale in­ sects, primarily) a n d t h u s of considerable value as biocontrol agents (New 1975). Species of Chrysoperla (fig. 5.15b) a r e even r e a r e d in insectaries to be b r o a d c a s t o n crops for this p u r p o s e . Mantispids (fig. 5.15a) a r e spider e g g p r e d a t o r s (Birabén 1960) o r bee parasites (Parker a n d S t a n g e 1965). Penny (1977) lists a p p r o x i m a t e l y 950 Latin A m e r i c a n species of N e u r o p t e r a . T h e s e are distributed a m o n g eleven fami­ lies, the largest a n d most c o m m o n of which a r e the ant lions ( M y r m e l e o n t i d a e ) , dusty wings (Coniopterygidae), a n d g r e e n lace­ wings (Chrysopidae). T h e r e a r e t h r e e n e u r o p t e r a n subfamilies f o u n d only in the Neotropical Region: the Platymantispinae, of u n c e r t a i n affinity b u t usually placed in the Mantispidae, are s t r a n g e s u b t e r r a n e a n p r e d a t o r s ( P a r k e r a n d S t a n g e 1965); t h e

figure 5.15 NEUROPTEROUS INSECTS, (a) Mantispid (Climaciella sp., Mantispidae). (b) Green ■Wwing (Chrysoperla sp., Chrysopidae). (c) Ant lion (Myrmeleon sp., Myrmeleontidae), larva in pit. »Ant lion (Glenurus peculiaris, Myrmeleontidae). (e) Owlfly (Corduleceris maclachlani, ""'aphidae).

NERVE-WINGED INSECTS

189

B r u c h e i s e r i n a e , two species of dusty wings f o u n d u n d e r rocks a n d possibly flightless; a n d finally t h e A l b a r d i n a e , c o n t a i n i n g o n e u n i q u e , highly modified owlfly species (Penny 1985).

References BIRABÉN, M. 1960. Mantispa parásita en el cocón de Metepeira. Neotropica 6: 61—64. EISNER, X , K. HICKS, M. EISNER, AND D. S.

ROBINSON.

1978. "Wolf-in-sheep's-clothing"

strategy of a predaceous insect larva. Science 199: 790-794. NEW, T. R. 1975. T h e biology of Chrysopidae and Hemerobiidae (Neuroptera), with refer­ ence to their usage as biocontrol agents: A review. Royal Entomol Soc. London Trans. 127: 115-140. NUÑEZ, E. 1989a. Chrysopidae (Neuroptera) del Perú y sus especies más comunes. Rev. Peruana Entomol. 31: 6 9 - 7 5 . NUÑEZ, E. 19896. Ciclo biológico y crianza de Chrysoperla externa y Ceraeochrysa cincta (Neur­ optera, Chrysopidae). Rev. Peruana Entomol. 31: 76-82. PARKER, F. D., AND L. A. STANGE. 1965. System­

atic and biological notes on the tribe Platymantispini and the description of a new species ol Plega from Mexico. Can. Entomol. 97: 604-612. PENNY, N. D. 1977. Lista de Megaloptera, Neuro­ ptera e Raphidioptera d o México, América Central, ilhas Caraibas e América do sul. Acta Amazónica 7 (4) suppl.: 1—61. PENNY, N. D. 1985 [1983]. Neuroptera of the Amazon Basin. Pt. 9. Albardiinae. Acta Ama­ zónica 13(3-4): 697-699. STANCE, L. A. 1970. Revision of the ant-lion tribe Brachynemurini of North America. Univ. Calif. Publ. Entomol. 55: 1-192.

Ant Lions Neuroptera, Myrmeleontidae. These are probably the best-known neuropterans, n o t because of t h e adults b u t b e ­ cause of t h e work of t h e larvae, t h e familiar, f u n n e l - s h a p e d a n t lion pits c o m m o n l y seen in fine sandy soil. T h e s e a r e c o n s t r u c t e d in places p r o t e c t e d from rain: u n d e r stair­ cases a n d by t h e edges of elevated build­ ings, b e n e a t h o v e r h a n g i n g rocks o r logs,

190

a n d n e a r t h e bases of trees. T h e y m a y be very n u m e r o u s a n d occur in large, concen­ trated g r o u p s at times (McClure 1976) Only larvae of t h e g e n u s Myrmeleon make these d e a t h t r a p s , into which ants o r other small, terrestrial insects fall (Wilson 1974) (Pit m a k i n g is f o u n d in a few o t h e r groups [Brachynemurus in A r g e n t i n a a n d Mexico] but tubes e x t e n d d o w n w a r d from t h e pits.) T h e escape of t h e hapless insect is pre­ vented by t h e constantly failing sloped sides of t h e pit. T h e a n t lion at t h e b o t t o m also flicks sand o n t o t h e s t r u g g l i n g prey to dislodge it a n d cause it to t u m b l e d o w n to the neck of t h e funnel. T h e larva waits t h e r e , b u r i e d , only its ice-tongjaws project­ ing above t h e surface. T h e s e formidable structures close o n t h e prey, t h e tips pierc­ ing its body, a n d take its blood through internal canals. T h e s e larvae only move backward, a trait s h a r e d by t h e giant Vella larvae that often prey o n t h e m (Stange pers. comra.). T h e m e a n d e r i n g subsurface bur­ rows of t h e latter c o m m o n Neotropical g e n u s a r e often evident in loose sandy areas.

large (BL 3 - 4 c m ) , with a very long, «lender a b d o m e n , sometimes twice as long as t h e wings in t h e male. T h e wing vena­ tion is very c o m p l e x : t h e r e is a very long c e ]l u n d e r t h e stigma spot in b o t h wings; the conspicuous vein forking in t h e m i d d l e of the wings n e a r t h e base looks identical in the fore wing a n d h i n d wing b u t actually involves totally different crossveins (Stange 1970); t h e subcostal a r e a of t h e wing is t h e only part lacking cross veins. T h e a n t e n n a e are short, t h i c k e n e d , a n d e x p a n d e d at t h e tip into a club. Males of m a n y species have claspers at t h e tip of t h e a b d o m e n a n d a peculiar b r u s h at t h e base of t h e h i n d wing. Most a r e dully m a r k e d with b r o w n a n d black speckling o n t h e b o d y a n d t r a n s p a r ­ ent wings, b u t a few have conspicuously marked wings, such as Morocordula apicalis from s o u t h e r n Mexico a n d species of Glenurus (fig. 5.15d) a n d Dimarini (Stange 1989).

O t h e r a n t lion larvae merely b u r r o w in sand o r loose soil, s e a r c h i n g for other s u b t e r r a n e a n insects to b e c a p t u r e d di­ rectly a n d eaten. Many c a n d i g rapidly in reverse by r e p e a t e d l y a r c h i n g t h e abdo­ m e n forward u n d e r t h e body. Some ant lions that live in d r y tree holes (e.g., some Glenurus; Miller a n d S t a n g e 1983) o r on tree bark o r bare rock surfaces (Navasoleon; see Miller a n d Stange 1985) have dimin­ ished o r lost their ability to d i g backward.

MCCLURE, M. S. 1976. Spatial distribution of pit-making ant lion larvae (Neuroptera: Myrmeleontidae): Density effects. Biotropica 8: 179-183.

A n t lion larvae (fig. 5.15c) a r e small (BL 5-10 m m ) , ovoid c r e a t u r e s with a boxlike h e a d a n d typical n e u r o p t e r a n sickle-shaped jaws. T h e y usually have en­ larged bristles at t h e r e a r e n d , employed in rapid digging, a n d have their eyes on stalks. T h e a b d o m e n is b u l b o u s a n d the a n t e r i o r p o r t i o n steep, so that t h e head a n d t h o r a x e m e r g e very low. A d u l t m y r m e l e o n t i d s a r e mostly fairly

ORTHOPTEROIDS AND OTHER ORDERS

Owlflies N e u r o p t e r a , Ascalaphidae. Looking somewhat like dragonflies b u t actually related to a n t lions, these a r e medium-sized to large (BL 2—4 cm, B W L to 10 cm) n e u r o p t e r a n s with reticulate wing venation. T h e wings a r e often in­ fused with b r o w n o r black, a l t h o u g h they are usually crystalline except for t h e small stigmal spot n e a r t h e a n t e r i o r e d g e of t h e tip. T h e y have large, many-faceted eyes, a n d t h e a n t e n n a e a r e characteristically long a n d filamentous, with a large, flat, terminal k n o b , s o m e w h a t like those of butterflies. T h e a n t e r i o r p a r t of t h e h e a d , t h o r a x , a n d legs is frequently hairy. Males of some species have a p r o n o u n c e d elon­ gate dorsal process o n t h e second a b d o m i ­ nal segment.

MILLER, R. B., AND L. A. STANCE. 1985. Descrip­

Owlfly larvae a r e n o n b u r r o w e r s , living openly o n leaves, o n tree t r u n k s , o r o n t h e g r o u n d . T h e y a r e similar to those of a n t lions b u t lack t h e e n l a r g e d d i g g i n g claw o n the hind leg. Almost all species have hairy, fingerlike processes o n t h e body, which a r e r a r e in a n t lions ( H e n r y 1976). T h e y a r e slow-moving a n d usually wait in a m b u s h for prey. T h e p u p a is p r o t e c t e d by a weak cocoon woven entirely of silk by t h e m a ­ t u r e larva. A n o t h e r peculiarity of t h e g r o u p is t h e laying of d e f o r m e d eggs (repagula) a r o u n d t h e fertile o n e s (New 1971).

tion of the antlion larva Navasoleon boliviana Banks with biological notes (Neuroptera; Myr­ meleontidae). Neuroptera Intl. 3: 119-126. PENNY, N. D. 1977. Lista de Megaloptera, Neuroptera e Raphidioptera do México, América Central, ilhas Caraibas e América do sul. Acta Amazónica 7(4) suppl.: 1-61. STANCE, L. A. 1970. Re vision of the ant-lion tribe Brachynemurini of North America. Univ. Calif. Publ. Entomol. 55: 1-192. STANCE, L. A. 1989. Review of the New World Dimarini with the description of a new genus from Peru (Neuroptera: Myrmeleontidae). Fla. Entomol. 72: 450-461. WILSON, D. S. 1974. Prey capture and competi­ tion in the ant lion. Biotropica 6: 187-193.

Biological i n f o r m a t i o n o n t h e family in the Neotropics is meager. A d u l t s of t h e g e n u s Corduleceris (fig. 5.15e), u n i q u e in having strongly d i m o r p h i c sexes, h a v e b e e n observed to a g g r e g a t e at t h e tips of tree b r a n c h e s o v e r h a n g i n g s t r e a m s (Covell 1989, H o g u e a n d Penny 1989). T h e p u r ­ pose of this behavior is u n k n o w n , a l t h o u g h the clusters m a y b e c o m p o s e d of sleeping individuals. Single adults often rest o n twigs, head d o w n w a r d , with wings a n d t h e a n t e n n a e held closely parallel to t h e sur­ face, t h e a b d o m e n sharply erect. T h e y a r e

T h e r e a r e 35 g e n e r a in Latin A m e r i c a n containing 2 2 4 species (Penny 1977).

References

MILLER, R. B., AND L. A. STANCE. 1983. T h e

antlions of Florida: Glenurus grains (Say) (Neuroptera: Myrmeleontidae). Fla. Dept. Agrie. Entomol. Circ. 251: 1—2.

NERVE-WINGED INSECTS

191

usually excellent fliers, a l t h o u g h they some­ times elect to catch o t h e r insects while foraging from leaf surfaces r a t h e r t h a n o n the wing. T h e r e a r e b o t h n o c t u r n a l a n d d i u r n a l species. T h e f o r m e r c o m e to artifi­ cial light occasionally. T h e family is fairly diverse in Latin America. Penny (1977) lists 94 species in 15 g e n e r a , t h e largest a n d most c o m m o n of which a r e Ululodes a n d Ameropterus. T h e r e are two large g r o u p s s e p a r a t e d by the form of the eye, those with split eyes, that is, with the eye partially divided by a transverse groove, a n d those with e n t i r e eyes. A t h i r d subfamily, r e p r e s e n t e d in the New World only by Albardia furcata in Brazil, is peculiar in the very s h o r t a n t e n n a e a n d woolly a b d o m e n of t h e a d u l t (Penny 1983).

References COVELL, JR., C. V. 1989. Aggregation behavior in a Neotropical owlfly, Cordulecerus maclachlani (Neuroptera: Ascalaphidae). Ento­ rno]. News 100: 155-156. HENRY, C. S. 1976. Some aspects of the external morphology of larval owlflies (Neuroptera: Ascalaphidae), with particular reference to Ululodes and Ascaloptynx. Psyche 83: 1—31.

6

AQUATIC ORDERS

HOGUE, C. L., AND N. D. PENNY. 1989. Aggrega­

tions of Amazonian owlflies (Neuroptera: Ascalaphidae: Cordulecerus). Acta Amazónica 18:359-361 NEW, T. R. 1971. Ovariolar dimorphism and repagula formation in some South American Ascalaphidae. J. Entomol. 46: 73-77. PENNY, N. D. 1977. Lista de Megaloptera Neuroptera e Raphidioptera do México, América Central, ilhas Caraibas e América do sul. Acta Amazónica 7(4) suppl.: 1-61. PENNY, N. D. 1983. Neuroptera of the Amazon Basin. Acta Amazónica 13: 697-699.

Aquatic insect life in Latin America is diverse a n d a b u n d a n t in b o t h r u n n i n g a n d standing inland waters a n d , to a limited extent, in the seas s u r r o u n d i n g the conti­ nental a n d island areas (see Ecosystems, chap. 2). All t h e major categories of waterdwelling insects a r e r e p r e s e n t e d a n d widely distributed e x c e p t in the n u t r i e n t - p o o r black water a n d clear water systems. Rela­ tive to those c a r r y i n g white waters, the major black water d r a i n a g e s of the G u i a n a (e.g., Rio N e g r o ) a n d Brazilian (e.g., Rio Tapajos) shields of S o u t h A m e r i c a a r e well known as faunistically d e p a u p e r a t e ("hun­ gry rivers") with r e g a r d to insects as well as vertebrates ( J u n k a n d Furch 1985: 15). Amazonia is still largely u n e x p l o r e d with respect to aquatic insects, as is most of the Andes. Many taxa f o u n d in streams in the southern p o r t i o n of South A m e r i c a a r e very ancient a n d show affinities to the faunas of the Australian region a n d south­ ern Africa ( G o n d w a n a l a n d a n d a m p h i n o t i c distributions) (lilies 1969). Families of insects in mostly terrestrial orders that h a v e a d a p t e d to life in fresh water are discussed e l s e w h e r e (see water bugs, c h a p . 8; water beetles, c h a p . 9; water midges, p u n k i e s , mosquitoes, etc., chap. 11). T h o s e o r d e r s totally a d a p t e d to life in fresh water a r e t h e subject of this chapter. Two of these o r d e r s , the E p h e m e r optera and O d o n a t a , form t h e Paleoptera, ^nsidered to be the stem g r o u p a n d the t ancient of the winged insects, from hich all h i g h e r insects evolved. Evidence

192

ORTHOPTEROIDS AND OTHER ORDERS

for this comes partly from t h e lack of a flexing m e c h a n i s m in the wing articulation a n d the reliance of the i m m a t u r e s on gills for respiration, not a t m o s p h e r i c air, the latter being a secondary a d a p t a t i o n in aquatic insects. Possibly the most primitive of the N e o p t e r a a r e the Plecoptera, as indicated by their generalized b o d y struc­ ture. T h e Megaloptera and Trichoptera are h i g h e r N e o p t e r a b u t are t h o u g h t to occupy positions basal to the phylogeny of the n e u r o p t e r o i d a n d p a n o r p o i d o r d e r s for the same reasons (see Evolution a n d Classification, c h a p . 1). T h e larvae a n d n y m p h s (sometimes called naiads) of these o r d e r s inhabit wa­ ters of all descriptions, i n c l u d i n g saline a n d t h e r m a l waters, b u t n o n e a r e m a r i n e as a r e some species of the aquatic families discussed elsewhere. T h e y a r e active but r e m a i n within the b o u n d a r i e s of their vastly varied microhabitats. Dobsonfly a n d caddis fly p u p a e a r e s u b m e r g e d , often in protective cases or cocoons, but may b e terrestrial a n d even active, having func­ tional muscles for m o v e m e n t of legs a n d m o u t h p a r t s . T h e adults r e m a i n close by the habitats of the i m m a t u r e s a n d a r e mostly p r e d a c e o u s o r d o not feed. Current and comprehensive introduc­ tions to the biology a n d z o o g e o g r a p h y of Latin A m e r i c a n aquatic insects, including extensive literature citations, a r e to be f o u n d in an i m p o r t a n t t h r e e - p a r t collabora­ tive t r e a t m e n t edited by H u r l b e r t a n d o t h e r s (1977, 1981, 1982); for i m p o r t a n t t r e a t m e n t s of local faunas, see also Vanzo-

193

lini (1964) for Brazil only a n d R o l d a n (1988) for Colombia. References HURLBERT, S. H. ed. 1977. Biota acuática de sudamérica austral. San Diego State Univ., San Diego. HURLBERT, S. H., G. RODRÍGUEZ, AND N. DÍAS

DOS SANTOS, eds. 1981. Aquatic biota of tropical South America. Pt 1. Arthropoda. San Diego State Univ., San Diego. HURLBERT, S. H., AND A. VILLALOBOS FIGUE-

ROA, eds. 1982. Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ, San Diego. ILLIES, J. 1969. Biogeography and ecology of Neotropical freshwater insects, especially those from running waters. In E. J. Fittkau, J. Illies, H. Klinge, G. H. Schwabe, and H. Sioli, eds., Biogeography and ecology in South America. 2: 685-708. Junk, T h e Hague. JUNK, W. J., AND K. FURCH. 1985. T h e physical

and chemical properties of Amazonian wa­ ters and the relationships with the biota. In G. T Prance and T E. Lovejoy, Amazonia. Pergamon, Oxford. Pp. 3—17. ROLDAN, G. 1988. Guía para el estudio de los macroinvertebrados acuáticos del Departa­ mento de Antioquia. Fondo Fen Colombia/ COLCIENCIAS, Univ. Antioquia, Bogotá. VANZOLINI, P. E., ed. 1964. Historia natural de organismos aquáticos do Brasil: Bibliografía comentada. Fund. Amparo Pesq. Est. Sao Paulo, Sao Paulo.

MAYFLIES Ephemeroptera (= Ephemerida). Portuguese: Siriruias (Brazil). T h i s is c o n s i d e r e d t h e most primitive of all the living o r d e r s of w i n g e d insects. Only mayflies u n d e r g o a molt after acquiring functional wings. T h e wings a r e incapable of being folded r e a r w a r d a n d often possess a very c o m p l e x , netlike venation, both ancestral c h a r a c t e r s indicative of early ori­ gin. T h e h i n d wing is always m u c h smaller t h a n t h e t r i a n g u l a r fore wing a n d in m a n y cases is lost altogether. A d u l t mayflies a r e f u r t h e r recognized by their strongly d e v e l o p e d eyes, particu­

194

AQUATIC ORDERS

larly in t h e males. T h e s e a r e often divided into u p p e r a n d lower p o r t i o n s of larger a n d smaller o m m a t i d i a . T h e a n t e n n a e are m e r e stylets, b u t t h e two o r t h r e e terminal sensory filaments (cerci plus m e d i a n cau­ dal filament) a r e very long, e x t e n d i n g from t h e tip of the a b d o m e n . M o u t h p a r t s are vestigial a n d t h e legs weak or reduced, or even vestigial in Campsurus (fig. 6.1a). Adults of most species a r e dull colored. T h o s e in t h e w i d e s p r e a d g e n u s Thraulodes (fig. 6.1c) may have striking p a t t e r n s ; male Tricorythodes a r e black with milky wings. T h e wings of t h e males in m a n y species have partially maculate wings. T h e biology of mayflies has b e e n exten­ sively studied by entomologists a n d limnologists (Flannagan a n d Marshall 1980). T h e body form of t h e aquatic n y m p h s differs m u c h a m o n g families b u t usually little from that of its own adult. Eyes, a n t e n n a e , and m o u t h p a r t s a r e well f o r m e d , as a r e t h e cerci a n d m e d i a n caudal filament. Conspicuous also a r e four to seven pairs of articulated, lateral, platelike gills (often d o u b l e plates) on most of t h e a b d o m i n a l segments. N y m p h s also display varied a n d sometimes bizarre shapes as specializations to their different s u b m e r g e d aquatic habitats. Slen­ der, cylindrical types a r e strong, "min­ nowlike" swimmers, occupying still water in p o n d s a n d stream pools a n d sometimes m o u n t a i n t o r r e n t s . T h e r e a r e flattened forms that lodge between a n d u n d e r rocks, a n d some have splayed legs that cling to exposed rock surfaces in fast c u r r e n t s (e.g., Thraulodes, fig. 6 . I d ) . O t h e r s a r e robust, with heavy, shovel-shaped, spiny legs and head used for b u r r o w i n g in b o t t o m muds or sand (e.g., Campsurus, fig. 6.1b). T h e n y m p h s of most species a r e herbi­ vores or scavengers, taking detritus and aquatic microorganisms, especially dia­ toms. A minority, such as Chiloporter and Chaquihua in Chile a n d A r g e n t i n a , are p r e d a t o r y o n o t h e r small aquatic inverte­ brates ( E d m u n d s pers. c o m m . ) . O n m a t u r i n g , t h e n y m p h s transform

Figure 6.1 MAYFLIES, (a) Legless mayfly (Campsurus albicans, Polymitarcyidae). (b) Legless mayfly (Campsurus sp.) nymph, (c) Tropical mayfly (Thraulodes sp., Leptophlebiidae). (d) Tropical mayfly (Thraulodes sp.) nymph. into the alate, flying, b u t sexually imma­ ture s u b i m a g o (the " d u n " ) . T h e transfor­ mation usually takes place at t h e water's surface b u t m a y also occur below t h e water or after t h e n y m p h h a s crawled o u t of t h e water o n t o s o m e object. T h i s stage soon m e t a m o r p h o s e s into a r e p r o d u c t i v e adult (imago). S u b i m a g o s have infúscate wings and t h e i n t e g u m e n t covered with microspines to distinguish t h e m from t h e glassy-winged, glossy-surfaced imagos (Ed­ munds 1988). Imagos a r e e p h e m e r a l , their active lives lasting only a few h o u r s or, at most, days. They d o n o t feed a n d s p e n d most of their short existence on t h e wing, m a t i n g a n d egg laying. T h e y generally r e m a i n n e a r their b r e e d i n g g r o u n d s w h e r e they a r e seen flying o r resting. Many species swarm, some (as in t h e g e n e r a Tortopus a n d Campsurus) in such great n u m b e r s as to constitute a p l a g u e , especially w h e n d r a w n to street lights o r h o u s e lights in u r b a n i z e d areas. T h o u s a n d s of individuals, mostly females, pile u p in t h e streets or press indoors to m a k e themselves a nuisance. Within these s w a r m s , t h e sexes copulate in flight, a n d females fall into t h e water as they release t h e i r eggs. Mayflies in all stages f o r m a large p a r t of the diet of freshwater fish a n d small ripar­ ian birds. T h e y a r e also e a t e n by m a n y types of c a r n i v o r o u s aquatic insects. T h e importance o f s o m e prolific species in food chains has c a u s e d t h e m to be r e f e r r e d to as "insect cattle" (e.g., Callibaetis).

Mayfly classification a n d evolutionary history a r e c o m p l e x ( E d m u n d s 1972). T h e r e a r e 13 families, p e r h a p s c o n t a i n i n g , w h e n all a r e discovered, a b o u t 500 o r m o r e species in all of Latin A m e r i c a , extrapolat­ ing from H u b b a r d ' s (1982) list of 300 currently n a m e d species. T h e i r p o o r abili­ ties to disperse m a k e t h e m useful as biogeographic indicators. A l t h o u g h s o m e c o m m o n g e n e r a in S o u t h A m e r i c a may e x t e n d to N o r t h America, m e m b e r s of several g r o u p s in s o u t h e r n S o u t h A m e r i c a show closer relationships with species from o t h e r austral areas (Australia, New Zea­ land, a n d s o u t h e r n Africa) t h a n with spe­ cies to t h e n o r t h . E x a m p l e s include t h e g e n e r a Metamonius, Chiloporter, Chaquihua, a n d Siphlonella of t h e S i p h l o n u r i d a e a n d Nousia of t h e L e p t o p h l e b i i d a e (Pescador a n d Peters 1982, 1985). For g e n e r a l r e ­ views of t h e o r d e r in Latin America, see H u b b a r d a n d Peters (1977, 1981) a n d E d m u n d s (1982). A n i m p o r t a n t t a x o n o m i c p a p e r is t h e review of t h e Neotropical Leptophlebiidae by Savage (1987).

References EDMUNDS, JR., G. F. 1972. Biogeography and

evolution of Ephemeroptera. Entomol. 17: 21-42.

Ann. Rev.

EDMUNDS, JR., G. F. 1982. Ephemeroptera. In

S. H. Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central Amer­ ica and the West Indies. San Diego State Univ., San Diego. Pp. 242-248. EDMUNDS, JR., G. F. 1988. T h e mayfly sub-

imago. Ann. Rev. Entomol. 33: 509-529.

MAYFLIES

195

FLANNAGAN, J. F., AND K. E. MARSHALL, eds.

1980. Advances in Ephemeroptera biology. Plenum, New York. HUBBARD, M. D. 1982. Catálogo abreviado de Ephemeroptera da América do sul. Pap. Avul. Zool. (Sao Paulo) 34(24): 257-282. HUBBARD, M. D., AND W. L. PETERS.

1977.

Ephemeroptera. In S. H. Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 165-169. HUBBARD, M. D., AND W. L. PETERS. 1981.

Ephemeroptera. In S. H. Hurlbert, G. Rodri­ guez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt 1. Arthropoda. San Diego State Univ., San Diego. Pp. 55-63. PESCADOR, M. L., AND W. L. PETERS. 1982. Four

new genera of Leptophlebiidae (Ephemer­ optera: Atalophlebiinae) from southern South America. Aquatic Ins. 4: 1 — 19. PESCADOR,

M. L., AND W. L. PETERS.

1985.

Biosystematics of the genus Nousia from southern South America (Ephemeroptera: Leptophlebiidae: Aetalophlebiinae). Kans. Entomol. Soc. J. 58: 9 1 - 1 2 3 . SAVAGE, H. M. 1987. Biogeographic classifica­ tion of the Neotropical Leptophlebiidae (Ephemeroptera) based upon geological cen­ ters of ancestral origin and ecology. Stud. Neotrop. Fauna Environ. 22: 199-222.

DRAGONFLIES AND DAMSELFLIES Odonata. T h e s e a r e familiar insects, always f o u n d n e a r water, a l t h o u g h t h e powerful a n d u n t i r i n g wings of dragonflies m a y take t h e m o n l o n g j o u r n e y s . T h e y a r e easily recognized by their e l o n g a t e bodies, four

similar, many-veined wings (with a dark spot o n t h e leading e d g e n e a r t h e tip, the pterostigma), a n d b u l b o u s eyes with an e n o r m o u s n u m b e r of m i n u t e ommatidia. Between t h e eyes arise t h e tiny, bristlelike a n t e n n a e . T h e thoracic s e g m e n t s a r e an­ gled obliquely so that their dorsal surfaces form a n incline. Dragonflies a r e distinguished from damselflies principally by their m o r e robust a n d usually larger bodies a n d their habit of e x t e n d i n g t h e wings o u t t o t h e sides when at rest. S o m e damselflies a r e also large, but they a r e always slender. Most damselflies fold their wings t o g e t h e r back over the a b d o m e n w h e n n o t in use. T h e wings are also abruptly n a r r o w e d a n d slender at the base in contrast to t h e broadly based drag­ onfly wings. T h e n y m p h s of dragonflies are also m o r e heavily built t h a n those of the damselflies. T h e f o r m e r have a broad, t a p e r i n g a b d o m e n , t i p p e d with short spinose processes (fig. 6.2b); their gills are located internally in folds of t h e rectum. Damselfly n y m p h s have elongate, slender a b d o m e n s , b e a r i n g t h r e e , conspicuous, finlike terminal gills (fig. 6.3b, e). T h e body a n d wings of o d o n a t e s are very often highly d e c o r a t e d with bright or gaudy colors. T h e tints of t h e body are transient a n d quickly d i s a p p e a r from dead specimens; b u t those of t h e wings persist as spot a n d b a n d p a t t e r n s o r b r o a d fields, most often over t h e basal half o r third of the wing. I n some, t h e entire wing may be colored, often in glossy r e d , o r a n g e , or

Figure 6.2 LIBELLULID DRAGONFLIES (LIBELLULIDAE). (a) Globetrotter (Pantala flavescens). (b) Globetrotter nymph, (c) Black wing (Diastatops dimidiata). (d) Amber wing (Perithemis indensa).

196

AQUATIC ORDERS

blue. T h e coloration of t h e male fre­ quently differs from t h a t of t h e female, and it is sometimes difficult to know they are of t h e s a m e species until they a r e seen copulating. D u r i n g t h e p a i r i n g process, t h e sexes are peculiarly j o i n e d in a t a n d e m configu­ ration u n i q u e to this o r d e r of insects. T h e male's c o p u l a t o r y s t r u c t u r e s (genital fossa) are situated o n t h e u n d e r s i d e s of t h e second a n d t h i r d a b d o m i n a l s e g m e n t s at a considerable distance from t h e t r u e sexual aperture at t h e tip of t h e a b d o m e n . Prior to mating, h e transfers s p e r m a t o z o a from the g o n o p o r e t o t h e penis in t h e fossa. During m a t i n g , t h e male grasps t h e female behind t h e h e a d (dragonflies) o r t h o r a x (damselflies) with his s t r o n g , tonglike cerci while s h e b e n d s h e r a b d o m e n forward to the fossa a n d receives t h e s p e r m from t h e penis. C o u p l e s fastened t o g e t h e r in this manner a r e c o m m o n l y seen resting o n vegetation by t h e water's e d g e , o r even in flight. Adult biology, such as flight p a t t e r n s and competition for prey a n d h u n t i n g space, in Latin A m e r i c a h a s n o t b e e n well studied b u t is generally t h e same as that observed elsewhere. S o m e special a d a p t a ­ tions for surviving d r y p e r i o d s have b e e n noted (Morton 1977). I m m a t u r e o d o n a t e s a r e all aquatic, t h e nymphs b e i n g f o u n d in all sorts of r u n n i n g and still water e n v i r o n m e n t s : p o n d s , shal­ low stream a n d lake m a r g i n s , s t r e a m pools, tree holes, a n d t a n k plants. T h e n y m p h s are insectivorous. T h e y c a p t u r e prey with a mantislike g r a b of t h e e n l a r g e d a n d elongated, extensile lower lip (labium). The n y m p h s show structural a d a p t a t i o n s in response t o t h e diverse ecological niches they occupy, b u t these a r e less e x t r e m e than those of mayflies. B u r r o w e r s tend to be short a n d b r o a d , b o t t o m sprawlers flat­ tened a n d long-legged (fig. 6.2b), a n d •wimmers slender a n d s t r e a m l i n e d . ProbaWy only a b o u t 15 p e r c e n t of t h e i m m a t u r e s Of the Latin A m e r i c a n species a r e k n o w n .

T h e literature o n Latin A m e r i c a n o d o ­ nates is reviewed by Dias d o s Santos (1981) a n d Paulson (1977, 1982). A species list is p r o v i d e d by Davies a n d Tobin ( 1 9 8 4 - 8 5 ) . References DAVIES, D. A. L., AND P. TOBIN. 1984-85. T h e

dragonflies of the world: A systematic list of the extant species of Odonata. 2 vols. Int. Soc. Odonatologia, Utrect. DÍAS DOS SANTOS, N. 1981. Odonata. In S. H.

Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 64-85. MORTON, E. S. 1977. Ecology and behavior of some Panamanian Odonata. Entomol. Soc. Wash. Proc. 79: 273. PAULSON, D. R. 1977. Odonata. In S. H.

Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 170-184. PAULSON, D. R. 1982. Odonata. In S. H.

Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 249-277.

DRAGONFLIES Anisoptera. Spanish: Zurcidores, caballitos del diablo, caballitos d e San P e d r o , matacaballos, libélulas, aguacilas (General); chispiaguas (Colombia); matapiojos (Chile). Portuguese: Cávalos d e cao, lavandeiras, lavabundas, pitos, cavalinhos d e j u d e u (Brazil). Dragonflies (Corbet 1962, 1980) a r e highly opportunistic p r e d a t o r s that h u n t singly o r sometimes in aggregations, taking o t h e r aquatic insects flying over still water p o n d s , m a r s h e s , a n d t h e m a r g i n s of lakes a n d streams (Young 1980). T h e i r aerial acrobat­ ics a r e spectacular; they c a n hover, d a r t forward instantly, a n d even fly backward with c o n s u m m a t e ease a n d speed, feats rivaled by few o t h e r aerial c r e a t u r e s . I n m a n y species, most of their daylight h o u r s of activity a r e spent o n t h e wing, a n d so highly modified is t h e body for flying that

DRAGONFLIES

197

they c o m e to rest only occasionally to p e r c h a n d to s p e n d t h e night in sleep. T h e legs are useless for walking, f o r m i n g instead a basketlike aerial sieve t o "strain" insects o u t of the air. Dragonflies usually oviposit while they a r e flying, d i p p i n g the tip of the a b d o m e n below t h e water's surface as t h e eggs a r e e x t r u d e d . T h e s u b m e r g e d eggs t h e n drift to the b o t t o m . S o m e h a v e m o r e specialized habits, including d r o p p i n g t h e eggs in t h e water from s o m e h e i g h t , o r directly plac­ ing t h e m o n t o leaves, rocks, o r m u d , o r inserting t h e m into p l a n t tissue (Paulson 1969). Dias dos Santos (1981) r e c o r d s 705 spe­ cies of dragonflies in 9 9 g e n e r a from t h e Neotropics, b u t m a n y u n d e s c r i b e d species a r e certain to exist. Most b e l o n g to t h e family Libellulidae. A few taxa in t h e s o u t h e r n parts of S o u t h A m e r i c a , such as Phenes raptor (Petaluridae), Gomphomacromia chilensis ( C o r d u l i i d a e ) , a n d t h e family Petaliidae, h a v e t h e i r closest relatives in o t h e r s o u t h e r n c o n t i n e n t s . Dragonflies a r e mainly tropical, however, a n d t h e g r o u p t e n d s to b e poorly r e p r e s e n t e d in t h e h i g h e r latitudes of Chile a n d A r g e n t i n a . T h e s t r o n g flight capabilities of t h e adults, s o m e of which a r e migratory, often carry t h e m g r e a t distances. A few species a r e c o m m o n l y f o u n d far o u t at sea, a n d some have colonized isolated oceanic islands. T h e cosmopolitan globetrotter (Pantala flavesceris) (fig. 6.2a), which is capable of flying for h o u r s at a t i m e , h a s b e e n col­ lected o n Coco Island a n d t h e mid-Atlantic Ilha T r i n d a d e . S o m e Latin A m e r i c a n dragonflies a r e distinctive a n d w i d e s p r e a d . T h e following e x a m p l e s all b e l o n g t o t h e family Li­ bellulidae. T h e r u b y tail {Libellula hercúlea) is a m e d i u m - s i z e d species (WS 8 cm) that stands o u t conspicuously against the vegeta­ tion o n which it is o f t e n seen resting because of its brilliant, almost glowing, scarlet a b d o m e n . T h e rest of t h e b o d y is

198

AQUATIC ORDERS

black. T h e sides of the t h o r a x a r e covered with a gray pruinosity. T h e ubiquitous f e r r u g i n o u s s k i m m e r (Orthemis ferruginea) is a dusky r e d - b o d i e d , m o d e r a t e l y large (WS 88 m m ) species. T h e " a m b e r wings" (Perithemis) (fig 6.2d) a r e small (WS 5 c m ) a n d h a v e par­ tially o r completely a m b e r - t i n t e d wings. T h e r e m a y also b e b r o w n cross b a n d s or spots o n t h e wings. Similar in size a r e the "black wings" (Diastatops; fig. 6.2c) that have solid black wings a n d a b r i g h t to dull red abdomen. A n o t h e r conspicuous g r o u p of small dragonflies a r e t h e "butterfly dragonflies" (Zenithoptera). T h e s e r e s e m b l e t h e black wings in size (WS 4 - 5 cm) a n d obscure wings, but t h e r e a r e clear streaks intruding from the a n t e r i o r wing m a r g i n j u s t beyond t h e halfway point which r u n transversely into t h e d a r k field. Also, it is t h e habit of the butterfly dragonflies t o slowly open a n d close t h e wings while p e r c h e d , in the m a n n e r of a s u n n i n g butterfly. References CORBET, P. S. 1962. A biology of dragonflies. Quadrangle, Chicago. CORBET, P. S. 1980. Biology of Odonata. Ann. Rev. Entomol. 25: 189-217. DÍAS DOS SANTOS, N. 1981. Odonata. In S. H.

Hurlbert, G. Rodriguez, and N. Dios dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 6 4 - 8 5 . PAULSON, D. R. 1969. Oviposition in the tropical dragonfly genus Micrathyria (Odonata, Li­ bellulidae). Tombo 12: 12-16. YOUNG, A. M. 1980. Observations on feeding aggregations of Orthemis ferruginea (Fabricius) in Costa Rica (Anisoptera: Libellulidae). Odonatologica 9: 325-328.

DAMSELFLIES Zygoptera. Spanish: Doncellas (General), chupajeringas (Peru).

Figure 6.3 DAMSELFLIES. (a) Ruby wing (Hetaerina sp., Calopterygidae), male, (b) Ruby wing (Hetaerina americana) nymph, (c) Giant damselfly (Magaloprepus coerulatus, Pseudostigmatidae). (d) Giant damselfly (Mecistogaster sp.), wing tips of male, (e) Common damselfly (Argia vivida, Coenagrionidae) nymph. very close to t h e habitats of their n y m p h s , flitting from p e r c h to p e r c h in search of small insects, mosquitoes, a n d o t h e r flies on which t o feed. T h e i r bodies a r e often multicolored, b u t t h e wings, with a few exceptions, a r e entirely t r a n s p a r e n t . T h e r e a r e 7 8 6 species of damselflies in 94 genera r e c o r d e d for t h e Neotropics by Dias dos Santos (1981). T h e actual n u m b e r will certainly g o h i g h e r after the faunas of Amazonia a n d o t h e r r e m o t e tropical areas are m o r e fully e x p l o r e d . T h e s e species belong to a d o z e n families, the largest a n d most c o m m o n by far b e i n g t h e Coena­ grionidae, d o m i n a n t g e n e r a of which a r e Argia (fig. 6.3e), Acanthagrion, a n d Telchasis. Hetaerina (Calopterygidae) is also c o m m o n and easily r e c o g n i z e d by brilliant, d e e p r e d wing bases ("ruby spots") in t h e male (fig. 6.3a, b) ( E b e r h a r d 1986, G a r r i s o n 1990, Williamson 1923). Four families a r e exclu­ sive to Latin A m e r i c a (Pseudostigmatidae, Polythoridae, Perilestidae, a n d Heliocharitidae). T h e r e a r e n o relictual taxa with amphinotic o r G o n d w a n a l a n d distribu­ tions, t h e P a t a g o n i a n species having m o r e northern relatives.

References DÍAS DOS SANTOS, N. 1981. Odonata. In S. H.

Damselflies a r e m u c h smaller a n d weaker fliers t h a n dragonflies. T h e y always stay

Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South

America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 64-85. EBERHARD, W. G. 1986. Behavioral ecology of the tropical damselfly Hetaerina macropus Selys (Zygoptera: Calopterygidae). Odonato­ logica 15: 51-60. GARRISON, R. W. 1990. A synopsis of the genus Hetaerina with descriptions of four new spe­ cies (Odonata: Calopterygidae). Amer. Ento­ mol. Soc. Trans. 116: 175-259. WILLIAMSON, E. B. 1923. Notes on the habitats of some tropical species of Hetaerina (Odo­ nata). Mus. Zool. Univ. Michigan Occ. Pap. 130: 1-45. Giant Damselflies Pseudostigmatidae, Megaloprepus a n d Mecistogaster. Spanish: H e l i c ó p t e r o s , chinchilejos (Peru). M e m b e r s of the small family Pseudostigma­ tidae inhabit m a t u r e forests from n o r t h e r n Mexico to s o u t h e r n Brazil. M e m b e r s of the g e n e r a Megaloprepus a n d Mecistogaster a r e very large a n d showy insects. T h e sight of o n e of these gossamer c r e a t u r e s slowly flying t h r o u g h the trees, its wings tracing a b l u r r e d whirl above its body, is o n e of the loveliest of t h e rain forest. S o m e natives believe t h e m to b e h u m a n spirits that have recently b e c o m e d i s e m b o d i e d a n d insist that they b e left u n m o l e s t e d (Klots a n d Klots 1959). T h e s e a r e e n o r m o u s d a m ­ selflies with s o m b e r b o d y colors b u t g a u d y

DAMSELFLIES

199

p a t t e r n s o n t h e wing tips. Megaloprepus contains only o n e species, M. coerulatus (fig. 6.3c), which h a s a g r e a t e r w i n g s p a n ( 6 0 - 7 5 m m ) t h a n most species of Mecistogaster (usually 6 0 - 6 5 m m , b u t u p to 170 m m ) b u t a s h o r t e r a b d o m e n (76—96 m m c o m p a r e d to u p to 135 m m ) . T h e wings of Megalo­ prepus a r e m a r k e d subapically with a b r o a d , d a r k p u r p l e b a n d a n d a r e suffused basally with white (in t h e males), while in Mecistogaster, o r a n g e , yellow, o r r e d color fields a r e p r e s e n t n e a r t h e wing tips (fig. 6.3d). W i n g coloration sometimes varies within a species d u e to a g e o r seasonal differences. Mecistogaster often h a s n o light-colored spot, b u t all t h e m e m b e r s of t h e family a r e c h a r a c t e r i z e d , without e x c e p ­ tion, by a multicelled p t e r o s t i g m a . Size has b e e n f o u n d to vary greatly in t h e adults owing to differences in t h e quality of t h e n y m p h a l habitats (Fincke 1984). T h e n y m p h s a r e a m o n g t h e few Zygoptera that e m p l o y b r o m e l i a d tanks a n d tree holes for d e v e l o p m e n t . Y o u n g (1981) has observed Megaloprepus ovipositing in a n accumulation of water in a d e p r e s s i o n in the t r u n k of a fallen t r e e . Female Mecisto­ gaster a r e k n o w n to t h r o w t h e i r eggs singly o n t o t h e water surface in such c o n t a i n e r habitats. T h e very long, s l e n d e r a b d o m e n a p p e a r s to b e a n a d a p t a t i o n to facilitate this p r o c e d u r e ( M a c h a d o a n d Martinez 1982). For food, t h e a d u l t s specialize o n kleptoparasitic s p i d e r s t h a t live in t h e webs of o r b weavers ( A r a n e i d a e such as Nephila a n d Gasteracantha) (Young 1980). I n flight, they a p p r o a c h t h e w e b a n d flutter b e f o r e it briefly p r i o r to d a r t i n g in to pluck o u t t h e prey, which they carry t o a n e a r b y p e r c h to devour. Males of t h e sexually d i m o r p h i c Megalo­ prepus hold m a t i n g territories a r o u n d water-filled tree holes for u p to two m o n t h s , d e f e n d i n g t h e m from conspecific males a n d p e r m i t t i n g only females with w h o m they h a v e m a t e d to e n t e r a n d oviposit in t h e holes (Fincke 1984).

200

AQUATIC ORDERS

References FINCKE, O. M. 1984. Giant damselnies in a tropical forest: Reproductive biology 0 f Megaloprepus coerulatus with notes on Mecisto­ gaster (Zygoptera: Pseudostigmatidae). Adv Odonatol. 2: 13-27. KLOTS, A. B., AND E. B. KLOTS. 1959. Living

insects of the world. Doubleday, Garden City. MACHADO, A. B. M., AND A. MARTÍNEZ.

1982.

Oviposition by egg-throwing in a zygopteran, Mecistogaster jocaste Hagen, 1869 (Pseudostigmatidae). Odonatologica 11: 15-22. YOUNG, A. M. 1980. Feeding and oviposition in the giant tropical damselfly Megaloprepus coerulatus (Drury) in Costa Rica. Biotropica 12: 237-239. YOUNG, A. M. 1981. Notes on the oviposition microhabitat of the giant tropical damselfly Megaloprepus coerulatus (Drury) (Zygoptera: Pseudostigmatidae). Tombo 23: 17-21.

STONEFLIES Plecoptera. T h i s is a small o r d e r of very ancient and primitive insects whose n y m p h s a r e typical inhabitants of streams. T h e adults remain n e a r t h e n y m p h a l habitat a n d a r e fre­ quently f o u n d resting on rocks o r boulders in m i d s t r e a m (hence their c o m m o n name) or on nearby vegetation a n d tree trunks. A d u l t stoneflies a r e soft bodied, small to medium-sized (most with length to tips of folded wings 1—2 cm, b u t s o m e to 4 - 5 cm), a n d r a t h e r elongate in overall form. Ex­ cept for a few with stubby o r n o wings, all possess two pairs of c o m p l e t e wings with a fairly complex venation: t h e fore wing is long, with parallel sides; t h e h i n d wing has a large, fanlike area posteriorly. Stoneflies are drably colored in b r o w n s a n d grays, except for some S o u t h A m e r i c a n Eustheniidae that have wings splashed with bright reds a n d yellows. T h e a n t e n n a e a n d cerci (when present) a r e b o t h filiform, the f o r m e r m u c h l o n g e r than t h e latter. Mouthparts a r e m a n d i b u l a t e b u t with weak ele­ m e n t s . Between t h e two s u b o r d e r s , the Arctoperlaria a n d A n t a r c t o p e r l a r i a , there is a basic difference in food habits corre-

Figure 6.4 DOBSONFLIES AND STONEFLIES. (a) Dobsonfly (Corydalus cornutus, Corydalidae). (b) Dobsonfly (Corydalus cornutus) larva, (c) Stonefly {Anacroneuria sp., Perlidae). (d) Stonefly [Anacroneuria sp.) nymph. lated with two m o u t h p a r t types. In t h e Arctoperlaria, t h e lobes of t h e labium a r e long a n d flexible a n d well s t r u c t u r e d for carnivory; in t h e A n t a r c t o p e r l a r i a , these lobes a r e r e d u c e d , relatively inflexible, a n d used for c h e w i n g plant tissues. ( T h e suborders Filipalpia a n d Setipalpia, used widely in earlier literature, a r e n o l o n g e r recognized as phylogenetically logical sub­ divisions of t h e order.) T h e i m m a t u r e s r e s e m b l e t h e adults, except that they lack wings (fig. 6.4d). They may h a v e c o n s p i c u o u s e x t e r n a l fila­ mentous gill tufts ventrally on various parts of t h e body, vestiges of which r e m a i n in adults a n d a r e of t a x o n o m i c utility. Nymphs of t h e South A m e r i c a n Pelurgoperla have long, sticky dorsal hairs in which debris becomes e n t a n g l e d , r e n d e r i n g t h e m inconspicuous amid t h e b o t t o m trash of their habitat. Except for a few limited studies (lilies 1964), t h e biology of Latin A m e r i c a n stone­ flies is not well investigated (Hynes 1976). Most species b r e e d only in cool, r u n n i n g water or cold m o u n t a i n lakes. Adults a r e sluggish a n d often difficult to see w h e n they are at rest o n their similarly colored sub­ strata. T h e y fly readily b u t slowly a n d feebly. T h e Latin A m e r i c a n stonefly f a u n a is fairly extensive a n d diverse ( B e n e d e t t o 1974). T h e r e a r e m o r e t h a n 170 species in 46 genera d i s t r i b u t e d a m o n g 7 families (Hlies 1966, Zwick 1973). Because of their considerable geologic a g e a n d significant

fossil r e c o r d , t h e o r d e r offers a wealth of facts for phylogenetic a n d z o o g e o g r a p h i c analysis o n a worldwide basis (lilies 1965). T h e m o r e primitive A n t a r c t o p e r l a r i a a r e mainly austral a n d exhibit an a m p h i n o t i c distribution, b e i n g f o u n d in s o u t h e r n South America as well as in Australia a n d New Zealand (Eustheniidae, G r i p o p t e r y gidae, A u s t r o p e r l i d a e ) . D i a m p h i p n o i d a e is restricted entirely to t h e s o u t h e r n A n d e s . T h e arctoperlarian Neonemura illiesi (Noton e m o u r i d a e ) is also s o u t h e r n , b u t its n e a r ­ est relatives a r e distributed m u c h like t h e p r e c e d i n g family a n d in S o u t h Africa ( G o n d w a n a l a n d distribution). T h e s e pat­ t e r n s indicate t h e origin a n d early diversifi­ cation of t h e Plecoptera in G o n d w a n a l a n d . A few boreal m e m b e r s of this s u b o r d e r have invaded Latin America, except t h e West Indies, by way of a s o u t h w a r d disper­ sion from N o r t h America. T h e s e a r e cer­ tain Perlidae, including t h e d o m i n a n t g e ­ nus Anacroneuria (74 species) (fig. 6.4c), which has m o v e d over t h e e n t i r e c o n t i n e n ­ tal area, a n d t h e g e n u s Amphinemura ( N e m o u r i d a e ) , which e x t e n d s only to cen­ tral Mexico. G e n e r a l information a n d lit­ e r a t u r e on t h e o r d e r in Latin A m e r i c a can be f o u n d in B a u m a n n (1982), lilies (1977), and Froehlich(1981). References BAUMANN, R. W. 1982. Plecoptera. In S. H.

Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 278-279.

STONEFLIES

201

BENEDETTO, L. 1974. Clave para la determina­ ción de los Plecópteros sudamericanos. Stud. Neotrop. Fauna9: 141-170. FROEHLICH, C. G. 1981. Plecoptera. In S. H.

Hurlbert, G. Rodríguez, and N. Días dos Santos, eds., Aquatic biota of tropical South America. San Diego State Univ., San Diego. Pp. 86-88. HYNES, H. B. N. 1976. Biology of the Plecop­ tera. Ann. Rev. Entomol. 21: 135-153. ILLIES, J. 1964. T h e invertebrate fauna of the Huallaga, a Peruvian tributary of the Amazon River, from the sources down to Tingo Maria. Int. Ver. Limnol. Verh. 15: 1077-1083. ILLIES, J. 1965. Phylogeny and zoogeography of the Plecoptera. Ann. Rev. Entomol. 10: 117-140. ILLIES, J. 1966. Katalog der rezenten Plecop­ tera. Das Tierreich 82: i-xxx, 1—632. ILLIES, J. 1977. Plecoptera. In S. H. Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 185-186. ZWICK, P. 1973. lnsecta: Plecoptera. Phylogenetisches System und Katalog. Das Tier­ reich 94: i—xxxii, 1—465.

DOBSONFLIES M e g a l o p t e r a . Spanish: P e r r o s del a g u a (larvae, Mexico). T h i s is a small o r d e r , related to t h e N e u r o p t e r a (Penny 1977, 1983). T h e r e a r e two families, t h e large C o r y d a l i d a e (BL 8— 12 cm, w i n g s p a n to 16 cm) a n d t h e m u c h smaller Sialidae (BL to 2 5 m m ) . T h e Cory­ dalidae ("dobsonflies") a r e w i d e s p r e a d a n d d o m i n a n t in m a i n l a n d Latin A m e r i c a (only a single Antillean species o n Dominica), with 7 g e n e r a a n d 47 species. Protochauliodes a n d Archichaulides a r e f o u n d in s o u t h ­ e r n Chile a n d have t h e i r c o n g e n e r s in Australia. T h e N o r t h A m e r i c a n Corydalus cornutus (fig. 6.4a) is k n o w n as far south as P a n a m a , from w h e r e a second, closely related species, C. armatus, c o n t i n u e s into S o u t h America. T h e Sialidae ("alderflies") a r e r e p r e s e n t e d by j u s t 7 localized species in a single g e n u s a n d a r e n o t discussed further here. Like

202

neuropterans,

dobsonflies

AQUATIC ORDERS

have

two pairs of m e m b r a n o u s wings with com­ plete venation, a l t h o u g h t h e multiplicity of s p u r i o u s veinlets a n d cross veins f o u n d in the N e u r o p t e r a is lacking. Except for mi­ n o r differences in t h e b r a n c h i n g of veins a n d being slightly narrower, t h e fore wings are similar to t h e h i n d wings. T h e body is cylindrical, elongate, a n d s o m e w h a t soft a n d flexible. Legs a r e short a n d similar on each thoracic segment. I n b o t h t h e adults a n d larvae, t h e m o u t h p a r t s a r e m a n d i b u late a n d with elements, especially t h e m a n ­ dibles, strongly d e v e l o p e d . Males of some species of Corydalus have extremely long, tusklike jaws (Glorioso 1981). All dob­ sonflies, except t h e l e m o n yellow Chloronia (Penny a n d Flint 1982), a r e b r o w n to dull gray or black. Males of Platyneuromus (= Doeringia) have a flattened lateral expan­ sion on t h e h e a d b e h i n d t h e c o m p o u n d eyes. T h e larvae (hellgrammites) (fig. 6.4b) a r e similar in body s t r u c t u r e to t h e adults b u t a r e slightly flattened a n d have con­ spicuous fingerlike gills laterally o n the a b d o m e n . T h e s e gills may be b a r e or fringed, a n d t h e r e may be a tuft of acces­ sory gill filaments at t h e base of most of the p r i m a r y a p p e n d a g e s . T h e last abdominal s e g m e n t bears a pair of large prolegs, each with a lateral filament a n d large claws at the tips. Dobsonflies a r e ubiquitous aquatic in­ sects, generally associated with clean, cold, m o u n t a i n streams. T h e adults r e m a i n near such streams a n d a r e rarely seen except w h e n they c o m e to lights at n i g h t o r when flying, as they d o occasionally o n cool, overcast days o r at dusk. Females attach their eggs in large, single-layered clusters to objects n e a r o r o v e r h a n g i n g t h e water. T h e s e white, flattened masses a r e some­ times conspicuous o n d a r k - c o l o r e d boul­ d e r s o r tree t r u n k s . T h e larvae a r e active p r e d a t o r s , catching a n d eating a variety of o t h e r aquatic insects that they find u n d e r a n d a r o u n d debris a n d rocks. T h e y leave the water to p u p a t e . P u p a e have free,

muscled legs a n d a r e able to walk a n d use their m a n d i b l e s to bite in defense. In t h e Peruvian A m a z o n region, dobsonfly larvae, d e v o u r e d raw or cooked, a r e collected by t h e natives for food ( M u r p h y pers. c o m m . ) . Called perros de agua, they a r e considered v e n o m o u s a n d m u c h feared in parts of Mexico. T h e l i t e r a t u r e of t h e Latin A m e r i c a n m e m b e r s of this o r d e r is reviewed by Flint (1977) a n d P e n n y (1981, 1982).

References FLINT, JR., O. S. 1977. Neuroptera. In S. H.

Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 187-188. GLORIOSO, M. J. 1981. Systematics of the dob-

sonfly subfamily Corydalinae (Megaloptera: Corydalidae). Syst. Entomol. 6: 253-290. PENNY, N. D. 1977. Lista de Megaloptera, Neuroptera e Raphidioptera do México, América Central, ilhas Caraibas e América do sul. Acta Amazónica 7(4) suppl.: 1-61. PENNY, N. D.

1981. Neuroptera. In

S. H.

Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 8 9 - 9 1 . PENNY, N. D.

1982. Neuroptera. In

S. H.

Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 280-282. PENNY, N. D. 1983. Neuroptera of the Amazon Basin. Pt. 7. Corydalidae. Acta Amazónica 12: 825-837.

a n d a r e virtually terrestrial ( H o l z e n t h a l 1988). T h e i m m a t u r e s a r e s o m e w h a t similar to l e p i d o p t e r o u s larvae b u t with t h e a b d o m i ­ nal prolegs restricted to t h e t e r m i n a l seg­ m e n t a n d b e a r i n g anal claws. All types p r o d u c e silk a n d a r e free-living, m a k e fixed retreats, o r construct p o r t a b l e cases for themselves. T h e r e t r e a t m a k e r s use silk for fabricating shelters a n d f o o d - t r a p p i n g nets. T h e case m a k e r s use silk as a m a t r i x to bind t o g e t h e r t h e material c o m p o s i n g their cases, such as leaf f r a g m e n t s , twigs, a n d sand grains. S o m e cases a r e con­ structed entirely of silk. A l t h o u g h some case types a r e very characteristic, usually of g e n e r a , o t h e r s , such as t h e c o m m o n t a p e r ­ ing, cylindrical, sand grain case, a r e m a d e by a n u m b e r of species in a variety of g e n e r a a n d families. Larvae of t h e leptocerid g e n u s Triplectides will pick u p a n d use e m p t y cases of o t h e r caddisflies o r m a k e their own by t u n n e l i n g a small stick. Most cases a r e tubular, b u t o t h e r s may have t h e form of a snail shell (Helicopsyche) o r may be somewhat flattened, four sided, o r purselike (Hydroptilidae). Dense masses of the long, tusk-shaped cases of Atanatolica a n d Grumichella (Leptoceridae) a n d Grumicha (Sericostomatidae) a r e often seen clinging to rock faces in waterfalls. T h e s e cases a r e m a d e entirely of silk o r have fine sand i n c o r p o r a t e d in their walls (Holzen­ thal 1988, Müller 1880).

PENNY, N. D., AND O. S. FLINT. 1982. A revision

of the genus Chloronia (Neuroptera: Corydali­ dae). Smithsonian Contrib. Zool. 348: 1—27.

CADDISFLIES Trichoptera. Caddisflies p o p u l a t e all types of freshwater habitats in Latin America, both in r u n n i n g waters (springs, s t r e a m s , waterfalls, seeps, rivers) a n d s t a n d i n g waters (lakes, p o n d s , marshes, pools). S o m e Phylloicus d e v e l o p in the leaf axils of b r o m e l i a d s . A few Atanatolica move o u t to t h e moist m a r g i n s of seeps

T h e p a r t of t h e larva p r o t e c t e d by t h e case is generally weakly sclerotized in case m a k e r s . T h e free-living types a r e well p i g m e n t e d a n d thick s k i n n e d t h r o u g h o u t . A m o n g still water species, t h e larval food consists primarily of algae, fungi, a n d decaying organic m a t t e r a n d occasionally living plant tissue. S o m e free-living larvae, especially those in fast water, a r e p r e d a ceous on o t h e r aquatic invertebrates. Adult caddisflies a r e mothlike, with body a n d wings clothed in short, easily d e t a c h e d , hairlike scales. At rest, they hold their fore wings rooflike over t h e body at a

CADDISFLIES

203

steep angle. T h e a n t e n n a e a r e usually very long a n d filamentous. T h i s stage lives n e a r t h e larval habitats, a n d t h e d i u r n a l species are often seen resting on rocks a n d vegeta­ tion by t h e water's e d g e . Many of t h e n o c t u r n a l species a r e attracted to artificial light. T h e o r d e r is well studied generally, b u t very little ecological o r n a t u r a l history i n f o r m a t i o n is available specifically on Neotropical r e p r e s e n t a t i v e s (Flint 1977, 1981; B u e n o - S o r i a a n d Santiago Fragoso 1982). McElravy a n d o t h e r s ( 1 9 8 1 , 1982) c o m p a r e d t h e diversity of species at a n o n s e a s o n a l site in P a n a m a to t h a t of Nearctic s t r e a m s a n d f o u n d it n o t signifi­ cantly h i g h e r b u t with relatively less varia­ tion over long time p e r i o d s . T h e o r d e r ' s t a x o n o m y in t h e region is likewise very i n c o m p l e t e . A c c o r d i n g to Flint (pers. c o m m . ) , t h e r e a r e probably between 3,000 a n d 4,000 species, a l t h o u g h only a b o u t 1,200 to 1,500 a r e n o w d e ­ scribed (Fischer 1 9 6 0 - 1 9 7 3 ) . Flint (1983) also p r o v i d e s a key t o t h e S o u t h A m e r i c a n families that is of g e n e r a l use for all of Latin A m e r i c a . T h r o u g h t h e lowlands, t h e most c o m m o n species a r e f o u n d in t h e families L e p t o c e r i d a e (e.g., t h e black-andwhite speckled Nectopsyche; fig. 6.5d, e) a n d H y d r o p s y c h i d a e (e.g., pale g r e e n Lepto­ nema albovirens; fig. 6.5a, b). In P a t a g o n i a n habitats, t h e r e a r e m a n y L i m n e p h i l i d a e (especially Dicosmoecinae). T h e d o m i n a n t

free-living, actively predatory, Holarctic g e n u s , Rhyacophila, is absent a n d replaced by t h e H y d r o b i o s i d a e (e.g., Atopsyche cal­ losa; fig. 6.5c). T h e family A n a m o l o psychidae is restricted to t h e far s o u t h e r n regions, as a r e t h e H e l i c o p h i d a e , Kokiriidae, P h i l o r h e i t h r i d a e , Stenopsychidae, Tasimiidae, a n d a n u m b e r of g e n e r a in o t h e r families (e.g., L e p t o c e r i d a e : Hudsonema, Notalina; Holzenthal 1986a, 19866). All show a distinct relationship to taxa in Australia a n d New Zealand. Lepidostomatidae a r e p r e s e n t in m o n t a n e areas of Mexico a n d Central A m e r i c a b u t absent from S o u t h America. In parts of Brazil, t h e larval cases (called curubixá o r grumixá) a r e a d m i r e d by the I n d i a n s as artworks of n a t u r e a n d used to a d o r n clothing a n d t h e body. S o m e water­ courses a r e even given these n a m e s be­ cause of t h e c o m m o n o c c u r r e n c e of caddis flies in t h e m ( I h e r i n g 1968: 328).

Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 221-226. FLINT, JR., O. S. 1983. Studies of Neotropical caddisflies. XXXIII. New species from aus­ tral South America (Trichoptera). Smithso­ nian Contrib. Zool. 377: 1-100. FISCHER, F. C. J. 1960-1973. Trichopterorum catalogus. 15 vols. Nederlandsche Entomol. Ver, Amsterdam. HOLZENTHAL, R. W. 1986a. Studies in Neo­ tropical Leptoceridae (Trichoptera). VI. Im­ mature states of Hudsonema flaminii (Navas) and the evolution and historical biogeography of Hudsonemini (Triplectidinae). Entomol. Soc. Wash. Proc. 88: 268-279. HOLZENTHAL, R. W. 19866. T h e Neotropical species of Notalina, a southern group of long-horned caddisflies (Trichoptera: Lepto­ ceridae). Syst. Entomol. 11: 6 1 - 7 3 . HOLZENTHAL, R. W. 1988. Studies in Neo­

tropical Leptoceridae (Trichoptera). VIII. The genera Atanatolica Mosely and Grumichella Müller (Triplectidinae: Grumichellini. Amer. Entomol. Soc. Trans. 114: 71 — 128. IHERING, R. VON. 1968. DICIONÁRIO DOS ANIMÁIS DO BRASIL. E D . UNIV. BRASILIA, SAO PAULO. MCELRAVV, E. P., V. H. RESH, H. WOLDA, AND O.

S. FLINT, J R 1981. Diversity of adult Trichop­ tera in a "non-seasonal" tropical environment. 3d Int. Symp. Trichoptera, Ser. Entomol., Proc. 20: 149-156. MCELRAVV, E. P., H. WOLDA, AND V. H. RESH.

1982. Seasonality and annual variability of caddisfly adults (Trichoptera) in a "nonseasonal" tropical environment. Arch. Hydrobiol. 94: 302-317. MÜLLER, F. 1880. Sobre as casas construidas pelas larvas de insetos Trichoptera da Provin­ cia de Santa Catharina. Mus. Nac. Rio de Janeiro Arch. 3: 99-134, 210-214, pis. 8 - 1 1 .

References BUENO-SORIA, J . , AND S. SANTIAGO FRAGOSO.

1982. Trichoptera. In S. H. Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West In­ dies. San Diego State Univ., San Diego. Pp. 398-400. FLINT, JR., O. S. 1977. Trichoptera. In S. H.

Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 249-253. FLINT, JR., O. S. 1981. Trichoptera. In S. H.

Hurlbert, G. Rodriguez, and N. Dias dos

Figure 6.5 CADDISFLIES. (a) Hydropsychid caddisfly (Leptonema sp., Hydropsychidae) larva. (b) Hydropsychid caddisfly (Leptonema albovirens). (c) Hydrobiosid caddisfly (Atopsyche callosa, Hydrobiosidae) larva, (d) Leptocerid caddisfly (Nectopsyche sp., Leptoceridae), larva in case. (e) Leptocerid caddisfly (Nectopsyche punctata).

204

AQUATIC ORDERS

CADDISFLIES

205

nonparasitic

í

ECTOPARASITIC ORDERS

E x t e r n a l parasitism is a pervasive a d a p t a ­ tion d e t e r m i n i n g s o m e major evolutionary lines; all m e m b e r s of t h r e e e n t i r e o r d e r s of insects a r e obligatory ectoparasites of verte­ b r a t e s (Askew 1 9 7 1 , Marshall 1981). T h e y live o n t h e host superficially, sometimes b u r r o w i n g into its skin o r p e n e t r a t i n g e x t e r i o r cavities b u t completely tied to it (Nelson et al. 1975). Such is t h e way of life of t h e M a l l o p h a g a (chewing lice), A n o p l u r a (sucking lice), a n d S i p h o n a p t e r a (fleas). T h i s c h a p t e r treats these o r d e r s i n d e p e n d e n t l y , s e p a r a t e from those fami­ lies a n d lesser a g g r e g a t i o n s of ectoparasitic insects a m o n g o t h e r o r d e r s which a r e dis­ cussed in their places n e x t to their freeliving relatives (see scabies mite, chiggers, etc., c h a p . 4 ; b e d b u g s , b a t b e d b u g s , etc., c h a p . 8; parasitic r o v e beetles, c h a p . 9; bat tick flies, louse flies, c h a p . 11). (For p h y l o g e n e t i c r e a s o n s , t h e louse o r d e r s a r e c o m b i n e d by s o m e a u t h o r s into o n e , t h e P h t h i r a p t e r a , o r e v e n j o i n e d with t h e Psocoptera; they a r e t r e a t e d h e r e as sepa­ rate, t h e m o s t familiar a r r a n g e m e n t . ) All these ectoparasites exhibit similar a d a p t i v e s t r u c t u r e s a n d ecological strate­ gies to life o n t h e skin a n d a m o n g t h e pelage o r p l u m a g e of v e r t e b r a t e animals. Most obvious is a flattening of t h e body, e i t h e r laterally (compression), as in t h e case of fleas, o r dorsoventrally ( d e p r e s ­ sion), as in lice, b e d b u g s , bat b e d b u g s , a n d o t h e r s . T h i s c o n d i t i o n gives t h e insect a m i n i m a l profile a n d s t r e a m l i n i n g to h i d e a n d m o v e over t h e b o d y u n h i n d e r e d by hairs o r feathers. C o r r e l a t e d with this gen­

206

eral body r e m o d e l i n g , in m a n y cases, is a r e d u c t i o n or c o m p l e t e loss of wings and eyes. Linear sets (combs) of flat structures that function like t h e barbs of an arrow­ h e a d , allowing rapid forward p r o g r e s s be­ tween obstructions b u t inhibiting motion backward as m i g h t be caused by preening by t h e host (Marshall 1980), a r e d e novo d e v e l o p m e n t s in these g r o u p s . Most also have holdfast devices of some kind, includ­ ing heavy g r a s p i n g clawlike tarsi, sticky secretions, o r even suction cups. T h e m o u t h p a r t s may b e a r r e c u r v e d hooks or spines to a n c h o r t h e m into t h e host during feeding. T h e food of ectoparasites consists of t h e blood, l y m p h , skin, hair, a n d other skin p r o d u c t s of t h e host. Because of t h e close a t t a c h m e n t of most ectoparasites to their host, their lack of wings, a n d scant mobility in most cases, dispersion is accomplished primarily by direct body contact b e t w e e n hosts. Phoresy is also occasionally practiced by several g r o u p s , especially by Mallophaga. They affix themselves by their m a n d i b l e s to the wing veins o r hairs of o t h e r winged insects (most often biting flies a n d louse flies) that are visiting their hosts a n d t h u s become t r a n s p o r t e d to a new host. Ectoparasites evidently all have evolved from free-living ancestors, as indicated by their body s t r u c t u r e a n d host associations. T h e sclerites of t h e adult flea thorax, for e x a m p l e , still b e a r t h e muscles a n d scle­ rotic braces of a winged insect, although in a vestigial a n d nonfunctional state. T h e larvae of fleas resemble those of primitive

D i p t e r a f o u n d in t h e nests of vertebrates. It is easy to i m a g i n e how a d e p e n d e n c e o n a w a r m - b l o o d e d host could have d e v e l o p e d from living in such close­ ness, followed by structural a d a p t a t i o n s . T h e lowland ectoparasite f a u n a of Mid­ dle a n d S o u t h A m e r i c a is r e m a r k a b l y uniform in t a x o n o m i c composition. M u c h greater e n d e m i s m is e x p e r i e n c e d by high­ land g r o u p s , except those using cosmopoli­ tan domestic animals as hosts. Locally domesticated hosts, however, may h a r b o r endemic species (Escalante 1981). Species of both categories a r e strongly isolated ecologically f r o m t h e i r N o r t h A m e r i c a n counterparts (Wenzel 1972) a n d show in­ d e p e n d e n t evolution. T h e systematics of ectoparasites h a s b e e n used to p r o v i d e clues to t h e relationships of host taxa. T h e study of p a t t e r n s of infestation with lice tends to confirm m o d e r n views on t h e affinities of S o u t h A m e r i c a n land m a m ­ mals (Vanzolini a n d G u i m a r á e s 1955). I m p o r t a n t regional studies of ectopara­ site faunas h a v e b e e n c o n d u c t e d for Pan­ ama (Wenzel a n d T i p t o n 1966) a n d Vene­ zuela (various a u t h o r s 1972, 1 9 7 5 - 1 9 7 6 ) .

References ASKEW, R. R. 1971. Parasitic insects. American Elsevier, New York. ESCALANTE, J. A. 1981. Ectoparásitos de ani­ males domésticos en el Cusco. Rev. Peruana Entomol. 24: 123-125. MARSHALL, A. G. 1980. The function of combs in ectoparasitic insects. In R. Traub and H. Starke, Fleas. Balkema, Rotterdam. Pp. 79-87. MARSHALL,

A.

G.

1981. T h e

ecology

of

ectoparasitic insects. Academic, London. NELSON, W. A., J. E. KEIRANS, J. F. BELL, A N D

C. M. CLIFFORD. 1975. Host-ectoparasite rela­ tionships.]. Med. Entomol. 12: 143-166. VANZOLINI, P., AND L. GUIMARÁES. 1955. Lice

and the history of South American land mammals. Rev. Brasil. Entomol. 3: 13-45. VARIOUS AUTHORS. 1972, 1975-76. Ectopara­

sites of Venezuela. A series of articles on ectoparasite groups published by Brigham Young Univ., Sci. Bull., Biol. Ser. Vols. 17 (1972) and 20 (1975-76).

WENZEL, R. L. 1972. Some observations on the zoogeography of Middle and South Ameri­ can ectoparasites. J. Med. Entomol. 9: 589. WENZEL, R. L., AND V J. TIPTON, eds.

1966.

Ectoparasites of Panama. Field Mus. Nat. Hist., Chicago.

CHEWING LICE Mallophaga. Feather lice, bird lice. As its n a m e implies, this o r d e r is character­ ized by having m a n d i b u l a t e m o u t h p a r t s . Chewing lice c o n s u m e feathers, hairs, a n d o t h e r c u t a n e o u s material, including blood, if accessible (dried from w o u n d s , etc.), a n d sebaceous secretions. T h e s e a r e m o r e heav­ ily sclerotized t h a n t h e sucking lice, with well-defined a b d o m i n a l sclerites a n d com­ paratively rigid body. I n t h e majority of species, t h e h e a d is relatively large, wider t h a n t h e p r o t h o r a x , a n d freely movable. Both of t h e two s u b o r d e r s a r e f o u n d in Latin America: t h e Ischnocera have fili­ form a n t e n n a e , e x p o s e d o n t h e sides of t h e h e a d , a n d vertically biting m a n d i b l e s b u t lack maxillary palpi; t h e Amblycera (Clay 1970) have short a n t e n n a e concealed in pockets o n t h e u n d e r s i d e s of t h e h e a d , mandibles that work laterally, a n d max­ illary palpi. Both bird- a n d mammal-infesting spe­ cies a r e f o u n d a m o n g t h e two g r o u p s . T h e y a r e often very host-specific, espe­ cially t h e Ischnocera, some even b e i n g confined to a particular area of t h e body of birds. A few hosts s u p p o r t a diversity of lice. T i n a m o u s , for e x a m p l e , carry n o less t h a n twelve m a l l o p h a g a n g e n e r a ( C a r r i k e r 1 9 5 3 - 1 9 6 2 ) . N o n e live o n bats, m a r i n e m a m m a l s , o r l a g o m o r p h s . T h e world's seven widespread families of Mallophaga, plus two e n d e m i c families, A b r o c o m o p h a g i d a e o n t h e r a t chinchilla ( E m e r s o n a n d Price 1976) a n d T r o c h i l i p h a g i d a e o n h u m m i n g b i r d s , a r e r e p r e s e n t e d in Latin America. Mammal-associated Amblycera are almost entirely confined to marsupials

CHEWING LICE

207

Figure 7.1 CHEWING LICE, (a) Cat louse (Felicola felis, Trichodectidae). (b) Bird louse (Paragoniocotes mirabilis, Philopteridae). (c) Oval guinea pig louse (Gyropus ovalis, Gyropidae). (d) Giant bird louse (Laemobothrion opisthocomi, Laemobothriidae). (e) Fowl louse (Menacanthus stramineus, Menoponidae). a n d r o d e n t s in t h e N e o t r o p i c s ; those of t h e Ischnocera infest placental m a m m a l s . A m o n g t h e I s c h n o c e r a , t h e Philopteri­ d a e is t h e largest family, with diverse spe­ cies o n birds, i n c l u d i n g t h e characteristicNeotropical p a r r o t s a n d macaws (Paragoniocotes, fig. 7.1b) a n d o t h e r s . M a m m a l s are t h e hosts of T r i c h o d e c t i d a e , i n c l u d i n g Felicola o n felines (fig. 7.1a) a n d Geomydoecus o n fossorial r o d e n t s (Werneck 1945). Sloths, m o n k e y s , kinkajous, coatis, a n d o t h e r uniquely N e o t r o p i c a l m a m m a l s have their o w n c h e w i n g lice as well (Lymeon, Cebidicola, Trichodectes, a n d Neotrichodectes, respectively). To d a t e , M a l l o p h a g a have not b e e n f o u n d o n t h e tapir, c a p y b a r a , anteater, or a r m a d i l l o . F r o m t h e Amblycera, t h e G y r o p i d a e a r e well d e v e l o p e d in Latin A m e r i c a ; t h e r e a r e several small g e n e r a o n wild pigs a n d r o d e n t s . Gyropus ovalis (fig. 7.1c) a n d Gliricola porcelli a r e t h e most c o m m o n of several g u i n e a pig lice. Bird lice of t h e family M e n o p o n i d a e a r e d i v e r s e in species a n d habits. Piagetiella bursaepelecani lives in t h e t h r o a t p o u c h of pelicans. L a e m o b o t h r i i d a e a r e typical of water b i r d s a n d birds of prey. A primitive species, Laemobothrion opistho­ comi (fig. 7 . I d ) , parasitizes t h e likewise primitive hoatzin. T h e Ricinidae parasitize s o n g b i r d s . T h e T r i m e n o p o n i d a e live o n marsupials a n d r o d e n t s . A few i n t r o d u c e d , c o s m o p o l i t a n species a r e pests of d o m e s t i c a n i m a l s . T h e s e in­ clude mainly t h e fowl lice, Menacanthus

208

ECTOPARASITIC ORDERS

stramineus (fig. 7.1e) a n d Menopon gallinae, (Ancona 19356); t h e ox, goat, a n d d o n k e y lice, Bovicola; a n d t h e pigeon louse, Columbicola columbae (Ancona 1935a). T h e latter affect their hosts only w h e n very n u m e r ­ ous, w h e n they cause aggravation from their persistent p r e s e n c e as well as skin irritation from feeding. Aside from host associations, little is k n o w n of t h e biology of chewing lice in Latin America. Many passerine bird hosts have b e e n observed i n d u l g i n g furiously in the habit of "anting" with m e m b e r s of the ant subfamily Formicinae. T h e y squat near an ant nest a n d passively allow t h e ants to crawl o n t o their p l u m a g e or place them t h e r e with their beaks. Agitation from p r e e n i n g m o v e m e n t s causes t h e ants to release formic acid vapors that apparently act as a repellent to any c h e w i n g lice present.

ANCONA, L. 19356. Contribución al conoci­ miento de los piojos de los animales de México. II. Menopon gallinae Linn. Inst. Biol. Univ. Nac. Aut. México Anal. 6: 53-62. £ AR RIKER, JR., M. A. 1953-62. Studies in Neotropical Mallophaga. XII. Lice of the tinamous. Pts. 1-2. Rev. Bras. Biol. 13: 2 0 9 224, 324-346; pts. 3 - 4 , Bol. Entomol. Venezolana 11: 3-30, 9 7 - 1 3 1 ; pts. 5 - 7 , Rev. Brasil. Biol. 2 1 : 205-216, 325-338, 3 7 3 384; 22: 433-448 (1962). CLAY, T. 1970. T h e Amblycera (Phthiraptera: Insecta). Brit. Mus. Nat. Hist. (Entomol.) Bull. 25: 7 3 - 9 8 . EMERSON, K. C , ed. 1967. Carriker on Mallo­ phaga. Posthumous papers, catalog of forms described as new, and bibliography. U.S. Nati. Mus. Bull. 248: 1-150. EMERSON, K. C , AND R. D. PRICE. 1976. Abro-

comophagidae (Mallophaga: Amblycera), a new family from Chile. Fla. Entomol. 59: 425-428. VON KÉLER, S. 1960. Bibliographic der Mallophagen. Zool. Mus. Berlin Mitt. 36: 146-403. WERNECK, F. L. 1945. Os tricodectideos dos

roedores (Mallophaga). Inst. Oswaldo Cruz Mem. 4 2 : 8 5 - 1 5 0 . WERNECK, F. L. 1948. Os malófagos de mamí­ feros. 2 vols. Ed. Inst. Oswaldo Cruz, Rio de Janeiro.

SUCKING LICE Anoplura. M a m m a l lice. Sucking lice r e s e m b l e biting lice b u t a r e immediately d i s t i n g u i s h e d by modification of the m o u t h p a r t s into stylets for sucking blood from t h e i r m a m m a l hosts. N o n e a r e

f o u n d o n birds. I n general, they also a r e less well sclerotized t h a n o t h e r lice, t h e a b d o ­ m e n having a n elastic cuticle, with small, dorsal a b d o m i n a l sclerites, allowing for con­ siderable distension d u r i n g feeding. Hosts include most major g r o u p s of placental m a m m a l s , excepting, most nota­ bly, t h e bats, anteaters, a n d aquatic g r o u p s , a l t h o u g h some highly specialized forms infest m a r i n e p i n n i p e d s . T h e latter a r e a d a p t e d for resisting cold a n d s u b m e r s i o n in water by a d e n s e covering of h y d r o phobic, scalelike bristles a n d spiracular valves for closing off t h e tracheal system w h e n the host dives. Host specificity is less rigid t h a n a m o n g t h e Mallophaga. Latin A m e r i c a n species n u m b e r a p p r o x i ­ mately 4 3 , classified in only a few of t h e 15 world families (Kim a n d L u d w i g 1978). Two a r e major pests of t h e h u m a n body, the h u m a n louse a n d crab louse (see b e ­ low). O t h e r s a r e w i d e s p r e a d forms that infest domestic animals, such as Haemato­ pinus, especially t h e h o g louse (Haemato­ pinus suis, fig. 7.2a); five species of Solenopotes which live o n cattle ( a n o t h e r o n d e e r ) ; andLinognathuspeddalis (Linognathidae), which parasitizes s h e e p in S o u t h America (Kim a n d Weisser 1974). Micro­ thoracicus mazzai (fig. 7.2c), M. praelongic.eps, a n d M. minor a r e f o u n d o n t h e llama; t h e last species is also a parasite of t h e alpaca, while t h e second also occurs o n t h e vicuña (Ferris 1951).

Worldwide, t h e n u m b e r of m a l l o p h a g a n species described exceeds 5,000, which is probably less t h a n 10 p e r c e n t of t h e species that await discovery. I n Latin America, t h e r e a r e several t h o u s a n d species to be n a m e d . I m p o r t a n t literature in t h e study of these lice is p r o v i d e d by E m e r s o n (1967), Werneck (1948), a n d v o n Kéler (1960).

References ANCONA, L. 1935a. Contribución al conoci­ miento de los piojos de los animales de México. I. Columbicola columbae. Inst. Biol. Univ. Nac. Auc. México Anal. 5: 342-351.

*

'

Rgure 7.2 SUCKING LICE, (a) Hog louse (Haematopinus suis, Haematopinidae). (b) Wild pig louse (Pecaroecus javalii, Haematoponidae). (c) Llama louse (Microthoracicus mazzai, Linognathidae). (d) Human louse (Pediculus humanus, Pediculidae). (e) Crab louse (Phthirus pubis, Pediculidae) and egg (nit).

SUCKING LICE

209

T h e r e m a i n d e r a r e f o u n d o n wild m a m m a l s of m a n y sorts. Pecaroecus (Haem a t o p i n i d a e , fig. 7.2b) lives o n peccaries (Babcock a n d Ewing 1938). T h e g e n e r a Hoplopleura a n d Polyplax ( H o p l o p l e u r i d a e ) are widespread on rodents.

References BABCOCK, O. G., AND H. E. EWING. 1938. A

new

genus and species of Anoplura from the pec­ cary. Entomol. Soc. Wash. Proc. 40: 197-210. FERRIS, G. F. 1951. The sucking lice. Pacific Goast Entomol. Soc. Mem. 1: 1—320. KIM, K.

C.,

AND H.

W.

LUDWIG.

1978.

The

family classification of the Anoplura. Syst. Entomol. 3: 249-252. KIM, K. C., AND C. F. WEISSER. 1974. Taxon­

omy of Solenopotes Enderlein, 1904, with redescription of Linognathus panamensis Ew­ ing (Linognathidae: Anoplura). Parasitology 69: 107-135.

PRIMATE LICE Human Louse Pediculidae, Pediculus humanus. Spanish: Piojo, l i e n d r e s (eggs). Náhuatl: Atémitl. Quechua: Usa. Portuguese: Piolho, piolho l a d r o , l é n d e a s (eggs). Tupi-Guaraní: M u q u i r a n a . Nits (eggs). T h e h u m a n louse ( B u x t o n 1947) has b e e n an intimate c o m p a n i o n of h u m a n s as long as we have existed, a n d they may have lived o n o u r s u b h u m a n p r o g e n i t o r s . Ewing (1926) recognized distinct subspecies of h u m a n louse (not n o w recognized, see below), associated with t h e N e g r o i d , Caucasoid, Asian, a n d A m e r i c a n I n d i a n host races. T h e last w e r e described from speci­ m e n s taken from the scalps of 4,000-yearold, p r e - C o l u m b i a n m u m m i e s f o u n d in Peru (see also Weiss 1932). T h e s e origi­ nally distinct types h a v e f o r m e d hybrids with t h e c o n t e m p o r a r y m e l d i n g of h u m a n ­ ity a n d a r e n o l o n g e r recognizable, b u t their existence indicates that the species is to be c o n s i d e r e d i n d i g e n o u s to all areas of h u m a n occupation.

210

ECTOPARASITIG ORDERS

T h e h u m a n louse is well established ¡ n Latin A m e r i c a n history ( H o e p p l i 1969). A louse plays a p a r t in the Popol-Vuh, the very famous book of Mayan mythology T h e story tells of I x m u c a n é , a goddess in the s h a p e of an old w o m a n w h o sends an i m p o r t a n t message by a louse to h e r grand­ sons. Chronicles of the Mexican conquest r e f e r r e d to bags of lice in the treasure houses of Moctezuma, b u t these may have been stocks of d r i e d cochineal bugs. Oviedo, speaking of the h o u s e h o l d of the Aztec e m p e r o r , m e n t i o n s priests who were wearing long hair full of lice that they were catching a n d eating while m u r m u r i n g prayers. Actual lice w e r e collected by the Inca r u l e r s of Peru as a tax that sufficed from the p o o r a n d destitute (de la Vega 1979). A curious belief persisted among early travelers to t h e New World with r e g a r d to lice. It was t h o u g h t that lice d i s a p p e a r e d a b o u t the l o n g i t u d e of the Azores on the voyage west a n d r e a p p e a r e d on the r e t u r n to E u r o p e ( H o g u e 1981). Two h u m a n louse subspecies a r e distin­ guished, based on the t e n d e n c y of popula­ tions to segregate either o n t h e head ("head lice," Pediculus humanus capitis) or body ("body lice," P. h. corporis). T h e y are scarsely distinguishable structurally, they hybridize readily, a n d m a n y even cross into opposite habitats a n d show only some be­ havioral differences so that they a r e obvi­ ously variants of o n e species. T h e species (fig. 7.2d) is small (BL 2 - 3 m m ) a n d gray to b r o w n ; b l o o d e d individu­ als usually show a large, d a r k globule in the a b d o m e n r e p r e s e n t i n g the coagulated last meal. T h e flattened body is elongate, the a b d o m e n elliptical in outline. T h e anten­ n a e are clearly visible projecting from the anterolateral m a r g i n s of the spherical h e a d . T h e single s h a r p pointed claws at the tips of the stout legs a r e also conspicuous. Infestations of t h e h u m a n louse (pedicu­ losis) are c o m m o n t h r o u g h o u t the world a n d r e m a i n an accepted way of life among peasants a n d indigents as well as peoples

whose c u l t u r e s d o n o t e m p h a s i z e bodily cleanliness. People normally feel e x t r e m e discomfort w h e n lice a r e biting, a n d the skin becomes sensitive to c o n t i n u e d feed­ ing» d e v e l o p i n g r e d p a p u l e s a n d rashes and possibly b e c o m i n g h a r d e n e d a n d deeply p i g m e n t e d ("vagabond's disease"). It is still a c o m m o n sight a m o n g I n d i a n s and in the A n d e s to see m o t h e r s g r o o m i n g the hair of t h e i r children a n d r e m o v i n g lice and nits a n d usually e a t i n g t h e m . Many other ways of killing or d i s c o u r a g i n g lice are practiced, using plant extracts (Lenko and P a p a v e r o 1979: 120f.). T h e y a r e also used in m a k i n g r e m e d i e s . T h e h u m a n louse is a vector of e p i d e m i c typhus {tabardillo in Mexico; classical ty­ phus), a very serious disease f o u n d t h r o u g h ­ out the world c a u s e d by Rickettsia prowazekii. In Latin A m e r i c a , it is most prevalent in colder climates w h e r e the b o d y louse, t h e primary carrier, is p r e s e n t . It is a povertyassociated disease a n d also a c o m m o n result of social d i s o r d e r d u r i n g wars or following catastrophe. It c u r r e n t l y has r e c e d e d but remains a potential t h r e a t to h u m a n health. This species of Pediculus is restricted to Homo sapiens. M e m b e r s of the s u b g e n u s Parapediculus live on New World spider monkeys (Áteles). H u m a n lice are distin­ guished by h a v i n g flat lateral plates on the abdomen instead of lobed as in the m o n ­ key lice a n d h a v e m u c h heavier setae a n d a more weakly sclerotized cuticle. H u m a n lice a r e t r a n s m i t t e d by close body contact, a l t h o u g h they cling to cloth­ ing and may leave the body a n d r a n g e over bedding a n d t h u s gain access to new indi­ viduals. Infestations are usually heaviest when p e r s o n s a r e c r o w d e d into sleeping quarters a n d w h e r e b a t h i n g a n d cleaning facilities are i n a d e q u a t e . In tropical cli­ mates, body lice t e n d to disappear, proba­ bly because of their sensitivity to high skin temperatures, b u t h e a d lice r e m a i n . T h e life cycle is simple. After h a t c h i n g , the n y m p h s begin sucking blood at once and feed frequently t h r o u g h o u t life, both

night a n d day. T h e y r e a c h m a t u r i t y after t h r e e molts in 16 to 18 days. Females m a t e a n d begin to attach their eggs to hairs a n d clothing a day or two after r e a c h i n g adult­ h o o d . If unfed, these lice very soon die.

References BUXTON, P. A. 1947. The louse: An account of the lice which infest man, their medical impor­ tance and control. 2d ed. Arnold, London. DE LA VEGA, G. 1979. The Incas: The royal commentaries of the Inca Garcilaso de la Vega. Translated by M. Jolas. Lib. A B C , Lima. EWING, H. E. 1926. A revision of the American lice of the genus Pediculus, together with a consideration of the significance of their geographical and host distribution. U.S. Nati. Mus. Proc. 68: 1-30. HOEPPLI, R. 1969. Parasitic diseases in Africa and the Western Hemisphere, early documen­ tation and transmission by the slave trade. Acta Trop. suppl. 10: 1-240. HOGUE, C. L. 1981. Commentaries in cultural entomology. 2. The myth of the louse line. Entomol. News 92: 53-55. LENKO, K., AND N. PAPAVERO. 1979.

Insetos no

folclore. Conselho Esl. Artes Cien. Hum., Sao Paulo. WEISS, P. 1932. Restos humanos de Cerro Colo­ rado. Rev. Mus. Nac. (Lima) 1(2): 90-102.

Crab Louse Pediculidae, Phthirus pubis. Spanish: Ladilla (General). Portuguese: C h a t o (Brazil). Pubic louse. T h i s bloodsucking louse (Payot 1920) in­ habits the h u m a n b o d y a l o n g with the h u m a n louse b u t seldom strays from the pubic region. It is distinct in form, crablike, with a short b r o a d body a n d very stout laterally projecting legs (fig. 7.2e). It is smaller than the h u m a n louse (BL 1.5—2 m m ) b u t about the same color. A l t h o u g h it infests t h e genital area pri­ marily, the c r a b louse occasionally migrates to the a r m p i t s or o t h e r hairy p a r t s of the b o d y — t h e eyelids, b e a r d , eyebrows. Indi­ viduals t e n d to be stationary, r e m a i n i n g attached for days at o n e p o i n t with the m o u t h p a r t s inserted in the skin. Contin-

PR1MATELICE

211

u e d defecation by t h e louse d u r i n g this time results in t h e a c c u m u l a t i o n of excre­ tory material a r o u n d t h e site. T h e life cycle (Nutall 1918) r e q u i r e s a p p r o x i m a t e l y a m o n t h . T h e female d e p o s ­ its eggs o n t h e c o a r s e r hairs of t h e host. N y m p h s a n d adults feed almost c o n t i n u ­ ously, causing skin rashes a n d intense itch­ ing. Transmission is by physical contact, most often sexual i n t e r c o u r s e , b u t t h e louse may s p r e a d on c o n t a m i n a t e d towels, blankets, a n d clothing.

References NUTALL, G. H. F. 1918. T h e biology of Phthirus pubis. Parasitology 10: 383-405. PAYOT, F. 1920. Contribution á l'étude du Phthirus pubis (Linné, Leach). Soc. Vaud. Sci Nat Bull. 53: 127-161.

FLEAS S i p h o n a p t e r a . Spanish a n d Portuguese: Pulgas. Náhuatl: T e c p i n t i n , sing, tecpin (Mexico). Quechua: P i q u e k u n a . Fleas a r e all too familiar insects. Several species infest pets a n d domestic animals a n d bite h u m a n s . As b l o o d s u c k e r s , they t r a n s m i t diseases, a m o n g t h e m o r e serious, m u r i n e t y p h u s (Rickettsia typhi) a n d p l a g u e (Yersinia pestis) (see Medical Entomology, c h a p . 3). T h e s e a r e small (BL 1—6 m m ) , d a r k b r o w n insects with sclerous bodies that a r e wingless a n d strongly c o m p r e s s e d laterally. T h e h e a d projects f o r w a r d as a r o u n d e d shield a n d bears short, t h r e e - s e g m e n t e d , clubbed a n t e n n a e a n d s o m e t i m e s d a r k ocu­ lar spots. T h e h e a d , t h o r a x , a n d a n t e r i o r a b d o m i n a l s e g m e n t s m a y also have rows of stiff, flat, scalelike bristles (ctenidia) that aid in mobility t h r o u g h hairs o r feathers. T h e legs a n d body h a v e p r o m i n e n t setae. T h e m o u t h p a r t s a r e e l o n g a t e d a n d su­ p r e m e l y a d a p t e d for b l o o d s u c k i n g with t h r e e saberlike stylets a n d associated palpi a n d c h a n n e l e d labium. T h e legs, especially

212

ECTOPARASITIC ORDERS

the posterior pair, a r e m u c h larger a n d powerfully constructed a n d give t h e fleas incredible leaping capabilities (Rothschild e t a l . 1973). Fleas a r e usually confined to animals that customarily b u r r o w a n d have a d e n , g r o u n d nest, or habitual resting place. Insectivores a n d r o d e n t s a r e c o m m o n hosts, but fleas a r e ectoparasites of owls, swallows, domestic fowl, r o d e n t s , bats, a n d m a n y o t h e r m a m m a l s (except aquatic ones). Host specificity varies, some flea species being catholic in their tastes, others highly restricted to o n e host species (da Costa Lima a n d H a t h a w a y 1946). Unlike t h e o t h e r ectoparasite o r d e r s discussed in this chapter, fleas have com­ plete m e t a m o r p h o s i s . T h e larval stage lives in the soil a n d o r g a n i c debris that collects in the host's nest, feeding on cast-off blood, feces, a n d scurf from t h e animals living t h e r e . After attaining full growth, the larva (fig. 7.3b) spins a cocoon in which it p u p a t e s (fig. 7.3e). T h e p u p a is inactive a n d passes a d e v e l o p m e n t a l p e r i o d of a few days to several m o n t h s while m e t a m o r ­ p h o s i n g into t h e a d u l t flea. Adults remain on t h e hosts most of t h e time b u t must go elsewhere, especially w h e n hosts leave the nest o r perish. Eggs a r e placed randomly, b u t t h e timing of e g g p r o d u c t i o n is depen­ d e n t on t h e r e p r o d u c t i v e cycle of t h e host. Copulation a n d oviposition occur most often p r i o r to nesting, e g g laying, o r partu­ rition of the vertebrates on which t h e fleas' life d e p e n d s . Blood is t h e only a d u l t food. When passing into the stomach, t h e blood courses t h r o u g h a check valve (proventriculus) lined with h u n d r e d s of promi­ n e n t spines. After t h e flea has engorged, the valve prevents a back flow of fluid into the foregut. T h i s s t r u c t u r e m a y become clogged with p l a g u e bacilli that develop after a blood meal o n an infected rat or h u m a n . T h e o r g a n i s m s a r e regurgitated forward d u r i n g s u b s e q u e n t feeding at­ t e m p t s a n d e n t e r t h e b l o o d s t r e a m of a

—■OB

Figure 7.3 FLEAS, (a) Cat flea {Ctenocephalides felis, Pulicidae). (b) Cat flea larva, (c) Oriental rat flea (Xenopsylla cheopis, Pulicidae). (d) Rodent flea (Dasypsyllus lasius, Dolichopsyllidae). (e) Cat flea, pupa in cocoon, (f) Chigoe (Tunga penetrans, Tungidae), male, (g) Chigoe, gravid female. new individual, w h e r e t h e disease can develop. In Latin A m e r i c a , t h e flea fauna is diverse a n d well d e v e l o p e d , with a total n u m b e r of species probably in excess of five h u n d r e d , b u t flea t a x o n o m y in this part of t h e world is i n c o m p l e t e ( J o h n s o n 1957, T r a u b 1950). T h e flea fauna of South A m e r i c a is notable for its high percentage of e n d e m i c h i g h e r taxa. Some, like the S t e p h a n o c i r c i d a e (helmeted fleas) and Pygiopsyllidae, f o u n d elsewhere only in the Australasian region, suggest that t h e original hosts of fleas w e r e marsupials (Wenzel 1972). O t h e r primarily South American flea families a r e Malacopsyllidae and Rhopalopsyllidae. T h e f o r m e r live on edentates a n d carnivores; they have en­ larged tarsal claws a n d a leathery integu­ ment; t h e latter a r e parasites of r o d e n t s and a variety of o t h e r m a m m a l s . As with o t h e r ectoparasites, fleas may be grouped a c c o r d i n g to those associated with household a n d h u s b a n d e d animals, some­ times secondarily attacking h u m a n s a n d probably mostly i n t r o d u c e d from the Old World, a n d i n d i g e n o u s species on wild mammals a n d wild birds. H u m a n s suffer mostly from t h e species found on animals k e p t in a n d a r o u n d homes, because of their s h a r e d a b o d e . T h e dog flea (Ctenocephalides canis) a n d cat flea (C. felis, fig. 7.3a) a r e p r i m a r y in this r e ­ spect. T h e O r i e n t a l rat flea (Xenopsylla cheopis, fig. 7.3c), a n o t h e r cosmopolitan species, r a n g e s freely from its hosts, often

biting people; it is t h e p r e m i e r vector of plague. O t h e r domestic r o d e n t fleas a r e t h e m o u s e flea (Leptopsylla segnis) a n d t h e n o r t h ­ e r n rat flea (Nosopsyllus fasciatus). T h e h u ­ m a n flea (Pulex irritans) has a wide r a n g e of hosts a m o n g animals whose lives a r e en­ c o u r a g e d by h u m a n activity a n d is a n o t h e r excellent t r a n s m i t t e r of p l a g u e , including two u n u s u a l types of t h e disease f o u n d in Ecuador, vesicular (virola pestosa) a n d tonsillar (angina pestosa) forms. T h e sticktight flea (Echidnophaga galliná­ cea) attaches to u n f e a t h e r e d skin on t h e h e a d of domestic fowl, causing consider­ able agitation. It is a relative of t h e b u r ­ rowing flea (see below) b u t rarely infests humans. Bat fleas form t h e family Ischnopsyllidae a n d mainly associate with insectivo­ r o u s c h i r o p t e r a n species. T h e i r d r o p p i n g s provide a better n u t r i e n t m e d i u m on roost floors t h a n those of the fruit-eating or carnivorous bats. Ceratophyllus (Ceratophyllidae) a n d Dasypsyllus (Dolichopsyllidae, fig. 7.3d) a r e c o m m o n g e n e r a mainly asso­ ciated with wild birds.

References DA COSTA LIMA, A. AND C. R. HATHAWAY.

1946.

Pulgas: Bibliographia, catálogo e animáis por elas sugados. Inst. Ozwaldo Cruz Monogr. 4: 1-522. JOHNSON, P. T. 1957. A classification

of the

Siphonaptera of South America. Entomol. Soc. Wash. Mem. 5: 1-299. ROTHSCHILD, M., Y. SCHLEIN, K. PARKER, C. NEVILLE, AND S. STERNBERG. 1973. T h e

FLEAS

213

flying leap of the flea. Sci. Amer. 229(5): 9 2 100. TRAUB, R. 1950. Siphonaptera from Central America and Mexico. Fieldiana Zool. Mem. 1: 1-127. WENZEL, R. L. 1972. Some observations on the zoogeography of Middle and South Ameri­ can ectoparasites. J. Med. Entomol. 9: 589.

BURROWING FLEA T u n g i d a e , Tunga penetrans. Spanish: Nigua, chique (General). Portuguese: Bicho d e pé, bicho d e p o r c o , j a t e c u b a (Brazil). Indian: Sika, t u n g a (Brazil). French: C h i q u e . C h i g o e , jigger, sand flea. Unlike o t h e r fleas, females of this species work their way into the skin of their hosts a n d encyst, e n l a r g i n g to several times their original size, u p to the size of a small pea ( 3 - 5 m m d i a m e t e r ) (fig. 7.3g). T h e male, which is mobile a n d active t h r o u g h o u t its life, copulates with the female after she has completely p e n e t r a t e d the skin. Females may e n t e r skin a n y w h e r e o n the body b u t most often b u r r o w u n d e r the toenails or into the soles of the feet of h u m a n s w h e r e the p r e s s u r e of their g r o w t h causes g r e a t discomfort. N o d u l a r , ulcerating swellings result. Persons g o i n g b a r e f o o t in infested areas (usually w h e r e pigs, the usual alter­ n a t e hosts, a r e c o m m o n ) a r e likeliest to pick u p the fleas in this way. T h e y are r e m o v e d only precariously with the tip of a sterile n e e d l e ; if the insect's body is r u p ­ t u r e d , infection a n d i m m u n e reactions with serious c o n s e q u e n c e s may follow, in­ c l u d i n g g a n g r e n e a n d loss of the a p p e n d ­ age. Many of t h e early settlers of Brazil lost their feet to this insect in a d r e a d f u l m a n n e r . To be infested is r e f e r r e d to as cambado in that country. T h e original h o m e of this flea a p p e a r s to be A m e r i c a ( H o e p p l i 1969: I69f.). T h i s is based on the e v i d e n c e of m u c h earlier accounts of the species f r o m tropical A m e r ­ ica (dating to 1526 by Oviedo) t h a n from

214

ECTOPARASITIC ORDERS

Africa, w h e r e it is not definitely recorded until the mid-1700s. Also, a n t h r o p o m o r ­ phic clay vessels from p r e - I n c a n Peru show feet unmistakably infested by the flea, A l t h o u g h t h e r e a r e earlier r e p o r t s of this flea in Africa, a k n o w n i n t r o d u c t i o n oc­ c u r r e d in 1872 by a British ship traveling from Rio d e J a n e i r o to Angola, whence it s p r e a d rapidly along the West African coast a n d subsequently disseminated to the interior. T h e early e x p l o r e r s of the Ama­ zon Basin found it a terrible nuisance: "It is a great h a p p i n e s s that its legs have not the elasticity with those of [other] fleas; for could this insect leap, every animal body would be filled with t h e m ; a n d , conse­ quently, both the b r u t e a n d h u m a n species be soon extirpated by the m u l t i t u d e of these insects" (de Ulloa 1758). T h e pest became so well k n o w n to the Brazilians that a body of folklore a n d poetry has evolved a r o u n d t h e m (Lenko a n d Papavero 1979).

nigs, o t h e r d o m e s t i c a n d wild animals serve as hosts (dogs, r o d e n t s , a n d b u r r o w ­ ing owls). All a r e fossorial a n d probably nick u p the fleas from t h e g r o u n d w h e n digging in it. T h e larvae a r e typically flealike in f o r m ; they may h a t c h within the sinus f o r m e d by t h e m o t h e r but usually drop to the g r o u n d to develop. T h e species is tropical, r a n g i n g only a few d e g r e e s n o r t h a n d s o u t h of t h e equator, on the c o n t i n e n t a n d in the West Indies. It may be f o u n d in a b u n d a n c e in dry, sandy places, in a n i m a l enclosures, a n d on ranches a n d farms, especially w h e r e pigs

a r e raised. O t h e r k n o w n species of Tunga a r e of n o medical significance.

References DE ULLOA, A. ] 758. Relación histórica del viaje a la América Meridional. 4 vols. Madrid. HOEPPLI, R. 1969. Parasitic diseases in Africa and the Western Hemisphere, early documen­ tation and transmission by the slave trade. Acta Trop. suppl. 10: 1-240. LENKO, K., AND N. PAPAVERO. 1979. O bicho-do-

pé—nasce fémea, morre macho. In K. Lenko and N. Papavero, Insetos no folclore. Cons. Est. Artes Cien. Hum., Sao Paulo. Pp. 4 7 9 498.

I n d i a n s have effective m e t h o d s for the extraction of encapsulated fleas. T h e most ancient t e c h n i q u e utilized fish spines, wood splinters, a n d plant spines to pry t h e m out physically. Later, p o i n t e d knives a n d nee­ dles were e m p l o y e d . Certain w o m e n and children in the interior of Brazil became a d e p t at r e m o v i n g the fleas a n d received considerable status for this specialized pro­ fession. Various o t h e r r e m e d i e s relying on the application of poultices of all kinds are ineffectual (ibid.). T h e b u r r o w i n g flea is a small (BL 1.0 mm), r e d d i s h - b r o w n flea with a somewhat stubby a p p e a r a n c e from the contracted thoracic segments (fig. 7.3f). T h e large h e a d a n d t h o r a x completely lack combs. T h e outline of the h e a d is angular, a n d the m o u t h p a r t s are overly large a n d stiff, with a f o u r - s e g m e n t e d palpus. T h e stiff elon­ gate mandibles are used to slash o p e n the skin to p e r m i t the e n t r y of the body when the female inserts herself into a new locus. Both males a n d females suck blood, but only the females b u r r o w into skin. Besides

BURROWING FLEA

215

O

In this suborder of bugs, the fore wing is composed of two distinct areas as de­ scribed above. The thickened basal portion may be subdivided further into elongate triangles, an anterior, basal corium, and a small, posterior clavus. The soft, papery membrane makes up a third or more of the apical part of the wing. Veins are apparent in the membrane but rarely in the other areas. Less apparent but highly characteristic of the suborder are repugnatorial or stink glands that are present in most families. In the adult, these open ventrally on the thorax, often through spoutlike processes,

Rgure 8.1 HETEROPTERANS. (a) Ant mimicking bug {Hyalymenus sp., Alydidae). (b) Ant model fofant mimicking bug (Camponotus sp., Formicidae). (c) Giant big-legged bug (Pachylis pharaonis, Coreidae). (d) Leaf-legged bug (Diactor bilineatus, Coreidae). (e) Rice stinkbug (Oebalus poecilus, l*6ntatomidae). (f) Conchuela (Chlorochroa ligata, Pentatomidae).

BUGS

Hemiptera. Spanish: Chinches. Portuguese: Percevejos. In English, the term "bug" has a precise meaning for insects of this order, in addi­ tion to its general application to all kinds of small insectoid creatures. The origin of the word is obscure. Theories suggest that it comes either from the Old English or Welsh bwg, meaning a goblin or ghost, or from the Arabic buk, a longstanding and widespread name for the infamous bedbug, Cimex lectularius (Usinger et al. 1966: 5). True bugs (Weber 1968) are all recogniz­ able by their mouthparts, which are modi­ fied for sucking fluids, such as plant sap, nectar, and insect or vertebrate blood. There are two pairs of sclerotized, flexible stylets (modified mandibles and maxillae) lying in a groove in a one- to foursegmented labium. Together, these struc­ tures form a proboscis, always arising on the front of the head but flexible and when not in use, projecting toward the posterior between the forelegs. There are two suborders, separated most evidently by wing structure. The first is the Heteroptera ("uneven wings"), with fore wing divided midway into two areas of radically different textures: a basal thick and rigid part and an apical papery and flexible portion. This type of wing is re­ ferred to as a hemielytron and is character­ istic of the heteropterans. The other suborder is the Homoptera ("uniform wings"), which have homogeneous, diapha­ nous, or parchmentlike fore wings. Be­

216

Hemiptera, Heteroptera.

near the base of the middle pair of legs. Scent glands also are present in the nymphs but open on the back of the abdomen. Although they may attract oth­ ers of the same species, having aggregating or alarm-calling functions, the products of these glands are usually foul smelling and repugnant. Their chemical composition varies greatly and is complex. In coreids and relatives (Aldrich and Yonke 1975), they contain varied aliphatic acids, alde­ hydes, alcohols, and esters. These chemi­ cals readily volatilize into the environment and seem to have a repellent effect on other insects and animals. They may also be rubbed or squirted on an assailant to burn the skin or other vulnerable tissues. In Trinidad and the Guianas, the "pepper flies" (Amnestus) may accidentally enter the eye, where they liberate a strong, caustic secretion that causes an agonizing, persis­ tent burning comparable to that of cay­ enne pepper (Myers 1934). Assassin bugs possess formidable bites in which they inject toxins in their saliva that can cause humans excruciating pain (Ryckman 1979, Ryckman and Bentley 1979). Two major heteropteran groups are medically significant as disease vectors or noxious bloodsuckers: the kissing bugs and the bedbugs. Most heteropterans are phytophagous,

cause it is a widespread habit of the Homoptera to produce waxy secretions from integumentary glands, I use the con­ venient and appropriate common name "wax bugs" here. Wings are partially or totally absent in many members of both suborders, especially among animal and plant ectoparasites or forms with secluded life-styles. References USINGER, R. L., J. CARAYON, N. T. DAVIS, N. UESHIMA, AND H. E. MCKEAN. 1966. Mono­

graph of the Cimicidae. Vol. 7. Entomol. Soc. Amer. (Thomas Say Foundation), College Park, Md. WEBER, H. 1968. Biologie der Hemipteren. Asher, Amsterdam.

HETEROPTERANS

sucking sap and tissue liquids from all parts of plants. Some kinds spend much of their lives exposed on tree trunks where their cryptic colors and rough integuments afford them camouflage against similarly appearing bark. Wetting and darkening of the substratum by rain would destroy this protection, but the cuticle of at least two species of bark bugs (Aradidae) absorb water and undergo corresponding color change, thus preserving this protective device (Silberglied and Aiello 1980). A recurring phenomenon in heterop­ teran families is mimicry of other insects, especially of ants (fig. 8.1a, b). Adults or nymphal stages may be involved. There are numerous examples. A species of Barberiella (Miridae) is a member of a Batesian complex of several different insects and a spider that mimics Camponotus planatus, an aggressive ant in Honduras (Jackson and Drummond 1974). The Brazilian bug Thaumastaneis montandoni (Pyrrhocoridae) (Hussey 1927), many Arhaphe (Largidae), and a Hyalymenus species (Alydidae) (da Costa Lima 1940: 90) have ant-shaped bodies, but the specific models are yet unknown. Some heteropterans are also myrmecophiles, termitophiles, or arachnophiles. In South America, nymphs of Arachnocoris albomaculatus (Nabidae) live in company

HETEROPTERANS

217

with certain spiders a n d generally have t h e a p p e a r a n c e of their host, with long slender legs a n d slightly b u l b o u s a b d o m e n (Myers 1925). A bizarre n y m p h a l mirid (possibly Paracarnus) is f o u n d in C u b a (China 1931). It bears a f o u r - p a r t , p e d u n c l e d p r o t h o r a c i c h o r n , giving it a r e s e m b l a n c e to certain t r e e h o p p e r s (genus Cyphonial), with which it may have s o m e association, possibly as a predator. A l t h o u g h primarily treating t h e Ecua­ d o r i a n fauna, F r o e s c h n e r (1981) provides a useful c o m p e n d i u m of t h e s u b o r d e r for Latin A m e r i c a .

References ALDRICH, J. R., AND T. R. YONKE. 1975. Natural

products of abdominal and metathoracic scent glands of coreoid bugs. Entomol. Soc. Amer. Ann. 68: 955-960. CHINA, W. E. 1931. A remarkable mirid larva from Cuba, apparently belonging to a new species of the genus Paracarnus, Dist. (Hemiptera, Miridae). Ann. Mag. Nat. Hist. (ser. 10) 8: 283-288. DA COSTA LIMA, A. 1940. Hemípteros, insectos

de Brasil. Vol. 4. Escuela Nac. Agron., Ser. Didac, Rio de Janeiro. FROESCHNER, R. C. 1981. Heteroptera or true bugs of Ecuador: A partial catalog. Smithso­ nian Contrib. Zool. 322: 1-147. HUSSEY, R. F. 1927. O n some American Pyrrhocoridae, Brooklyn Entomol. Soc. Bull. 22:227-235. JACKSON, J. F , AND B. A. DRUMMOND 111. 1974.

A Batesian ant-mimicry complex from the mountain pine ridge of British Honduras, with an example of transformational mim­ icry. Amer. Midi. Nat. 91: 248-251. MYERS, J. G. 1925. Biological notes on Arachnocoris albomaculatus Scott (Hemiptera; Nabidae). N.Y. Entomol. Soc. J. 33: 136-146, pi. 1. MYERS, J. G. 1934. Field observations on some Guiana insects of medical and veterinary interest. Trop. Agrie. 11: 279-283. RYCKMAN, R. E. 1979. Host reactions to bug bites (Hemiptera, Homoptera), a literature review and annotated bibliography. Calif. Vect. Views 26(1-2): 1-24. RYCKMAN, R. E., AND D. G. BENTLEY. 1979. Host

reactions to bug bites (Hemiptera, Homop­ tera): A literature review and annotated bibli­ ography. Calif. Vect. Views 26(3-4): 25-30.

218

BUGS

SILBERGLIED, R., AND A. AIELLO. 1980. Camou.

flage by integumentary wetting in bark bugs Science 207: 773-775.

BIG-LEGGED BUGS C o r e i d a e . Squash bugs. Big-legged bugs a r e c o m m o n a n d typical H e t e r o p t e r a , generally similar to stink bugs a n d seed bugs b u t distinguished by the n u m e r o u s c o m p l e x veins in t h e mem­ b r a n o u s portion of t h e fore wing which the o t h e r s lack. A l t h o u g h m a n y have all slen­ d e r legs, t h e h i n d legs a r e often enlarged a n d at times very stout, especially the femur, which may b e greatly swollen; the tibia also is sometimes wide a n d flattened, as in t h e so-called leaf-legged bugs. Be­ cause of this c o m m o n h y p e r t r o p h y of the legs a n d their diverse food, I have coined the c o m m o n n a m e used h e r e which seems m o r e widely applicable a n d descriptive t h a n "squash bugs," better restricted to a few g e n e r a such as Anasa. Most a r e small to medium-sized bugs (BL 1 0 - 1 5 m m ) , b u t some a r e large and robust, such as Pachylis (fig. 8.1c), which may be over 30 millimeters long. They usually a r e dull b r o w n o r g r e e n i s h , but quite a few display r e d o r yellow lines on the fore wings, a n d t h e d o r s u m of the a b d o m e n is often colored brilliantly in red or yellow. T h e s e parts of t h e body are displayed w h e n t h e b u g is molested in what a r e believed to b e w a r n i n g signals of the foul tastes a n d smells m a n y of t h e m pos­ sess. T h e n y m p h s also form aggregations, which e n h a n c e t h e noxious effect. Nymphs of Thasus acutangulus, for e x a m p l e , are vivid o r a n g e , yellow, a n d black a n d group t o g e t h e r o n twigs of Pithecellobium trees in Central America. W h e n a l a r m e d , they pul­ sate violently, spray j e t s of anal fluid, and e x u d e a noxious secretion over t h e back of the a b d o m e n . Paryphes blandus n y m p h s re­ act similarly, giving off a p u n g e n t odor. A few coreids a r e p r e d a c e o u s , b u t most

feed o n plants, many, such as t h e injurious genus Anasa, o n vines of Cucurbitaceae (Brailovsky 1985). A c o m m o n pest of t h e nassion vine (Passiflora) is Diactor bilineatus (fig- 8.Id), a leaf-legged b u g (percevejo do maracuja). It is colorfully m a r k e d with yellow s p o t t e d black leg e x p a n s i o n s a n d a green d o r s u m with a n o r a n g e c h e v r o n ; t h e nymphs a r e bizarre, with black legs a n d wing b u d s , white collar, a n d o r a n g e a b d o ­ men with l a r g e black b u t t o n s . An estimate of t h e n u m b e r of Neo­ tropical coreid species c a n n o t b e m a d e at present; t h e r e a r e at least o n e h u n d r e d genera.

References ALDRICH,

J.

R.,

AND M.

S.

BLUM.

1978.

Aposematic aggregation of a bug (Hemip­ tera: Coreidae): T h e defensive display and formation of aggregations. Biotropica 10: 58-61. BRAILOVSKY, H. 1985. Revisión del género Anasa Amyot-Serville (Hemiptera-Heteroptera-Coreidae-Coreinae-Coreini). lnst. Biol. Univ. Nac. Aut. Mexico Monogr. 2: 1-266. YOUNG, A. M. 1980. Notes on the interaction of the Neotropical bug Paryphes blandus Horvath (Hemiptera: Coreidae) with the vine Anguria warscewiczii Hook F. (Curcurbitaceae). Brenesia 17: 27-42.

STINKBUGS Pentatomidae. Spanish: C o n c h u e l a s (General). Portuguese: Persevejos d o mato, Marias fétidas (Brazil). As their n a m e implies, stinkbugs have welldeveloped a n d very effective r e p u g n a torial glands a n d a r e k n o w n as m u c h by their smell as by their physical attributes. Among t h e latter a r e a broadly oval o r shield-shaped body with a large, t r i a n g u l a r central plate (scutellum) in t h e m i d d l e of the back. T h e lateral c o r n e r s of t h e prothorax a n d posterolateral c o r n e r s of the a b d o m i n a l s e g m e n t s a r e often p r o ­ longed o r s h a r p p o i n t e d . T h e legs a r e always simple a n d similar in size a n d

length. Most a r e plain colored, g r e e n o r b r o w n , b u t some, especially in t h e i m m a ­ t u r e stages, a r e p a i n t e d in b r i g h t h u e s of r e d , blue, a n d yellow (pi. I d ) . Both t h e n y m p h s a n d adults suck s a p a n d m a y be very a b u n d a n t o n their host plants (Young 1984). T h e latter a r e found a m o n g diverse taxa. Because of their h a b ­ its of feeding o n cultivated plants, m a n y species a r e pests. Such is t h e rice stinkbug (Oebalus poecilus; fig. 8.1e), t h e b a n e of rice in many areas (pulga d'anta, chupador, tamanjuá). O t h e r injurious species a r e t h e conchuela (Chlorochroa = Petidia ligata; fig. 8.If) a n d several m e m b e r s of t h e g e n u s Euschistus a n d Mormidea. S o m e , however, a r e considered beneficial because they at­ tack o t h e r insects which may b e e n e m i e s of cultigens. N y m p h s , especially t h e y o u n g e r instars, are often g r e g a r i o u s . Subsocial b e h a v i o r is exhibited by t h e C o l o m b i a n Antiteuchus tripterus (Discocephalinae). Its females g u a r d b o t h their eggs a n d first instar n y m p h s against p r e d a t o r s a n d parasitoid wasps of the family Scelionidae ( E b e r h a r d 1975). Species of P e n t a t o m i d a e (jumiles) a r e sold in t h e Mexican villages of C u a u t l a a n d Taxco for h u m a n c o n s u m p t i o n . M a r k e t e d live in p a p e r cones in h a n d f u l lots, they have a p i q u a n t taste a n d a r e believed to alleviate liver, kidney, a n d stomach ail­ m e n t s w h e n ingested (orig. obs., A n c o n a 1932, 1933). T h e r e a r e m o r e t h a n a t h o u s a n d species of stinkbugs in some ninety g e n e r a in South America alone.

References ANCONA, L. 1932. Los jumiles de Taxco (Gro.), Atizies taxcoensis, spec. nov. Inst. Biol. Univ. Mexico Anal. 4: 149-162. ANCONA, L. 1933. Los jumiles de Cuautla, Euschistus zopilotensis Distant, lnst. Biol. Univ. Mexico Anal. 4: 103-108. EBERHARD, W. G. 1975. T h e ecology and behav­ ior of a subsocial pentatomid bug and two scelionid wasps: Strategy and counter strat­ egy in a host and its parasites. Smithsonian Contrib. Zool. 205: 1-39.

STINKBUGS

219

YOUNG, A. M. 1984. Phenological patterns in reproduction oí Senna fructuosa (Mill.) Irwin and Barneby (Caesalpinaceae) and a pod associate, Pellaea sticta (Dallas) (Heteroptera: Pentatomidae), in Costa Rican tropical rain forest. Kans. Entomol. Soc. J. 57: 413-422.

SEED BUGS Lygaeidae. T h i s family is c o m p o s e d of e l o n g a t e , small to medium-sized b u g s (BL 3 - 1 5 m m ) char­ acterized by a wing m e m b r a n e with only four o r five veins. T h e a n t e n n a e a n d beak are four s e g m e n t e d . Most a r e d r a b , b u t m a n y display r e d s , yellows, a n d o t h e r bright colors t h a t a r e doubtlessly a p o sematic in function (Sillén-Tullberg et al. 1982). P o l y m o r p h i s m in wing size occurs; usually t h e h i n d pair is a t r o p h i e d o r lost, a n d t h e m e m b r a n o u s tip of t h e fore wing may b e r e d u c e d . M e m b e r s of this family feed largely o n the m a t u r e seeds of various plants, a large n u m b e r from figs {Ficus) (Slater 1972). T h e o r a n g e - a n d b l a c k - b a n d e d milkweed bugs, Oncopeltus, form a conspicuous g r o u p (Ojeda 1973). T h e s e infest milk­ weed plants, especially t h e w i d e s p r e a d weed Asclepias curassavica, o n the seed p o d s of which they m a y b e f o u n d in clusters (Root a n d C h a p l i n 1976). T h e s e a n d Lygaeus ( a n d probably others) feed o n m a n y asclepiadaceous plants, which con­ tain cardiac glycosides. T h e bugs probably

sequester these toxic alkaloids for their own protection. Oncopeltus fasciatw (fig 8.2a) is widely c u l t u r e d as a laboratory animal for research in physiology and o t h e r areas (Feir 1974). Several Central A m e r i c a n lygaeids ex­ hibit very s t r o n g resemblances to beetles (coleoptery), with shell-like, coreaceous hemielytra, which meet evenly along the midline (Slater 1985). T h e functional sig­ nificance o f t h e r e s e m b l a n c e is not known. A few species a r e c r o p pests. T h e infamous chinch b u g (Blissus leucopterusfig. 8.2b) causes considerable d a m a g e to rice in Peru. O t h e r s a r e beneficial. T h e big-eyed b u g (Geocoris punctipes) is a n effi­ cient p r e d a t o r of r e d s p i d e r mites on cotton plantations. T h e r e a r e about 4 5 0 species of this family in Latin America (Slater 1964).

References FEIR, D. 1974. Oncopeltus fasciatus: A research animal. Ann. Rev. Entomol. 19: 81-96. OJEDA, D. 1973. Contribución al estudio del género Oncopeltus Stál (Hemiptera: Lygaei­ dae). Rev. Peruana Entomol. 16: 88-94. ROOT, R. B., AND S. J. CHAPLIN. 1976. T h e life­

styles of tropical milkweed bugs, Oncopeltus (Hemiptera: Lygaeidae), utilizing the same hosts. Ecology 57: 132-140. SlLLÉN-TULLBERG,

B . , C . WlKLUND,

AND L.

JARVI. 1982. Aposematic coloration in adults and larvae of Lygaeus equestris and its bearing on Müllerian mimicry: An experimental study on predation on living bugs by the great tit Parus major. Oikos 39: 131-136.

Figure 8.2 HETEROPTERANS. (a) Large milkweed bug (Oncopeltus fasciatus, Lygaeidae). (b) Chinch bug (Blissus leucopterus, Lygaeidae). (c) Tarnished plant bug (Lygus lineolaris, Miridae). (d) Lunate flat bug (Dysodius lunatus, Aradidae). (e) Cotton lace bug (Corythucha gossypii, Tingidae).

220

BUGS

SLATER, J- A.

1964. A catalogue

of the

Lygaeidae of the world. 2 vols. Univ. Conn. Storrs. SLATER, J- A. 1972. Lygaeid bugs (Hemiptera: lygaeidae) as seed predators of figs. Biotropica4: 145-151. SLATER, J- A. 1985. A remarkable new coleopteroid lygaeid from Colombia (Hemiptera: Heteroptera). Int. J. Entomol. 27: 229-234.

PLANT BUGS Miridae. Leaf b u g s . The best-known plant b u g s a r e t h e m a n y species that cause direct h a r m from their feeding. D a m a g e often takes t h e form of rough, h a r d , gall-like lesions (stigmonosis) on the surface o f valued leaves (especially on tobacco), fruit (commonly cacao), o r tender stems. Chief a m o n g t h e offending genera a r e Lygus, Engytatus, a n d Monalonion. A l t h o u g h m e m b e r s of the family a r e widely recognized for their d e p r e d a t i o n s among field c r o p s , m a n y a r e beneficial predators of a p h i d s a n d o t h e r injurious insects. A w i d e s p r e a d c r o p pest is t h e tarnished p l a n t b u g (Lygus lineolaris, fig. 8.2c). For the most p a r t , these a r e small (BL less than 10 m m in most), e l o n g a t e bugs with the hemielytra characteristically " b r o k e n " or angled sharply d o w n w a r d at the base of the m e m b r a n e . A small n o t c h (cunneus) at this point, only o n e o r two cells in t h e veins of the hemielytral m e m b r a n e , a n d lack of ocelli a r e also family characteristics. T h e i r bodies a r e slightly flattened, soft, a n d fre­ quently have brightly colored, elongate stripes or h a r l e q u i n p a t t e r n s . Many species, especially in t h e subfamily Mirinae, a r e ant mimics (see H e t e r o p t e r a n s , above).

Rio de |aneiro Arch., Pt. 1, 44: 1-158; Pt. 2, 45:1-216; Pt. 3, 47: 1-161; Pt. 4, 48: 1-384; Pt. 5,51-194.

LUNATE FLAT BUG A r a d i d a e , Mezirinae, Dysodius

lunatus.

T h e extraordinarily d e p r e s s e d s h a p e of this b u g (fig. 8.2d) a n d its family relatives constitute a n a d a p t a t i o n primarily for life o n o r u n d e r the bark of d e a d trees. T h e s e bugs also possess t r e m e n d o u s l y long (max­ illary) stylets for sucking the juices of fungi that grow well in their habitat. T h e s e structures a r e coiled u p within t h e h e a d when not in use. T h e i n t e g u m e n t is fluted, scored, tuberculate, a n d very coarsely r o u g h e n e d , which, with its s o m b e r colors, gives t h e insect such a resemblance to t h e substrata of lichens, e r o d e d wood, a n d such mottled surfaces that it is n o t easily seen. Speci­ m e n s a r e usually found e x p o s e d o n t h e bark of trees b u t a r e detected with diffi­ culty because of their cryptic f o r m a n d color (Silberglied a n d Aiello 1980). T h i s species is the best-known r e p r e s e n ­ tative of 472 species of A r a d i d a e in t h e Neo­ tropics (Kormilev a n d F r o e s c h n e r 1987, Usinger a n d M a t s u d a 1959). It is very large by family s t a n d a r d s (BL to 15 m m ) a n d has scallops o n the m a r g i n s of the a b d o m e n a n d the sides of the p r o t h o r a x , the latter greatly p r o d u c e d anteriorly as b r o a d , platelike lobes, r o u n d e d o n t h e o u t e r m a r g i n s a n d e x t e n d i n g forward well b e y o n d the level of the eyes. Adults h a v e c o m p l e t e wings.

References KORMILEV, N. A., AND R. C. FROESCHNER. 1987.

Flat bugs of the world: A synonymic list (Heteroptera: Aradidae). Entomography, Sacramento.

This is t h e largest family o f h e t e r o p ­ terans, with several t h o u s a n d species in the Neotropics (Carvalho 1 9 5 7 - 1 9 6 0 ) .

SILBERGLIED, R., AND A. AIELLO. 1980. Camou­

References

USINGER, R. L., AND R. MATSUDA. 1959. Classifi­

CARVALHO, J. C M . 1957-1960. A catalogue of Miridae of the world (Hemiptera). Mus. Nac.

flage by integumentary wetting in bark bugs. Science 207: 773-775. cation of the Aradidae (Hemiptera-Heteroptera). Brit. Mus. Nat. Hist., London.

LUNATE FLAT BUG

221

LACE BUGS

LEONARD, M. D., AND A. S. MILLS. 1931. Obser­

vations on the bean lace bug in Puerto Ri c o Dept. Agrie. Puerto Rico J. 15: 309-323.

Tingidae. In this family ( D r a k e a n d Davis 1960), t h e e n t i r e dorsal surface, i n c l u d i n g t h e wings, has taken o n a n alveolar o r reticulate a p ­ p e a r a n c e . Like tiny p a n e s of glass, t r a n s p a r ­ e n t m e m b r a n e s enclose t h e spaces between a c o m p l e x lacework of cells. T h e s e m a y form inflated sacs o r b r o a d , winged e x p a n ­ sions o n t h e sides a n d r e a r of the p r o t h o r a x , the latter e x t e n d i n g f o r w a r d over t h e h e a d . T h e surface of some m a y b e profusely spined in a d d i t i o n . Because they a r e g e n e r ­ ally small (BL usually less t h a n 5 m m ) , o n e may a p p r e c i a t e this s t r u c t u r e only w h e n t h e b u g is b e n e a t h t h e microscope. Because of its e c o n o m i c i m p o r t a n c e , t h e family h a s received c o n s i d e r a b l e taxonomic a t t e n t i o n (Drake a n d Ruhoff 1960). All of t h e a p p r o x i m a t e l y 615 Latin A m e r i ­ can species ( D r a k e a n d R u h o f f 1965) live o n t h e foliage of trees a n d s h r u b s . O r d i ­ narily, they c o n g r e g a t e o n t h e u n d e r surfaces of leaves, a n d their sap-sucking, especially by m e m b e r s of t h e large g e n u s Corythucha, occasionally causes h a r m to c r o p s such as cotton. C. gossypii (fig. 8.2e) is a widely distributed injurious species with m a n y hosts a m o n g t h e cultivated plants ( L e o n a r d a n d Mills 1931). Blind, beetlelike m e m b e r s of t h e sub­ family V i a n a i d i n a e a r e atypically s m o o t h , only t h e surface b e i n g lightly p u n c t a t e . T h e y live symbiotically with ants u n d e r ­ g r o u n d ( D r a k e a n d Davis 1960). References DRAKE C. J., AND N. T. DAVIS.

1960.

The

morphology, phylogeny, and higher classifica­ tion of the family Tingidae, including the description of a new genus and species of the subfamily Vianaidinae (Hemiptera: Heteroptera). Entomol. Amer. 39: 1-100. DRAKE, C. J., AND R. A. RUHOFF. 1960. Lace-bug

genera of the world (Hemiptera: Tingidae). U.S. Nati. Mus. Bull. 112: 1-105. DRAKE, C. J., AND R. A. RUHOFF. 1965. Lace bugs

of the world: A catalog (Hemiptera: Tingi­ dae). U.S. Nati. Mus. Bull. 213: 1-634.

222

BUGS

ASSASSIN BUGS Reduviidae. T h e s e a r e mostly m e d i u m to large b u g s of varied form b u t always with a narrowly attached, elongate h e a d that bears a flexi­ ble, s e g m e n t e d beak a n d fairly long anten­ nae. A t rest, t h e beak fits neatly into a p r o n o u n c e d furrow o n t h e u n d e r s i d e of the h e a d which h a s microscopically visible cross striations o n its i n n e r surface. In m a n y species, t h e forelegs a r e modified for g r a s p i n g small insect prey. T h e p r o t h o r a x is trapezoidal a n d often b e a r s s h a r p spines at its o u t e r c o r n e r s . T h e a b d o m e n is usu­ ally b r o a d a n d e x p a n d s m a r k e d l y beyond the m a r g i n s of t h e folded wings. Typical e x a m p l e s a r e found in t h e widespread g e n e r a Apio•merus (fig. 8.3a) a n d Arilus. T h e w i d e s p r e a d cogwheel b u g {chinche crestada, Arilus carinatus) is large (BL 20 m m ) and easily recognized by a p r o m i n e n t , vertical, semicircular crest o n t h e back of t h e thorax which is m a r g i n e d with coarse teeth (fig. 8.3b). M e m b e r s of t h e subfamily Emesinae, called t h r e a d - l e g g e d bugs, a r e sticklike (fig. 8.4c) a n d often mistaken for phasmids (Wygodzinsky 1966). S o m e of these are k n o w n to steal prey c a u g h t in spiders' webs. Many assassin bugs a r e brightly colored, with r e d o r yellow legs a n d spots that a p p e a r o n t h e back of t h e a b d o m e n when the wings a r e raised. T h e brightly colored, spiny n y m p h s a r e a c o m m o n sight o n low vegetation in forests. T h e i r brilliant mark­ ings ( p i . 3d) obviously w a r n enemies of their v e n o m o u s a n d painful bite (aposematic). Many such species participate in Müllerian mimicry complexes with other insects. R e m a r k a b l e e x a m p l e s a r e certain Spiniger that closely resemble tarantula

Figure 8.3 ASSASSIN BUGS (REDUVIIDAE). (a) Assassin bug (Apiomerus lanipes). (b) Cog­ wheel bug (Arilus carinatus). (c) Tarantula hawk model for tarantula hawk-mimicking assassin bug (Pepsis sp., Pompilidae). (d) Tarantula hawk-mimicking assassin bug (Spiniger sp.). (e) Trashgathering assassin bug (Salyavata variegata). hawks (spider wasps of t h e g e n u s Pepsis) (fig. 8.3c, d) as well as Hiranetix a n d Graptocleptes, which a r e colored a n d b e h a v e like b a n d - w i n g e d , i c h n e u m o n i d wasps. Notocyrtus vesiculostis mimics t h e stingless bee, Trígona fulviventris, both species hav­ ing a black h e a d , t h o r a x , a n d legs, a n d orange-yellow a b d o m e n (Jackson 1973). Apiomerus pictipes ( J o h n s o n 1983) h a s b e e n observed in Mexico feeding o n Trígona bees, which it resembles a n d which it seems to be attracted by some agent, possibly a chemical l u r e (Weaver et al. 1975). A species with a n o t h e r deceptive h u n t ­ ing strategy is Salyavata variegata (fig. 8.3e), whose n y m p h lives in n a s u t e t e r m i t e nests in Costa Rica. For camouflage, it scratches off bits of nest a n d plasters t h e m to hairlike projections o n its back. It also "fishes" for termites, using t h e e m p t y car­ casses of p r e v i o u s p r e y as a lure. W o r k e r s are attracted t o their o w n d e a d as a source of protein; w h e n a t e r m i t e takes t h e "bait," the b u g d r o p s it a n d grabs t h e fresh p r e y (McMahan, 1982, 1983). Assassin bugs, a s their n a m e suggests, are fiercely p r e d a c e o u s . All feed o n blood from o t h e r insects. ( T h o s e of t h e subfamily Triatominae suck v e r t e b r a t e blood; see Kissing B u g s , below.) T h e y attack merci­ lessly, u s i n g their raptorial forelegs to grasp a n d h o l d their victims while they stab with their m o u t h stylets. Chemicals con­ tained in t h e saliva immobilize their cap­ tives a n d can cause c o n s i d e r a b l e pain w h e n

used as a defensive "sting" against verte­ brates a n d h u m a n s . Because of their insec­ tivorous habits, assassin bugs a r e i m p o r ­ tant controllers of c r o p pest a n d n a t u r a l insect populations. J u s t over a t h o u s a n d species of R e d u ­ viidae a r e r e c o r d e d from Latin A m e r i c a (Wygodzinsky 1949). References JACKSON, J. F. 1973. Mimicry oí Trígona bees by a reduviid (Hemiptera) from British Hondu­ ras. Fla. Entomol. 56: 200-202. JOHNSON, L. K. 1983. Apiomerus pictipes (reduvio, chinche asesina, assassin bug). In D. H. Janzen, ed., Costa Rican natural history. Univ. Chicago Press, Chicago. Pp. 684-687. MCMAHAN, E. A. 1982. Bait-and-capture strat­ egy of a termite-eating assassin bug. Ins. Soc. 29: 346-351. MCMAHAN, E. A. 1983. Bugs angle for termites. Nat. Hist. 92(5): 40-47. WEAVER, E. C , E. T. CLARKE, AND N. WEAVER.

1975. Attractiveness of an assassin bug to sting­ less bees. Kans. Entomol. Soc. J. 48: 17—18. WYGODZINSKY, P. 1949. Elenco sistemático de los Reduviiformes Americanos. Univ. Nac. Tucumán Publ. 473, Monogr. 1: 1-102. WYGODZINSKY, P. 1966. A monograph of the

Emesinae (Reduviidae, Hemiptera). Amer. Mus. Nat. Hist. Bull. 133: 1-614.

Kissing Bugs Reduviidae, T r i a t o m i n a e , Panstrongylus, Triatoma, a n d relatives. Spa?iish: Chirimachas (Peru), vinchucas (Argentina, Chile), pitos (Venezuela), chinches voladoras (Mexico), chinches

ASSASSIN BUGS

223

y u r ú p u c ú (Paraguay). Portuguese: Bichos d e p a r e d e , c h u p a n c a s , p i n c h a d o r e s , b a r b e i r o s , fincoes (Brazil). Conenose bugs. T h e s e R e d u v i i d a e a r e of major medical i m p o r t a n c e in t h e New W o r l d tropics b e ­ cause of t h e role of several species as vectors of C h a g a s ' disease (Zeledón a n d Rabinovich 1981). T h i s is a very debilitat­ ing a n d frequently fatal s y n d r o m e caused by t h e p r o t o z o a n Trypanosoma (Schizotrypanum) cruzi. T h e p a t h o g e n s a r e intro­ d u c e d by t h e b u g w h e n it feeds, n o t directly t h r o u g h t h e bite b u t in liquid fecal material that it habitually releases follow­ ing e n g o r g e m e n t . T h e m i c r o o r g a n i s m s in these d r o p p i n g s e n t e r t h r o u g h t h e p e r f o r a ­ tion m a d e by t h e bite o r t h r o u g h t h e m u c o u s m e m b r a n e s . T h e site of inocula­ tion m a y b e a n y w h e r e o n t h e skin, b u t infection c o m m o n l y occurs at t h e o u t e r c o r n e r of t h e eye. T h e parasites multiply rapidly a n d localize eventually in vital inter­ nal o r g a n s ; after years of c h r o n i c disease, the patient often succumbs. C h a g a s ' dis­ ease is restricted to t h e New World ("Ameri­ can trypanosomiasis"), a n d a l t h o u g h t h e p a t h o g e n occurs in its b u g a n d i n t e r m e d i ­ ate wild m a m m a l hosts over a m u c h wider area, it is p r e v a l e n t principally in d r y areas of m a r g i n a l a g r i c u l t u r e t h r o u g h o u t south­ e r n Mexico, a r o u n d t h e p e r i p h e r y of t h e A m a z o n Basin (but a p p a r e n t l y n o t within, even t h o u g h vectors a r e p r e s e n t ; Barbosa

d e Almeida 1971), a n d across t h e middle of South America. It is estimated that 13 to 14 million p e o p l e in South A m e r i c a suffer from t h e disease at this time. In 1909, Carlos C h a g a s first discovered the flagellate in t h e t r i a t o m i n e b u g , Panstrongylus megistus (fig. 8.4a). Since that time, several o t h e r species in this subfamily of bloodsucking bugs have b e e n incrimi­ nated as transmitters. T h e most effective vector seems to be t h e widely distributed, often domestic, Triatoma infestans (Rabino­ vich 1972). O t h e r i m p o r t a n t g e n e r a l vec­ tors a r e T. dimidiata in C e n t r a l America, Ecuador, a n d Peru a n d Rhodnius pallescens in P a n a m a . Several species of Panstrongylus (Lent a n d J u r b e r g 1975) a n d Triatoma and Rhodnius prolixus (fig. 8.4b) a r e localized vectors in different parts of Latin America (Lent a n d Wygodzinsky 1979: 135f.). Many m o r e triatomines have b e e n f o u n d natu­ rally infected with t h e parasite. Kissing b u g s a r e mostly m e d i u m to fairly large reduviids (BL 10—45 m m ) . T h e h e a d is cylindrical, a b o u t t h r e e times as long as wide, a n d h a s a constricted, cone-shaped a n t e r i o r p o r t i o n from which the t h r e e - s e g m e n t e d beak arises. A mem­ b r a n o u s connection between t h e second a n d third rostral segments, permitting flexure of t h e third s e g m e n t d u r i n g blood­ sucking, is a u n i q u e condition among bugs. T h e a b d o m e n is concave dorsally a n d widely e x p a n d e d laterally. I n many species, these expansions a r e conspicu-

Figure 8.4 HETEROPTERANS. (a) Kissing bug (Panstrongylus megistus, Reduviidae). (b) Kissing bug {Rhodnius prolixus, Reduviidae). (c) Thread-legged bug (Empicoris rubromaculatus, Reduviidae). (d) Bat bug (Hesperoctenes sp., Polyctenidae). (e) Bedbug (Cimex lectularius, Cimicidae).

224

BUGS

b a n d e d alternately with light a n d ¿ a r k colors. T h e general b o d y color is black o r b r o w n , with well-defined harle­ quin o r variegated p a t t e r n s of light yellow, orange, o r r e d . T h e m a i n biological f e a t u r e of kissing bugs is their obligate, v e r t e b r a t e bloodfeeding habits. All species n e e d this blood to complete t h e i r life cycle. T h e i r p r i m a r y habitat is t h e nest of their hosts a n d n e a r b y confined refuges ( u n d e r b a r k , rock crev­ ices, crowns of palms, etc.) w h e r e they h i d e during t h e day, e m e r g i n g at n i g h t to feed on v e r t e b r a t e s visiting these niches. Many species a r e domestic, o c c u r r i n g in poultry pens a n d stock corrals a n d habitations, especially m u d o r a d o b e h u t s .

a n d B l a n k e n s h i p 1984; Ryckman a n d Zackrison 1987). Rhodnius prolixus colonies a r e now widely used as e x p e r i m e n t a l animals in m a n y labo­ ratories as a result of t h e p i o n e e r i n g work of insect physiologist V. B . Wigglesworth.

Adults rarely fly b u t occasionally a r e attracted to lights at night. Most have a long life cycle, 3 0 0 days o n t h e average from e g g to a d u l t ; s o m e two years. O t h e r s complete d e v e l o p m e n t in less t h a n a year and may h a v e two to t h r e e g e n e r a t i o n s p e r a n n u m . T h e r e a r e five n y m p h a l stages. When d i s t u r b e d , m a n y species release a pungent o d o r from m e t a t h o r a c i c glands (Schofield a n d U p t o n 1978). T h e active chemical is butyric acid, which is r e p u g ­ nant t o h u m a n s a n d o t h e r animals, includ­ ing ants. O t h e r aspects of t h e behavior of Triatominae a r e reviewed by Schofield (1979).

LENT, H., AND P. WYGODZINSKY. 1979. Revision

o U S |y

Because of t h e i r voracious appetites for h u m a n blood, kissing b u g s w e r e well known to t h e i n d i g e n o u s A m e r i c a n s a n d have attracted attention historically since early colonial days. Fray Reginaldo d e Lizárraga was t h e first t o describe t h e b u g s in 1590 (Abalos a n d Wygodzinsky 1951). T h e subfamily contains five tribes with 13 g e n e r a a n d 111 species in t h e Western H e m i s p h e r e , whose vector i m p o r t a n c e , classification, a n d s t r u c t u r e h a v e b e e n thor­ oughly reviewed (Lent a n d Wygodzinsky 1979, U s i n g e r et al. 1966). Bibliographies of Chagas' disease a n d lists of species of t h e kissing b u g s o f Latin A m e r i c a a r e also available (Ryckman 1984, 1986; R y c k m a n

References ABALOS, J. W., AND P. WYGODZINSKY. 1951. La

vinchuca: Folklore y antecedentes históricos. Cien. Invest. 7: 472-475. BARBOSA DE ALMEIDA, F. 1971. Triatomíneos da

Amazonia. Acta Amazónica 1: 8 9 - 9 3 . LENT, H., AND J. JURBERG.

1975. O

género

Panstrongylus Berg, 1879, com um estudo sobre a genitalia externa das especies (Hemiptera, Reduviidae, Triatominae). Rev. Brasil. Biol. 35: 379-438. of the Triatominae (Hemiptera, Reduviidae), and their significance as vectors of Chagas' disease. Amer. Mus. Nat. Hist. Bull. 163: 123-520. RABINOVICH, J. E. 1972. Vital statistics of Triatominae (Hemiptera: Reduviidae) under laboratory conditions. I. Triatoma infestans Klug. J. Med. Entomol. 9:351-370. RYCKMAN, R. E. 1984. T h e Triatominae of

North and Central America and the West Indies: A checklist with synonymy (Hemip­ tera: Reduviidae, Triatominae). Soc. Vector Ecol. Bull. 9: 71-83. RYCKMAN, R. E. 1986. T h e Triatominae of

South America: A checklist with synonymy (Hemiptera: Reduviidae: Triatominae). Soc. Vector Ecol. Bull. 11: 199-208. RYCKMAN, R. E., AND C. M. BLANKENSHIP. 1984.

The Triatominae and Triatominae-borne trypanosomes of North and Central America and the West Indies: A bibliography with index. Soc. Vector Ecol. Bull. 9: 112-430. RYCKMAN, R. E., AND J. L. ZACKRISON. 1987. A

bibliography to Chagas' disease, the Triato­ minae and Triatominae-borne trypanosomes of South America (Hemiptera: Reduviidae: Triatominae). Soc. Vector Ecol. Bull. 12: 1-464. SCHOFIELD,

C. J.

1979. T h e behavior

of

Triatominae (Hemiptera: Reduviidae): A Re­ view. Bull. Entomol. Res. 69: 363-379. SCHOFIELD,

C. J.,

AND C. P. UPTON.

1978.

Brindley's scent-glands and the metasternal scent-glands oí Panstrongylus megistus (Hemip­ tera, Reduviidae, Triatominae). Rev. Brasil. Biol. 38: 665-678.

ASSASSIN BUGS

225

USINGER, R. L., P. WYGODZINSKY, AND R. E.

RYCKMAN. 1966. The biosystematics of Triatominae. Ann. Rev. Entomol. 11: 309-330. ZELEDÓN,

R.,

AND J.

E.

RABINOVICH.

1981.

Chagas' disease: An ecological appraisal with special emphasis on its insect vectors. Ann. Rev. Entomol. 26: 101-133.

Parasitic Bugs Bugs in two h e t e r o p t e r a n families have lost their wings a n d b e c o m e totally a d a p t e d for a parasitic existence o n t h e bodies, o r in the nests, of birds a n d m a m m a l s . A m o n g these, the Polyctenidae, o r bat bugs, a r e the m o r e e x t r e m e l y modified structurally a n d biologically, for life o n bats. T h e y are small (BL 3 to 5 m m ) , lack eyes, a r e flattened, a n d have comblike rows of flattened spines o n t h e b o d y (fig. 8.4d). T h e females a r e viviparous (Ryckman a n d Casdin 1977). A single g e n u s , Hesperoctenes, is p r e s e n t in t h e New World ( U e s h i m a 1972). T h e cimicids, o r b e d b u g s , a r e m u c h better k n o w n because of t h e two species that associate closely with m a n k i n d . T h e y are n o t well a d a p t e d to cling to f u r o r leathers b u t a r e t e m p o r a r y visitors t o t h e body of the host, n o r m a l l y bats a n d various birds. Most c o m m o n a n d best k n o w n a r e the i n t r o d u c e d domestic b e d b u g s (Cimex lec­ tularius a n d C. hemipterus) o n bats, chickens, a n d h u m a n s t h r o u g h o u t A m e r i c a (see b e ­ low). Also of e c o n o m i c i m p o r t a n c e a r e t h e p a r r o t b e d b u g (Psitticimex), o n p a r r o t s in n o r t h e r n A r g e n t i n a , t h e Mexican chicken b u g (Haematosiphon inodorus), o n chickens in Mexico (Lee 1955), a n d t h e Brazilian chicken b u g (Ornithocoris toledoi), which, p r i o r to its control with o r g a n i c insecticides, was a pest o n chickens in s o u t h e a s t e r n South America. Twelve g e n e r a of cimicids a r e f o u n d in Latin America. T h e absence of native spe­ cies in C e n t r a l A m e r i c a , w h e r e only t h e domestic pests a r e r e p r e s e n t e d , is a curious distribution p a t t e r n a m o n g New World b e d b u g s (Ryckman et al. 1981, U s i n g e r et al. 1966).

226

BUGS

Cimicids are larger t h a n polyctenids (BL 4—5 m m ) , m o r e o r less oval, a n d slightly flattened. T h e a b d o m e n is soft a n d capable of e n o r m o u s e n l a r g e m e n t with blood at the time of e n g o r g e m e n t . T h e y a r e flight­ less, b u t padlike, vestigial fore wings per­ sist in most species. A u n i q u e d e v e l o p m e n t in this family i s the presence of a secondary copulatory receptacle (spermalege) in the female, situ­ ated o n o n e side, at t h e base of the a b d o m e n . T h e male genitalia a r e asymmet­ rical a n d possess a formidable, swordlike penis. Males p e r f o r a t e t h e i n t e g u m e n t with this i n t r o m i t t e n t o r g a n in o n e or m o r e places, usually t h r o u g h a n external fold in a n intersegmental m e m b r a n e over the receptacle. T h e s p e r m is t h e n deliv­ e r e d to a n u n d e r l y i n g pocket that c o m m u ­ nicates with t h e genital ducts. Copulation with t h e female's n o r m a l terminal geni­ talia never occurs.

References LEE, R. D. 1955. T h e biology of the Mexican chicken bug, Haematosiphon inodorus (Duges) (Hemiptera: Cimicidae). Pan-Pacific Ento­ mol. 31: 4 7 - 6 1 . RYCKMAN, R. E., D. G. BENTLEY, AND E. F. ARCH-

BOLD. 1981. The Cimicidae of the Americas and oceanic islands: A checklist and bibliogra­ phy. Soc. Vector Ecol. Bull. 6: 93-142. RYCKMAN, R. E., AND M. A. CASDIN. 1977. The

Polyctenidae of the world, a checklist with bibliography. Calif. Vector Views 24: 25-31. UESHIMA, N. 1972. New World Polyctenidae (Hemiptera), with special reference to Vene­ zuelan species. Brigham Young Univ. Sci. Bull. (Biol. Ser.) 17: 13-21. USINGER, R. L., J. CARAYON, N. T. DAVIS, N. UESHIMA, AND H. E. MCKEAN. 1966. Mono­

graph of Cimicidae. Vol. 7. Entomol. Soc. Amer. (Thomas Say Foundation), College Park, Md.

BEDBUGS Cimicidae, Cimicinae, Cimex lectularius a n d C. hemipterus. Spanish: C h i n c h e s de cama (General). Portuguese: Percevejos

das camas (Brazil). Náhuatl: Texcantin, sing- texcan (Mexico). Wall lice, r e d coats. H u m a n b e d b u g s a r e truly cosmopolitan insects, h a v i n g followed h u m a n s over most of t h e world ( A n o n . 1973). T h e two anthrophilic species have b e e n well k n o w n since antiquity in t h e O l d World, w h e r e their original hosts were probably bats. They first took to m a n in s o u t h e r n E u r o p e or t h e M i d d l e East in t h e case of t h e common b e d b u g (C. lectularius) o r t h e Ori­ ent in t h e case of t h e tropical b e d b u g (C. hemipterus) a n d surely s p r e a d to t h e New World with t h e first e x p l o r e r s a n d immi­ grants. Early published m e n t i o n s of t h e species a r e always suspect o w n i n g to t h e chroniclers' use of chinche a n d Aztec texcan not only for c o m m o n b e d b u g s b u t for other a r t h r o p o d s ( H o e p p l i 1969: 183). These bugs are ravenous nocturnal blood seekers cohabiting with m a n , h i d i n g in cracks, mattresses, a n d so o n , a n d b e ­ hind f u r n i t u r e d u r i n g t h e day. T h e y emerge at n i g h t to j o i n us in o u r b e d s a n d to fill t h e i r g u t s with o u r life fluids. I n u n k e m p t p r e m i s e s , h u n d r e d s m a y be found in hideaways, a n d their o b n o x i o u s , sickeningly sweet "nest o d o r " can be over­ whelming. T h e chemicals responsible for the o d o r m a y have alternative attractant and repellent functions t o w a r d o t h e r bed­ bugs o r animals (Levinson a n d Barllan 1971). B e d b u g bites cause varied reactions in humans. I n s o m e , t h e r e is m a r k e d swelling and irritation; in o t h e r s , n o effect. Al­ though m a n y e x p e r i m e n t s have b e e n con­ ducted to i n c r i m i n a t e these two species as natural disease vectors, t h e results have virtually always b e e n negative. T h e y can transmit several p a t h o g e n s u n d e r labora­ tory conditions, however. Bedbugs a r e small, ovate, a n d m u c h flattened e x c e p t w h e n e n g o r g e d , a n d t h e n they a r e almost spherical. A d u l t s a r e reddish-brown a n d covered with m i n u t e

bristles. T h e i r legs a r e slender a n d fairly long. T h e lateral e x p a n s i o n s of t h e p r o t h o r a x of t h e c o m m o n b e d b u g (fig. 8.4e) are m o r e extensive a n d flattened t h a n those of the tropical b e d b u g , a n d its color is lighter. T h e f o r m e r also t e n d s to be m o r e u r b a n a n d e x t e n d s into h i g h e r latitudes t h a n the latter, which is a rural species a n d restricted to t h e m i d d l e latitudes in t h e Americas. I n f o r m a t i o n o n all aspects of the n a t u r a l history of these species is s u m m a r i z e d in the masterful m o n o g r a p h by U s i n g e r et al. (1966).

References ANONYMOUS. 1973. The bed-bug. 8th ed. Brit. Mus. Nat. Hist. Econ. Ser. 5: 1-17. HOEPPLI, R. 1969. Parasitic diseases in Africa and the Western Hemisphere, early documen­ tation and transmission by the slave trade. Acta Trop. suppl. 10: 1-240. LEVINSON, H. Z., AND A. R. BARLLAN. 1971.

Assembling the alerting scents produced by the bedbug Cimex lectularius L. Experientia 27: 102-103. USINGER, R. L., J. CARAYON, N. T. DAVIS, N. UESHIMA, AND H. E. MCKEAN. 1966. Mono­

graph of the Cimicidae. Vol. 7. Entomol. Soc. Amer. (Thomas Say Foundation), College Park, Md.

Water Bugs As h a s o c c u r r e d secondarily in t h e evolu­ tion of Coleóptera a n d o t h e r o r d e r s , sev­ eral families of H e t e r o p t e r a have a d o p t e d aquatic habits a n d a r e modified for swim­ ming, diving, respiring, a n d f e e d i n g while s u b m e r g e d . Two basic life-styles have evolved: the "semiaquatic b u g s " ( A n d e r s e n 1982) live o n t h e surface film a n d a r e generally less a d a p t e d to the m e d i u m t h a n the truly "aquatic b u g s " that s p e n d t h e majority of their time b e n e a t h t h e surface. T h e s e are insects of g r e a t i m p o r t a n c e in the ecology of all watery habitats, from small trickles, h i g h l a n d streams, a n d p o n d s to the margins of the seas. A l t h o u g h specifically treating t h e Guy-

BEDBUGS

227

ana region, t h e review by Nieser (1975) is widely applicable. T h e g e n e r a l Latin A m e r i ­ can literature is c o m p i l e d in t h e bibliogra­ phies of B a c h m a n (1977), Nieser (1981), a n d P o l h e m u s (1982).

References ANDERSEN, N. M. 1982. T h e semiaquatic bugs (Hemiptera, Gerromorpha), phylogeny, adap­ tations, biogeography and classification. Entomonograph 3: 1—455. BACHMANN, A. O. 1977. Heteroptera. In S. H.

Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 189-212. NIESER, N. 1975. T h e water bugs (Heteroptera: Nepomorpha) of the Guyana Region. Stud. Fauna Suriname Guyanas 59: 1—310. NIESER, N. 1981. Hemiptera. In S. H. Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 100-128. POLHEMUS, J. T

1982. Hemiptera. In S. H.

Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 288-292.

GIANT WATER BUGS Belostomatidae, L e t h o c e r i n a e , Lethocerus. Spanish: C u c a r a c h a s d e l a g u a (General). Portuguese: Baratas d ' a g u a (Brazil). Electric light b u g s , toe biters.

Belonging to a cosmopolitan family, this ge­ nus has speciated prolifically in Latin A m e r ­ ica. T h e r e a r e nearly twenty distinct spe­ cies, mostly in t h e g e n u s Lethocerus (Menke 1963). T h e two species L. maximus (fig. 8.5a) a n d L. granáis a r e a m o n g t h e largest of in­ sects with a b o d y length u p to 11.5 centime­ ters a n d weight of 15 to 25 g r a m s . All a r e m u c h flattened, ellipsoid in out­ line, a n d shiny d a r k b r o w n . T h e h e a d is rigidly fixed to t h e t h o r a x a n d d o e s not rotate; its frontal p o r t i o n projects strongly forward between t h e eyes. Straplike, retrac­ tile respiratory a p p e n d a g e s a r e b o r n e at the tip of t h e a b d o m e n . T h e latter pene­ trate t h e surface film w h e n t h e b u g as­ cends, r e a r w a r d , to take in air. Most of the store is carried u n d e r t h e wings in a cavity created by t h e d e p r e s s e d a b d o m e n . T h e s e a r e rapacious p r e d a t o r s , catching all sorts of o t h e r aquatic invertebrates and even small vertebrates like tadpoles and fish with their raptorial forelegs, then killing t h e m with a vicious stab of the m o u t h stylets. T h e r o s t r u m acts like a h y p o d e r m i c syringe, injecting saliva that b o t h immobilizes a n d digests t h e organ­ isms o n which they feed (Picado 1937). T h e bugs wait patiently in a m b u s h a m o n g plants or debris in t h e water, relying on r e m a i n i n g motionless a n d o n their cryptic coloration to escape detection by their

prey. Female giant water b u g s lay their e ggs o n e m e r g e n t aquatic vegetation, not on t h e back o f males as is characteristic of other g e n e r a in t h e family. Specimens m a y b e c o m m o n at times near their well-vegetated, m a r s h y p o n d and lakeshore habitats. O n w a r m evenings d u r i n g their dispersal season, they some­ times a c c u m u l a t e u n d e r electric lights a n d attract a g r e a t deal of attention (Lanzer 1975). T h e biology of L. maximus h a s b e e n investigated in s o m e detail in T r i n i d a d (Cullen 1969) a n d it has b e e n m a i n t a i n e d in the l a b o r a t o r y for studies o n t h e physiol­ ogy of its flight muscles (Barros 1973).

References BARROS S., M. C. 1973. Mantencáo da barata d'agua gigante (gen. Lethocerus) no labora­ torio. Univ. Sao Paulo, lnst. Biol. Mar., Bol. Zool. Biol. Mar. (Nov. Ser.) 30: 613-623. CULLEN, M. J. 1969. T h e biology of giant water bugs (Hemiptera: Belostomatidae) in Trini­ dad. Royal Entomol. Soc. London Proc. A 44: 123-136. LANZER, M. E. B. 1975. Nota previa sobre o comportamento de Belostoma Latreille, 1807 e Lethocerus Mayr, 1853 em aquário e no meio ambiente. Iheringia (Ser. Div.) 4: 47-50. MENKE, A. S. 1963. A review of the genus Lethocerus in North and Central America, including the West Indies (Hemiptera: Belo­ stomatidae). Entomol. Soc. Amer. Ann. 56: 261-267. PICADO T , C , 1937. Estudo experimental sobre o veneno de Lethocerus del-pontei (DeCarlo) (Hemiptera-Belostomidae). lnst. Butantan (Sao Paulo) Mem. 10: 303-310, figs. 1-3.

BACK SWIMMERS Notonectidae.

Figure 8.5 WATER BUGS, (a) Giant water bug (Lethocerus maximus, Belostomatidae). (b) Back swimmer (Buenoa pallens, Notonectidae). (c) Salt marsh water boatman (Trichocorixa reticulata, Corixidae). (d) Common water strider (Gerris remigis, Gerridae). (e) Sea strider (Halobates micans, Gerridae).

228

BUGS

Back s w i m m e r s a r e so n a m e d because of their habit of s w i m m i n g u p s i d e d o w n . They a r e also recognized by their long, oarlike h i n d legs t h a t pull in u n i s o n w h e n the insect swims. T h e b o d y is slender, boat shaped, a n d often colored with white, r e d , or greenish areas, a l t h o u g h it is usually all dull brown. T h e fore tarsus is visibly two

s e g m e n t e d , slender, a n d b a r e . Like their relatives, t h e giant water bugs, they have large eyes a n d a s h a r p beak with p o t e n t powers; their bite is very painful a n d often is t h e salvation of a specimen in t h e collec­ tor's clutches. Also like their relatives, they are p r e d a c e o u s , b u t t h e size of their prey is smaller, consisting of fly a n d beetle larvae, fish fry, a n d crustácea. T h e r e a r e between 300 a n d 4 0 0 species, mostly in t h e d o m i n a n t g e n e r a Notonecta, Martarega, a n d Buenoa. T h e s e a r e distrib­ u t e d almost e v e r y w h e r e w h e r e suitable sluggish water habitats a r e available b u t show considerable selectivity to specific niches d e t e r m i n e d by water conditions a n d prey availability (Gittleman 1975). Notonecta a r e t h e largest (BL of most 1 2 15 m m ) a n d must rise frequently to take o n oxygen from t h e a t m o s p h e r e ; males a r e silent. Buenoa a r e smaller ( m a x i m u m BL 11 m m ) a n d possess h e m o g l o b i n c o n t a i n i n g cells that store oxygen a n d assist t h e m in r e m a i n i n g s u b m e r g e d for longer p e r i o d s t h a n o t h e r notonectids; males stridulate by m e a n s of a r i d g e d p r o t u b e r a n c e o n t h e inside of the fore tibia. B. pallens (fig. 8.5b) is a widespread species. Martarega a r e t h e smallest ( m a x i m u m B L 10 m m ) a n d a r e most c o m m o n l y f o u n d at water's e d g e in slow-moving rivers a n d d o n o t stridulate. Since earliest times, Mexicans a r o u n d Lake Texcoco a n d Chalco n e a r Mexico City have collected Notonecta unifasciata (axayácatl) a n d associated water b o a t m e n , which they c o n s u m e in various forms (Ancona 1933). T h e eggs (ahuautli, axayácatl, etc.) are cooked into a kind of bread (hautle); adults (ahuatle, bledo delagua) may be g r o u n d a n d mixed with saltpeter to m a k e a salty h a s h ( B o d e n h e i m e r 1951). Refer to B a c h m a n n (1977), Nieser (1981), a n d P o l h e m u s (1982) for bibliogra­ phies o n t h e regional fauna.

References ANCONA, L. 1933. El ahuatle de Texcoco. lnst. Biol. Univ. Nac. Mexico Ann. 4: 51-69.

BACK SWIMMERS

229

BACHMANN, A. O. 1977. Notonectidae. In S. H.

Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State University, San Di­ ego. Pp. 193-195. BODENHEIMER, F. S. 1951. Insects as human food. Junk, T h e Hague. GITTLEMAN, S. H. 1975. T h e ecology of some Costa Rican backswimmers (Hemiptera: Noto­ nectidae). Entomol. Soc. Amer. Ann. 68: 511-518. NIESER,

N.

1981.

Notonectidae.

In

S.

H.

Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State University, San Diego. Pp. 115-117. POLHEMUS, J. T

1982. Notonectidae. In S. H.

Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univer­ sity, San Diego. Pp. 306-308.

WATER BOATMEN Corixidae. Water b o a t m e n a r e superficially similar to back s w i m m e r s b u t always swim r i g h t side u p a n d differ in m a n y anatomical details: the b o d y is cylindrical o r slightly flattened, the m o u t h p a r t s a r e short, c o n e s h a p e d , a n d u n s e g m e n t e d , a n d t h e male genitalia are asymmetrical. T h e fore wings a r e often m a r k e d with fine, t r a n s v e r s e zigzagging d a r k lines. T h e tarsus of t h e foreleg is c o m p o s e d of a single b r o a d , hairy seg­ m e n t . T h e y vary considerably in size (BL 3-10 mm). T h e s e water b u g s c h o o s e habitats like the back s w i m m e r s , a l t h o u g h they t e n d to p r e f e r m o r e s t a g n a n t or t o r p i d waters w h e r e they s p e n d most of their time at t h e b o t t o m . S o m e , such as Trichocorixa reticulata (fig. 8.5c), tolerate saline c o n d i t i o n s a n d may be e x t r e m e l y a b u n d a n t in inland salt pools o r estuaries o n t h e c o n t i n e n t a n d in the Galápagos Islands. Corixid food a n d f e e d i n g habits a r e varied. A c o m m o n m e t h o d of feeding is by sieving edible particles from b o t t o m d e ­ bris. T h e y also m a y ingest o t h e r living o r

230

BUGS

d e a d aquatic invertebrates if small e n o u g h or in a d e c o m p o s e d state. Males chirp by r a s p i n g areas of small pegs on the base of t h e fore f e m u r against the s h a r p e d g e of t h e m o u t h beak. T h e family is diverse in t h e Neotropics, with a p p r o x i m a t e l y 117 species in 14 gen­ era. As with notonectids (see above), some corixids (Corisella) a r e f o u n d in Mexican markets. Tons of these d r i e d insects are s h i p p e d a b r o a d as bird o r fish fodder, and they a r e g a t h e r e d also for h u m a n use, both as eggs or adults (mosco, moschitos). Eggs are laid in e n o r m o u s n u m b e r s o n r e e d s placed in t h e water by the g a t h e r e r s . Like those of Notonecta, they a r e m a d e into a fishflavored cake, also called huatlé o r ahuahutl ( B o d e n h e i m e r 1951: 295f.). Refer to B a c h m a n n (1977), Nieser (1981), a n d P o l h e m u s (1982) for bibliogra­ phies on the Latin A m e r i c a n fauna.

References BACHMANN, A. O. 1977. Corixidae. In S. H.

Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 191-193. BODENHEIMER, F. S. 1951. Insects as human food. Junk, The Hague. NEISER, N. 1981. Corixidae. In S. H. Hurlbert, G. Rodriguez, and N. Dios dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 117-119. POLHEMUS, J. T. 1982. Corixidae. In S. H.

Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 308-310.

WATER STRIDERS G e r r i d a e . Spanish: Zapateros (Argentina). Pond skaters. T h e s e a r e oval to elongate b u g s with fours e g m e n t e d a n t e n n a e a n d large, globular eyes. T h e i r o t h e r distinctive features are slender spiderlike legs, with tarsal claws

inserted b e f o r e t h e tip of t h e last segment. The body is covered with a velvety pile of hairs that a r e h y g r o p h o b i c a n d m a k e t h e insect resistant to wetting s h o u l d it sub­ merge. T h e i r colors r a n g e from black to brown with occasional silvery m a r k i n g s . Wings m a y be absent, abbreviated, o r fully developed. Water striders ( A n d e r s e n a n d P o l h e m u s 1976) all a r e "semiaquatic bugs," s p e n d i n g their entire lives m o v i n g jerkily to a n d fro on the surface film. T h e y feed on t h e blood of insects a n d invertebrates that fall onto t h e surface film o r reside o n it. A common species is Gerris remigis (fig. 8.5d). O n e g r o u p of striders has colonized t h e open ocean. T h e best k n o w n of these, t h e "sea skaters," belong to t h e g e n e r a Halobates ( H e r r i n g 1 9 6 1 , C h e n g 1985) a n d Rheumatobates ( C h e n g a n d Lewin 1971), which generally r e s e m b l e freshwater strid­ ers except for a grossly r e d u c e d a b d o m e n and c o m p l e t e absence of wings. J u s t a few species live o n t h e waters of t h e ocean off both coasts of A m e r i c a a n d a m o n g associ­ ated oceanic islands. Halobates micans (fig. 8.5e) a n d H. robustus a r e c o m m o n inhabit­ ants of small bays in t h e Galápagos Islands. Little is k n o w n of their biology. Coastal species lay their eggs on rocks; pelagic species may oviposit o n floating objects (wood, feathers) a n d a r e even k n o w n to attach eggs to birds that h a v e b e e n resting on the waves. T h e i r food consists of pelagic, surface-dwelling animals like jellyfish. Refer to B a c h m a n n (1977), Nieser (1981), a n d P o l h e m u s (1982) for bibliogra­ phies on t h e r e g i o n a l fauna.

References ANDERSEN, N. M., AND J. T. POLHEMUS.

1976.

Water-striders (Hemiptera: Gerridae, Veliidae, etc.). In L. Cheng, ed., Marine insects. North-Holland, Amsterdam. Pp. 187-224. BACHMANN, A. O. 1977. Gerridae. In S. H.

Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 204-206. CHENG, L. 1985. Biology of Halobates (Heter-

optera: Gerridae). Ann. Rev. Entomol. 30: 11-135. CHENG, L., AND R. A. LEWIN. 1971. An interest­

ing marine insect, Rheumatobates aestuarius (Heteroptera: Gerridae), from Baja Califor­ nia, Mexico. Pacific Ins. 13: 333-341. HERRING, J. L. 1961. T h e genus Halobates (Hemiptera: Gerridae). Pacific Ins. 3: 223— 305. NIESER, N. 1981. Gerridae. In S. H. Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 125-128. POLHEMUS, J. T.

1982. Gerridae. In S. H.

Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ. San Diego. Pp. 319-323.

HOMOPTERANS H e m i p t e r a , H o m o p t e r a . Wax bugs. T h i s s u b o r d e r is a large o n e with m a n y families, several of which a r e strictly Neotropical. O p i n i o n s differ as to its inter­ nal classification because of t h e diversity of basic body forms a n d e x t r e m e biological a n d anatomical a d a p t a t i o n s . S o m e unify­ ing characteristics a r e sucking m o u t h p a r t s similar to those of h e t e r o p t e r a n s b u t with the elements usually consolidated into a short beak set so far back o n t h e h e a d as to a p p e a r to arise from b e h i n d t h e forelegs a n d piercing stylets often very long a n d coiled in t h e body w h e n n o t in use. Wings, when present, a r e h o m o g e n o u s in t e x t u r e , usually m e m b r a n o u s b u t often thickened a n d fairly rigid a n d with few veins (al­ t h o u g h some have a reticulate p a t t e r n ) . T h e r e a r e two pairs of wings except for male scale insects, which have only t h e fore pair d e v e l o p e d . I use t h e n a m e "wax b u g s " to refer to the s u b o r d e r because of the almost univer­ sal p r e s e n c e of w a x - p r o d u c i n g glands in its constituent families. T h e s e glands a r e vari­ ously developed o n different p a r t s of t h e body a n d always o p e n exteriorly o n t h e cuticle. T h e i r secretions a r e e x t r u d e d to

HOMOPTERANS

231

the outside to take many forms for diverse functions, some well defined, such as the protective scale covering of armored scale insects, and others with unclear signifi­ cance, such as the massive plumes trailing from the abdomen of some large fulgorids. Silk- and lac-producing glands are also present. All are sap feeders and often very spe­ cific in their preferences for the sap of certain plants (Johnson and Foster 1986). They remove this fluid, sometimes in such quantity, because of enormous populations that may develop, that they kill or seriously injure their hosts. For this reason, and also because many are very efficient carriers of plant pathogens, wax bugs are plant pests of prime importance. Injurious species are found in almost all families but especially among the aphids, scale insects, and leafhoppers. Unlike their heteropteran relatives, no wax bugs have adapted to feeding on vertebrate blood or have be­ come aquatic. A mutualistic relationship exists be­ tween some hymenopterans and many wax bugs, especially aphids and scale insects (Letourneau and Choe 1987). The latter secrete "honeydew," a carbohydrate-rich overflow from the alimentary canal or integumentary glands which is greedily consumed and even specifically solicited, especially by ants. T h e ants attend the bugs, protecting them from predators, dis­ persing them, and even building shelters for them (Way 1953).

References JOHNSON, L. K., AND R. B. FOSTER. 1986. Associa­

tions of large Homoptera (Fulgoridae and Cicadidae) and trees in a tropical forest. Kans. Entomol. Soc.J. 59: 415-422. LETOURNEAU, D.

K., AND J. C. CHOE.

1987.

Homopteran attendance by wasps and ants: The stochastic nature of interactions. Psyche 94: 81-91. WAY, J. T. 1953. Mutualism between ants and honeydew producing Homoptera. Ann. Rev. Entomol. 8: 307-344.

232

BUGS

CICADAS Cicadidae. Spanish: Chicharras (General); coyuyos, cigarras (Argentina). Portuguese: Cigarras. Among homopterans, cicadas are famous for their sound-producing abilities. (A few others have vocal organs but none so well developed or capable as those of Cicadi­ dae, and nothing is known of their func­ tion in Neotropical forms.) On the back of the first abdominal segment there is a pair of exposed (or protected by a fold of the body wall), taut membranes (tymbals) that may be made to rapidly vibrate by well-developed oblique muscles. The tymbals vibrate extremely fast, producing a loud, sometimes strident, steady or pulsating buzz or siren scream that varies in pitch, pulse, intensity, dura­ tion, or other acoustic quality according to species. One type (tentatively determined as the very widespread Quesada gigas; fig. 8.6a) in park trees in Caracas emits a fairly deafening, throbbing wail. Some species are silent or produce only interrupted clicks. Both sexes have auditory tympani anteri­ orly on the underside of the abdomen, concealed beneath large protective plates (not to be confused with the tymbals). Males often congregate on trees in forests and join their voices in intense synchro­ nized chorusing that acts as a call to assem­ ble females for mating. This aggregative behavior seems to be exhibited in secon­ dary habitats and on trees that are the nymphal hosts (Young 1980). The latter are varied hardwoods and palms (Young 1973). Peaks of singing intensity occur in many species at dawn and dusk (Young 19816). On trees, females insert masses of eggs in twigs or fronds with an ovipositor adapted for slitting bark. In many species, oviposition is limited to dead trees and fronds in the forest understory, while oth­ ers put their eggs into dead grasses near

Figure 8.6 WAX BUGS, (a) Giant cicada (Quesada gigas, Cicadidae). (b) Spotted cicada (Zammara smaragdina, Cicadidae). (c) Cicada nymph (Cicadidae). (d) Ground pearls (Margarodes formicarum, Margarodidae), females, (e) Axin (Uaveia axin, Margarodidae).

the ground. T h e nymphs drop to the soil and burrow deeply. They commonly spend two (but some, several to many) years underground, feeding on sap from the host's roots. On maturity, they crawl out of the ground and up onto tree trunks and limbs to transform. Fixed to such surfaces, their cast skins, split widely down the back, are a common sight. The nymphs of at least one Amazonian cicada (Fidicina chlorogena; Ginzberger 1934) constructs tall (2030 cm), hollow, mud chimneys in which to pass their final days before becoming adults. The imagos are presumed to be short­ lived, perhaps existing from a few weeks to a few months. They imbibe fluids from the xylem tissue of their hosts with a sturdy, jointed proboscis. Most are active in the forest canopy, although some species fre­ quent the trunks of trees close to the ground. Few additional details of tropical cicadan biology are available, and these are mostly from species in Costa Rica (Young 1981a). Others (Bartholomew and Barnhart 1984) have studied flight metabolism in the Central American F. manijera. It can fly at a body temperature of 22° C , but takeoff must be preceded by warming up by body movements. During active flight, the body heats up to 33° C. They do not jump but are easily excited and frightened into flight. Adults are easily distinguished as a group by their robust form large size (BL °f most 20-50 mm), and membranous

wings, the anterior pair of which are al­ most twice as long as the posterior and extending well beyond the end of the body when folded. A transversely ridged swell­ ing on the face and the singing organs of the male also are unique features of the family. The lateral margins of the thorax have winglike flanges in some, for exam­ ple, Zammara (fig. 8.6b). The bodies of most cicadas are marked with green or white against a black background. T h e wing veins are usually dark, the color usually staining only the adjoining mem­ brane, although the usually transparent membranous areas may be splotched with brown patterns as well. Nymphs (fig. 8.6c) are stout-bodied, brownish, molelike creatures with outsized, powerful fore tibiae and otherwise heavy legs adapted for digging. The abdo­ men is stubby and curved downward. Cicadas seldom cause economic damage to plants with their feeding and ovi­ position. They have been recorded molest­ ing fruit trees and coffee in Brazil (da Fonseca 1945). A complete bibliography on the family to 1956 is available (Metcalf 1962). The species are cataloged by Metcalf (1963a, 19636, 1963c) and Duffels and van der Laan (1985). The number of described Latin American species is roughly esti­ mated at over 800, in some 80 or so genera; the former figure may represent as few as 60 percent of those that are to be found (Moore pers. comm.).

CICADAS

233

References BARTHOLOMEW, G. A., AND M. C. BARNHART.

1984. Tracheal gases, respiratory gas ex­ change, body temperature and flight in some tropical cicadas. J. Exper. Biol. I l l : 131 -144. DA FONSECA, J. P. 1945. As cigarras do cafeeiro e seu combate. Bol. Agr. 46: 297-304. DUFFELS, J. P., AND P. A. VAN DER LAAN.

1985.

Catalogue of the Cicadoidea (Homoptera, Auchenorhyncha) 1956-1980. Junk (Series Entomológica), T h e Hague. GINZBERGER, A. 1934. Die Bauten der Larve der Singzikade Fidicina chlorogena Wlk. Anz. Akad. Wiss. W i e n 7 1 : 55. METCALF, Z. P. 1962. A bibliography of the

Cicadoidea (Homoptera: Auchenorhyncha). Entomol. Dept., N.C. Agrie. Exper. Sta. Pap. 1373: 1-229. METCALF, Z. P. 1963a. Cicadoidea, Cicadidae, Tibiceninae. In Z. P. Metcalf, General cata­ logue of the Homoptera. N.C. State Coll., Raleigh. Fase. 8, pt. 1, sec. 1. METCALF, Z. P. 19636. Cicadoidea, Cicadidae, Gaeaninae and Cicadinae. In Z. P. Metcalf, General catalogue of the Homoptera. N.C. State Coll., Raleigh. Fase. 8, pt. 1, sec. 2. METCALF, Z. P. 1963c. Cicadoidea, Tibicenidae. In Z. P. Metcalf, General catalogue of the Homoptera. N.C. State Coll., Raleigh. Fase. 8, pt. 2. YOUNG, A. M. 1973. Cicada populations on palms in tropical rain forest. Principes Palm Soc.J. 17: 3 - 9 . YOUNG, A. M. 1980. Observations on the aggre­ gation of adult cicadas (Homoptera; Cica­ didae) in tropical forests. Can. J. Zool. 58: 711-722. YOUNG, A. M. 1981a. Notes on the population ecology of cicadas (Homoptera: Cicadidae) in the Cuesta Angel forest ravine of northeast­ ern Costa Rica. Psyche 88: 175-195. YOUNG, A. M. 19816. Temporal selection for communicatory optimization: The dawn-dusk chorus as an adaptation in tropical cicadas. Amer. Nat. 117: 826-829.

SCALE INSECTS Coccoidea. Spanish: Cochinillas, coccídeos (General). Portuguese: C o c h o n i l h a s , coccídeos (Brazil). Wax g l a n d s a r e especially well d e v e l o p e d in these usually m i n u t e to small (BL 1—2 m m ) h o m o p t e r a n s a n d chiefly function to

234

BUGS

secrete a protective shell o r covering for the insect. T h i s takes a scalelike form in m a n y g r o u p s . All stages a n d both sexes have legs with s e g m e n t e d tarsi t i p p e d with a single claw. Females a r e quite unlike t h e males in form a n d life-style. T h e latter a r e active, midgelike, free-living forms with o n e pair of wings a n d no feeding beak. Most, but not all, females live their entire adult lives as a m o r p h o u s bags of tissue without func­ tional legs o r wings, r e m a i n i n g attached to their food plants by their incredibly long, hairlike feeding stylets, usually u n d e r a protective scale or i n t e r r e d in a h a r d enc r u s t m e n t . T h e y lay their eggs u n d e r or within their casements. T h e eggs hatch into tiny mobile crawling n y m p h s (crawl­ ers) that molt a n d either settle d o w n to a sessile life as females o r r e m a i n active and develop into flying males. T h e r e a r e several different subgroups of scale insects, each with its o w n structural a n d secretory characteristics. Female "gi­ a n t scale insects" {cochinillas de cola, Margarodidae) have well-formed legs a n d a n t e n n a e a n d cover their bodies in white, waxy, o v e r l a p p i n g plates that build u p to form large (to 2 cm), h a r d e n e d s p h e r e s . T h e insects a r e s u b t e r r a n e a n , feeding o n roots a r o u n d which they form grapelike clusters. T h e s e cystlike structures ( " g r o u n d pearls") at o n e time were collected a n d m a d e into bead necklaces in parts of t h e Caribbean, especially those of Margarodes formicarum (fig. 8.6d), a species first f o u n d associated with ants in t h e Lesser Antilles (Guilding 1830). Llaveia axin (fig. 8.6e) is a n o t h e r margarodid type from acacias a n d t h e hog-plum tree (Spondias purpurea) in Yucatán which were used for centuries by Mesoamerican I n d i a n s . T h e bright o r a n g e females, each almost 3 centimeters long, were crushed a n d k n e a d e d t o g e t h e r into a wax ball that f o r m e d t h e base for cosmetics a n d medici­ náis a n d an i n g r e d i e n t of h a r d waxy fin­ ishes applied to early Mayan a n d Aztec

pottery a n d g o u r d s . N a m e s used for this insect by natives of the a r e a in t h e past include ni-in, nije, axin, oji, a n d tuch-cuy (Edwards 1970, J e n k i n s 1970). Giant scale insects a r e also notoriously hardy a n d l o n g lived. T h e published rec­ ord for vitality in an insect is for specimens of Margarodes vitium that r e m a i n e d alive without f e e d i n g for seventeen years (Ferris 1919). T h e species is a pest o n g r a p e s in dry areas of S o u t h America. Apparently, this ability to lie d o r m a n t for long p e r i o d s is an a d a p t a t i o n to dryness, t h e adults emerging n o r m a l l y after t h e first rains. O t h e r very large coccoids a r e the leath­ ery "tortoise scales" (Coccidae), which have a convex s h a p e a n d a r e covered with a thin layer of wax b u t n o scale (BL 1—20 m m ) . An e x a m p l e is Neolecanium sallei (Wheeler 1913), f o u n d o n coral trees (Erythrina) in Central A m e r i c a . I n c l u d e d in this g r o u p a r e t h e "fluted scales," so called because of t h e r i d g e d form of the wax accumulations that build up b e n e a t h t h e body. T h e most well k n o w n representative is t h e cottony cushion scale (Icerya purchasi; fig. 8 . 7 a - c ) , a major e n e m y of citrus t h r o u g h o u t America. Female " a r m o r e d scales" {escamas—Diaspididae) typically live u n d e r a flattened, disklike scale of wax a n d cast n y m p h a l skins. T h i s is t h e largest a n d probably most injurious g r o u p , p o p u l a t i o n s often explod­ ing on their hosts; m a n y a r e n o t o r i o u s

pests of o r c h a r d a n d s h a d e trees a n d o t h e r cultivars. Particularly b a d a r e t h e Califor­ nia red scale {Aonidiella aurantii; fig. 8.7d) a n d San Jose scale {Quadraspidiotus perniciosus) (Marín 1987). Mealybugs (Pseudococcidae a n d Eriococcidae) a r e a b e r r a n t scale insects. T h e y a r e motile a n d without a r m o r , being covered with a thick, crusty o r flaky, white layer of wax instead. T h e y also have long, taillike filaments projecting from t h e p o s t e r i o r (fig. 8.7f). A serious a n d w i d e s p r e a d pest of p i n e a p p l e is the p i n e a p p l e m e a l y b u g (Dysmicoccus brevipes). A n o t h e r g r o u p is t h e "cochineal bugs," discussed below. Many scale insects p r o d u c e copious quantities of honeydew, a source of polysaccharide for m u l t i t u d e s of insects n e a r the base of food chains in ecosystems (Salas a n d J i r ó n 1977).

References EDWARDS, J. G. 1970. Giant margarodid scales from Yucatán. Pan-Pacific Entomol. 46: 68. FERRIS, G. F. 1919. A remarkable case of longev­ ity in insects (Hem., Horn). Entomol. News 30: 27-28. GUILDING, L. 1830. An account oí Margarodes, a new genus of insects found in the neighbor­ hood of ants nests. Linnaean Soc. London Trans. 9:912-914. JENKINS, K. D. 1970. T h e fat-yielding coccid, Llaveia, a monophlebine of the Margarodidae. Pan-Pacific Entomol. 46: 7 9 - 8 1 . MARÍN, L. R. 1987. Biología y morfología de la "escama de San José" Quadraspidiatus perni-

m*

ftL

—

A N D

A

-

H.

EHRLICH.

1982.

Lizard predation on tropical butterflies. J. Lepidop. Soc. 36: 148-152. BLANCO, J. L. 1961. Representaciones de la

mariposa en Mesoamérica. El México Anti­ guo 9: 195-244. GILBERT, L. E., AND M. C. SINGER. 1975. Butter­

fly ecology. Ann. Rev. Ecol. Syst. 6: 365-397. HlNTON, H. E. 1951. Myrmecophilous Lycaenidae and other Lepidoptera—A summary. Proc Trans. So. London Entomol. Nat. Hist. Soc. (1949-50): 111-175. HOFFMAN, C. C. 1918. Las mariposas entre los antiguos mexicanos. Cosmos 1. LAMAS, G. 1977. Bibliografía de catálogos y listas regionales de mariposas (Rhopalocera) de América Latina. Soc. Mexicana Lepidop. Publ. Esp. Pp. 1-44. LAMAS, G. 1978. Adiciones a la bibliografía de catálogos y listas regionales de mariposas de América Latina (Rhopalocera). Soc. Mexi­ cana Lepidop. Bol. Inf. 4(5): 1-14. LAMAS, G. 1983. Mariposas atraídas por hormi­ gas legionarias en la Reserva de Tambopata, Perú. Rev. Soc. Mexicana Lepidop. 8(2): 49-51. LAMAS, G. 1986. Drinking crocodile tears. An­ tenna 10(4): 162. LEGG, G. 1978. A note on the diversity of world Lepidoptera. Biol. J. Linnean Soc. 10: 343-347. ; MAUCKY, H. 1970. New aspects on the associa­ tion between lycaenid larvae (Lycaenidae) and ants (Formicidae, Hymenoptera). J. Lepidop. Soc. 24: 190-202. MAZOKHIN-PORSHNYAKOV, G. A. 1957. Reflect­

ing properties of butterfly wings and role of ultra-violet rays in the vision of insects. Bio­ physics 2: 352-362. NORMS, W. E. 1955. Dawn adventure. Cham­ bers Journal, London. PWEN, D. F. 1974. Trade threat to butterflies. Oryx 12: 479-483. [»CE, N. E. 1987. T h e evolution and biot S^graphy of associations between lycaenid butterflies and ants. Oxford Surv. Evol. Biol. 4:49-116.

PLISKE, T E. 1975. Courtship behavior and use of chemical communication by males of cer­ tain species of ithomiine butterflies (Nym­ phalidae: Lepidoptera). Entomol. Soc. Amer. Ann. 68: 935-942. PYLE, R. M. 1976. T h e ecogeographic basis for lepidoptera conservation. Ph.D. diss., Yale Univ., New Haven. RAY, T

S., AND C. C. ANDREWS.

1980.

Ant

butterflies: Butterflies that follow army ants to feed on antbird droppings. Science 210: 1147-1148. REMINGTON, C. L. 1973. Ultraviolet reflectance in mimicry and sexual signals in the Lepi­ doptera. New York Entomol. Soc. J. 81: 124. RILEY, N. D. 1975. A field guide to the butter­ flies of the West Indies. Collins, London. ROBBINS, R. K. 1982. How many butterfly species? Lepidop. Soc. News (1982): 4 0 - 4 1 . SWIHART, S. L. 1967. Hearing in butterflies (Nymphalidae: Heliconius, Ageronia). J. Ins. Physiol. 13:469-476. SWIHART, S. L. 1972. Modelling the butterfly visual pathway. J. Ins. Physiol. 18: 1915-1928. TURNER, J. R. G. 1977. Butterfly mimicry: T h e genetical evolution of an adaptation. Evol. Biol. 10: 1636-206. WELLING, E. C. 1959. Notes on butterfly migra­ tions in the peninsula of Yucatán. J. Lepidop. Soc. 13: 62-64. WESLEY, D. J., AND T

C. EMMEL. 1975.

The

chromosomes of Neotropical butterflies from Trinidad and Tobago. Biotropica 7: 2 4 - 3 1 . YOUNG, A. M. 1980a. Evolutionary responses by butterflies to patchy spatial distributions of resources in tropical environments. Acta Biotheor. 29: 37-64. YOUNG, A. M. 19806. The interaction of preda­ tors and "eyespot butterflies" feeding on rotting fruits and soupy fungi in tropical forests; variations on a theme developed by the Muyshondts and Arthur M. Shapiro. Entomol. Rec. J. Var. 92: 63-69. YOUNG, A. M. 1984. Ithomiine butterflies associ­ ated with non-antbird droppings in a Costa Rican tropical rain forest. J. Lepidop. Soc. 38: 61-63.

SWALLOWTAILS Papilionidae. L a r g e (WS of most 7 - 1 1 cm), colorful, a n d graceful in flight, t h e swallowtail b u t t e r ­ flies (so-called from t h e taillike extensions

SWALLOWTAILS

337

Figure 10.15. SWALLOWTAIL BUTTERFLIES (PAPILIONIDAE). (a) Kite (Eurytides bellerophon) (b) Kite (Eurytides philolaus), larva, (c) Giant swallowtail, pupa, (d) Giant swallowtail {Papilio toaos) (e) Giant swallowtail, larva, (f) Aristolochia swallowtail (Parides iphidamas). (g) Aristolochia swallowtail, larva. of t h e apices of t h e h i n d wings of s o m e g e n e r a ) a d o r n t h e Neotropics. Color pat­ t e r n s vary, b u t t h e r e is frequently a r e d tinted, eyelike spot at t h e i n n e r notch of the h i n d wing. T h e i r caterpillars a r e all n a k e d , w i t h o u t spines o r visible hairs b u t sometimes with tubercles, a n d a r e some­ what club s h a p e d , with t h e t h o r a x e n ­ larged. W h e n d i s t u r b e d , t h e larvae arch their backs a n d e v e r t a n o d o r i f e r o u s forked o r g a n ( o s m e t e r i u m ) from b e h i n d the h e a d which is a d e t e r r e n t device (Eisner et al. 1 9 7 1 , L ó p e z a n d Q u e s n e l 1970, Young et al. 1986). T h e chrysalids mimic wood f r a g m e n t s with their a n g u l a r form (two projecting points o n t h e h e a d ) and rough brown or greenish integument. T h e y rest u p r i g h t , fastened to a t e r m i n a l silk b u t t o n a n d l e a n i n g back into a silken girdle. T h e r e a r e t h r e e m a j o r types of swallow­ tails based o n f o r m a n d habits ( H a n c o c k 1983). T h e kites (pages, zebras, swordtails, Leptocircini, e.g., Eurytides; fig. 10.15a) a r e smaller t h a n most (WS 7 c m ) a n d have pale, thinly scaled, often white wings, with thin, transverse, black stripes; t h e tails a r e extra long a n d flexible. T h e larvae (fig. 10.15b) a r e usually g r e e n a n d s m o o t h , a n d they feed o n species of t h e c u s t a r d a p p l e family ( A n n o n a c e a e ) . T h e s e a r e forest dwellers, a n d t h e males a r e c o m m o n par­ ticipants in t h e clouds of butterflies seen d r i n k i n g from wet sand along water­ courses.

338

MOTHS AND BUTTERFLIES

T h e sun-loving t r u e swallowtails (Papilionini, Papilio; fig. 10.15d) a r e t h e largest of t h e g r o u p (WS to 12 cm) a n d variously m a r k e d , a l t h o u g h most a r e black with b r o a d bright yellow bars t h r o u g h the mid­ dle of t h e wings a n d crescent-shaped spots b o r d e r i n g t h e o u t e r m a r g i n of t h e hind wing. A few a r e "tiger m a r k e d " a n d mimic similarly colored heliconian a n d ithomiine models. Tails a r e lacking in t h e latter but are nearly always p r e s e n t in t r u e swallow­ tails, a l t h o u g h short. T h e r a r e Papilio homerus of Jamaica is t h e largest tailed swal­ lowtail in t h e world (WS t o 15 cm). T h e larvae (fig. 10.15e) a r e mottled brown and c r e a m streaked, simulating bird drop­ pings, a n d often c o n g r e g a t e for mutual protection. Food plants a r e varied but often a r e of t h e p e p p e r (Piperaceae) and citrus families (Rutaceae), t h e latter includ­ ing o r a n g e , l e m o n , a n d lime. O n citrus, those of certain species {Papilio cresphontes, P. andraemon, P. anchisiades) sometimes con­ stitute pests a n d a r e k n o w n to fruit grow­ ers as " o r a n g e p u p p i e s " o r " o r a n g e dogs" (Lawrence 1972). Papilio chrysalids often mimic b r o k e n twigs (fig. 10.15c). Aristolochias ("poison eaters," pharmac o p h a g o u s swallowtails, Troidini) a r e mod­ erate-sized (WS 7—9 cm), mostly without tails, a n d inky black with r e d o r magenta color fields in t h e c e n t e r of t h e hind wing a n d g r e e n (blue o r yellow) areas near the base of t h e fore wing {Parides; fig. 10.150 or dull, greenish-black with splotches oi

vellow mostly o n h i n d wings {Battus). These swallowtails a r e partial to s h a d e a n d 0 ¡ s t u r e a n d a r e c o n s u m m a t e forest in­ l e t s w h e r e t h e larvae (fig. 10.15g) feed o n their p i p e v i n e {Aristolochia, Aristolochiaceae) hosts. T h e s e plants contain toxic alkaloids s e q u e s t e r e d by t h e caterpillars and t r a n s m i t t e d to t h e adults, m a k i n g them u n p a l a t a b l e . Many serve as models in Batesian a n d Müllerian mimicry com­ plexes (Young 1971). T h e t u b e r c u l a t e lar­ vae are also p r o t e c t e d by these chemicals, a fact they seem to advertise with r e d - o r a n g e or yellow streaks, c o n s p i c u o u s against a n otherwise completely black body. T h e p u ­ pae are flared o u t laterally along t h e edges of the wing cases (Young 1977). T h e r e a r e m o r e t h a n ninety swallowtail species in Latin A m e r i c a ( d ' A b r e r a 1981, d'Almeida 1966); t h e family occurs as far south as central Chile a n d n o r t h e r n Patago­ nia (Slansky 1973). T h e biologies of most are still u n k n o w n in spite of t h e attention this attractive g r o u p h a s received from collectors a n d hobbyists. S o m e information is available on t h e several species, including Papilio homerus, above, c o n s i d e r e d possibly in danger of extinction by t h e I n t e r n a t i o n a l Union for t h e C o n s e r v a t i o n of N a t u r e (Col­ lins a n d Morris 1985).

References COLLINS, N. M., AND M. G. MORRIS.

1985.

Threatened swallowtail butterflies of the world: T h e IUCN Red Data Book. IUCN, Gland, Switzerland. D'ABRERA, B. 1981. Papilionidae. In B. d'Abrera, Butterflies of the Neotropical Region. Pt. I. Papilionidae and Pieridae. Lansdowne, East Melbourne, Australia. D'ALMEIDA, R. F. 1966. Catálogo dos Papi­ lionidae americanos. Soc. Brasil. Entomol., Sao Paulo. EISNER, T., A. F. KLUGE, M. I. IKEDA, Y. C. MEINWALD, AND J. MEINWALD. 1971. Sesqui-

terpenes in the osmeterial secretion of a papilionid butterfly, Battus polydamus. J. Ins. Physiol. 17: 245-250. HANCOCK, D. L. 1983. Classification of the

Papilionidae (Lepidoptera): A phylogenetic

approach. Smithsersia (Nat. Mus. Mon. Zim­ babwe) 2: 1-48. LAWRENCE, P. O. 1972. The Jamaican "orange dog," Papilio andraemon (Lepidoptera: Papi­ lionidae). Fla. Entomol. 55: 243-246. LÓPEZ, A., AND V. C. QUESNEL. 1970. Defensive

secretions of some papilionid caterpillars. Carib.J. Sci. 10: 5 - 7 . SLANSKY, JR., F. 1973 [1972]. Latitudinal gradi­ ents in species diversity of the New World swallowtail butterflies. J. Res. Lepidop. 11: 210-217. YOUNG, A. M. 1971. Mimetic associations in natural populations of tropical papilionid butterflies (Lepidoptera: Papilionidae). New York Entomol. Soc. J. 79: 210-224. YOUNG, A. M. 1977. Studies on the biology of Parides iphidamas (Papilionidae: Troidini) in Costa Rica. J. Lepidop. Soc. 31: 100-108. YOUNG, A. M., M. S. BLUM, H. M. FALES, AND

Z. BIAN. 1986. Natural history and ecological chemistry of the Neotropical butterfly Papilio anchisiades (Papilionidae). J. Lepidop. Soc. 40: 36-53.

METALMARKS Lycaenidae, Riodininae ( = Erycinidae, Nemeobiidae). Such a large p r o p o r t i o n of t h e species of this very diverse subfamily (over 9 0 % of the world's 1,500 o r so species, s e p a r a t e d into 150 genera) live in t h e Neotropics that they could b e c o n s i d e r e d t h e most characteristic butterflies of t h e r e g i o n . Most a r e small (WS 2 0 - 3 5 m m ) a n d delicately built, a n d their colors a n d shapes a r e so varied as to defy descrip­ tion. Many a r e vividly a n d e x t r e m e l y b e a u ­ tifully h u e d with metallic blues, d e e p scar­ let, g r e e n , white, a n d o t h e r colors, often in complex combinations a n d designs (e.g., Amarynthis menaria; fig. 10.16d). Many have metallic gold o r silver flecks o n the u n d e r s i d e s , a n d a few have eyespots. Some take t h e p a t t e r n s of distasteful m o d ­ els a m o n g t h e o t h e r L e p i d o p t e r a . T h e wings may b e r o u n d e d o r a n g u l a r a n d in some cases flamboyantly multitailed {Helicopis; fig. 10.16a), r e s e m b l i n g hairstreaks,

METALMARKS

339

or with single, long "swallowtails" (Chlo­ rinea; fig. 10.16c). A d u l t s a r e fast fliers generally b u t often settle r e p e a t e d l y o n the s a m e p e r c h , with their r e s p l e n d e n t wings o u t s t r e t c h e d . S o m e display a "false h e a d " posteriorly o n t h e u n d e r s i d e s of t h e wings, as d o t h e h a i r s t r e a k s (see hairstreaks, below) (Robbins 1985). T h e early stages of few a r e r e c o r d e d in the scientific l i t e r a t u r e . I n these, food plants a r e varied, a n d t h e larvae a r e elon­ gate a n d sluglike with small lateral e x p a n ­ sions held closely a p p r e s s e d to t h e substra­ t u m . T h e d o r s u m of t h e larva's p r o t h o r a x is thickened a n d rigid a n d usually b r o w n in contrast to t h e rest of t h e b o d y a n d often with h o r n l i k e projections o r large bristles o n either side. Most a r e greenish o r pale with m a n y short body hairs; s o m e have brilliantly colored p r o t u b e r a n c e s m a r k i n g e x t e r n a l glands that p r o d u c e e x u d a t e s av­ idly s o u g h t by ants (De Vries 1989, Ross 1985). S o m e m e t a l m a r k larvae a r e participants in mutualistic associations with ants (as in Juditha molpe; fig. 10.16b), obtaining their protection in e x c h a n g e for these sub­ stances ( B o u l a r d 1 9 8 1 , C a l l a g h a n 1977, Horvitz et al. 1987). It is suspected that t h e p h e n o m e n o n is w i d e s p r e a d t h r o u g h t h e subfamily. A n t s actually build shelters for o n e species (Ross 1966). T h e caterpillar of o n e species (Thisbe irenea) not only feeds o n leaf tissue b u t also d r i n k s t h e extrafloral nectar of its host plant, creating a conflict between plant a n d h e r b i v o r e for t h e atten­ tions of ants (De Vries a n d B a k e r 1989).

References BOULARD, M. 1981. Nouveaux documents sur

les chenilles de lycénes tropicaux. Alexanor 12: 135-140. CALLAGHAN, C. J.

1977. Studies on

restinga

butterflies. 1. Life cycle and immature biology of Menander felsina (Riodinidae), a myrmecophilous metalmark. J. Lepidop. Soc. 31: 173-182. DE

340

VRIES, P. 1989. T h e ant associated larval

MOTHS AND BUTTERFLIES

organs of Thisbe irenea (Riodinidae) and their effects on attending ants. Zool. J. Linnean Soc. 94: 379-393. DE VRIES, R J., AND 1. BAKER. 1989. Butterflv

exploitation of an ant-plant mutualism: Add ing insult to herbivory. New York Entornol Soc.J. 97: 332-340. HORVITZ, C. C , C. TURNBULL, AND D. J. HAR-

VEY. 1987. Biology of immature Eurybia elvinn (Lepidoptera: Riodinidae), a myrmecophilous metalmark butterfly. Entornol. Soc Amer. Ann. 80: 513-519. ROBBINS, R. K. 1985. Independent evolution of

"false head" behavior in Riodinidae 1 Lepidop. Soc. 39: 224-225. Ross, G. N. 1966. Life-history studies on Mexi­ can butterflies. IV T h e ecology and ethology of Anatole rossi, a myrmecophilous metalmark (Lepidoptera: Riodinidae). Entornol. Soc Amer. Ann. 59: 985-1004. Ross, G. N. 1985. T h e case of the vanishing caterpillar. Nat. Hist. 94(11): 48-55.

HAIRSTREAKS Lycaenidae, Lycaeninae. T h e Lycaeninae a r e poorly k n o w n in Latin America c o m p a r e d to their sister group, the m e t a l m a r k s . Currently, about 1,000 species have been discovered. W h e n all are described, it is estimated that their number will exceed t h e Riodininae. In t h e Neotropics, well over 90 percent of this subfamily of small butterflies (WS 1 5 - 4 0 m m ) is c o m p r i s e d of o n e tribe, the Eumaeini (Eliot 1973). T h e s e a r e the hairstreaks, typically with short hairlike a p p e n d a g e s e x t e n d i n g from a lobe at the r e a r of the h i n d wing, at t h e base of which are conspicuous, eyelike spots. T h e y rest with t h e h i n d wings a p p r e s s e d tightly over the back which they characteristically rub together, setting these "tails" in motion so that they resemble waving a n t e n n a e . T h e action is t h o u g h t to divert t h e attacks of p r e d a t o r s away from the t r u e head to these e x p e n d a b l e wing structures. T h e validity of t h e hypothesis has b e e n tested o n the c o m m o n "false h e a d hairstreak," Arawacus

pigure 10.16 LYCAENID BUTTERFLIES (LYCAENIDAE). (a) Multitailed metalmark (Helicopis ac/s). (b) Metalmark (Juditha molpe), larva being attended by ants of the genus Hypoclinea. (c) Longtailed metalmark (Chlorinea faunus). (d) Metalmark (Amarynthis menaria). (e) False head hairstreak ¡Arawacus aetolus).

oetolus (fig- 10.16e) (Robbins 1980, 1981). On the u p p e r sides, they a r e solid colored and plain, a l t h o u g h m a n y o t h e r s display iridescent blues a n d g r e e n a n d o t h e r s a r e vividly p a t t e r n e d . T h e early stages of t h e majority of t h e species a r e u n k n o w n . T h e few larvae that are known a r e mostly sluglike with a small retracted h e a d a n d a r e s o m e w h a t flattened, some very m u c h so (Callaghan 1982). Most feed on a variety of d i c o t y l e d o n e o u s plants, often the flowers, a n d fruit, b u t a few (e.g., (Miaría) may h a v e specialized food prefer­ ences a m o n g o t h e r plants, such as orchids. Some are associated with ants symbiotically and have a t h i c k e n e d , t o u g h cuticle, p r e ­ sumably to p r o t e c t t h e m from attack w h e n entering a n t nests to feed o n their larvae o r other guests, such as coccids. T h e s e also exude substances from special integu­ mentary glands to attract ants for protec­ tion o r to entice t h e m to carry t h e m to their nest. Some a r e agricultural pests, such as the pineapple h a i r s t r e a k (Tmolus basilides) whose larvae e a t t h e flowers a n d b o r e into the developing fruit of p i n e a p p l e ( H a r r i s 1927), b u t h a i r s t r e a k s a r e largely benign

ELIOT, J. N. 1973. T h e higher classification of

insects.

Like their relatives, t h e swallowtails, they possess a fully functional pair of front legs, a n d t h e p u p a is attached at t h e tip of the a b d o m e n a n d held u p r i g h t by a silken girdle passing a r o u n d t h e t h o r a x . T h e larvae a r e largely s m o o t h except for mi­ n u t e papillae in t h e i n t e g u m e n t .

References .CALLAGHAN, C. J. 1982. Notes on the immature

biology of two myrmecophilous Lycaenidae: Juditha molpe (Riodininae) and Panthiades billas (Lycaeninae). J. Res. Lepidop. 20: 36-42.

the Lycaenidae (Lepidoptera): A tentative arrangement. Brit. Mus. Nat. Hist. Entornol. Bull. 28: 373-505, pis. 1-6. HARRIS, W. V 1927. On a lycaenid

butterfly

attacking pineapples in Trinidad, B.W.I. Bull. Entornol. Res. 18: 183-188, pis. 7 - 8 . ROBBINS, R. K. 1980. T h e lycaenid "false head" hypothesis: Historical review and quantitative analysis. J. Lepidop. Soc. 34: 194-208. ROBBINS, R. K. 1981. T h e "false head" hypothe­ sis: Predation and wing pattern variation of ly­ caenid butterflies. Amer. Nat. 118: 770-775.

WHITES AND SULFURS Pieridae. Spanish: Isocas (General, larvae), pirpintos (Argentina). In these familiar butterflies, yellow, o r a n g e , a n d white a r e t h e p r e d o m i n a t i n g colors. T h e y f r e q u e n t flowers of o p e n glades a n d clearings a n d a r e a m o n g t h e sun-loving insect t h r o n g s that t e n d the blossoms of t h e forest canopy. T h e y also r a n g e widely from the coastal deserts a n d h u m i d s w a m p l a n d to well above tree line in t h e p á r a m o s a n d rocky slopes of s n o w c a p p e d peaks.

WHITES AND SULFURS

341

c Figure 10.17 BUTTERFLIES, (a) Great southern white (Ascia monuste, Pieridae). (b) Cloudless sulfur (Phoebis sennae, Pieridae), larva, (c) Cloudless sulfur, pupa, (d) Cloudless sulfur, adult. (e) Alpine pierid (Catasticta semiramis, Pieridae). (f) Silver-winged Butterfly (Argyrophorus argenteus, Nymphalidae), larva, (g) Silver-winged butterfly, adult. Several species, especially t h e cloudless sulfurs, Phoebis ( d ' A l m e i d a 1940), partici­ p a t e in m i g r a t o r y swarms. T h e males a r e avid p u d d l e r s . Many mimics also a r e found in t h e family. T h e larvae of a few (gusanos de la col, isocas del repollo, lagartas da hortalica, lagartas da couve) a r e n o t o r i o u s pests, for e x a m p l e , the w i d e s p r e a d , native g r e a t s o u t h e r n white (Ascia monuste; fig. 10.17a) a n d , in Chile only w h e r e it was accidentally intro­ d u c e d , t h e E u r o p e a n c a b b a g e butterfly (Pieris brassicae) ( G a r d i n e r 1974). T h e lar­ vae of Colias lesbia (isoca de la alfalfa, cuncuna) destroy alfalfa in A r g e n t i n a . Adults of t h e g e n u s Catasticta (fig. 10.17e) a r e atypical pierids, h a v i n g a check­ e r e d wing p a t t e r n a n d o c c u r r i n g only at high elevations from Mexico to t h e A n d e s , w h e r e t h e greatest n u m b e r of species a r e f o u n d . T h e larvae feed o n L o r a n t h a c e a e ; they a r e g r e g a r i o u s a n d r e s p o n d by h e a d r e a r i n g w h e n molested. T h e r e a r e a p p r o x i m a t e l y 400 species of pierids in Latin A m e r i c a ( d ' A b r e r a 1981).

References D'ABRERA, B. 1981. Pieridae. In B. d'Abrera, Butterflies of the Neotropical Region. Pt. I. Papilionidae and Pieridae. Lansdowne, East Melbourne, Australia. Pp. 80—165. D'ALMEIDA, R. F. 1940. Revisáo do género Phoebis Hiibn. (Lepidopt Pieridae). Arq. Zool. Est. Sao Paulo 1: 67-148. GARDINER, B. O. C. 1974. Pieris brassicae L. established in Chile; another palaearctic pest

342

MOTHS AND BUTTERFLIES

crosses the Atlantic (Pieridae). J. Lepidop Soc. 28: 269-277.

Cloudless Sulfur Pieridae, Pierinae, Coliadini, Phoebis sennae. T h i s is probably t h e most c o m m o n and widespread of the Neotropical pierids. A moderately large butterfly (WS 6 cm), its intense greenish-yellow wings, punctuated o n t h e u n d e r s i d e s with scattered patches or lines of r e d d i s h - b r o w n scales, immedi­ ately identify it (fig. 10.17d). T h e u p p e r sides of t h e male wings a r e immaculate; those of the female a r e b o r d e r e d by a b r o k e n black m a r g i n a n d an irregular black spot midway n e a r t h e leading edge of the fore wing. T h e species is strongly migratory. Vast clouds m o v i n g t h r o u g h m a n y parts of South a n d Central A m e r i c a a n d even over the o p e n sea in the Caribbean are of frequent occurrence. T h e i m m a t u r e s a r e well known, the larva (fig. 10.17b) being almost a pest on l e g u m i n o u s o r n a m e n t a l s in t h e genus Cas­ sia. It feeds on Calliandra a n d Inga as well. It grows to 35 to 40 millimeters a n d is elongate (slightly t a p e r e d at both ends), a n d its skin is transversely wrinkled or r i d g e d . It is generally g r e e n to yellowg r e e n with a lateral yellow line. T h e upper half of the body is speckled with small purplish dots in each of which is a minute black wart b e a r i n g fine white hairs. T h e

p U pa (fig- 10.17c) is w e d g e s h a p e d , with greatly e x p a n d e d wing cases that form a sharp heel w h e r e they j o i n from each side. The very d e e p wing cases give it an a r c h e d backed a p p e a r a n c e . A l t h o u g h pink p u p a e are known, this stage is typically g r e e n with a whitish-yellow longitudinal lateral stripe a n d d a r k m i d d o r s a l line.

BRUSH-FOOTED BUTTERFLIES Nymphalidae. Butterflies classified into this diverse fam­ ily, which is c o n s i d e r e d h e r e in a b r o a d sense, include all of t h e succeeding sub­ families, which s o m e a u t h o r s consider sepa­ rate families.

Satyrs Nymphalidae, Satyrinae. Satyrs a r e almost all d r a b d e n i z e n s of t h e forest floor. M a n y d o have bright colors o n the hind wings a n d eyespot p a t t e r n s ventrally. A few a r e t r a n s p a r e n t , like "glassywings" (Haeterini). T h e wings a r e also soft and thinly scaled a n d with few exceptions have the bases of t h e veins of t h e fore wing inflated to f o r m a conspicuous swelling. The host plants of almost all species a r e grasses a n d b a m b o o s . Silver-winged Butterfly Nymphalidae, Satyrinae, Argyrophorus argenteus. Spanish: M a r i p o s a plateada, cinta plateada. This is an exceedingly beautiful, m e d i u m sized (WS 4 cm) butterfly of t h e lower slopes of t h e P a t a g o n i a n A n d e s , t h e u p p e r Wing surfaces b e i n g solid, s h i n i n g silver (fig. 10.17g). I t is a difficult butterfly to catch because of its erratic flight over its brushland habitat. T h e i m m a t u r e s a r e Poorly k n o w n ; t h e larva (fig. 10.17f) is ught yellowish-brown with longitudinal •tapes a n d a p p a r e n t l y feeds o n t h e grass

Stipa (coirón), o n which females have b e e n observed to oviposit (Shapiro 1982).

Reference SHAPIRO, A. M. 1982. Notas sobre los estados inmaduros de la mariposa plateada, Argyro­ phorus argenteus Blanchard (Lepidoptera: Satyridae). Soc. Mexicana Lepidop. Rev. 7: 29-31.

Monarch Butterflies N y m p h a l i d a e , D a n a i n a e , Danaus. Monarcas.

Spanish:

Like ithomiines a n d s o m e o t h e r tropical L e p i d o p t e r a , male m o n a r c h butterflies seek organic c o m p o u n d s , called pyrrolizid i n e alkaloids, from w i t h e r e d plants, most c o m m o n l y of t h e g e n e r a Heliotropium, Eupatorium, Senecio, a n d Tournefortia. T h e s e substances a r e imbibed with t h e t o n g u e a n d form substrates in t h e butterfly's b o d y from which a r e synthesized special a p h r o disiacal p e r f u m e s that h e l p attract females a n d e n s u r e success in c o u r t s h i p a n d mat­ ing. T h e chemicals (di-hydro-pyrrolizidines) a r e secreted by p o u c h e d glands n e a r t h e c e n t e r of t h e h i n d wings from which they a r e first picked u p a n d t h e n disseminated into t h e air by p r o t r u s i b l e fan-shaped hair b r u s h e s located at t h e tip of t h e a b d o m e n (Brower a n d J o n e s 1965). T h e role of these chemicals in t h e court­ ship process is still n o t clear. Individuals of this g e n u s also s e q u e s t e r cardiac glycosides in their bodies ( B o p p r é 1978). T h e s e c o m p o u n d s a r e a c q u i r e d by the larvae w h e n feeding o n their asclepiad host plants a n d a r e toxins capable of i n d u c ­ ing severe intestinal complaints in birds o r o t h e r animals that swallow t h e m (Roeske et al. 1976). T h u s , t h e adults gain protection for themselves a n d also serve as a m o d e l for a n u m b e r of o t h e r mimetic butterflies a n d m o t h s . N o t every milkweed (Asclepias) food plant provides cardiac glycosides, a n d butterfly individuals that eat these a r e n o t poisonous. T h e r e a r e four Danaus species r a n g i n g

BRUSH-FOOTED BUTTERFLIES

343

References ACKERY, P. R-, AND R. 1. VANE-WRIGHT. 1984.

Milkweed butterflies, their cladistics and biol­ ogy. Brit. Mus. Nat. Hist., London. See esp. pp. 106-110. BLAKLEY, N. R., AND H. DINGLE. 1978. Competi­

Figure 10.18 NYMPHALID BUTTERFLIES (NYMPHALIDAE). (a) Monarch (Danaus plexippus). (b) Monarch, larva, (c) Monarch, pupa, (d) Cracker (Hamadryas feronia). (e) Head-for-tail (Coloburá dirce). (f) Malachite green (Siproeta stelenes).

tion: Butterflies eliminate milkweed bugs from a Caribbean Island. Oecologia 37: 133-136. goppRÉ, M. 1978. Chemical communication, plant relationships, and mimicry in the evolu­ tion of the danaid butterflies. Entomol. Exper. Appl. 24: 64-77. BROWER, L. P. 1985. New perspectives on the

t h r o u g h o u t t h e Neotropics, t h e most wide­ s p r e a d b e i n g t h e " t r u e " m o n a r c h (D. plex­ ippus; fig. 10.18a), which exists as various subspecific a n d local forms (the s o u t h e r n / ) . erippus is sometimes c o n s i d e r e d as a sepa­ rate species). D. gilippus a n d relatives a r e t h e so-called q u e e n s , recognized by d a r k , brownish g r o u n d color in t h e wings. T h e w i d e s p r e a d soldier (D. eresimus) has found its way to several islands of t h e Antilles; t h e J a m a i c a n m o n a r c h (D. deophile) is restricted to Jamaica a n d Hispaniola. All m o n a r c h butterflies (Ackery a n d Vane-Wright 1984) a r e frost-sensitive a n d essentially tropical butterflies (Young 1982). T h e n o r t h e r n subspecies D. plexippus plexippus p e n e t r a t e s t h e t e m p e r a t e latitudes of N o r t h A m e r i c a (to s o u t h e r n C a n a d a ) b u t only as a s u m m e r visitor. I n t h e fall m o n t h s , its n o r t h e r n p o p u l a t i o n s m i g r a t e south­ ward over two major r o u t e s , a Pacific flyway to t h e west of t h e Rocky M o u n t a i n s a n d a continental flyway t o t h e east (Brower 1985). T h e o v e r w i n t e r i n g sites of most of the latter h a v e only b e e n recently discov­ e r e d in a r e m o t e m o u n t a i n r a n g e o n t h e Mexican plateau (Calvert a n d B r o w e r 1986, U r q u h a r t a n d U r q u h a r t 1976, B r o w e r et al. 1977). T h i s roosting p h e n o m e n o n is so spectacular t h a t t h e butterflies a r e n o w p r o t e c t e d by presidential d e c r e e (López Portillo 1980, N o r m a n 1986). A civic g r o u p , Pro M o n a r c a , has even b e e n f o r m e d which is d e d i c a t e d t o t h e conservation of t h e o v e r w i n t e r i n g sites ( O g a r r i o 1984). T h e p h e n o m e n o n of r o o s t i n g m o n a r c h s h a s

344

MOTHS AND BUTTERFLIES

b e e n k n o w n for a long time to t h e local residents, w h o refer to t h e m as palomas de los Santos. T h e butterflies a r e believed to arrive o n t h e Day of t h e Dead (All Soul's Day, N o v e m b e r 2). Some of t h e N o r t h A m e r i c a n migrants of D. plexippus overwinter in t h e West Indies w h e r e t h e larval host is t h e universal red-flowered weed, A. curassavica, a n d Calotropis procera, a n African i m m i g r a n t . On Barbados, t h e larvae have c o n s u m e d all traces of t h e former, which is also t h e food plant of t h e milkweed bugs (Oncopeltus). T h e bugs, as a c o n s e q u e n c e , a r e n o longer f o u n d t h e r e (Blakley a n d Dingle 1978). T h e o t h e r Danaus species, a n d t h e south­ e r n subspecies of D. plexippus, a r e funda­ mentally sedentary. Most Danaus a r e rich rusty brown in general color with black wing veins. T h e wing a p e x a n d m a r g i n s a r e also black, containing small white spots. T h e related large tiger (Lycorea cleobaea, formerly L. ceres) a n d L. ilione, however, deviate strongly from this typical p a t t e r n , having tiger colors a n d a clear wing p a t t e r n , respectively, like the heliconians, ithomiines, a n d other L e p i d o p t e r a that they j o i n in Miillerian mimicry complexes. T h e larvae a r e colored in narrow m u l t i h u e d circular b a n d s a n d bear fleshy tentacular structures o n various segments (fig. 10.18b). T h e p u p a e a r e compact, smooth, a n d of various colors; that of D. plexippus, e m e r a l d g r e e n with golden spots, is well k n o w n for its beauty (fig. 10.18c).

migration biology of the monarch butterfly, Danaus plexippus L. In M. A. Rankin, Migra­ tion: Mechanisms and adaptive significance. Contrib. Mar. Sci. Suppl. 27:748-785. BROWER, L. P., W. H. CALVERT, L. E. HENDRICK,

AND J. CHRISTIAN. 1977. Biological observa­ tions on an overwintering colony of monarch butterflies (Danaus plexippus, Danaidae) in Mexico. J. Lepidop. Soc. 31: 232-242. BROWER, L. P., AND M. A . J O N E S . 1965. Precourt-

ship interaction of wing and abdominal sex glands in male Danaus butterflies. Royal Entomol. Soc. London Proc. 40: 147-151. CALVERT, W. H., AND L. P. BROWER. 1986. T h e

location of monarch butterfly (Danaus plex­ ippus L.) overwintering colonies in Mexico in relation to topography and climate. J. Lepi­ dop. Soc. 40: 164-187. LÓPEZ PORTILLO, J. 1980. Decreto por el que

por causa de utilidad pública se establece zona de reserva y refugio silvestre los lugares donde la mariposa conocida con el nombre de "Monarca" hiberna y se reproduce. Diario Oficial (México) 9 April 1980. Pp. 7 - 8 . NORMAN, C 1986. Mexico acts to protect over­ wintering monarchs. Science 233: 1252-1253. OGARRIO, R. 1984 [1981]. Development of the civic group, Pro Monarca, A . O , for the pro­ tection of the monarch butterfly over­ wintering grounds in the Republic of México. Átala 9: 11-13. ROESKE, C M., J. M. SEIBER, L. P. BROWER, AND

C. M. MOFFITT. 1976. Milkweed cardenolides and their comparative processing by mon­ arch butterflies. Rec. Adv. Phytochem. 10: 93-167. URQUHART, F. A., AND M. R. URQUHART. 1976.

The overwintering site of the eastern popula­ tion of the monarch butterfly (Danaus p. plexippus: Danaidae) in southern Mexico. J. Lepidop. Soc. 30: 153-158. YOUNG, A. M. 1982. An evolutionary-ecological

model of the evolution of migratory behavior

in the monarch butterfly and its absence in the queen butterfly. Acta Biotheor. 31: 219-237.

Nymphalines Nymphalidae, Nymphalinae. M e m b e r s of this subfamily a r e varied, small t o m e d i u m size, with r e d u c e d , brushlike forelegs a n d strongly clubbed a n t e n n a e . T h e central cell of t h e h i n d wing venation is always o p e n , a n d t h e wing has a c u p p e d depression o n t h e i n n e r m a r g i n t o accommodate the abdomen when the wings a r e closed. T h e larvae a n d p u p a e a r e likewise diverse, t h e f o r m e r usually spiny. Crackers Nymphalidae, Nymphalinae, Ageroniini, Hamadryas. Spanish: Calicoes (General), t r o n a d o r a s (Mexico), tabletas (Peru), cascabeles (Venezuela), s o ñ a d o r a s (Costa Rica), gritonas (Colombia). Portuguese: Assentas p á u , matracas, angolinhas, etc. (Brazil). T h e s e swift-flying a n d p u g n a c i o u s b u t t e r ­ flies (Perry 1964) invariably rest o n tree t r u n k s 1 to 10 m e t e r s above t h e g r o u n d , h e a d d o w n , wings s p r e a d , their colors m e r g i n g completely with t h e mottled b a r k b a c k g r o u n d . T h e edges of t h e wings a r e pressed flat to t h e b a r k so that n o s h a d o w is t h r o w n . O n t h e a p p r o a c h of a n o t h e r individual, a male cracker lurches from its p e r c h a n d fights off t h e i n t r u d e r . Such e n c o u n t e r s are usually p u n c t u a t e d by crackling o r clicking noises that t h e butterflies t h e m ­ selves emit. T h e s o u n d s a r e audible t o t h e h u m a n ear, in still air, u p t o several m e t e r s away. Both sexes i n d u l g e in these fast aerial pursuits a n d place o t h e r kinds of butterflies a n d even h u m a n s (orig. obs.) u n d e r "attack." O n e a u t h o r (Ross 1963) thinks that this e x t r e m e wariness m a y b e wholly a m e a n s of escaping p r e d a t o r s a n d not a n instance of t r u e territorial behavior. T h e m e c h a n i s m of clicking is n o t well u n d e r s t o o d . S h a r p contact b e t w e e n certain thoracic sclerites d u r i n g i r r e g u l a r wing

BRUSH-FOOTED BUTTERFLIES

345

beats a p p e a r s to emit t h e actual s o u n d ( H a n n e m a n n 1956, Swihart 1967). T h e butterflies often choose p e r c h e s o n w o u n d e d trees oozing f e r m e n t i n g s a p , on which they feed. T h e y a r e also fond of juices e x u d i n g from r o t t i n g food, carrion, a n d o t h e r moist o r g a n i c matter. T h e wings of c r a c k e r s a r e m a r k e d on the u p p e r sides with a l t e r n a t i n g , zigzag lines of b r o w n , o r b l u e a n d c r e a m crossed by t h e d a r k wing veins, t o g e t h e r p r o d u c i n g an intricate, irregular, mosaic o r checker­ b o a r d p a t t e r n (fig. 10.18d). T h i s is b r o k e n by a s u b m a r g i n a l row of spots, which a r e smaller anteriorly o n t h e fore wing a n d gradually increase to fairly conspicuous eyespots posteriorly o n t h e h i n d wing. T h e twenty species a r e all m e d i u m - s i z e d (WS 5—6 cm) a n d fairly c o m m o n t h r o u g h o u t the Neotropics in d r y to wet forest habitats ( J e n k i n s 1983). C r a c k e r caterpillars mostly feed on toxic vines in t h e e u p h o r b i a family, r e c o r d e d spe­ cies b e i n g Dalechampia scandens a n d Tragia volubilis. T h e y grow to 30 to 35 millimeters (BL) a n d a r e s p i n e d all o v e r like m a n y o t h e r n y m p h a l i d larvae, b u t t h e m a i n shaft of each spine process is fine a n d t h e lateral barbs l o n g a n d arising n e a r t h e base so that the o r g a n a p p e a r s b r a n c h e d in stellate fash­ ion. T h e p a i r e d h e a d spines a r e e x t r a long a n d decidedly c l u b b e d . B r i g h t colors deco­ r a t e t h e skin; c r e a m , r e d , g r e e n , o r a n g e , a n d d a r k lines r u n a l o n g t h e back a n d sides. T h e chrysalids mimic d e a d leaves, t h e h e a d manifesting a p a i r of very p r o m i n e n t flat­ t e n e d flange-shaped e x t e n s i o n s with scal­ loped e d g e s . T h e y h a n g straight d o w n from its t e r m i n a l s u p p o r t a n d w h e n tou­ c h e d a r e capable of violent, s n a p p i n g move­ m e n t s ( M u y s h o n d t a n d M u y s h o n d t 1975a, 19756, 1975c; Y o u n g 1974).

References HANNEMANN, H. J. 1956. Über pterotarsale Stridulation und einige andere Arten der Lauterzeugung bei Lepidopteren. Deutche Entomol. Zeit., N.F., 3(1): 14-27.

346

MOTHS AND BUTTERFLIES

JENKINS, D. W. 1983. Neotropical Nymphalidae 1. Revision of Hamadryas. Allyn Mus. Bull 811-146. ' : MUYSHONDT, A., AND A. MUYSHONDT, JR. 1975.

Notes on the life cycle and natural history of butterflies of El Salvador. IB. Hamadryas februn (Nymphalidae-Hamadryadine). New York En tomol. Soc.J. 83: 157-169. MUYSHONDT,

A.,

AND A.

MUYSHONDT,

I»

19756. Notes on the life cycle and natural history of butterflies of El Salvador. Hjj Hamadryas guatemalena Bates (NymphalidaeHamadryadinae). New York Entomol. Soc I 83: 170-180. 'J' MUYSHONDT, A., AND A. MUYSHONDT, JR. 1975c

Notes on the life cycle and natural history of butterflies of El Salvador. I1IB. Hamadryas amphinome L. (Nymphalidae-Hamadryadinae) New York Entomol. Soc.J. 83: 181-191. PERRY, R. 1964. Notes on the genus Ageronia and the butterflies of Poponté. Entomologist 97: 140-141. Ross, G. M. 1963. Evidence for lack of territoriality in two species of Hamadryas (Nymphal­ idae). J. Res. Lepidop. 2: 241-246. SWIHART, S. L. 1967. Hearing in butterflies (Nymphalidae: Heliconius, Ageronia). J. Ins. Physiol. 13: 469-476. YOUNG, A. M. 1974. On the biology of Hama­ dryas februa (Lepidoptera: Nymphalidae) in Guanacaste, Costa Rica. Zeit. Angewan. Entomol. 76: 380-393. Head-for-Tail Butterfly N y m p h a l i d a e , N y m p h a l i n a e , Nymphalini, Colobura dirce. Zebra (Trinidad). T h i s butterfly is a lover of oozing, ferment­ ing fruit a n d sap a n d s p e n d s m u c h of its time in w o o d e d areas, sitting on food that has fallen to t h e g r o u n d o r p e r c h i n g on the t r u n k s of t h e trees, h e a d d o w n w a r d with wings tightly closed. I n t h e latter position, the full confusing effect of t h e pattern on the u n d e r s i d e s of t h e wings can be seen (fig. 10.18e). T h e illusion of t h e head and tail reversed (tergiversation) is produced by an e n l a r g e m e n t of t h e tip of the hind wing with a central ocellate spot, forming a false h e a d a n d eye; also, vertical bars of black at t h e base of t h e wing toward the t r u e h e a d simulate a b d o m i n a l segments. Presumably, this p a t t e r n is a deception directing t h e strikes of bird beaks away

from t h e vital h e a d a n d body. T h e butter­ fly also takes flight at an instant, scurrying through t h e a i r a n d m a k i n g a rustling noise with its wings. T h u s , its behavior as well as its f o r m a n d color afford it protec­ tion from v e r t e b r a t e p r e d a t o r s . T h e solid d a r k b r o w n of t h e u p p e r surfaces is b r o k e n only by a conspicuous yellow b a r crossing t h e m i d d l e of t h e fore wing obliquely. It is a m e d i u m - s i z e d butter­ fly (WS 1 1 - 1 2 cm). T h e life cycle is well k n o w n (Beebe 1952, M u y s h o n d t a n d M u y s h o n d t 1976). The m a t u r e larva (BL 36 m m ) , of typical nymphalid f o r m , is velvety black o r g r e e n with contrasting yellow areas a r o u n d t h e spiracles a n d white to bright yellow stellately b r a n c h e d spines. It has an eversible fingerlike g l a n d o n t h e u n d e r s i d e of the neck, whose function is u n k n o w n . T h e leaves of t h e cecropia tree a r e its food. T h e chrysalid (BL 3 cm) resembles a frag­ mented wood chip with its r o u g h e n e d exterior, j a g g e d d o r s u m , a n d light b r o w n color. It h a n g s from twigs by a fastening only at the tip of t h e a b d o m e n .

References BEEBE, W. 1952. A contribution to the life history of Colobura (Gynaecia auct.) dirce dirce (Linnaeus) butterfly. Zoológica 37: 199-202.

b a r of light g r e e n a n d small circular spots of t h e same color o n t h e o u t e r half of t h e h i n d wings (fig. 10.18f). T h i s coloration is strikingly similar to that of t h e u n r e l a t e d heliconian Philaethria dido (fig. 10.21a). T h e u n d e r s i d e s a r e richly m a r b l e d with g r e e n a n d b r o w n blotches, p u n c t u a t e d with black lines a n d ovals. Adults feed o n a variety of flowers b u t also c o m m o n l y take liquids from rotten fruit a n d even fresh e q u i n e o r bovine d u n g . T h e m a t u r e larva is dull black, of m o d e r ­ ate size (BL 50—53 m m ) , a n d liberally spined; t h e h e a d spines a r e decidedly longer t h a n those of t h e body a n d clubbed. Food plants a r e diverse species of the family Acanthaceae, often small semiwoody h e r b s (Justicia, Ruellia, Blechum) g r o w i n g as weeds in disturbed places such as wood lots, clear­ ings, a n d coffee plantations. T h e p u p a is a b o u t 30 millimeters long, f r e e - h a n g i n g from a black t e r m i n a l stalk, a n d generally translucent light g r e e n (and often covered with a white bloom). T h e r e a r e short, paired h e a d projections, short spines on t h e back of t h e a n t e r i o r a b d o m i ­ nal segments, a n d sparse black specks o n the skin. T h e species has g o n e u n d e r t h e n a m e "Metamorpha" o r "Victorina" steneles [sic] in the earlier literature.

MUYSHONDT, JR., A., AND A. MUYSHONDT. 1976.

Notes on the life cycle and natural history of butterflies of El Salvador. IC. Colobura dirce L. (Nymphalidae-Coloburinae). New York Ento­ mol. Soc. J. 84: 2 3 - 3 3 . Malachite Green Nymphalidae, N y m p h a l i n a e , N y m p h a l i n i , Siproeta stelenes. B a m b o o p a g e

(Trinidad). This is a n o t h e r w i d e s p r e a d a n d c o m m o n butterfly associated with forest clearings throughout C e n t r a l A m e r i c a a n d m u c h of northern S o u t h A m e r i c a (Young a n d Muyshondt 1973). It is m o d e r a t e l y large (WS 75 m m ) , with t h e u p p e r wing surfaces Mack except for a b r o a d , b r o k e n m e d i a n

Reference YOUNG,

A.

M.,

AND A.

MUYSHONDT.

1973.

Ecological studies of the butterfly Victorina stelenes (Lepidoptera: Nymphalidae) in Costa Rica and El Salvador. Stud. Neotrop. Fauna 8: 155-176. Number Butterflies Nymphalidae, Nymphalinae, C a t a g r a m m i n i , Diaethria a n d relatives. Spanish: O c h e n t a y ochos, o c h e n t a y nueves (General). Portuguese: Oitenta-eoitos, oitenta-e-noves, cruzeiros d o sul (Brazil). Eighty-eights, eighty-niners. While varied in details, t h e p a t t e r n s of t h e u n d e r s i d e s o n t h e h i n d wings of this famil-

BRUSH-FOOTED BUTTERFLIES

347

common a n d c o n s p i c u o u s d i u r n a l Lepi­ doptera e n c o u n t e r e d in the New World

Figure 10.19 NYMPHALID BUTTERFLIES (NYMPHALIDAE). (a) "89" (Diaethria clymena). (b) Red anartia (Anartia amathea). (c) Leaf butterfly (Memphis arachne). (d) Cecropia butterfly (Historis odius), larva, (e) Cecropia butterfly, pupa, (f) Cecropia butterfly, adult. iar g r o u p of m e d i u m - s i z e d (WS 3 - 5 cm), entirely Neotropical n y m p h a l i d s always consists of concentric d a r k rings o n a gray to yellow b a c k g r o u n d . T h e i n n e r m o s t of these are s e p a r a t e d into two figure eights (although o n e p u p i l of t h e a n t e r i o r figure may b e fused with t h e other, f o r m i n g m o r e of a "9"). With a little imagination, o n e can read 88, 89, 80, o r o t h e r n u m b e r s from the wing (fig. 10.19a), a l t h o u g h m a n y see let­ ters instead ("BD"). O n the u p p e r sides, the black b a c k g r o u n d color of the wings is i n t e r r u p t e d by a d i a g o n a l b a r of metallic blue or g r e e n a n d / o r b r o a d fields of these colors plus r e d or yellow. T h e r e a r e s o m e fifty species in this g r o u p (Dillon 1948), d i s t r i b u t e d widely t h r o u g h o u t the r e g i o n , mainly in forest habitats. A few species in the related a n d generally s i m i l a r - a p p e a r i n g g e n e r a Callicore, Paulo gramma, Perisama, Dynamine, Callidula, a n d Catacore h a v e numerological h i n d wings as well, b u t m o r e often, they sport b a r r e d , leaflike, o r o t h e r p a t t e r n s . In the past, m a n y of these w e r e l u m p e d into the catch-all g e n u s "Catagramma." T h e early stages of a few species a r e k n o w n ( M u y s h o n d t 1975). T h e m a t u r e larva of Diaethria asíala is m o r e o r less typical; it grows to m o d e r a t e size (BL 2 5 27 m m ) a n d is u n a r m e d except for the h e a d spines, which a r e very long (a t h i r d of the body length) a n d h a v e s t r o n g lateral b r a n c h e s . T h e b o d y is light g r e e n a n d covered with tiny white warts a n d longitudi­ nal rows of yellow tubercles. It exhibits an

348

MOTHS AND BUTTERFLIES

active defense behavior w h e n disturbed r e a r i n g its h e a d a n d striking with its horns. Food plants a r e vines in the family Sapindaceae (Serjania, Cardiospermum, Urvillea, etc.), a l t h o u g h the e u p h o r b s Sapium a n d Dalechampia are r e c o r d e d for Dyna­ mine. T h e s e plants have p o i s o n o u s proper­ ties that they i m p a r t to the larvae feeding on them. T h e chrysalid is g r e e n with dark, veinlike m a r k s on the wing cases and lateral, light g r e e n lines. It attaches to the u n d e r s i d e of a leaf, only by t h e tail, but hangs closely a p p r e s s e d to the surface. It may p r o d u c e a faint c r e a k i n g s o u n d by wiggling sideways or c o n t r a c t i n g its abdo­ m e n accordionlike.

References DILLON, L. S. 1948. The tribe Catagrammini (Lepidoptera: Nymphalidae). Pt. I. The ge­ nus Catagramma and allies. Read. Pub. Mus. Art Gal. Sci. Pub. 8: 1-v, 1-113. MUYSHONDT, A. 1975. Notes on the life cycle and natural history of butterflies of El Salvador. VIA. Diaethria astala Guérin. (Nymphalidae— Callicorinae). New York Entomol. Soc. J. 83: 10-18. Peacock Butterflies N y m p h a l i d a e , N y m p h a l i n a e , Nymphalini, Anartia. Any of the t h r e e continental species in this g e n u s (Silberglied et al. 1979) might be f o u n d in a wayside flower p a t c h o r dis­ t u r b e d sunny clearing a n y w h e r e in the h u m i d lowlands. T h e y a r e a m o n g the most

tropics. T h e fatima (A. fatima) (Young a n d Stein 1976), r a n g e s from Mexico to P a n a m a , where it is r e p l a c e d by its close relative, the similar r e d anartia (also called t h e coolie or t o m a t o , A. amathea; fig. 10.19b), whose distribution c o n t i n u e s through South A m e r i c a to n o r t h e r n A r g e n t i n a . Two r a r e r species, A. chrysopelea a n d A. lytrea, are of local o c c u r r e n c e in t h e larger islands of the West Indies. T h e fatima a n d r e d anartia have the same basic coloration, t h e u p p e r wing surface basically d a r k b r o w n with trans­ verse pale b r o w n a n d r e d b a n d s . T h e essential difference c o n c e r n s the extent of a red area inside t h e o u t e r m o s t light b a n d : here t h e r e a r e j u s t a few spots in the h i n d wing in the fatima, b u t this is a b r o a d field extending across b o t h fore a n d h i n d wings in the r e d a n a r t i a . T h e o u t e r b a r may be white or yellow in the f o r m e r species, a dimorphic characteristic a p p a r e n t l y deter­ mined genetically a n d with possible (but not probable?) implications for m a t e selec­ tion a n d survival ( E m m e l 1972, Silberglied e t a l . 1979). T h e white peacock (biscuit, A. jatrophae) has the widest distribution of the peacocks, occurring t h r o u g h o u t t h e Neotropics, in­ cluding the Antilles b u t e x c e p t i n g the high Andes a n d Chile. Its wings a r e off-white to gray, with c o m p l e x zigzag lines crossing the wings a n d t h r e e d a r k spots j u s t b e y o n d the center (one o n t h e fore wing, two o n the hind wing). All peacocks a r e m e d i u m - s i z e d b u t t e r ­ flies (WS 4 cm) a n d generally similar in shape, having t r i a n g u l a r wings with scal­ loped b o r d e r s . T h e y a r e u n p a l a t a b l e to vertebrates a n d p r e s u m a b l y rely on this and disruptive wing colors for survival, although at least o n e field study does not support the latter idea (Silberglied et al. 1980). Larvae of all a r e similar, with coarse

longitudinal stripes usually on a black back­ g r o u n d a n d with rows of elongate spines that are thickly bristled b u t not b r a n c h e d . T h e spines of the h e a d a r e clubbed. Food plants are varied but a r e often water-loving herbs of the families S c r o p h u l a r i a c e a e (Bacopa monnieri or water hyssop) a n d Labiaceae for the white peacock or Acanthaceae (Blechum) for the o t h e r species. T h e p u p a e a r e s u s p e n d e d terminally a n d are smooth a n d of simple s h a p e . T h e y a r e usually j a d e g r e e n with small black spots, a l t h o u g h occasional individuals may be black. Adults of the red a n d brown species have a jaunty, erratic flight, while the white species is an inveterate glider. At rest, those of all species habitually orient with their wings o p e n a n d bask in t h e sun (Fosdick 1973).

References EMMEL, T. C. 1972. Mate selection and balanced polymorphism in the tropical nymphalid but­ terfly, Anartia fatima. Evolution 26: 96-107. FOSDICK, M. K. 1973 [1972]. A population study of the Neotropical nymphalid butterfly, Anartia amalthea [sic], in Ecuador. J. Res. Lepidop. 11: 65-80. SILBERGLIED, R. E., A. AIELLO, AND G. LAMAS.

1979. Neotropical butterflies of the genus Anartia: Systematics, life histories and general biology (Lepidoptera: Nymphalidae). Psyche 86: 219-260. SILBERGLIED,

R.

E.,

A.

AIELLO,

AND D.

M.

WINDSOR. 1980. Disruptive coloration in but­ terflies: Lack of support in Anartia fatima. Science 209: 617-619. YOUNG, A. M., AND D. STEIN. 1976.

Studies on

the evolutionary biology of the Neotropical nymphalid butterfly Anartia fatima in Costa Rica. Milwaukee Pub. Mus. Contrib. Biol. Geol. 8: 1-29. Leaf Butterflies Nymphalidae, Nymphalinae, Charaxini, Memphis (formerly Anaea). T h e s e are medium-sized (WS 4 - 5 cm) butterflies having thin wings with a n g u l a r m a r g i n s , a s h a r p t i p p e d fore wing a n d a

BRUSH-FOOTED BUTTERFLIES

349

short-tailed h i n d wing. Memphis arachne (fig. 10.19c) is a n e x a m p l e . Above, the wings a r e brightly colored, b u t the lower surfaces a r e gray to b r o w n a n d mottled in a way closely r e s e m b l i n g a d e a d leaf, even to the details of the veins, imitated by d a r k streaks a n d m o l d spots a n d o t h e r imperfec­ tions faked by clear spots. T h e d e c e p t i o n is e n h a n c e d by their habit of resting with wings tightly closed o n tree t r u n k s a n d a m o n g d e a d foliage o n t h e g r o u n d . T h e y a r e c o m m o n forest dwellers a n d n u m e r o u s in species (Comstock 1961). T h e i r larvae feed o n the leaves of various small trees a n d s h r u b s in t h e families L a u r a c e a e a n d Flacourtiaceae. T h e y a r e cylindrical a n d lacking spines o r o t h e r elaborations. T h e cuticle possesses only n u m e r o u s small beadlike g r a n u l e s , each b e a r i n g a short, white hair (Young 1981).

References COMSTOCK, W. P. 1961. Butterflies of the Ameri­ can tropics: The genus Anaea, Lepidoptera, Nymphalidae. Amer. Mus. Nat. Hist., New York. YOUNG, A. M. 1981. Notes on the seasonal distribution of Anaea butterflies (Nympha­ lidae) in tropical dry forests. Acta Oecologica 2: 17-30. Cecropia Butterfly Nymphalidae, Nymphalinae, Nymphalini, Historis odius. Spanish: Pescadito (Costa Rica, p u p a ) . T h i s well-known butterfly (De Vries 1983) r a n g e s widely over Latin America, includ­ ing the C a r i b b e a n Islands a n d even iso­ lated Coco Island, w h e r e it is the only resident butterfly. It is a m o d e r a t e l y large (WS 10 cm) r o b u s t a n d powerful flier in the c a n o p y ; but its f o n d n e s s for the juices of r o t t i n g fruit often b r i n g it to the g r o u n d . T h e fore wings a r e b o r d e r e d broadly in black o n the u p p e r sides a r o u n d a basal m e d i a n field of dull o r a n g e . T h e h i n d wings a r e almost entirely black (fig. 10.19f).

350

MOTHS AND BUTTERFLIES

T h e large, m a t u r e larvae (BL to 75 nxn\ b e a r m a n y b r a n c h e d spines a n d are mot­ tled black, b r o w n , o r a n g e , a n d blue (fig 10.19d). T h e y a r e often seen resting singly o n the apical g r o w i n g stems of the host which is the cecropia tree. Young larvae feed in g r o u p s o n the u n d e r s i d e s of the leaves. T h e chrysalid has fine, fork-tipped dorsal a b d o m i n a l spines a n d c u r v e d head h o r n s (fig. 10.19e).

Reference DE VRIES, P. J. 1983. Historis odius (Orion). ¡n D. H. Janzen, ed., Costa Rican natural his­ tory. Univ. Chicago Press, Chicago. Pp 731 732.

Ithomiines Nymphalidae, lthomiinae. Ithomiines (Fox 1956, 1960, 1967; Fox and Real 1971) superficially resemble heliconiines, with the same a t t e n u a t e antennal clubs, wing shapes, flight behavior, and color p a t t e r n s (except for the "transpar­ ent" type, which is lacking in that sub­ family a n d very c o m m o n in this, e.g., Oleria a n d Hypoleria; fig. 10.20a). T h e y also are p r e s u m e d u n p a l a t a b l e forest dwellers that serve as models in Müllerian mimicry com­ plexes with heliconians a n d o t h e r Lepi­ d o p t e r a . Structural details distinguishing the family a r e a basally s p u r r e d anal vein in the fore wing a n d a weak s p u r only at t h e base of t h e h u m e r a l vein of the hind wing; they also lack the eversible hair tufts on the a b d o m e n which the heliconiines possess a n d seem to be less complex in their behavior a n d life history, although this may be d u e to the relatively poorer e x t e n t to which they have b e e n studied. Like their m o n a r c h butterfly relatives, male ithomiines are attracted to dead plants of the g e n u s Heliotropium (Boraginaceae) containing certain chemicals of decomposi­ tion called pyrrolizidine alkaloids (Lamas a n d Pérez 1983). Dried inflorescences, emit­ ting volatile c o m p o n e n t s of these alkaloids, a r e the most attractive p a r t of the plant. On

W5S1

Figure 10.20 NYMPHALID BUTTERFLIES (NYMPHALIDAE). (a) Glassy wing ithomiine (Hypoleria andromica). (b) Heliconius-mimicking ithomiine (Mechanitis polymnia). (c) Ithomiine (Mechanitis sp.), pupa, (d) Ithomiine (Mechanitis sp.), larva, (e) Gulf fritillary (Agraulis vanillae). (f) Julia (Dryas lulia). arriving at a moist plant, the butterflies drink surface d r o p l e t s ; or if the plant is dry, they r e g u r g i t a t e liquids a n d s p r e a d it over twigs a n d r e i m b i b e . I n the process, they obtain quantities of these alkaloids in solu­ tion. While feeding, they b e c o m e docile a n d can even be picked off a n d r e t u r n e d with­ out causing t h e m to fly. Males use these substances as metabolic substrates to p r o ­ duce a sexual secretion t h a t is disseminated from hair tufts o n t h e costal m a r g i n s of the hind wings (Pliske et al. 1976) a n d protect them from s p i d e r p r e d a t o r s (Brown 1984). They often g r o u p t o g e t h e r d u r i n g these displays, a m o n g t h e few butterflies to ex­ hibit a p p a r e n t lek behavior. D u r i n g n o r m a l behavior, females c o m e to males emitting this p h e r o m o n e from erect hair pencils (Piiske 1975). T h i s m a l e a p h r o d i s i a c of o n e species may p e r v a d e a territory also occu­ pied by males of a second a n d lead to cross mating, t h o u g h t to have significance in the family's evolution (Vasconcellos Neto a n d Brown 1982). Mechanitis females (and those of a few Other genera, e.g., Hypothyris) lay their eggs in clusters o n thick, pilose or spiny (trichomes), p o i s o n o u s s o l a n u m , a n d a p o cyanaceous (Echites G r o u p ) types a n d the larvae live gregariously o n silk p a d s that "ley spin over the coarse leaf surfaces (Rathcke a n d Poole 1975; Young a n d Moffett 1979a, 19796). However, they d o not •eem to sequester from t h e m t h e distaste­ ful or poisonous chemicals that r e n d e r

t h e m largely i m m u n e from v e r t e b r a t e p r e dation (Brown 1984). S o m e birds have even learned to feed selectively o n the a b d o m i n a l contents of a few species t h a t may lack these chemicals (Brown a n d Vasconcellos Neto 1976). By contrast, most g e n e r a (Greta, formerly Hymenitis, etc.) dwell in u n d e r s t o r y habitats w h e r e they exploit nonpilose, thin, p a p e r y leaves, oviposit singly, a n d have solitary larvae. T h e larvae of o n e species have b e e n observed interacting with ants in a n a p p a r ­ ently rudimentary, mutualistic way (Young 1978). Adults may follow a r m y a n t swarms, feeding o n the d r o p p i n g s of a n t birds (Ray a n d A n d r e w s 1980), b u t may take n o u r i s h ­ m e n t from any liquid bird feces (Young 1984). Externally, the larvae (fig. 10.20d) a r e simple, slender, t a p e r i n g at b o t h e n d s , a n d s m o o t h , o r with s p a r s e short hairs, (al­ t h o u g h some have elongate, fleshy lateral protuberances). T h e y are unicolorous or b a n d e d with various colors. T h e silver chrysalids (fig. 10.20c) a r e s u s p e n d e d only from the tip of the a b d o m e n a n d a r e strongly a r c h e d dorsally, t h e h e a d re­ c u r v e d ; the t h o r a x is e n l a r g e d so that the a p e x of the wing covers p r o t r u d e strongly. T h e widespread and common members of the g e n e r a Melinaea a n d Mechanitis (fig. 10.20b) have been implicated as p r i m e movers in the evolution of mimicry rings (Brown 1977). T h e subfamily contains 300 species

BRUSH-FOOTED BUTTERFLIES

351

( d ' A l m e i d a 1978, Mielke a n d B r o w n 1979, Lamas p e r s . c o m m . ) .

tionary race continues: Butterfly larval adan tation to plant trichomes. Science 187- 175 176. RAY, T. S., AND C. C. ANDREWS.

References BROWN, JR., K. S. 1977. Geographical patterns of evolution in Neotropical Lepidoptera: Dif­ ferentiation of the species of Melinaea and Mechanitis (Nymphalidae, Ithomiinae). Syst. Entomol. 1: 161-197. BROWN, J R , K. S. 1984. Adult-obtained pyr-

rolizidine alkaloids defend ithomiine butter­ flies against a spider predator. Nature 309: 707-709. BROWN, J R , K. S., A N D J . VASCONCELLOS NETO.

1976. Predation on aposematic ithomiine but­ terflies by tanagers (Pipraeidea melanonota). Biotropica8: 136-141. D'ALMEIDA, R. F. 1978. Catálogo dos Ithomiidae Americanos (Lepidoptera). Cons. Nac. Desenv. Cien. Teen., Curitiba. Fox, R. M. 1956. A monograph of the Ithomiidae (Lepidoptera). Pt. 1. Amer. Mus. Nat. Hist. Bull. I l l : 1-75. Fox, R. M. 1960. A monograph of the Ithomiidae (Lepidoptera). Pt. II. T h e Tribe Melinaeini Clark. Amer. Entomol. Soc. Trans. 86: 109-171. Fox, R. M. 1967. A monograph of the Ithomiidae (Lepidoptera). Pt. III. T h e tribe Mechanitini Fox. Amer. Entomol. Soc. Mem. 22: 1-90. Fox, R. M., AND H. G. REAL. 1971. A mono­

graph of the Ithomiidae (Lepidoptera). Pt. IV. T h e tribe Napeogenini Fox. Amer. Entomol. Inst. Mem. 15: 1-368.

1980. Ant

butterflies: Butterflies that follow army ants to feed on ant bird droppings. Science 2101147-1148. VASCONCELLOS NETO, J., AND K. S. BROWN, ID

1982. Interspecific hybridization in Mechanitis butterflies (Ithomiidae): A novel pathway for the breakdown of isolating mechanisms Biotropica 14:288-294. YOUNG, A. M. 1978. Possible evolution of mu­ tualism between Mechanitis caterpillars and an ant in northeastern Costa Rica. Biotropica 1077-78. YOUNG, A. M. 1984. Ithomiine butterflies associ­ ated with non-antbird droppings in Costa Rican tropical rain forest. J. Lepidop. Soc 3861-63. YOUNG, A. M., AND M. W. MOFFETT. 1979a.

Behavioral regulatory mechanisms in popula­ tions of the butterfly Mechanitis isthmia in Costa Rica: Adaptations to host plants in secondary and agricultural habitats (Lepidoptera: Nym­ phalidae: Ithomiidae). Deutsche Entomol Zeit.,N.F, 26: 21-38. YOUNG, A. M., AND M. W. MOFFETT.

1979ft.

Studies on the population biology of the tropical butterfly Mechanitis isthmia in Costa Rica. Amer. Midi. Nat. 101: 309-319.

Passion Vine Butterflies N y m p h a l i d a e , Heliconiinae, Heliconius a n d relatives. Heliconians.

LAMAS, G., AND J. E. PÉREZ. 1983. Danainae e

Ithomiinae (Lepidoptera, Nymphalidae) atraí­ dos por Heliotropium (Boraginaceae) en Madre de Dios, Perú. Rev. Peruana Entomol. 24: 5 9 62. MIELKE, O. H. H., AND K. S. BROWN, J R 1979.

Suplemento ao catalogo do Ithomiidae Ameri­ canos (Lepidoptera) de Romualdo Ferreira DAlmeida. Cons. Nac. Desenv. Cien. Tecn., Curitiba. PLISKE, T. E. 1975. Courtship behavior and use of chemical communication by males of cer­ tain species of ithomiine butterflies (Nym­ phalidae: Lepidoptera). Entomol. Soc. Amer. Ann. 68: 935-942. PLISKE,

T.

E., J.

A.

EDGAR,

AND C.

C. J.

CULVENOR 1976. T h e chemical basis of attrac­ tion of ithomiine butterflies to plants contain­ ing pyrrolizidine alkaloids. J. Chem. Ecol. 2: 255-262. RATHCKE, B. J., AND R. W. POOLE. 1975. R e v o l u ­

352

MOTHS AND BUTTERFLIES

Passion vine butterflies a r e a successful g r o u p , j u d g i n g from t h e i r a b u n d a n c e in low to mid-elevation forest habitats, the fairly large n u m b e r of species (65, mostly Heliconius), a n d their m a n y highly devel­ o p e d behavioral a n d physical adaptations (Brown 1981). T h e s e uniquely Neotropical butterflies have attracted n u m e r o u s bio­ logical studies since t h e days of their first appreciation by H e n r y Walter Bates (1862) in Amazonia. O f all taxa, it seems best to have fulfilled Bates's o w n prediction that "the study of butterflies will some day be valued as o n e of t h e most important b r a n c h e s of biological science." Bates himself p r o p o s e d a n d demon­ strated t h e principle of simple mimicry

from his A m a z o n i a n observations, mainly of ithomiines a n d heliconians. T h e p h e ­ nomenon is n o w recognized as c o m m o n among insects, fish, a n d even plants. Later, working in s o u t h e r n Brazil, Fritz Müller extended t h e c o n c e p t by discovering that two o r m o r e distasteful species benefit by jjjplaying t h e same p a t t e r n (see mimicry,

chap- 2). Substantiating conclusions from bird, reptile, plant, a n d o t h e r butterfly distribu­ tions, detailed investigation oí Heliconius in Central a n d South A m e r i c a h a s h e l p e d reveal t h e existence a n d locations of p r e ­ sumed Q u a t e r n a r y ice a g e forest refugia (see life zones, c h a p . 2) (Brown et al. 1974, Benson 1982). Localized speciation of heli­ conians is a c o n t i n u i n g d y n a m i c process, linked to p r e s e n t - d a y ecological factors and historical fluctuations in vegetation (Brown a n d B e n s o n 1977, Descimon a n d M a s t d e M a e g h t 1984). Much of t h e heliconians' significance in evolutionary biology derives from their association with two types o f plants, pas­ sion vines (Passiflora) a n d t h e cucurbit ge­ nus Psiguria (formerly Anguria) a n d close relatives (Gilbert 1975). T h e f o r m e r is t h e larval food a n d contains n o x i o u s chemicals (cyanogenic glucosides a n d alkaloids) that deter most o t h e r herbivores b u t from which t h e butterflies probably derive a n inedibility t h a t d e t e r m i n e s t h e i r role as mimicry models. T h e vine also possesses extrafloral nectaries that m a i n t a i n a d e ' fense force of p u g n a c i o u s ants. T h e latter's presence strongly influences m a n y aspects of the butterfly's utilization of t h e plant as a host. ,

Curiously, s o m e plants also d e f e n d t h e m «elves against f e e d i n g by heliconians by producing small, oval, yellow growths o n the leaves a n d tendrils, which mimic t h e ¡butterfly's eggs. Females refrain from Ovipositing o n plants already occupied by and a r e t h u s d i s c o u r a g e d from d o i n g on parts with these s t r u c t u r e s (Gilbert 1982).

T h e b r i g h t o r a n g e flowers of Psiguria are p r o d u c e d in progressively m a t u r i n g p e d u n c l e s that attract r e g u l a r visitations of adult heliconians. T h e butterflies derive from these flowers n o t only nectar b u t also pollen, which clings to their proboscises, t o be later dissolved by r e g u r g i t a t e d digestive fluids a n d ingested (Gilbert 1972). T h i s rich p r o t e i n food enables heliconians t o live very long for butterflies, possibly u p to n i n e m o n t h s , d u r i n g which time e g g p r o ­ duction is also p r o l o n g e d . O t h e r flowers may be visited at r a n d o m . L o n g life has enabled such sophisticated behaviors to evolve in these butterflies. T h e y also form a g g r e g a t i o n s to pass t h e night, h a n g i n g from vegetation in g r o u p s of m a n y individuals (Mallet 1986). Roosts are habitually used by t h e same individuals. T h e y have d e v e l o p e d highly sensitive a n d efficient n e u r a l m e c h a n i s m s that give t h e m exceptional sight a n d color p a t t e r n recognition, p r e s u m a b l y because of t h e role played by such in releasing c o u r t s h i p behavior (Swihart 1967). Females attract mates also with s t r o n g sexual p h e r o m o n e s . Multiple males often arrive simultaneously a n d elicit r a p i d copulation (often b e f o r e the female completely escapes t h e chrysalid case). I n t e r f e r e n c e by rival suitors is discouraged with a postulating sexual s u p ­ pression p h e r o m o n e (so-called olfactory chastity belt, T u r n e r 1973) i m p a r t e d t o t h e female by t h e successful male (Gilbert 1976). Heliconian adults a r e long winged, with a t t e n u a t e a n t e n n a l clubs. I n t h e a b d o m e n , females have yellow eversible glands with hair pencils o n t h e tips. T h e r e is a basically simple anal vein in t h e fore wing (i.e., n o s p u r ) a n d a strongly r e c u r v e d h u m e r a l vein at t h e base of t h e h i n d wing. T h e i r color p a t t e r n s vary greatly within t h e basic types (except for t h e "clear" type, which is not r e p r e s e n t e d in this subfamily; see but­ terflies a n d m o t h s , above) a n d a r e con­ trolled by a simple set of genetic loci (Nijhout a n d Gilbert 1990). T h e r e a r e

BRUSH-FOOTED BUTTERFLIES

353

BROWN, JR, K-. S., AND W. W. BENSON.

1977.

Fvolution in modern Amazonian non-forest islands: Heliconius hermathena. Biotropica 9: 95-117BROWN, J R . K. S., O. H. H. MIELKE, AND H.

EBERT. 1974. Quaternary refugia in tropical America: Evidence from race formation on Heliconius butterflies. Royal Soc. London Proc. B 187: 369-378. Figure 10.21 HELICONIINE BUTTERFLIES (NYMPHALIDAE). (a) Green heliconius (Philaethria dido), (b) Tiger heliconius (Heliconius melanops). (c) Black heliconius (Heliconius erato), pupa (d) Black heliconius, adult, (e) Black heliconius, larva, (f) Zebra butterfly (Heliconius charitonius).

COOK, L. M., AND L. P. BROWER 1969. Observa­

tions on polymorphism in two species of heliconiine butterflies from Trinidad, West Indies. Entomologist 102: 125-128. COOK, L. M.,

some u n u s u a l p a t t e r n s , as in Philaethria dido (fig. 10.21a), which has a light g r e e n g r o u n d color o n t h e u p p e r side a n d black bars a n d spots. T h i s p a t t e r n is mimicked by an edible n y m p h a l i d (Siproeta [formerly Victorino] stelenes; fig. 10.18f) (Young 1974). Two very c o m m o n species often seen in g a r d e n s a n d weed fields a r e t h e zebra butterfly (Heliconius charitonius; fig. 10.21f), which is black with oblique yellow wing bars (Cook et al. 1976, Y o u n g 1976), a n d t h e julia (Dryas iulia [formerly spelled julia], fig. 10.20f), which is all dull o r a n g e except for a single subapical black wing b a r (Young 1978, M u y s h o n d t 1973). T h e latter species has b e e n o b s e r v e d d r i n k i n g tears from t h e eyes of caimans a n d turtles in Peru ( T u r n e r 1986). Because t h e wings of m e m b e r s of t h e g e n u s Agraulis (fig. 10.20e) a r e silver spot­ ted o n t h e u n d e r s i d e s like t h e Holarctic butterflies called "fritillaries," they b e a r that n a m e also. A c o m m o n species with "tiger" m a r k i n g s is Heliconius melanops (fig. 10.21b); a p r e d o m i n a n t l y b l a c k - m a r k e d species is H. erato (fig. 10.2 I d ) . T h e larvae (fig. 10.2le) a r e usually c r e a m colored o r pale with d a r k spots o r b a n d s a n d covered with fringed, single, n o n u r t i c a t i n g spines, i n c l u d i n g a pair d o r sally from t h e oversized h e a d . Chrysalids (fig. 10.21c) h a n g from t h e t e r m i n u s of t h e a b d o m e n with a n irregularly s h a p e d body a n d a pair of wide-set, flat h e a d projec­ tions. T h i s s h a p e a n d t h e i r b r o w n i s h o r grayish color give t h e m t h e a p p e a r a n c e of

354

MOTHS AND BUTTERFLIES

a d e a d leaf. Silver o r white streaks also are frequently present, imitating holes and imperfections in t h e fake leaf. A few spe­ cies, such as Laparus doris, have gregarious, n o n s p i n y larvae that may form e n o r m o u s clusters o n t h e leaves of t h e host vinesp u p a e of these h a n g in great numbers from t h e u n d e r s i d e s of leaves a n d stems of associated trees a n d s h r u b s . Such groups arise from eggs that a r e g r o u p e d , even those of different females (Cook and B r o w e r 1969). Because of their great diversity and plastic wing p a t t e r n s , t h e t a x o n o m y of the g r o u p is complicated (Emsley 1965, Brown 1976, Michener 1942). It is clear that m e m b e r s of t h e same morphological series can display widely d i v e r g e n t mimetic pat­ terns, a n d , conversely, similar species are often u n r e l a t e d ( T u r n e r 1976).

References BATES, H. W. 1862. Contributions to an insect fauna of the Amazon valley (Lepidoptera: Heliconidae). Linnean Soc. London Trans. 2 3 : 4 9 5 - 5 6 1 . 6 , pi. 55-56. BENSON, W. S. 1982. Alternative models for infrageneric diversification in the humid trop­ ics: Tests with passion vine butterflies. In G. T. Prance, ed., Biological diversification in the tropics. Columbia Univ. Press, New York. Pp. 608-640. BROWN, JR., K. S. 1976. An illustrated key to the

silvaniform Heliconius (Lepidoptera: Nymphalidae) with descriptions of new subspecies. Amer. Entomol. Soc. Trans. 102: 373-484. BROWN, J R , K. S. 1981. T h e biology of Heli­ conius and related genera. Ann. Rev. Ento­ mol. 26: 427-456.

E. W. THOMASON, AND A.

M.

YOUNG. 1976. Population structure, dynamics and dispersal of the tropical butterfly Heli­ conius charitonius. J. Anim. Ecol. 45: 851-863. DESCIMON, H., AND J. MAST DE MAEGHT.

1984.

Semispecies relationships between Heliconius erato cyrbia Godt. and H. himera Hew in southwestern Ecuador. J. Res. Lepidop. 22: 229-237. EMSLEY, M. G. 1965. Speciation in Heliconius (Lep., Nymphalidae): Morphology and geo­ graphic distribution. Zoológica 50: 191-254. GILBERT, L. E. 1972. Pollen feeding and repro­ ductive biology of Heliconius butterflies. Nat. Acad. Sci. Proc. 69: 1403-1407. GILBERT, L. E. 1975. Ecological consequences of a coevolved mutualism between butterflies and plants. In L. E. Gilbert and P. H. Raven, ed., Coevolution of animals and plants. Univ. Texas Press, Austin. Pp. 210-240. GILBERT, L. E. 1976. Postmating female odor in Heliconius butterflies: A male-contributed antiaphrodisiac? Science 193: 419-420. GILBERT, L. E. 1982. T h e coevolution of a

butterfly and a vine. Sci. Amer. 247: 15, 1 1 0 114, 116, 119-121. MALLET, J. 1986. Gregarious roosting and home range in Heliconius butterflies. Nati. Geogr. Res. 2(2): 198-215. MICHENER, C. D. 1942. A generic revision of the Heliconiinae (Lepidoptera, Nymphalidae). Amer. Mus. Nov. 197: 1-8. MUYSHONDT, A. 1973. Some observations on Dryas iulia iulia (Heliconiidae). J. Lepidop. Soc. 27: 302-303. NIJHOUT, H. F., AND L. E. GILBERT. 1990.

An

analysis of the phenotypic effects of certain colour pattern genes in Heliconius (Lepi­ doptera: Nymphalidae). Biol. J. Linnean Soc. 40: 357-372. SWIHART, S. L. 1967. Neural adaptations in the visual pathway of certain heliconiine butter­ flies, and related forms, to variations in wing coloration. Zoológica 52: 1-14.

TURNER, J. R. G. 1973. Passion flower butter­ flies. Animals 15: 15-17, 19-21. TURNER, J. R. G. 1976. Adaptive radiation and convergence in subdivisions of the butterfly genus Heliconius (Lepidoptera: Nymphali­ dae). Zool. J. Linnean Soc. 58: 297-308. TURNER, J. R. G. 1986. Drinking crocodile tears: The only use for a butterfly? Antenna 10: 119-120. YOUNG, A. M. 1974. Further observations on the natural history of Philaethria dido dido (Lepi­ doptera: Nymphalidae: Heliconiinae). New York Entomol. Soc. J. 82: 3 0 - 4 1 . YOUNG, A. M. 1976. Studies on the biology of Heliconius charitonius L. in Costa Rica (Nym­ phalidae: Heliconiinae). Pan-Pacific Entomol. 52: 291-303. YOUNG, A. M. 1978. Spatial properties of niche separation among Eueides and Dryas butter­ flies (Lepidoptera: Nymphalidae: Heliconi­ inae) in Costa Rica. New York Entomol. Soc. J. 86: 2-19. Morphos N y m p h a l i d a e , M o r p h i n a e . Spanish: Morfos. Portuguese: Azulonas, azulas. R e s p l e n d e n t with their great, metallic b l u e wings, t h e m o r p h o s a r e t h e flying jewels of the forests. T h i s effulgent blue is caused by the spectral reflectance of light passing t h r o u g h microscopic lamellae in t h e wing scales (Hirata a n d O h s a k o 1966, Pillai 1968). As it is a physical color, it n e v e r fades. Its function is not u n d e r s t o o d , b u t it may startle a n d confuse attackers o r func­ tion in sexual recognition (Young 197 \b). T h e subfamily is uniquely tropical A m e r i c a n a n d contains a b o u t twenty-five species, most in t h e typical g e n u s Morpho (LeMoult a n d Real 1962). (Many m o r e have been n a m e d by overly zealous a m a ­ t e u r lepidopterists, b u t all a r e n o t consid­ e r e d valid entities.) T h e sight of o n e of these magnifi­ cent iridescent butterflies lazily flapping t h r o u g h t h e v e r d a n t g r e e n is e n o u g h to dazzle t h e perceptions a n d elicit exclama­ tions from traveler a n d scientist alike. Natives in A m a z o n i a n Peru tell that t h e m o r p h o is o n e form a s s u m e d by t h e forest

BRUSH-FOOTED BUTTERFLIES

355

spirits, o r chullachaquis, w h o lead followers into t h e j u n g l e to b e c o m e lost forever. M o r p h o s a r e t h e favorites of collectors a n d o n c e w e r e prizes s o u g h t by the liberes from Devil's Island p e n a l colony in French G u i a n a d u r i n g its i n f a m o u s history. Today, t h e tradition of m a k i n g trays a n d decou­ pages from their wings still persists for t h e tourist t r a d e in Rio d e J a n e i r o (Carvalho a n d Mielke 1972). S o m e species a r e p r o ­ tected by federal law, b u t o t h e r s a r e still c a u g h t wild without a p p a r e n t d a m a g e to their p o p u l a t i o n s since primarily t h e males a r e t a k e n (Kesselring 1975). O t h e r s a r e r e a r e d artificially for commercial p u r ­ poses ( O t e r o 1971). B e r m ú d e z (1966) sug­ gested o n e species (M. peleides) as t h e national butterfly for Venezuela. It is a l e g e n d that t h e first o n e of these beautiful butterflies to b e t a k e n to E u r o p e was by Sir Walter Raleigh w h o p r e s e n t e d it to Q u e e n Elizabeth (ibid.) S h e is said to have a d o r n e d h e r hair with it o n e night at a ball. O n seeing t h e h o r r o r of s o m e naturalists also p r e s e n t , s h e gave it to t h e o n e w h o later used it as t h e type specimen of M. peleides. T h e i r f o n d n e s s for t h e e x u d a t e s of fer­ m e n t i n g fruit m a k e s m o r p h o s easy p r e y to h u n t e r s using the fruit as bait (Young a n d T h o m a s o n 1974). A piece of b l u e p a p e r o r a d e a d s p e c i m e n also t e m p t s their curiosity a n d will d r a w specimens into n e t r a n g e . T h e y m a y also b e t a k e n while sleeping in loose a g g r e g a t i o n s (Young 1971a).

Two types of b l u e m o r p h o s m a y be recognized according to basic wing pattern: solid blue (e.g., M. rhetenor) a n d blue only in a b r o a d medial field b o r d e r e d by black (e.g., M. achillaena; fig. 10.22a, pi. 2f). I n addition, t h e r e are all white species (e.g., M. polyphetnus) a n d a single very l a r g e species that is generally o r a n g e with black mark­ ings (M. hecuba; fig. 10.22e). Each g r o u p r e p r e s e n t s different adaptive strategies with distinctive behavioral a n d ecological traits (Young 1982, Young a n d M u y s h o n d t 1972). T h e u n d e r s i d e s of t h e wings bear eyespots o r a r e otherwise m a r k e d with dead leaf p a t t e r n s to camouflage t h e closedwinged resting butterfly (fig. 10.22c). Eyespots t e n d to be m o r e conspicuous in low-flying, g r o u n d - f e e d i n g types (Young 1980). T h e larvae, which feed mainly o n many species of legumes, e u p h o r b s , a n d grasses (Carvalho a n d Mielke 1972), a r e very color­ ful, with multicolored, serial designs on the d o r s u m (fig. 10.22b). T h e y a r e also beset with hair tufts whose color a n d arrange­ m e n t s vary a c c o r d i n g to species. All have very hairy, oversized h e a d s . Species are generalized in their choice of hosts, and larval colors vary o n different plants, a possible system of automimicry to protect palatable individuals o n low-toxicity hosts (Young a n d M u y s h o n d t 1973). T h e y are c r e p u s c u l a r feeders. T h e light, leaf green chrysalids (fig. 10.22d) a r e obovoid and h a n g from hooks o n a short, t e r m i n a l stalk.

Some of t h e m have spot p a t t e r n s reminis­ cent of a simian face.

References BERMÚDEZ, P. J. 1966. El Morpho peleides podría ser la mariposa nacional. Natura (Caracas) 31:20. CARVALHO, J. C. M., AND O. H. H. MIELKE.

1972. The trade of butterfly wings in Brazil and its effects upon the survival of the species. 9th Int. Cong. Entomol. 1: 486-488. HIRATA, K., AND N. OHSAKO. 1966. Studies on

the structure of scales and hairs of insects. IV. Microstructure of scales of the butterfly Morpho menelaus nakaharai Le Moult. Kagoshima Univ. Sci. Rpt. 15: 4 9 - 6 1 . KESSELRING, J. 1975. Are morphos endan­ gered? Atala 3(2): 31. LEMOULT, E., AND P. REAL. 1962. Les Morpho

d'Amérique du Sud et Central. Ed. LeMoult, Paris. OTERO, L. S. 1971. Instrucóes para criacáo de borboleta "capitáo-do-mato" (Morpho achil­ laena) e outras especies do género Morpho ("azul-seda," "bóia," "azuláo-branco," "praiagrande"). Inst. Brasil. Desenv. Flor, Rio de Janeiro. PILLAI, P K. C. 1968. Spectral reflection charac­ teristics of Morpho butterfly wing. Optical Soc. Amer.J. 58: 1019-1022. YOUNG, A. M. 1971a. Notes on gregarious roosting in tropical butterflies of the genus Morpho.). Lepidop. Soc. 25: 223-234. YOUNG, A. M. 19716. Wing coloration and reflectance in Morpho butterflies as related to reproductive behavior and escape from avian predators. Oecologia 7: 209-222. YOUNG, A. M. 1980. The interaction of preda­ tors and "eyespot butterflies" feeding on rotting fruits and soupy fungi in tropical forests: Variations on a theme developed by the Muyshondts and Arthur M. Shapiro. Entomol. Rec. 90: 63-69. YOUNG, A. M. 1982. Notes on the natural history of Morpho granadensis polybaptus Butler (Lepidoptera: Nymphalidae: Morphinae), and its relation to that oí Morpho peleides límpida But­ ler. New York Entomol. Soc. J. 90: 35-54. YOUNG, A. M., AND A. MUYSHONDT. 1972. Geo­

graphical and ecological expansion in tropical butterflies of the genus Morpho in evolutionary time. Rev. Biol. Trop. 20: 231-263. YOUNG, A. M., AND A. MUYSHONDT. 1973. Notes

Figure 10.22 MORPHO BUTTERFLIES (NYMPHALIDAE). (a) Achilles morpho (Morpho achillaena). (b) Achilles morpho, larva, (c) Achilles morpho, undersides, (d) Morpho (Morpho sp.). pupa, (e) Hecuba (Morpho hecuba).

356

MOTHS AND BUTTERFLIES

on the biology of Morpho peleides in Central America. Carib.J. Sci. 13: 1-49. YOUNG, A. M., AND J. H. THOMASON. 1974. T h e

demography of a confined population of the butterfly Morpho peleides during a tropical dry season. Stud. Neotrop. Fauna 9: 1-34.

Owl Butterflies N y m p h a l i d a e , Brassolinae, Caligo. Spanish: Buhitos p a r d o s (Costa Rica), mariposas d e Muzo (Colombia), vaquitas n e g r a s (Ecuador, larvae). T h e s e are familiar m e m b e r s of the family, easily recognized by their great size (WS 1 2 - 1 5 cm) a n d large, c o n s p i c u o u s eyespot n e a r t h e center of t h e u n d e r s i d e of t h e h i n d wing (fig. 10.23a). T h e latter, in some species, is s u r r o u n d e d by a n e l o n g a t e , d a r k field, creating t h e illusion of t h e h e a d profile of a lizard o r frog. A secondary spot, o n t h e leading e d g e of t h e wing, resembles t h e t y m p a n u m of such verte­ brates. T h i s is t h o u g h t to b e a false w a r n ­ ing device that affords the butterfly protec­ tion w h e n it is at rest with wings folded (Stradling 1976). T h e u p p e r sides a r e d a r k blue, p u r p l e , o r b r o w n o n t h e i n n e r half o r m o r e a n d black o n t h e o u t e r half o r d a r k b o r d e r e d . T h e male a b d o m e n bears con­ spicuous r e d to yellow scale-covered scent glands o n each side opposite t h e a n d r o conia of the wings. Volatile chemicals from these glands may play a role in the territo­ rial behavior exhibited in this sex (Blandin a n d Descimon 1975). T h e adults a r e n o t k n o w n to take nectar from flowers b u t avidly suck liquids from r o t t i n g fruit. T h e y are c r e p u s c u l a r b u t a r e often frightened into flight—an eratic, u n d u l a t i n g motion close to t h e g r o u n d — d u r i n g t h e daylight h o u r s from their perches in forest glades. Individuals m a y be spotted resting o n vegetation n e a r t h e g r o u n d . T h e y a r e most f r e q u e n t in t h e vicinity of t h e larval food plants, which most c o m m o n l y belong to t h e b a n a n a fam­ ily a n d its relatives: Heliconia a n d b a n a n a (Musaceae), Canna (Cannaceae), a n d Calathea (Marantaceae). S o m e plants in t h e ginger (Hedychium), grass (Pennisetum), a n d

BRUSH-FOOTED BUTTERFLIES

357

Figure 10.23 BUTTERFLIES AND SKIPPERS, (a) Owl butterfly (Caligo eurilochus, Nymphalidae), undersides, (b) Owl butterfly, larva, (c) Darius (Dynastor darius, Nymphalidae). (d) Darius, pupa. (e) Blue banded skipper (Elbella polyzona, Hesperiidae). (f) Long-tailed skipper (Urbanus proteus, Hesperiidae). (g) Canna skipper (Calpodes ethlius, Hesperiidae). palm families (Cocos) a r e also utilized (Carvalho a n d Mielke 1972). At times, t h e larvae a r e even a b u n d a n t e n o u g h o n ba­ n a n a to b e c o n s i d e r e d a pest ( H a r r i s o n 1963, Malo a n d Willis 1961). B a m b o o a n d m a n i o c a r e also r e c o r d e d hosts b u t a r e probably substitutes for t h e a f o r e m e n ­ tioned. Larvae feed at all times of t h e day (Condie 1976). M a t u r e larvae a r e large (BL 10—15 cm), mottled b r o w n , a n d fusiform (fig. 10.23b, pi. 2g). T h e h e a d h a s a pair of welld e v e l o p e d clubbed " h o r n s " projecting from its u p p e r o u t e r c o r n e r s ( a n d some­ times a second pair laterally) a n d short, s h a r p spines situated o n t h e back of each of t h e m i d d l e s e g m e n t s . T h e terminally s u s p e n d e d p u p a e a r e also large (BL 4—5 cm) a n d have t h e a p p e a r a n c e of a curled, dead leaf. N u m e r o u s m i n u t e prickly spines a r e p r e s e n t o n t h e back of t h e a b d o m e n (Young a n d M u y s h o n d t 1985). S o m e seventeen species a r e recognized in t h e g e n u s .

References BLANDIN, P., AND H. DESCIMON. 1975. Contribu­

tion a la connaissance des Lépidoptéres de l'Equateur: Les Brassolinae (Nymphalidae). Soc. Entomol. France Ann. (N.S.) 11: 3 - 2 8 . CARVALHO, J. C. M., AND O. H. H. MIELKE.

1972. T h e trade of butterfly wings in Brazil and its effects upon the survival of the species. 9th Int. Cong. Entomol. 1: 486-488. CONDIE, S. 1976. Some notes on the biology and behavior of three species of Lepidoptera

358

MOTHS AND BUTTERFLIES

(Satyridae: Brassolinae) on non-economic plants in Costa Rica. Tebiwa 3: 1-28. HARRISON, J. O. 1963. T h e natural enemies of some banana insect pests in Costa Rica. J. Econ. Entomol. 56: 282-285.

n a r y in its r e s e m b l a n c e to t h e h e a d of a snake (fig. 10.23d). It is large (4 c m long) and b r o w n o n t h e d o r s u m , beige a n d whitish o n t h e u n d e r s i d e s . Because it hangs in an incline position, its colors a r e reversed in position so that t h e p u p a is darker below, this s h a d e i n v a d i n g o n each light u p p e r p o r t i o n , giving t h e impression of t h e constriction b e h i n d a viper's h e a d . Reticulations in t h e p a t t e r n also r e s e m b l e reptilian scales, a n d t h e eyes a r e accentu­ ated.

Reference AIELLO, A., AND R. E. SILBERGLIED. 1978. Life

history oí Dynastor darius (Lepidoptera: Nym­ phalidae: Brassolinae) in Panama. Psyche 85: 331-345.

MALO, E, AND E. R. WILLIS. 1961. Life history

and biological control of Caligo eurilochus, a pest of banana.J. Econ. Entomol. 54: 530-536. STRADLING, D. J. 1976. T h e nature of the mi­

metic patterns of the brassolid genera Caligo and Eryphanis. Ecol. Entomol. 1: 135-138. YOUNG, A. M., AND A. MUYSHONDT. 1985. Notes

on Caligo memnon Felder and Caligo atreus Kollar (Lepidoptera: Nymphalidae: Bras­ solinae) in Costa Rica and El Salvador. J. Res. Lepidop. 24: 154-175.

The Darius N y m p h a l i d a e , Brassolinae, Dynastor darius. T h i s is a very large (WS 9 5 mm), c r e p u s c u l a r species (fig. 10.23c) found from G u a t e m a l a south into Brazil (Aiello a n d Silberglied 1978). Its larva (BL 6 cm) feeds o n bromeliads a n d is a bizarre crea­ t u r e with a s m o o t h fusiform body, a head with a c o r o n a of club-shaped h o r n s , and a forked tail. T h e cuticle is longitudinally pin-striped with d a r k b r o w n a n d dark g r e e n o n a n e u t r a l g r o u n d color. A con­ spicuous, oval, black spot, with a reticulate g r e e n c e n t e r a n d concentric o u t e r rings of g r e e n a n d black, m a r k s t h e back between a b d o m i n a l segments 3 a n d 4 (a second, similar, smaller spot is situated behind, s p a n n i n g a b d o m i n a l s e g m e n t s 5 a n d 6). It is t h e chrysalid, however, that is extraordi-

SKIPPERS Hesperioidea. Spanish: G u s a n o s cabezones (Central A m e r i c a , larvae). Skippers a r e m o t h l i k e butterflies with r o ­ bust bodies a n d d r a b colors for t h e most part, a l t h o u g h s o m e deviate from d r a b n e s s with metallic blue, r e d , o r o t h e r bright color fields o n t h e u p p e r wing surfaces. They a r e o t h e r w i s e closely related to t h e true butterflies. T h e y r a n g e in size from small (WS 17 mm) to large (WS 5 cm) a n d possess wideset a n t e n n a e with t h e knobs uniquely curved, s o m e even h o o k e d , at t h e tip. All six legs a r e completely d e v e l o p e d . Skip­ pers a r e n a m e d for t h e i r n e r v o u s , flitting flight. T h e i r larvae a r e n a k e d , often g r e e n , cream, o r yellow, a n d covered with a fine white p o w d e r o r bloom. T h e y h a v e over­

sized, r o u n d , brightly colored h e a d s at­ tached to a distinct s o m e w h a t constricted "neck" ( = first thoracic s e g m e n t ) . T h e i r tastes a r e extremely varied, b u t m a n y food plants a r e monocots, often grasses. Many have t h e habit of rolling themselves u p in t h e leaves of t h e i r hosts, tying t h e cut edges of their shelters d o w n with silk. H e r e they p u p a t e in cocoonlike s t r u c t u r e s o r o n t h e ground. T h i s is a large g r o u p with s o m e 1,800 t o 2,000 Neotropical species. R e p r e s e n t a t i v e species a r e t h e c a n n a skipper, Calpodes ethlius (fig. 10.23g), a n d fiery skipper, Hylephila phyleus (Young 1982). Many in t h e g e n e r a Urbanus (fig. 10.23f) a n d Chioides a r e tailed; these a n d t h e larger, m o r e brilliantly m a r k e d species belong t o t h e subfamily Pyrginae. Black bars o n a pale bluish-white b a c k g r o u n d is a n oftenr e p e a t e d p a t t e r n in several u n r e l a t e d spe­ cies, for e x a m p l e , Elbella polyzona (fig. 10.23e), Phocides thermus, Tarsoctenus papias, a n d Jamadia gnetus. Adults a r e avid nectar feeders a n d act as i m p o r t a n t pollinators of flowering plants, some even having greatly elon­ gated t o n g u e s for p r o b i n g d e e p - t h r o a t e d blossoms of plant tubes to which they may be mutualistically associated ( E m m e l 1971).

References EMMEL, T C. 1971. Symbiotic relationship of an Ecuadorian skipper (Hesperiidae) and MaxMaria orchids. J. Lepidop. Soc. 25: 20—22. YOUNG, A. M. 1982. Notes on the interaction of the skipper butterfly Calpodes ethlius (Lepi­ doptera: Hesperiidae) with its larval host plant Canna edulis (Cannaceae) in Mazatlán, State of Sinaloa, Mexico. New York Entomol. Soc.J. 90: 99-114.

SKIPPERS

359

11 FLIES AND MIDGES Díptera. Spanish: Moscas (General); z a n c u d o s (midges, long-legged flies, G e n e r a l ) ; mosquillas (midges, G e n e r a l ) . Portuguese: Moscas, p e r n i l o n g o s (Brazil, long-legged flies); m o s q u i n h a s (Brazil, midges). Tupi-Guaraní: M b e r ú , c a r a p a n á (Brazil, long-legged flies); b i r o n h a s ( = m b e r u - o b i ) (Brazil, biting flies). Quechua: C h u s p i (fly). Náhuatl: Zayolmeh (sing, zayolin) (Mexico). Except for a relatively few wingless forms, all t h e m e m b e r s of this o r d e r (Oldroyd 1964) a r e c h a r a c t e r i z e d by a single pair of wings arising from t h e m e s o t h o r a x ; t h e m e t a t h o r a c i c wings h a v e evolved into spe­ cial c l u b - s h a p e d , sensory o r g a n s (halteres), i m p o r t a n t to t h e insect's flight equilib­ r i u m . Flies a r e very efficient a e r o n a u t s , a n d m a n y a r e capable of powerful, sus­ tained flight, a n ability p r o v i d e d by an oversized t h o r a x , a c c o m m o d a t i n g h y p e r d e v e l o p e d flight muscles to move t h e wings. No o t h e r o r d e r o f insects h a s this particular a r r a n g e m e n t o f flight o r g a n s . All a d u l t flies feed o n liquids, which they imbibe with s i p h o n i n g t u b u l a r o r spongelike m o u t h p a r t s . Vertebrate blood is a n i m p o r t a n t food, b u t o t h e r n a t u r a l liquors a r e c o n s u m e d as well: blood of o t h e r insects, plant e x u d a t e s , nectar, ani­ mal secretions, h o n e y d e w , d e c o m p o s i n g o r g a n i c substances, feces, a n d so o n . T h e larvae feed o n , a n d live in, a still g r e a t e r variety of substances, b o t h liquid a n d solid. Many live in water a n d a r e filter feeders or browsers of algae a n d d i a t o m s . P u p a e of m a n y aquatic species a r e active swimmers.

360

T h e formal h i g h e r classification of the Diptera is controversial (Griffiths 1972, 1981; Steyskal 1974). For convenience, the o r d e r may be divided into t h r e e main g r o u p s : (1) N e m a t o c e r a ("midges"), in which t h e adults have long, many-seg­ m e n t e d a n t e n n a e , t h e larvae have welldeveloped h e a d s , a n d p u p a e a r e free; (2) Brachycera ("straight-seamed flies"), in which t h e r e a r e abbreviated a n t e n n a e com­ posed of several unlike segments, larval heads a r e r e d u c e d , a n d t h e r e a r e free p u p a e from which t h e adult escapes t h r o u g h a T - s h a p e d or straight opening; a n d (3) C y c l o r r h a p h a ("muscoid flies"), whose adults have very short a n t e n n a e , usually with only two, very small, basal segments a n d an oval, large third segment with a sensory bristle, whose larvae are maggotlike, completely without a sclerotized h e a d capsule, a n d whose p u p a e are contained in t h e persistent last larval skins that h a r d e n a n d form a capsule called a " p u p a r i u m " (fig. 11.8d). T h e last g r o u p is f u r t h e r divided into small muscoid flies without calypters, e n l a r g e d lobes at the base of t h e wings ("acalypterate muscoids"), a n d larger muscoid flies with welld e v e l o p e d calypters ("calypterate muscoids"). (A few families d o n o t fit into this dichotomy a n d a r e set off in a special g r o u p , t h e Aschiza.) T h e t e r m s "lower" or "primitive" flies, in contrast to "higher" flies, are also often applied informally to refer respectively to t h e N e m a t o c e r a ver­ sus Brachycera a n d muscoids. T h e Neotropics s u p p o r t a very large d i p t e r a n fauna, mostly poorly known.

Most families h a v e b e e n cataloged (Papavero 1 9 6 6 - ) , a n d t h e n u m b e r s of species found in each as given in t h e following sections a r e t a k e n from this publication. T h e r e a r e m a n y economically i m p o r ­ tant d i p t e r a n g r o u p s . Adults carry h u m a n and animal diseases (see flies a n d disease, below), a n d larval feeding d a m a g e s fruits, vegetables, t u b e r s , a n d o t h e r c r o p s . Flies, however, also i n c l u d e m a n y parasites use­ ful in biological control of insect pests. Certain Neotropical tachinid flies, t h e socalled C u b a n fly (Lixophaga diatraeae) a n d the A m a z o n fly (Metagonistylum mínense), have even b e e n e x p o r t e d to t h e Oriental tropics to r e d u c e p o p u l a t i o n s of lepidopterous stem b o r e r s of rice, s u g a r c a n e , a n d other g r a m i n e o u s c r o p s (although with only partial success; K a m r a n 1973). It is c u r i o u s that, a l o n g with t h e llama and o t h e r animals, flies {chuspi) w e r e held in some r e v e r e n c e by t h e ancient A n d e a n peoples. A c c o r d i n g to Rivero a n d von Tschudi (1857) flies w e r e sacrificed to t h e sun by t h e followers of t h e Incas in ancient Peru a n d a r e still used as a design motif in that c o u n t r y (see fig. 1.8).

References GRIFFITHS, G. C. D. 1972. T h e phylogenetic

classification of Diptera Cyclorrhapha, with special reference to the structure of the male postabdomen. W. Junk, T h e Hague. GRIFFITHS, G. C. D. 1981. [Book review of] Manual of Nearctic Diptera. Vol. 1. J. F. McAlpine, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, D. M. Wood, eds. Minister of Supply and Services. Entomol. Soc. Can. Bull. 13: 49-55. KAMRAN, M. A. 1973. Introduction of Neotrop­ ical tachinids into Southeast Asia for biological control of stem borers of graminaceous crops. Entomol. Soc. Amer. Bull. 19: 143-146. OLDROYD, H. 1964. T h e natural history of flies. Norton, New York. PAPAVERO, N., ed. 1966-. A catalogue of the Diptera of the Americas south of the United States. Dept. Zool., Sec. Agrie. Sao Paulo, Sao Paulo. Fase. 1-103. RIVERO, M. E., AND J. J. VON TSCHUDI.

1857.

Peruvian antiquities. Putnam, New York.

Translated from Spanish to English by F. L. Hawks. STEYSKAL, G. C. 1974. Recent advances in the primary classification of the Diptera. Ento­ mol. Soc. Amer. Ann. 67: 513-517.

FLIES AND DISEASE Exceptionally good flying a n d dispersal abilities, along with t h e bloodsucking, p r e daceous, unsavory habits of n u m e r o u s spe­ cies, have m a d e t h e D i p t e r a t h e single most i m p o r t a n t insect o r d e r from t h e medicalveterinary s t a n d p o i n t ( Z u m p t 1973). Most of t h e world's insect-borne diseases of h u m a n s a n d animals a r e t r a n s m i t t e d by biting a n d filth flies, a n d several serious pathological conditions a r e caused by flies themselves. Very large s u m s of m o n e y a r e spent from t h e public health b u d g e t s of nations e v e r y w h e r e , especially in t h e tropi­ cal portions of Latin America, in efforts to control diseases effected by d i p t e r a n s .

Reference ZUMPT, F. 1973. Diptera parasitic on vertebrates in Africa south of the Sahara and in South America, and their medical significance. In B.J. Meggers, E. S. Ayensu, and W. D. Duckworth, eds., Tropical forest ecosystems in Africa and South America: A comparative review. Smithsonian Insl. Press, Washington, D.C. Pp. 197-205.

Biting Flies as Disease Vectors D i p t e r a n m o u t h p a r t s , which a r e basically s t r u c t u r e d for a liquid diet, have b e e n f u r t h e r modified in several families for piercing o r slashing t h e skin of v e r t e b r a t e s a n d withdrawing blood. Blood is rich in p r o t e i n a n d provides t h e n u t r i e n t s neces­ sary for e g g d e v e l o p m e n t in these flies, which is why only t h e females bite. Attrac­ tion of t h e fly to its host is still n o t fully u n d e r s t o o d . Substances e m a n a t i n g from the host, such as c a r b o n dioxide, p e r s p i r a ­ tion, a n d body o d o r s , as well as skin color

FLIES AND DISEASE

361

a n d b o d y size a r e some factors that stimu­ late biting, b u t o t h e r s , still u n k n o w n , a r e probably involved. E n v i r o n m e n t a l condi­ tions a r e also i m p o r t a n t ; man-biting activ­ ity is largely d e p e n d e n t o n a m b i e n t tem­ p e r a t u r e a n d water v a p o r p r e s s u r e (Read e t a l . 1978). Many disease m i c r o o r g a n i s m s of birds a n d m a m m a l s , i n c l u d i n g h u m a n s , have evolved b l o o d - i n h a b i t i n g stages in their life cycles, t a k i n g a d v a n t a g e of t h e blood­ letting habits of m a n y insects, even devel­ o p i n g a n obligatory d e p e n d e n c e o n t h e m for t h e d e v e l o p m e n t of critical phases in the r e p r o d u c t i o n of t h e i r species. T h e s e a d a p t a t i o n s , a n d t h e fact that D i p t e r a fly, p r o m o t e t h e dissemination of these o r g a n ­ isms a n d m a k e m i d g e s a n d flies t h e most effective k n o w n biological vectors of parasites. A m o n g t h e blood f e e d e r s , t h e mosqui­ toes (Culicidae) a r e t h e best k n o w n a n d t h e most active s p r e a d e r s of p a t h o g e n s to b o t h m a n a n d animals. T h e a l r e a d y lengthy list of diseases that only mosquitoes t r a n s m i t p r o m i s e s to g r o w even l o n g e r as new ones, especially o n e s caused by viruses, a r e dis­ covered (Aitken et al. I 9 6 0 , Causey et al. 1961). I n t h e New World tropics, t h e most serious medical a n d v e t e r i n a r y p r o b l e m s a r e mostly m o s q u i t o related a n d include the malarias, yellow fever, encephalitides, d e n g u e , filariases, a n d a host of o t h e r poorly u n d e r s t o o d infections. O t h e r g r o u p s of b l o o d - f e e d i n g D i p t e r a with species also involved in t h e s p r e a d of disease a r e t h e blackflies (Simuliidae), which t r a n s m i t onchocerciasis, sand flies (Psychodidae, Lutzomyia), which carry cuta­ n e o u s leishmaniasis a n d oroya fever, p u n kies ( C e r a t o p o g o n i d a e , Culkoides), and horseflies a n d deerflies (Tabanidae). T h e latter two families a r e b o t h involved as vectors in several filarial, bacterial, a n d protozoal afflictions. T h e s e g r o u p s a r e all (except T a b a n i d a e [Brachycera]) classified in t h e N e m a t o c e r a , but several isolated species of Cyclor-

362

FLIES AND MIDGES

r h a p h a , such as t h e stable fly (Stornoxxs calcitrans) a n d h o r n fly (Haematobia irritans), also feed o n blood a n d act as vectors (of these, both males a n d females bite) ( T h e infamous tsetses [Glossina], carriers of sleeping sickness in Africa, a r e not found in t h e New World.) T h e direct effects of t h e bites of flies can be as grave as t h e infections they bring Mosquitoes a n d o t h e r biting flies are a constant nuisance in t h e e n v i r o n m e n t of I n d i a n s , w h o may be forced to live in a cloud of smoke from a wood fire to survive parts of t h e day. Every account of travel along t h e great rivers tells of t h e merciless attacks of these insects. E x p l o r e r s Alexan­ d e r von H u m b o l d t a n d A i m é Bonpland wrote, People who have not navigated the great rivers of equinoctal America . . . can scarcely conceive how, at every instant, with­ out intermission, you may be tormented by . . . mosquitoes, zancudos, jejenes, and tempraneros, that cover the face and hands, pierce the clothes with their long needleformed suckers, and getting into the mouth and nostrils, occasion coughing and sneez­ ing whenever any attempt is made to speak in the open air. In the missions of the Orinoco . . . the plague of the mosquitoes af­ fords an inexhaustible subject of conversa­ tion. When two people meet in the morning, the first questions they address to each other are: "How did you find the zancudos dur­ ing the night? How are we today for the mosquitoes?" (1852: 2:273) T h e last message received from the illfated 1925 expedition of Col. P. H . Fawcett, seeking lost cities in t h e Brazilian h i n t e r l a n d , tells how noxious insects con­ tributed to its failure: The attempt to write is fraught with much difficulty owing to the legions of flies that pester one from dawn till dark—and some­ times all through the night! The worst are the tiny ones smaller than a pinhead, almost

invisible, but stinging like a mosquito. Clouds of them are always present. . . . The stinging horrors get all over one's hands, and madden. Even the head nets won't keep them out. As for mosquito nets, the pests fly through them! ( 1 9 5 3 : 300) Habitation in s o m e areas is impossible even today because of t h e bloodthirsty onslaughts of h o r d e s of blackflies a n d sand flies. Bites often cause severe allergic reac­ tions a n d m a y p r o d u c e local sores that resist healing a n d b e c o m e septic. Not all species of t h e biting fly g r o u p s are equally offensive; those of p r i m a r y importance will be n o t e d below.

References AITKEN, T. H. G., W. G. DOWNS, C. R. ANDER­

SON, AND L. SPENCE. 1960. Mayaro virus isolated from a Trinidadian mosquito, Mansonia venezuelensis. Science 131: 986. CAUSEY, O. R., C. E. CAUSEY, O. M. MAROJA, AND

D. G. MACEDO. 1961. T h e isolation of arthro­ pod-borne viruses, including members of two hitherto undescribed serological groups, in the Amazon region of Brazil. Amer. J. Trop. Med. Hyg. 10: 227-249. FAWCETT, P. H. 1953. Lost trails, lost cities. Funk & Wagnalls, New York. READ, R. G., A. J. ADAMES, AND P. GALINDO.

1978. A model of microenvironment and man-biting tropical insects. Environ. Entotnol. 7: 547-552. VON HUMBOLDT, A. AND A. BONPLAND.

1852

[1814-1825]. Personal narrative of travels to the equinoctial regions of America, during the year 1799-1804. 3 vols. Henry Bohn, London. Translated by Thomasina Ross.

certain flesh flies, acalypterates, such as t h e eye gnats (Hippelates), a n d m a n y o t h e r s (Lindsay a n d S c u d d e r 1956). Because of the p r o p e n s i t y of these D i p t e r a to pick u p bacteria, viruses, amoebic cysts, a n d o t h e r p a t h o g e n s while f r e q u e n t i n g feces, gar­ bage, a n d carrion, a n d t h e capability to mechanically transmit t h e m to t h e food of p e o p l e a n d domestic animals, o r directly o n t o lips, eyes, a n d fingers, filth flies m u s t be considered of public health a n d veteri­ nary i m p o r t a n c e (Alcivar a n d C a m p o s 1946, G r e e n b e r g a n d B o r n s t e i n 1964). T h e evidence for fly involvement in t h e per­ petuation of m o r e t h a n sixty diseases is a b u n d a n t , t h e most convincing b e i n g that for enteric d i s o r d e r s o r dysenteries caused by Shigella a n d Salmonella bacteria. While not d e m o n s t r a b l y involved as p r i m a r y vec­ tors of these p a t h o g e n s , flies m u s t always be considered to often have considerable p o ­ tential epidemiological i m p o r t a n c e d u r i n g o u t b r e a k s of polio, hepatitis, conjunctivitis, staphylococcal infections, cholera, pinta, brucellosis, a n d s u r r a , especially u n d e r very u n s a n i t a r y conditions such as prevail in times of warfare a n d famine, following disasters, o r w h e n o t h e r d i s r u p t i o n s of social o r d e r a n d hygiene occur, such as t h e d e v e l o p m e n t of slums. A c o m p r e h e n s i v e list of t h e filth fly species a n d t h e diseases b o r n e by t h e m in the vast area of Latin A m e r i c a is b e y o n d the scope of this book; t h e subject is surveyed in some detail by G r e e n b e r g (1971, 1973).

Filth Flies Numerous species of flies live in close association with h u m a n s ("synanthropic" o r "domestic" flies), their larvae b r e e d i n g in wastes a n d a d u l t s f r e q u e n t i n g all sorts of organic m a t t e r c o n t a m i n a t e d with h u m a n disease o r g a n i s m s . T h e s e a r e t h e filth flies, a name describing t h e foul habits of t h e notorious housefly ( C o u t i n h o et al. 1957) *nd its relatives a m o n g t h e muscoid flies, ">e green blowfly a n d o t h e r c a r r i o n flies,

References ALCÍVAR, C , AND F. CAMPOS. 1946. Las moscas,

como agentes vectores de enfermedades en­ téricas en Guayaquil. Rev. Ecuatoriana Hig. Med. Trop. 3: 3-14. C O U T I N H O , } . O., A. DE E. TAUNAY, AND L. P DE

CARVALHO LIMA. 1957. Importancia de Musca

domestica como vector de agentes patogénicos para o homen. Rev. Inst. Adolpho Lutz Sao Paulo 17: 5 - 2 3 . GREENBERG, B. 1971, 1973. Flies and disease. 2 vols. Princeton Univ. Press, Princeton.

FLIES AND DISEASE

363

GREENBERG, B., AND A. A. BORNSTEIN. 1964. Fly

dispersion from a rural Mexican slaughter­ house. Amer. J. Trop. Med. Hyg. 13: 881-886. LINDSAY, D. R., AND H.

I. SCUDDER.

1956.

Nonbiting flies and disease. Ann. Rev. Entomol. 1: 323-346.

Myiasis T h e larvae of m a n y k i n d s of flies find their way into living a n i m a l s a n d feed o n healthy o r diseased tissues, a condition called myiasis ( G u i m a r á e s et al. 1983). T h e host acquires t h e infection by swallow­ ing fly eggs o r larvae in its food o r from eggs ( a n d s o m e t i m e s y o u n g larvae) p u r ­ posefully placed by t h e fly o n its b o d y o r o n s o m e s u b s t r a t u m f r o m which it will b e taken u p (Dove 1937). Many fly g r o u p s may b e involved ( J a m e s 1947), e i t h e r as (1) accidental invaders of t h e host, seldom causing a n y lasting effects a n d d y i n g b e ­ fore r e a c h i n g maturity, (2) facultative p a r a ­ sites usually b r e e d i n g in d e c o m p o s i n g o r ­ ganic m a t t e r b u t occasionally feeding within t h e body, o r (3) specific myiasis p r o d u c e r s , b e l o n g i n g t o species having obligatory d e v e l o p m e n t in living flesh. From the clinical s t a n d p o i n t , these infes­ tations a r e classified a c c o r d i n g to t h e site at which t h e larvae feed: (1) in t h e skin (cutaneous myiasis), i n c l u d i n g c r e e p i n g dis­ eases such as p r o d u c e d by early larvae of h o r s e bots, (2) in sores a n d w o u n d s (trau­ matic myiasis) by calliphorids, especially screwworms, (3) in cavities of t h e body (sinus myiasis), (4) in i n t e r n a l o r g a n s o r passages (organic a n d e n t e r i c myiasis) by almost any species that is accidentally swal­ lowed o r a t t r a c t e d t o orifices of the diges­ tive o r u r o g e n i t a l tracts, a n d (5) in t h e eye (ocular myiasis) caused by various types, a m o n g t h e m t h e s h e e p bots whose first instar larvae i n v a d e t h e conjunctiva. N u ­ m e r o u s publications contain case histories of all types of myiasis in h u m a n (Isola a n d O s i m a n i 1944, D o n o s o 1947, T o b a r a n d H o n o r a t o 1947) a n d a n i m a l (Lacey a n d

364

FLIES AND MIDGES

G e o r g e 1981) hosts in m a n y p a r t s of Latin America (Mazza a n d J ó r g 1939, T h o m a s 1987). T h e incidence of myiasis is apparently h i g h e r in t h e h u m i d tropics t h a n in cooler t e m p e r a t e regions (Méndez 1981). This may be d u e to t h e p o o r e r hygiene and faster d e v e l o p m e n t of larvae in the warmer climes. Myiasis lesions a r e seldom benign a n d should always b e c o n s i d e r e d as poten­ tially serious a n d treated promptly. T h e results of u n t r e a t e d cases in h u m a n s range from mild discomfort, with n o p e r m a n e n t d a m a g e , to gross destruction of vital organs a n d d r e a d f u l external disfigurements with associated psychological disturbances. Bot­ flies ( C u t e r e b r i d a e , O e s t r i d a e , a n d Gasterophilidae), carrion o r blowflies (Calliphoridae), flesh flies (Sarcophagidae), a n d cer­ tain o t h e r muscoid flies a r e most often the cause of myiasis, but almost any species with s a p r o p h a g o u s larvae m a y be involved, at least accidentally.

References DOVE, W. E. 1937. Myiasis of man. J. Econ. Entomol. 30: 39-49. DONOSO, R. B. 1947. Miasis humana en Chile y consideraciones clínicas y epidemiológicas. Rev. Chilena Hig. Med. Prevent. 1947: 1-54. GUIMARÁES, J. H., N. PAPAVERO, AND A. P. Do

PRADO. 1983. As miíases na regiáo Neotropi­ cal (identificacáo, biología, bibliografía). Rev. Brasil. Zool. 1: 239-416. ISOLA, W., AND J. OSIMANI.

1944. Un nuevo

Y los dípteros que las producen. Rev. Med. Panamá 6: 146-159. THOMAS, JR-, D. B. 1987. Incidence of screwworm (Díptera: Calliphoridae) myiasis on the Yucatán Peninsula oí Mexico. J. Med. Ento­ mol. 24: 498-502. TOBAR, G., AND A. HONORATO. 1947. Anota­

ciones acerca de una epidemia de miasis humana. Hospital Viña del Mar (Chile) Publ. Trimes. 3(1): 5-14.

CRANE FLIES Tipulidae. Extremely long, frail, easily d e t a c h e d legs and elongate body a n d wings immediately identify these flies (fig. 11.1c). T h e y a r e mistaken for mosquitoes by m a n y p e o p l e , but they n e v e r bite a n d have n o scaly vestiture. T h e i r size r a n g e is very great, from tiny (BL 5 - 7 m m ) to large (BL to 50 mm). Adults a r e c o m m o n p l a c e in d a m p situa­ tions, usually only n e a r water, which is a frequent larval m e d i u m , a l t h o u g h a few species occur over o p e n water o r r e m a i n continuously s u b m e r g e d (Byers 1977, 1981, 1982). Bromeliad tanks, tree holes, and seashore rock crevices a r e specialized aquatic habitats. Many o t h e r s d e v e l o p in soil, in leaf mold, a n d in o t h e r purely terrestrial sites. T h e larvae a r e mostly cylin­ drical, with a leathery i n t e g u m e n t a n d fleshy lobes s u r r o u n d i n g large b r e a t h i n g pores o n t h e b l u n t t e r m i n u s of the abdo­

m e n (fig. 11.Id). I n t h e forest, c r a n e flies often a g g r e g a t e in protected, moist places like hollow r o t t e n tree s t u m p s o r b e t w e e n t h e buttresses of l a r g e trees a n d " d a n c e . " T h e y gyrate u p a n d d o w n a n d create a ghostly illusion with their d i a p h a n o u s wings a n d w i d e s p r e a d legs. T h e function of this behavior is n o t k n o w n . Many also vibrate rapidly o n their leg tips over t h e substratum, another unexplained phe­ n o m e n o n . Mimetic relationships b e t w e e n c r a n e flies a n d i c h n e u m o n i d wasps a r e k n o w n (Slobodchikoff 1974). T h i s is t h e largest family of flies in t h e Neotropics. Practically n o t h i n g h a s b e e n written r e g a r d i n g t h e biology of t h e well over 3,400 Latin A m e r i c a n species, which certainly r e p r e s e n t s only a m o d e r a t e frac­ tion of t h e total n u m b e r yet to be f o u n d ( A l e x a n d e r a n d A l e x a n d e r 1970, B r u c h 1939).

References ALEXANDER, C. P., AND M. M. ALEXANDER. 1970.

Family Tipulidae. In N. Papavero, ed., 1967-, A catalogue of the Díptera of the Americas south of the United States. Mus. Zool., Univ. Sao Paulo, Sao Paulo. Pp. 4.1-4.259. BRUCH, C. 1939. Contribución al conocimiento de los tipúlidos argentinos (Díptera). Physis 17: 3-28. BYERS, G. W. 1977. Tipulidae. In S. H. Hurlbert, ed., Biota acuática de sudamérica aus­ tral. San Diego State Univ., San Diego. Pp. 259-265. BYERS, G. W. 1981. Tipulidae. In S. H. Hurlbert, G. Rodríguez, and N. Dias dos Santos,

caso de oftalmiasis conjuntival producida por Oestris ovis L. en el Uruguay. Arch. Uruguay Med. 25: 260-264. JAMES, M. T. 1947. The flies that cause myiasis in man. U.S. Dept. Agrie. Misc. Publ. 631: 1175. LACEY, L. A., AND T. K. GEORGE. 1981. Myiasis

in an Amazonian porcupine. Entomol. News 92: 79-80. MAZZA, S., AND M. E. JÓRG. 1939. Cochíiomyia

hominivorax americana C. y P., estudio de sus larvas y consideraciones sobre miasis. Univ. Buenos Aires, Mis. Estud. Pat. Reg. Argen­ tina, Publ. 41: 3-46. MÉNDEZ, E. 1981. Las miasis centroamericanas

Figure 11.1 FLIES (MIDGES), (a) Water midge (Chironomus sp., Chironomidae). (b) Water midge, larva, (c) Crane fly (Típula sp., Tipulidae). (d) Crane fly, larva, (e) Bathroom fly (Clogmia albipunctata, Psychodidae). (f) Bathroom fly, larva.

CRANE FLIES

365

eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 231-241. BYERS, G. W. 1982. Tipulidae. In S. H. Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 407-414. SLOBODCHIKOFF, C. N. 1974. Behavioral and

morphological mimicry in a cranefly and an ichneumonid. Pan-Pacific Entomol. 50: 155— 159.

WATER MIDGES C h i r o n o m i d a e . Spanish: Sayules (when s w a r m i n g , from N á h u a t l , zayollan = a b u n d a n c e of flies) (Nicaragua). A l t h o u g h they r e s e m b l e mosquitoes with their e l o n g a t e bodies, l o n g legs, a n d plu­ mose m a l e a n t e n n a e , water m i d g e s d o n o t have biting m o u t h p a r t s a n d lack scales o n any p a r t of t h e b o d y (fig. 11.1a) (Saether 1971). A few larvae o c c u r in wet decaying matter, b u t t h e majority a r e truly aquatic, leading s u b m e r g e d lives in all types of freshwater, brackish water, a n d m a r i n e coastal habitats. W h e n a b u n d a n t , they con­ stitute a n i m p o r t a n t food item for o t h e r aquatic animals, especially fish. Larval d e n ­ sities o f 50,000 p e r s q u a r e m e t e r a r e n o t u n u s u a l . Particularly noticeable a r e those of t h e g e n u s Chironomus t h a t inhabit lakes, w h e r e their n u m b e r s build e n o r m o u s l y d u r i n g t h e r e p r o d u c t i v e cycle. T h e s e a r e often b r i g h t r e d , from h e m o g l o b i n in their blood ("blood worms") (fig. 11.1b), which aids in r e s p i r a t i o n o n t h e oxygen-deficient b o t t o m waters in t h e t u b e houses they construct in t h e b o t t o m ooze. I n some places d u r i n g t h e d r y season, periodic mass e m e r g e n c e s of a d u l t s from these p o p u l a t i o n s take place, filling t h e air with clouds of gnats a n d c r e a t i n g a public nui­ sance. T h i s p h e n o m e n o n is well d o c u ­

366

FLIES AND MIDGES

m e n t e d at Lake N i c a r a g u a (Bay 1964) w h e r e t h e offending species is Siolimyia amazónica (Wirth 1979). T h e larvae of a few species associate as c o m m e n s a l s with o t h e r animals in t h e wa­ ter; some attach to o t h e r aquatic insects (Epler 1986) o r even t h e skin of bottomdwelling catfishes in t h e A m a z o n River (Fittkau 1974, Freihofer a n d Neil 1967). Except for a few c a r n i v o r o u s , p h y t o p h a ­ gous, a n d parasitic forms, c h i r o n o m i d lar­ vae a r e m i c r o p h a g e s o n o r g a n i c d e t r i t u s . T h e t a x o n o m y of Neotropical water midges is in a n infant state. At least 1,500 species a r e t h o u g h t to exist, m a n y of these yet u n n a m e d (Fittkau a n d Reiss 1979, Fittkau 1971). T h e S o u t h A m e r i c a n fauna is divided into a w a r m - a d a p t e d species complex, occupying t h e Guiana-Brazilian area, a n d m o r e o r less cold-loving groups, limited to t h e A n d e a n - P a t a g o n i a n regions (Fittkau 1978). O f t h e seven subfamilies, only t h e T a n y p o d i n a e a n d C h i r o n o m i n a e a r e a b u n d a n t in t h e region. Most of the water midges living in forest s t r e a m s of Central A m a z o n i a belong to t h e latter. T h e n u m b e r of species in a n aquatic habitat often accounts for m o r e t h a n 50 p e r c e n t of t h e total i n v e r t e b r a t e species present. Larvae of virtually all build some sort of a case o n o r within t h e substrate in which they live. T h e T e l m a t o g e t o n i n a e a r e largely a ma­ rine g r o u p , found in rocky intertidal areas along t h e entire A m e r i c a n coast. T h e i r larvae a r e physiologically a d a p t e d to salt water (Oliver 1971). T h e special c o n t r i b u t i o n of certain aquatic m i d g e g r o u p s to o u r u n d e r s t a n d ­ ing of t h e b i o g e o g r a p h y of southern S o u t h America h a s b e e n r e f e r r e d to else­ w h e r e (see physiographic regions, chap. 2). L i t e r a t u r e o n t h e Neotropical m e m b e r s of t h e family is reviewed by Reiss (1977, 1981, 1982); world bibliographies are also available (Fittkau et al. 1976, Hoffrichter a n d Reiss 1981).

Deferences BAY, E. C. 1964. An analysis of the "sayule" (Díptera: Chironomidae) nuisance at San Car­ los, Nicaragua, and recommendations for its alleviation. World Health Org./EBL20, WHO/ Vector Control 86: 1-19. EPLER, J- H. 1986. A novel new Neotropical Nanocladius (Díptera: Chironomidae), symphoretic on Traverella (Ephemeroptera: Leptophlebiidae). Fla. Entomol. 69: 319-327. FITTKAU, E. J. 1971. Distribution and ecology of Amazonian chironomids (Díptera). Can. Ento­ mol. 103:407-413. FITTKAU, E. J. 1974. Ichthyocladius n. gen., eine neotropische Gattung der Orthocladiinae (Chironomidae, Díptera) deren Larven epizoische auf Welsen (Astroblepidae und Loricariidae) leben. Entomol. Tidskr. 95: 91 — 106. FITTKAU, E. J. 1978. Sich abzeichnende Verbreitungsmuster in die neotropische-nearktischen Chironomidenfauna. Deutsch. Ges. Allg. Angewan. Entomol. Mitt. 1: 7 7 - 8 1 . FITTKAU, E. J., AND F. REISS. 1979. Die zoogeo-

graphische Sonderstellung der neotropischen Chironomiden. Spixiana 2: 273-280. FITTKAU, E. J., F. REISS, AND O. HOFFRICHTER

1976. A bibliography of the Chironomidae. Gunnera26: 1-177. FREIHOFER, W. C , AND E. H. NEIL. 1967. Com-

mensalism between midge larvae (Díptera: Chironomidae) and catfishes of the families Astroblepidae and Loricariidae. Copeia 1967: 39-45. HOFFRICHTER, O., AND R. REISS. 1981. Supple­

ment 1 to "A bibliography of the Chiro­ nomidae." Gunnera 37: 1—68. OLIVER, D. R. 1971. Life history of the Chiro­ nomidae. Ann. Rev. Entomol. 16: 211-230. REISS, F. 1977. Chironomidae. In S. H. Hurl­ bert, ed., Biota acuática de sudamérica aus­ tral. San Diego State Univ., San Diego. Pp. 277-280. REISS, F. 1981. Chironomidae. In S. H. Hurl­ bert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 261-268. REISS, F. 1982. Chironomidae. In S. H. Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 433-438. SAETHER, O. A. 1971. Notes on general morphol­ ogy and terminology of the Chironomidae (Diptera). Can. Entomol. 103: 1237-1260.

WIRTH, W. W. 1979. Siolimyia amazónica Fittkau, an aquatic midge new to Florida with nui­ sance potential. Fla. Entomol. 62: 134-135.

MOTH FLIES Psychodidae. Owl midges. So called because of a d e n s e body covering of elongate scales, which gives t h e m a l e p i d o p t e r a n a p p e a r a n c e (fig. 11. le), m o t h flies a r e dwellers in d a m p n e s s . T h e larvae of most a r e aquatic o r r e q u i r e considerable m o i s t u r e in their habitats. T h e s e r a n g e from wet soil a n d rotting leaf litter to p e r m a n e n t l y moist o r s u b m e r g e d s t r e a m a n d p o n d m a r g i n s ; t h e lance-winged midges {Maruina) live o n s m o o t h rocks u n d e r clean d r i p p i n g o r spray water a n d even totally s u b m e r g e d in torrential water in streams ( H o g u e 1973). Larvae a r e elongate a n d cylindrical (fig. 11.1 F), except those of Maruina, which a r e flattened a n d have ventral suckers. Each s e g m e n t possesses distinct, transverse, d o r ­ sal sclerotized plates a n d well-developed setae. T h e head capsule is small b u t com­ plete, a n d t h e r e is a short terminal b r e a t h ­ ing tube. Adults a r e all small (BL 1-4 m m ) a n d densely covered with e l o n g a t e scales. T h e wings a r e elliptical in outline (lanceolate in Maruina), with only longitu­ dinal veins a p p a r e n t a n d , w h e n at rest, are held rooflike over t h e a b d o m e n . T h e n u m b e r of species currently k n o w n ( a p ­ proximately 430 in 28 g e n e r a ) is probably far short of t h e total living in Latin America (Duckhouse 1973). A few well-known domestic species, such as Clogmia (formerly Telmatoscopus) albipunctata (fig. 11.le) (Sebastiani 1978) a n d Psychoda alternata, b r e e d in the foul o r g a n i c m a t t e r that accumulates in sink d r a i n t r a p s , sewage filters, septic tanks, a n d similar places. Adults of these a r e often seen in­ d o o r s , especially in b a t h r o o m s a n d lavato­ ries, w h e r e they are a m i n o r nuisance.

MOTH FLIES

367

Save for these few, little is k n o w n of Neotropical m o t h flies (see t h e l i t e r a t u r e reviews in D u c k h o u s e 1977, 1 9 8 1 , 1982). Sand flies of t h e subfamily P h l e b o t o m i n a e (see below), however, a r e b l o o d s u c k e r s a n d t r a n s m i t several h u m a n a n d animal dis­ eases a n d a r e well s t u d i e d .

References DUCKHOUSE, D. A. 1973. Family Psychodidae. In N. Papavero, ed., 1967—, A catalogue of the Díptera of the Americas south of the United States. Mus. Zool., Univ. Sao Paulo, Sao Paulo. Pp. 6A.1-6A.29. DUCKHOUSE, D. A. 1977. Psychodidae. In S. H.

Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 266-267. DUCKHOUSE, D. A. 1981. Psychodidae. In S. H.

Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pi. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 241-245. DUCKHOUSE, D. A. 1982. Psychodidae. In S. H.

Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 414-416. HOGUE, C. L. 1973. A taxonomic review of the genus Maruina (Díptera: Psychodidae). Los Angeles Co. Mus. Nat. Hist. Bull. 17: 1-69. SEBASTIANI, F. L. 1978. Ciclo biológico de Telmaloscopus albipunctatus (Williston, 1893) (Díp­ tera, Psychodidae). I. Comportamento sex­ ual. Cien. Cult. 30(6): 719-722.

SAND FLIES Psychodidae, P h l e b o t o m i n a e , Lutzomyia. Spanish: Manta-blancas (General), moscas m o r r a n o s (Colombia), chitras ( P a n a m a ) , titiras (Peru). Portuguese: A r r u p i a d o s , a s a - d u r a s , mosquitos palha (Brazil). Tupi-Guaraní: Birigüi. Females of several g e n e r a o f P s y c h o d i d a e (once c o m b i n e d with t h e O l d World spe­ cies as t h e single g e n u s Phlebotomus; Lewis et al. 1977, T h e o d o r 1965) feed o n verte­ b r a t e blood a n d a r e i m p o r t a n t vectors of h u m a n a n d animal diseases (Lewis 1974). In Latin America, species of Lutzomyia

368

FLIES AND MIDGES

transmit several types of leishmaniases (Kil lick-Kendrick 1978, W a r d 1977) from Mex­ ico to A r g e n t i n a (but not, strangely ¡ n Chile a n d U r u g u a y ) . T h e most prevalent a r e so-called c u t a n e o u s leishmaniases (y»a. palomoyo, espundia, uta, pian bois, bubón de Vélez, bubón de Aleppo, úlcera de Chinácota etc.), resulting from infections by various species of Leishmania, L. brasiliensis, L. peru­ viana, a n d L. mexicana. I n t h e n e a r past o n e variety of t h e last species (Leishmania m. mexicana) was c o m m o n a m o n g chicle g a t h e r e r s in Mexico, G u a t e m a l a , a n d Be­ lize, whose ulcerated ears, acquired during forays in t h e forest w h e r e t h e disease was rife a n d sand flies c o m m o n , became the hallmark of their profession (chiclero's ulcer). Visceral leishmaniasis (caused by Leishmania donovani chagasi) is also widely distributed in S o u t h A m e r i c a where a p r o v e n vector is Lutzomyia longipalpis. Sand flies also t r a n s m i t Peruvian ve­ r r u g a (Oroya fever, carrion's disease), which is restricted to certain A n d e a n val­ leys in Peru, Colombia, a n d Ecuador. This is an often deadly rickettsial disease, mani­ fested at o n e stage by ugly skin warts. It is transmitted t h r o u g h t h e bite of Lutzomyia verrucarum a n d L. colombiana. Pizarro's t r o o p s w e r e d e c i m a t e d by this disease d u r i n g military expeditions in 1533—1537 in Peru, a n d t h e r e is evidence that v e r r u g a was t h e cause of d e a t h of the Inca H u a y n a C a p a c shortly before that time (Patrón 1896). I n t h e 1870s, 7,000 d e a t h s were r e p o r t e d from t h e affliction in t h e Rimac Valley of Peru, when the construction of t h e T r a n s - A n d e a n railway was i m p e d e d by this sickness suffered by t h o u s a n d s of laborers (Noguchi et al. 1928). In addition to leishmanias, wild sand flies h a r b o r t r y p a n o s o m a t i d flagel­ lates (Christensen a n d H e r r e r 1976) and many viruses of u n k n o w n effects on their hosts (e.g., pacui virus in rice rats; Aitken e t a l . 1975). Sand flies (Martins et al. 1971) a r e small, hairy psychodids (BL 1.5-4 m m ) , grayish-

Mure 11-2 BITING MIDGES, (a) Sand fly {Lutzomyia sp., Psychodidae). (b) Sand fly, larva. JtjPunkie (Culicoides sp., Ceratopogonidae). (d) Punkie, larva, (e) Blackfly (Simulium sp., SJrnuliidae). (f) Blackfly, larva, (g) Blackfly, pupa in cocoon.

yellow to b r o w n , with e l o n g a t e wings that are held obliquely away from t h e body when at rest (fig. 11.2a). T h e y have a slender body a n d a set of e l o n g a t e biting mouthparts (Lewis 1975). T h e i r flight is also characteristic: a series of short, erratic hops, in which t h e fly seldom moves m o r e than a short distance. Adults often rest in sheltered places (Chaniotis et al. 1972), soil a n d rock crev­ ices, between t r e e buttresses, u n d e r loose tree bark, in caves (Williams 1976), tree holes, animal b u r r o w s , a n d t e r m i t e nests, and sometimes in h u m a n dwellings. T h e majority of t h e 260 o r m o r e Neotropical species a r e forest dwellers (Porter a n d De Folian 1 9 8 1 , Young 1979), a n d females normally suck t h e blood of sylvan m a m ­ mals, birds, a n d reptiles. B o t h sexes also take plant sugars in t h e form of nectar, honeydew, a n d t h e like. T h e larvae (fig. 11.2b) a r e s e p a r a t e d from those of o t h e r psychodids by a d o u ­ ble pair of l o n g anal spines. T h e y live in soil, feeding o n o r g a n i c debris such as excrement, d e a d insects, d e c a y i n g plant matter, a n d fungi.

sites of phlebotomine sandflies in a Panaman­ ian tropical forest, j . Med. Entomol. 9: 9 1 - 9 8 . CHRISTENSEN, H.

A.,

AND A.

HERRER.

LEWIS, D. J., D. G. YOUNG, G. B. FAIRCHILD, AND

D. M. MINTER. 1977. Proposals for a stable classification of the Phlebotomine sandflies (Diptera: Psychodidae). Syst. Entomol. 2: 319-332. MARTINS, A. V, D. J. LEWIS, AND O. THEODOR.

1971. Phlebotomine sandflies. World Health Org. Vector Biol. Contr. Unit. Pap. 71.255: 1-23. NOGUCHI, H., R. C. SHANNON, E. B. TILDEN, AND

J. R. TYPER. 1928. Phlebotomus and Oroya fever and verruga peruana. Science 68: 493-495. PATRÓN, P. 1896. Apuntes históricos sobre la verruga americana. Soc. Geogr. Lima Bol. 5(4): 435-445. PORTER, C. H., AND G. R. D E FOLIART.

References AITKEN, T. H. G., J. P. WOODALL, A. H. P. DE ANDRADE, G. BENSABATH, AND R. E. SHOPE.

1975. Pacui virus, phlebotomine flies, and small mammals in Brazil: An epidemiological study. Amer. J. Trop. Med. Hyg. 24: 358-368. CHANIOTIS, B. N., R. B. TESH, M. A. CORREA,

AND K. M. JOHNSON. 1972. Diurnal resting

1976.

Neotropical sand flies (Díptera: Psychodidae), invertebrate hosts of Endotrypanum schaudinni (Kinetoplastida: Trypanosomatidae). J. Med. Entomol. 13: 299-303. KILLICK-KENDRICK, R. 1978. Recent advances and outstanding problems in the biology of phlebotomine sandflies. Acta Trópica 35: 297-313. LEWIS, D. J. 1974. T h e biology of Phlebotomidae in relation to Leishmaniasis. Ann. Rev. Entomol. 19: 363-384. LEWIS, D. J. 1975. Functional morphology of the mouth parts in New World phlebotomine sandflies (Diptera: Psychodidae). Royal Ento­ mol. Soc. London Trans. 126: 497-532.

1981.

The man-biting activity of phlebotomine sandflies (Diptera: Psychodidae) in a tropical wet forest in Colombia. Arq. Zool. (Sao Paulo) 30:81-158. THEODOR, O. 1965. On the classification

of

American Phlebotominae. J. Med. Entomol. 2: 171-197. WARD, R. D. 1977. New World leishmaniasis: A

SAND FLIES

369

review of the epidemiological changes in the last three decades. 15th Int. Cong. Entomol. Proc. Pp. 505-522. WILLIAMS, P. 1976. The phlebotomine sandflies (Díptera, Psychodidae) of caves in Belize, Central America. Bull. Entomol. Res. 65: 601-614. YOUNG, D. G. 1979. A review of the blood­ sucking psychodid flies of Colombia (Diptera: Phlebotominae and Sycorocinae). Univ. Fla. Insl. Food Agrie. Sci. Agrie. Esp. Sta. Bull. 806: 1-266.

PUNKIES C e r a t o p o g o n i d a e . Spanish: Jejenes (var. ihenni), majes, plagas, m i m e s (General). Portuguese: M a r u i n s , mosquitos pólvora, b e m b é s (Brazil). French: Bigailles (Haiti). No-see-ums. T h e smallest of t h e biting flies, p u n k i e s m e a s u r e only 1 to 4 millimeters in body length. T h e y a r e gnatlike, s o m e w h a t r e s e m ­ bling blackflies, b u t their wings a r e elon­ gate, with few veins, often d a r k p i g m e n t e d with clear spots, a n d folded flat over t h e a b d o m e n w h e n at rest. T h e legs a r e slen­ d e r b u t short. Punkies have m o u t h p a r t s a d a p t e d for liquid food like t h o s e of o t h e r biting midges, b u t they feed o n a wider variety of a n i m a l hosts a n d plants as well. A few Atrichopogon, Pterobosca, a n d Forcipomyia a r e k n o w n to be ectoparasites o n a d u l t a n d larval l e p i d o p t e r a n s , o r t h o p t e r a n s , a n d o t h e r insects (Wirth 1956). Culicoides, Lasiohelea, a n d Leptoconops species suck blood from birds, m a m m a l s , including h u m a n s , a n d frogs. Many g e n e r a only take nectar from flowers (Dasyheled). S o m e species of Dasyhelea, of Forcipomyia, and small, hairy m e m b e r s of o t h e r g e n e r a , a r e pollinators of Para r u b b e r ( W a r m k e 1952), cacao (Bystrak a n d Wirth 1978, Y o u n g 1983), a n d o t h e r tropical c r o p s ( W i n d e r 1978). Many of t h e b l o o d - f e e d i n g Culicoides (fig. 11.2c) (Forattini 1957) a r e n o t o r i o u s pests a n d bite intensely, seemingly well beyond t h e capacities o f such m i n u t e crea­

370

FLIES AND MIDGES

tures (Aréan a n d Fox 1955, Kettle 1977\ T h e y often swarm in intolerable numbe in m a n g r o v e swamps a n d along r ¡ V e b a n k s o r m a r s h e s , m a k i n g such piar uninhabitable. Like blackflies a n d mosqu' toes, they a r e m e n t i o n e d in every account of tropical travel a m o n g t h e greatest haz ards encountered. The tiny black ihenni of the Bolivian part of the Amazon watershed, has a bite like the burn of a cigarette and travels millions strong. It has an uncanny ability to get un­ der or through any net. It belongs to no union and knows no hours—it attacks inces­ santly every minute of the day and night. Several travelers through this region have died, not directly from the bites of the in­ sects, but because of the impossibility of get­ ting a moment's rest. (Price 1952: 143) T h e i r bites often i n d u c e severe allergic reactions. Skin reactions r a n g e from mild itching to blistering a n d o p e n lesions often complicated by secondary infections. In some species of t e m p e r a t e areas, the larger females often eat t h e smaller males d u r i n g mating. T h i s habit has not been observed for tropical species but is likely to occur, since m a n y of t h e same or related g e n e r a , such as Bezzia a n d Palpomyia, in­ habit both regions (Downes 1978). Culicoides furens, f o u n d t h r o u g h o u t the C a r i b b e a n a n d along t h e Middle American coasts to E c u a d o r a n d Brazil, is the most widespread biter. It a n d related species are vectors of filarial h e a r t w o r m (Mansonella ozzardi) between a n d a m o n g h u m a n s and dogs. T h e g e n u s also h a r b o r s a variety of h u m a n pathological viruses (Linley et al. 1983). In tropical America, Culicoides trans­ mit onchocerciasis, blue t o n g u e , a n d other sicknesses of horses, s h e e p , a n d cattle, as well as p r o t o z o a n infections of birds (Kettle 1965). Curiously, t h e g e n u s is virtually ab­ sent from South America south of the a p p r o x i m a t e latitude of Santiago, Chile. Most i m m a t u r e punkies a r e aquatic or subaquatic, occupying m a n y habitats, usu-

«I m u d o r wet sand o n lake, p o n d , river, o r tream m a r g i n s , b u t they also live in water , c u m u l a t i o n s in w a t e r - h o l d i n g plants (Helignia Calathea inflorescences, b r o m e l i a d leaf axils; Wirth a n d d e J. Soria 1981). Marshes a n d l a n d c r a b b u r r o w s a r e favored breeding sites as well. O t h e r species a r e semiterrestrial, their larvae d e v e l o p i n g in damp situations u n d e r b a r k or in r o t t i n g wood, fruit, cactus stems, a n d b a n a n a stalks, or in d e c o m p o s i n g leaves. T h e larvae a r e very long a n d slender /jjL 1—3 m m ) , white, a n d practically hair­ less. T h e eellike freely aquatic species (fig. 11.3d) a r e recognized by their s e r p e n t i n e swimming m o v e m e n t s . T h e family h a s a b o u t 7 0 0 Neotropical species g r o u p e d into over 30 g e n e r a (Wirth 1974). Many species certainly r e m a i n u n d i s ­ covered. T h e l i t e r a t u r e o n t h e Neotropical members of t h e g r o u p has been well in­ dexed (Atchley et al. 1981; Wirth a n d Cavalieri 1977; Wirth 1 9 8 1 , 1982).

References ARÉAN, V. M., AND 1. Fox. 1955. Dermal alter­ ations in severe reaction to the bite of the sandfly, Culicoides furens. Amer. J. Clin. Path. 25: 1359-1366. ATCHLEY, W. R., W. W. W I R T H , C. T. GASKINS,

AND S. L. STRAUSS. 1981. A bibliography and

keyword index of the biting midges (Diptera: Ceratopogonidae). U.S. Depl. Agrie. Bibliog. Lit. Agrie. 13: 1-544. List of all publications on the family from 1758 to 1978. BYSTRAK, P. G., AND W. W. W I R T H . 1978.

The

North American species oí Forcipomyia, subgenus Euprojoannisia (Diptera: Ceratopogoni­ dae). U.S. Dept. Agrie. Tech. Bull. 1591: 1-51. DOWNES, J. A. 1978. Feeding and mating in the insectivorous Ceratopogonidae (Diptera). En­ tomol. Soc. Can. Mem., 104: 1-62. FORATTINI, O. P. 1957. Culicoides da regiáo Neotropical (Diptera, Ceratopogonidae). Fac. Hig. Saud. Púb. Univ. Sao Paulo Arq. 11: 161-526. KETTLE, D. S. 1965. Biting ceratopogonids as Vectors of human and animal diseases. Acta Trop. 22: 356-562. KETTLE, D. S. 1977. Biology and bionomics of bloodsucking ceratopogonids. Ann. Rev. En­ tomol. 22: 3 3 - 5 1 .

LINLEY, J. R., A. L. H O C H , AND F. P. PINHEIRO.

1983. Biting midges (Diptera: Ceratopogo­ nidae) and human health. J. Med. Entomol. 209: 347-364. PRICE, W. 1952. The amazing Amazon. John Day, New York. WARMKE, H. E. 1952. Studies on natural pollina­ tion of Hevea brasiliemis in Brazil. Science 116: 474-475. WINDER, J. A. 1978. Cocoa flower Diptera, their identity, pollinating activity, and breeding sites. Pest Art. News Sum. 24: 5 - 1 8 . WIRTH, W. W. 1956. New species and records of biting midges ectoparasitic on insects (Dip­ tera, Heleidae). Entomol. Soc. Amer. Ann. 49: 356-364. WIRTH, W. W. 1974. Family Ceratopogonidae. In N. Papavero, ed., 1967-, A catalogue of the Diptera of the Americas south of the United States. No. 14. Mus. Zool. Univ. Sao Paulo, Sao Paulo. Pp. 14.1-14.89. WIRTH, W. W. 1981. Ceratopogonidae. In S. H.

Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 268-275. WIRTH, W. W. 1982. Ceratopogonidae. In S. H.

Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 438-442. WIRTH, W. W., AND F. CAVALIERI. 1977. Cerato­

pogonidae. In S. H. Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 280-283. WIRTH, W. W., AND S. DE J. SORIA. 1981.

Two

Culicoides biting midges reared from inflores­ cences of Calathea in Brazil and Colombia, and a key to the species of the discrepans group (Diptera: Ceratopogonidae). Rev. Theobroma 11: 107-117. YOUNG, A. M. 1983. Seasonal differences in abundance and distribution of cocoa-pollina­ ting midges in relation to flowering and fruit sel between shaded and sunny habitats of the La Lola cocoa farm in Costa Rica. J. Appl. Ecol. 10: 801-831.

BLACKFLIES Simuliidae. Spanish: Mosquitos pelones, alazanes, r o d e d o r e s , poicos, tabardillos (General); majes, perrujas, moscas d e café (Costa Rica); b a r r i g o n e s , jerjeles (Venezuela, Peru). Portuguese:

BLACKFLIF:S

371

B o r r a c h u d o s , p i u n s (Brazil). K a b o u r a flies (Guianas). Buffalo gnats, coffee flies. T h e s e d i u r n a l biting gnats (Darsie 1982; Vulcano 1977, 1981) a r e similar to p u n k i e s but are generally l a r g e r (BL 1-5 m m ) , often with a black body, a l t h o u g h m a n y a r e gray o r yellow with silvery o r g o l d e n p u b e s ­ cence. T h e i r a n t e n n a e a n d legs are short, a n d their wings a r e i m m a c u l a t e a n d broadly triangular. T h e females suck blood from warm­ blooded vertebrates. Near fast-flowing rocky s t r e a m s a n d rivers w h e r e their larvae develop, they m a y o c c u r in such n u m b e r s a n d bite with such persistence as to drive off h u m a n s a n d animals alike a n d r e q u i r e d e t e r m i n e d control (Laird 1981). T h e bite is not b o t h e r s o m e at first b u t causes a b u r n i n g sensation that m a y last for days; the bite of some species is m a r k e d by a small, persistent blood spot. A m a z o n naturalist-explorer Alfred Russell Wallace ( 1 8 5 3 : 310) r e c o r d e d his e x p e r i e n c e with blackflies t h u s : "My feet were so thickly covered with t h e little blood-spots p r o ­ d u c e d by their bites, as to be of a d a r k p u r p l i s h - r e d colour, a n d m u c h swelled a n d inflamed. . . . T h e only m e a n s of taking a little rest in t h e day was by w r a p p i n g u p h a n d s a n d feet in a blanket." T h e blackfly f a u n a of Latin A m e r i c a is e x t r e m e l y rich, with over 300 species dis­ tributed in 10 g e n e r a , o c c u r r i n g in practi­ cally all regions a n d r u n n i n g water habitats from high A n d e a n melt waters a n d out­ flows of hot springs to the g r e a t cataracts of the major rivers (Coscaron 1987). T h e i r distribution includes most of the major C a r i b b e a n islands, b u t they a r e absent from the Galápagos a n d Malvinas; mysteri­ ously, they a r e k n o w n o n the island of M a r g a r i t a off the Venezuelan coast w h e r e there are no streams. Species in t h e g e n u s Simulium (fig. 11.2e) bear a n u m b e r of serious diseases affecting their v e r t e b r a t e hosts (Coscaron 1984). T h e

372

FLIES AND MIDGES

most i m p o r t a n t of these is onchocerciasis (enfermedad de robles), caused by the filarial w o r m Onchocerca volvulus, which, before the conquest, was absent from America. These filarial w o r m s u n d o u b t e d l y were intro­ d u c e d with slaves from Africa a n d became p e r m a n e n t l y established in coffee-growing areas of G u a t e m a l a (Dalmat 1955), neigh­ b o r i n g El Salvador, a n d t h e Mexican states of Chiapas a n d Oaxaca. T h e r e are now secondary foci in Venezuela, e x t r e m e north­ e r n Brazil, Colombia, a n d Ecuador. T h e affliction causes blindness in m a n y people in these locales w h e n t h e w o r m s situate in ocular tissues. Major vector species in Guate­ mala a r e S. ochraceum, S. metallicum, a n d S. callidum ( A n o n . 1 9 7 1 , G a r m s 1975). T h e t a x o n o m y of o t h e r vectors is still unsettled (Tidwell et al. 1981). Blackflies are also suspected of transmitting t h e hemorrhagic s y n d r o m e s of Altamira a n d black fever of Lábrea, recently recognized diseases of un­ known etiology discovered along the TransA m a z o n i a n Highway in Brazil (Anon. 1966, Pinheiro et al. 1974). T h e d o g h e a r t worm (Mansonella ozzardi) is also transmitted by Simulium amazonicum a n d has been found to affect a significant p a r t of the h u m a n popu­ lation, especially indigenes, in parts of Cen­ tral America, the West Indies, a n d the A m a z o n Basin (Shelley et al. 1982). These findings haved precipitated new interest in studies o n Neotropical blackflies (Lacey and C h a r l w o o d 1980). T h e r e a r e sixty-one spe­ cies k n o w n to bite h u m a n s in the Neotropics (Travis et al. 1974). Simuliid larvae (fig. 11.2f) a r e obliga­ tory inhabitants of swift streams a n d river rapids w h e r e they cling to rocks a n d other h a r d substrate surfaces, h e a d trailing in the c u r r e n t , in g r o u p s of a few to hun­ d r e d s of individuals (Takaoka 1981). They are elongate with a swollen posterior and a well-developed h e a d ; e x p a n d e d mouthb r u s h e s projecting from the u p p e r side of the head catch drifting p l a n k t o n a n d detri­ tus, closing intermittently to transfer the accumulated catch to the m o u t h . Around

an anal sucker a n d o n a small lobe b e n e a t h the head a r e fine hooks that, with the sucker, a n c h o r larvae to a silk m a t that they spin with modified salivary glands. If dis­ turbed, larvae loop t h e h e a d e n d into the current a n d m a y escape in u n i q u e fashion by releasing their hold a n d allowing t h e m ­ selves to be swept away into t h e c u r r e n t . Simultaneously, they let o u t a silk a n c h o r line with which they reel themselves back to their f e e d i n g site w h e n t h e d a n g e r has passed. Pupation usually occurs in a silk cocoon, by whose s h a p e species m a y often be identi­ fied. T h e r e s p i r a t o r y o r g a n s of the p u p a (fig. 11-2g) itself a r e also diagnostic in shape a n d b r a n c h i n g p a t t e r n . T h e cocoon is attached to s u b m e r g e d rocks so that the adult must later e m e r g e from some d e p t h . After leaving t h e p u p a l case a n d cocoon, the imago rises to the surface in an air bubble, from which it " p o p s " into flight.

References ANONYMOUS. 1966. Studies on the Amazonian fever (febre de Lábrea; Febre Negra; etc.). Belem Virus Lab. Ann. Rpt. App. Pp. 139— 157. ANONYMOUS. 1971. Blackflies in the Americas. World Health Org. Vector Biol. Control Unit Pap. 71.283 4:1-24. COSCARON, S. 1984. Simúlidos sudamericanos: Diptera-lnsecta. 9th Cong. Latinoamer. Zool. (Arequipa) Info. Final. Pp. 165-168. COSCARON, S. 1987. El género Simulium Latreille en la Región Neotropical: Análisis de los grupos sujarespedficos, especies que los integran y distribución geográfica (Simuliidae, Díptera). Mus. Paraense Emilio Goeldi, Belem. DALMAT, H. T. 1955. T h e black flies (Díptera: Simuliidae) of Guatemala and their role as vectors of onchocerciasis. Smithsonian Misc. Coll. 125: 1-425.

transmission of Onchocerca volvulus. Tropenmed. Parasit. 26: 169-182. LACEY, L. A., AND J. D. CHARLWOOD. 1980.

On

the biting activities of some anthropophilic Amazonian Simuliidae (Díptera). Bull. Entomol. Res. 70: 495-509. LAIRD, M., ed. 1981. Blackflies: The future for biological methods in integrated control. Aca­ demic, London. PINHEIRO, F. P., D. COSTA, Z. C. LINS, G. BENSABATH. O. M. MAROJA, AND A. H. P.

ANDRADE. 1974. Haemorrhagic syndrome of Altamira. Lancet 1: 639-642. SHELLEY, A. J.,

R.

R. PINGER, AND M. A.

P.

MORAES. 1982. The taxonomy, biology and medical importance of Simulium amazonicum Goeldi (Díptera: Simuliidae), with a review of related species. Brit. Mus. Nat. Hist. Bull. Ser. Entomol. 44: 1-29. TAKAOKA, H. 1981. Seasonal occurrence of Simulium ochraceum, the principal vector of Onchocerca volvulus in the southeastern en­ demic area of Guatemala. Amer. J. Trop. Med. Hyg. 30: 1121-1132. TIDWELL, M. A., B. V PETERSON, f. RAMÍREZ, M. DE TIDWELL, AND L. A. LACEY. 1981. Notas y

claves preliminares de los jejenes Simulium amazonicum y S. sanguineum (Díptera: Simulii­ dae) incluyendo los vectores de Onchocerca volvulus y Mansonella ozzardi. Dir. Malar. San. Amb. Bol. 21: 79-89. TRAVIS, B. V, M. VARGAS, AND f. C. SWARTZ-

WELDER. 1974. Bionomics of black flies (Díp­ tera: Simuliidae) in Costa Rica. 1. Species biting man, with an epidemiological summary for the Western Hemisphere. Rev. Biol. Trop. 22: 187-200. VULCANO, M. A. 1977. Simuliidae. In S. H.

Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 285-293. VULCANO. M. A. 1981. Simuliidae. In S. H.

Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 275-285. WALLACE, A. R. 1853. A narrative of travels on the Amazon and Rio Negro. Reeve, London.

DARSIE, JR., R. p. 1982. Simuliidae. In S. H.

Hurlberl and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 443-449. «ARMS, R. 1975. Observations on filarial infec­ tions and parous rates of anthropophilk blackflies in Guatemala, with reference to the

MOSQUITOES Culicidae. Spanish: Mosquitos, z a n c u d o s (General). Náhuatl: M o y o m e h (sing. moyotl) (Mexico). Portuguese:

MOSQUITOES

373

Mosquitos, pernilongos, muricocas (Brazil). Tupi-Guaraní: Carapaná. For many, images of hordes of blood­ thirsty mosquitoes arise when the word "tropics" is mentioned. Indeed, humid lowland swamps, teeming jungles, and mosquitoes are almost synonymous to those without direct experience in tropical biology. In truth, although there are more mosquito species in the lower latitudes than in the arcto-temperate zones, they seldom appear in such great numbers, and then only locally, certainly never like the vast multitudes spawned by the sum­ mer thaw on the Alaskan and Siberian tundras. Yet sometimes their attacks be­ come sufficiently persistent to kill. Pedro Teixeira, while journeying through Mex­ ico in 1600, wrote, "Almost all along this road [Acapulco to San Juan] is a plague of mosquitoes, so terrible and grievous that no defence avails against them, and they stung my best slave to death" (Gillett 1971). Mosquito diversity in Latin America is phenomenal, however; almost a third of the world's more than 3,000 species, in 18 genera, are found here (Knight 1978, Knight and Stone 1977, Mattingly 1971), as many as in any other single zoogeographic region. The best known are the medically important genera, especially Anopheles, Aedes, and Culex. Some dominant groups are the subgenus Melanoconion of Culex and the sabethines, especially Wyeomyia. Exploi­ tation of the innumerable available breed­ ing sites probably has engendered their variety. Some classic, general works on the taxonomy of the family in the region are those of Dyar (1928) and Howard, Dyar, and Knab (1912-1917). Although histori­ cally important, they are outdated and are now replaced by a multitude of modern contributions on specific groups. But as yet, there is no comprehensive work on the fauna in general. This is greatly needed. Since mosquitoes are the best-known

374

FLIES AND MIDGES

large family of organisms, much can be said about the developmental stages and other aspects of their biology (Bates 1949 Horsfall 1955, Machado-Allison 19801982). It is a mistake to think only 0 f stagnant swamps as the mosquito's do­ main, although larvae do not survive in currents flowing much faster than a few centimeters per second. Neither is fresh water a requirement. Many species breed in coastal marshes and mangrove swamps (Aedes taeniorhynchus,

Anopheles

Figure 11.3 MOSQUITOES (CULICIDAE). (a) Malaria mosquito (Anopheles darlingi). (b) Malaria mosquito, pupa, (c) Malaria mosquito, larva, (d) Giant mosquito (Toxorhynchites sp.). (e) Yellow fever mosquito (Aedes aegypti). (f) Yellow fever mosquito, larva.

albimanus)

limestone solution holes (Culex opisthopus) land crab burrows (Deinocerites), and even coral rock holes (Culex bahamensis), where brackish, salty, or sometimes even supersaline aquatic conditions prevail. Larvae that live in these situations have special physio­ logical mechanisms for maintaining inter­ nal water balance; for example, the rectal gills, organs normally used to absorb salts, are greatly reduced. Mosquito populations tend to build dur­ ing the early part of the rainy season when the number of available breeding sites increases. Even minor rainfall fluctuations in nonseasonal areas may effect changes in numbers (Wolda and Galindo 1981). The most common habitats of the immatures are freshwater accumulations on the ground, especially where there is abun­ dant vegetation, such as in swamps, in marshes, or along the overgrown margins of lakes, ponds, and rivers. This is the home of the ubiquitous Anopheles, Culex, and Aedes mosquitoes, but others typically breed here, such as Uranotaenia. The amount of water need not be great; small puddles and pools and even small niches on the surfaces of rocks or animal foot­ prints may hold enough water to support mosquito life. However, because the nutrient-poor groundwaters of central Amazonia support little food, the region is relatively free of mosquitoes, save those breeding in containers and epiphytes. Larvae and pupae can thrive in incredi­ bly small amounts of water, as is often

found in various natural and artificial containers. T h e former includes the hol­ lowed or cupped parts of plants in which rainwater collects, such as rot holes in trees, limbs, and logs (Aedes, Toxorhynchites), on fallen leaves, especially large, broad, thick types that drop from trees like Cecropia and aroids (Aedes, Wyeomyia), and in fruit husks. Trichoprosopon digitatum larvae form extremely dense populations in the fetid liquid accumulating in hollow cacao pods that have had their contents eaten away by scavengers. Certain types of New World tropical plants also collect water: bromeliads in their leaf axils, bamboo in hollow stems, Heliconia in flower bracts, and pitcher plants (Heliamphora). These are the niches occupied by specialized genera with numerous species, often with highly local distributions (Wyeomyia, Sabethes).

Larval mosquitoes are elongate with swollen thorax and well-developed, pigmented head (fig. 11.3f). The mouth has brushlike elements (developed from the labrum), kept almost constantly in motion to sweep suspended microorganisms and organic particles into the mandibles and mouth opening. In some predaceous forms (Toxorhynchites), the filaments of the mouth brushes are heavy blades used to grasp and hold prey, which is then chewed and swallowed in bits. Projecting from the last abdominal seg­ ment is a sclerotized breathing tube contain"•g the major trachea and entry pores for

air at its apex. The larva relies on atmo­ spheric oxygen and must come to the sur­ face often to take in fresh air. A few types (Mansonia, Coquillettidea) have a breathing tube modified for piercing the air-filled vessels of aquatic plants and never need to break the surface film to breathe. Larvae swim by rapid sideways jerking movements of the body, for which they are called "wrigglers"; pupae are also active and progress similarly, except that the action of the abdomen is forward under the large, oval thorax, and propulsion is aided by a pair of overlapping, rounded flat paddles at the terminus of the abdo­ men (fig. 11.3b). Rapidly moving pupae often roll over and over, a motion earning them the name "tumblers." The outer body cuticle of both of these stages is set with numerous, often complex hairs whose configurations are extensively used in clas­ sification and identification. The shape of the "trumpets," breathing organs project­ ing from the anterior part of the thorax of the pupa, is also of diagnostic value. Mosquitoes also live varied and involved lives as adults. The females of nearly all but those belonging to the species with preda­ ceous larvae (Toxorhynchites) take terrestrial vertebrate blood as their principal protein food source. The female bites wherever she can penetrate surface capillaries, through exposed skin almost anywhere on the bod­ ies of mammals and amphibians, through the feet and base of the beaks of birds, and between the scales of reptiles. Blood meals

MOSQUITOES

375

may n o t be r e q u i r e d by some (autogenous) strains, which e x p l a i n s h o w species may survive in areas t e m p o r a r i l y without a nor­ mal food source. T h e adult is distinguishable from Díp­ tera in g e n e r a l a n d o t h e r biting midges by having scales o n e l o n g a t e wings a n d a very long proboscis. T h e legs a r e also very long a n d fragile, a n d t h e eyes a r e well-devel­ o p e d , large h e m i s p h e r e s . I n most, males may be distinguished by densely p l u m o s e a n t e n n a e ; female a n t e n n a e a r e only lightly haired. L a r g e swarms of males emit ultrasonic whines with their vibrating wings to attract females for m a t i n g . Most mosquitoes follow this p a t t e r n . Deinocerites, however, never swarm, a n d males seek females directly in crab b u r r o w s (see c r a b h o l e mosquitoes, below). Mosquito females lay their eggs either directly o n o r n e a r water. Eggs m a y be placed singly o r in small floating g r o u p s of a h u n d r e d o r m o r e (egg rafts). Eggs placed out of water a r e d r o u g h t resistant a n d may r e m a i n viable for m o n t h s b e f o r e i n d u c e d to h a t c h by rising w a t e r in t h e i r habitat. A d u l t mosquitoes a r e by far t h e most i m p o r t a n t insect vectors of diseases to h u m a n s a n d o t h e r v e r t e b r a t e s in t h e N e o tropics. Historically, malaria, b o r n e by Anopheles species, h a s b e e n t h e greatest killer a n d debilitator of h u m a n k i n d . Its incidence was r e d u c e d to a low level as a result of t h e w i d e s p r e a d control c a m p a i g n s waged with D D T following World W a r I I . However, t h e d e v e l o p m e n t of insecticide resistance by t h e vectors, e c o n o m i c r e ­ straints, d r u g resistance by t h e plasmodia, a n d g o v e r n m e n t a p a t h y t o w a r d control p r o g r a m s have allowed a d r a m a t i c resur­ gence of t h e disease in t h e 1960s a n d 1970s (Agarwal 1978, B r o w n et al. 1976, C h a p i n a n d Wasserstrom 1981). A l t h o u g h all four species of h u m a n malaria a r e found in t h e New World tropics, Plasmodium falciparum is d o m i n a n t a n d t h e most serious. Yellow fever is second to malaria in

376

FLIES AND MIDGES

prevalence, but it is a m o r e virulent HU ease. It is t h o u g h t that t h e u r b a n vector Aedes aegypti, s p r e a d from Africa a n d wa< responsible for epidemics d u r i n g the coin. nization of t h e Americas, although the j u n g l e form, m a i n t a i n e d by native Haerna gogus, Aedes, a n d o t h e r vectors, may have existed h e r e in p r e - C o l u m b i a n times. T h e disastrous effect of t h e disease on the construction of t h e P a n a m a Canal is \v e || k n o w n . Like malaria, yellow fever is ¡n. creasing in o c c u r r e n c e , following a decline resulting from t e m p o r a r i l y successful con­ trol efforts in t h e first half of t h e century Many other, probably indigenous, viral infections a r e k n o w n to have mosquito vectors in t h e Neotropics, including den­ gue, transmitted by Aedes aegypti, and Saint Louis, Eastern, Venezuelan, and o t h e r encephalitides t r a n s m i t t e d by Culex a n d Aedes species (McLintock 1978). Bancroftian filariasis is n o n i n d i g e n o u s but has become established in limited foci, un­ d o u b t e d l y by i n t r o d u c t i o n from Africa d u r i n g t h e slave days. T h e s e points of infection a r e scattered t h r o u g h o u t the Antilles a n d n o r t h e a s t e r n coastal a n d subcoastal South America w h e r e t h e vectors are mainly house mosquitoes (Culex quinquefasciatus). While t h e seriousness of these diseases is widely recognized, t h e direct effects of the bites of mosquitoes should n o t be dis­ c o u n t e d . Saliva injected with t h e bite con­ tains proteins foreign to t h e h u m a n physiol­ ogy a n d often elicits s t r o n g sensitization reactions. Allergic reactions manifest on the skin as mild to severe urticaria, forma­ tion of s u b d e r m a l tubercles, o r production of large, water-filled blisters; systemic symp­ toms a r e fever a n d nausea. Anthropophilic species often show p r e f e r e n c e for attacking different hosts a n d areas of the body. Stud­ ies to d e t e r m i n e t h e reasons for these preferences a r e inconclusive, but some gen­ eralities a r e evident. Persons with lighter skin p i g m e n t a t i o n seem t o be m o r e attrac­ tive t h a n d a r k e r - s k i n n e d individuals. Ex-

ed carbon d i o x i d e is a major attractant, though o t h e r o d o r s e m a n a t i n g from t h e "ju a lso d r a w mosquitoes. Favored feeding ots a r e t h e ankles, elbows, ears, a n d back «f the h a n d s ; w a r m skin is p r e f e r r e d over cold o r hot. T i g h t l y w o r n clothing is n o t a -nod protection since most mosquitoes can Wte t h r o u g h t h e weave. M o d e r n chemical 'impellents (such as N, N-diethyl-m-toluajiiide, D E E T ) a r e effective if s t r o n g solu­ tions a r e e m p l o y e d a n d m a i n t a i n e d o n t h e

skin. T h e effects of a few bites a r e usually -ggligible, b u t t h e persistence of mosqui­ l e s a n d their a b u n d a n c e in s o m e places lenders t h e m f o r m i d a b l e pests. Tales from the colonial p e r i o d of e x p l o r a t i o n tell of mosquito attacks a n d t h e i r curious effects j©n people. S e v e n t e e n t h - c e n t u r y F r e n c h aturalist Mouffet said, " T h e gnats in ¡America d o so slash a n d cut, that they will pierce t h r o u g h very thick clothing; so that is excellent s p o r t to b e h o l d h o w ridicuusly the b a r b a r o u s p e o p l e , w h e n they a r e bitten will skip a n d frisk, a n d slap with eir hands their thighs, buttocks, shoul­ ders, arms a n d sides, even as a c a r t e r d o t h horses." v o n H u m b o l d t a n d B o n p l a n d 1852, 2: 274) f o u n d d u r i n g their South aerican travels in 1 7 9 9 - 1 8 0 4 that "he­ neen t h e little h a r b o r of H i g u e r o t e a n d mouth of t h e Rio U n a r e [in coastal 'enezuela n e a r Caracas, to avoid mosqui], t h e w r e t c h e d inhabitants a r e accusmed to . . . pass t h e night b u r i e d in t h e d . . . leaving o u t t h e h e a d only, which y cover with a handkerchief." arences CARWAL, A. 1978. Malaria makes a comeback. NewScient. 77: 274-277. TES, M. 1949. The natural history of mosqui­ toes. Macmillan, New York. OWN, A. W. A., J. HAWORTH, AND A. R.

ZOHAR. 1976. Malaria eradication and control from a global standpoint. I. Med. Entomol. 13: 1-25 PIN, G., AND R. WASSERSTROM. 1981. Agri­

cultural production and malaria resurgence

in central America and India. Nature 293: 181-185. CLEMENTS, A. N. 1963. T h e physiology of

mosquitoes. Pergamon, Oxford. DYAR, H. G. 1928. T h e mosquitoes of the

Americas. Carnegie lnst., Washington, D.C., Publ. 387. FOOTE, R. H., AND D. R. COOK. 1959. Mosqui­

toes of medical importance. U.S. Dept. Agrie. Agrie. Handbk. 152; 1-158. FORATTINI, O. P. 1962, 1965. Entomología médi­ ca. 1. Parte geral, Díptera, Anophelini; 2. Culicini. Culex, Aedes e Psorophora; 3. Culicini: Haemagogus, Mansonia, Culiseta, Sabethini. Toxorhynchitini, Arboviroses, Filariose bancroftiana, Genética. Fac. Hig. Saúde Pub., Univ. Sao Paulo, Sao Paulo. GILLETT, J. D. 1971. Mosquitoes. Weidenfeld and Nicolson, London. GILLIES, M. T. 1980. The role of carbon dioxide in host finding by mosquitoes (Díptera: Culicidae): A review. Bull. Entomol. Res. 70: 525-532. HARBACH, R. F.., AND K. L. KNIGHT.

1980.

Taxonomists' glossary of mosquito anatomy. Plexus, Marlton, N.J. HORSFALL, W. R. 1955. Mosquitoes—their bio­ nomics and relation to disease. Ronald, New York. HOWARD, L. O., H. G. DYAR, AND F. KNAB.

1912-1917. T h e mosquitoes of North and Central America and the West Indies. Carne­ gie lnst., Washington, D.C., Publ. 159. JONES, J. C. 1978. T h e feeding behavior of mosquitoes. Sci. Amer. 238(6): 138-140, 1 4 3 144, 146, 148. KNIGHT, K. L. 1978. Supplement to A catalog of the mosquitoes of the world (Diptera: Culicidae). Entomol. Soc. America, College Park, Md. KNIGHT, K. L., AND A. STONE. 1977. A catalog

of the mosquitoes of the world. 2 ed. Entomol. Soc. America (Thomas Say Found.), College Park, Md., 6: 1-611. LAIRD, M. 1988. T h e natural history of larval mosquito habitats. Academic, New York. MACHADO-ALLISON, C. E. 1980-1982. Ecología de los mosquitos (Culicidae). Acta. Biol. Vene­ z u e l a 10:303-371; 11:51-129, 133-237. MCLINTOCK, J. 1978. Mosquito-virus relation­ ships of American encephalitides. Ann. Rev. Entomol. 23: 17-37. MATTINGLY, P. F. 1969. The biology of mosquitoborne disease. Sci. Biol. Ser. 1: 1 — 184. MATTINGLY, P. F. 1971. Illustrated key to the

genera of mosquitoes. Amer. Entomol. lnst. Contrib. 7(4): 1-84.

MOSQUITOES

377

SEIFERT, R. P. 1980. Mosquito fauna of Heliconia áurea. J. Anim. Ecol. 49: 687-697. SERVICE, M. W. 1976. Mosquito ecology: Field sampling methods. Applied Science, London. VON HUMBOLDT, A . AND A . BONPLAND.

1852

[1814—1825]. Personal narrative of travels to the equinoctial regions of America, during the years 1799-1804. 3 vols. Henry Bohn, London. Translated by Thomasina Ross. WARD, R. A. 1977. Culicidae. In S. H. Hurlbert, ed., Biota acuática de sudamérica austral. San Diego State Univ., San Diego. Pp. 268-274. WARD, R. A. 1981. Culicidae. In S. H. Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 245-256. WARD, R. A. 1982. Culicidae. In S. H. Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 417-429. WOLDA, H., AND P. GALINDO. 1981.

Population

fluctuations of mosquitoes in the non-sea­ sonal tropics. Ecol. Entomol. 6: 99-106.

Malaria Mosquitoes Culicidae, A n o p h e l i n a e , Anopheles. Adults in this g e n u s a r e characterized by an evenly r o u n d e d , r a t h e r t h a n trilobed, scutellum a n d the usual absence of scales o n the a b d o m e n . Biting specimens can be identified by their " h e a d - s t a n d i n g " pos­ t u r e ; the h e a d is r o t a t e d forward o n the t h o r a x m o r e t h a n in o t h e r mosquitoes, forcing t h e m o s q u i t o to elevate the body w h e n w o r k i n g its m o u t h p a r t s into t h e skin. Larvae lack a r e s p i r a t o r y t u b e a n d exhibit flat, p a l m a t e setae o n the d o r s u m of most a b d o m i n a l s e g m e n t s . With n o t c h e d , tho­ racic lobes, their spiracular flaps, a n d these p a l m a t e setae, they a n c h o r themselves to the u n d e r s i d e of t h e surface film to feed a n d r e s p i r e (fig. 11.3c). (Four species in the related g e n u s Chagasia have a trilobed scutellum a n d larvae with the a n t e r i o r spiracular flap p r o d u c e d into a long, spine­ like process.) T h e p u p a is typical of most mosquitoes (fig. 11.3b). T h e r e are seventy-eight species, r e p r e ­ senting all the world s u b g e n e r a in the

378

FL1F.S AND MIDGES

Neotropical Region except Cellia. Most of the h u m a n malaria vectors are in [k s u b g e n e r a Nysorrhynchus (Faran 1980) and Kerteszia (Zavortink 1973). T h e s e are mainly the following: A. pseudopunctipennis widely in South America; A. bellator, Trinid a d a n d Brazil, a b r o m e l i a d breeder (Downs a n d P i t t e n d r i g h 1946); A. cruzii Brazil, also in b r o m e l i a d s ; A. albimanus Caribbean, Mexico, a n d C e n t r a l Americaa n d A. darlingi (fig. 11.3a), widespread and the best vector (Mendes dos Santos et al 1981). Many o t h e r s act as reservoirs for the malaria parasite (Plasmodium) in the wild. T h r o u g h o u t all of history, malaria has rightfully been c o n s i d e r e d o n e of the worst scourges of m a n k i n d . Extremely debilitat­ ing a n d widespread, it has probably caused m o r e severe suffering t h a n any o t h e r single a r t h r o p o d - b o r n e disease. T h e r e is no proof, however, of its presence in America prior to the Spanish Conquest, a n d it was probably not c o m m o n for fifty years or m o r e after its i n t r o d u c t i o n , possibly first via o n e of Columbus's own crews. It surely c a m e with m a n y s u b s e q u e n t trans-Atlantic expeditions. To p l u n d e r the rich West In­ dian cities, Sir Francis D r a k e left Plymouth in 1585 with twenty-nine ships carrying 1,500 s e a m e n a n d 800 soldiers. H e picked u p malaria in the C a p e Verde Islands and t r a n s p o r t e d it to the C a r i b b e a n and Central America w h e r e his m e n became so afflicted a n d so m a n y h u n d r e d s d i e d that he was forced to a b a n d o n the mission (Keevil 1957: 9 2 - 9 4 ) . Fortunately, the primary Af­ rican vector species, Anopheles gambiae, accidently i n t r o d u c e d to the n o r t h e a s t coast of Brazil a b o u t 1930, was contained and even­ tually eradicated (Soper a n d Wilson 1943).

References DOWNS, W. G.,

AND C. S. PITTENDRIGH.

1946.

Bromel iad malaria in Trinidad, British West Indies. Amer. J. Trop. Med. Hyg. 26: 46-66. FARAN, M. E. 1980. Mosquito studies (Diptera, Culicidae). XXXIV. A revision of the Albi­ manus Section of the subgenus Ny.ssorhyiuhus

y/_ E. KERR 1981. Biología de anofelinos amazónicos. 1. Ciclo biológico, postura e estadios larvais de Anopheles darlingi. Root 1926 (Diptera: Culicidae) da Rodovia Manaus-Boa Vista. Acta Amazónica 11: 789-797. SoPER. F. L., AND D. B. WILSON. 1943. Anopheles gambiae in Brazil, 1930 to 1940. Rockefeller Foundation, New York. ZAVORTINK, T. J. 1973. Mosquito studies (Dip­ tera, Culicidae). XXIX. A review of the subge­ nus Kerteszia of Anopheles. Amer. Entomol. Inst. Contrib. 9(3): 1-54.

t r u n k s or vegetation n e a r the larval b r e e d ­ ing sites. Giant mosquitoes are used to detect d e n g u e in h u m a n s suspected of having the disease (xenodiagnosis). S e r u m from a pa­ tient is injected into the t h o r a x of the a d u l t or head of the male a n d allowed to incu­ bate for a prescribed n u m b e r of days. Fluorescence of brain tissue u n d e r ultravio­ let light is positive for the disease (Ramalingam pers. c o m m . ) . T h e r e a r e sixteen regional species, all belonging to the s u b g e n u s Lynchiella. T h e most c o m m o n a n d w i d e s p r e a d are T. haemorrhoidalis a n d T. theobaldi.

Giant Mosquitoes

References

0f

Anopheles. Amer. Entomol. Inst. Contrib. 15(7): 1-215. KEEVIL. J- I- 1957. Medicine and the navy 1200-1900. Vol. 1. E. and S. Livingstone, Edinburgh. MENDES DOS SANTOS, J. M., E. P. B. CONTEL, AND

Culicidae, T o x o r h y n c h i t i n a e , Toxorhynchites. E l e p h a n t mosquitoes. These are very large, n o n b i t i n g mosqui­ toes (fig. 11.3d) seen only in d e e p forest (Steffan a n d E v e n h u i s 1981). T h e p r o b o s ­ cis of the adults of b o t h sexes is long a n d ventrally r e c u r v e d , c o n t r a r y to the straight or upward b e n d of o t h e r mosquitoes, a n d used to s i p h o n n e c t a r a n d plant e x u d a t e s , instead of blood. T h e body is usually d e e p metallic blue o r violaceous, a n d males of­ ten display b r i g h t r e d scale patches at the tip of the a b d o m e n . T h e larvae are also large a n d prey o n active aquatic invertebrates, usually o t h e r mosquito larvae, which they search o u t a n d catch with t h e i r p r e h e n s i l e , flat-bladed mouthparts. Because of this habit, they are considered potential biological control ' agents against h a r m f u l mosquitoes (Stef. Ian 1975). However, their aggressiveness and p e n c h a n t for cannibalism diminishes their effective application in this area. T h e i m m a t u r e s d e v e l o p in n a t u r a l con­ tainers, such as cut b a m b o o , b r o m e l i a d s , *nd tree holes. T h e y also d e v e l o p in artificial c o n t a i n e r s d i s c a r d e d in or n e a r forested areas, most commonly, metal ®ns, barrels, e a r t h e n pots, a n d old a u t o ­ mobile tires. A d u l t s generally rest on tree

STEFFAN, W. A. 1975. Systematics and biological control potential of Toxorhynchites (Diptera: Culicidae). Mosq. Syst. 7: 59-67. STEFFAN, W.

A.,

AND N.

L.

EVENHUIS.

1981.

Biology of Toxorhynchites. Ann. Rev. Entomol. 26: 159-181.

Aedes Mosquitoes Culicidae, Culicinae, Aedini, Aedes. A d u l t Aedes mosquitoes are usually a d o r n e d with brilliant white, silvery, or g o l d e n scale patches on the t h o r a x . Details identifying t h e m are a lack of setae in the spiracular area a n d the presence of bristles b e h i n d the spiracle a n d o n the a n t e r i o r sides of the t h o r a x . T h e larvae have a n o r m a l s i p h o n with lateral teeth basally, a n anal s e g m e n t with a n incomplete sclerotization, a n d a single pair of tufted setae placed n e a r t h e middle. T h i s is a large a n d diversified g e n u s in the Neotropics, with a b o u t 120 species. Several s u b g e n e r a are r e p r e s e n t e d in Latin America, some found n o w h e r e else (Arnell 1976, Berlin 1969, Schick 1969, Zavortink 1972). A very w i d e s p r e a d , often t r o u b l e s o m e , species in this g e n u s is the salt m a r s h mos­ quito {Aedes taeniorhynchus), found a l o n g both of the Central a n d South A m e r i c a n coasts a n d the inland saline areas as well as

MOSQUITOES

379

in t h e Antilles a n d G a l á p a g o s Islands. Adults may e m e r g e in e n o r m o u s n u m b e r s to m a k e territory b o r d e r i n g salt m a r s h e s a n d swamps u n i n h a b i t a b l e .

References ARNELL, J. H. 1976. Mosquito studies (Díptera, Culicidae). XXXIII. A revision of the Scapularis Group of Aedes (Ochlerotatus). Amer. Entomol. Inst. Contrib. 13(3): 12-144. BERLIN, D. G. W. 1969. Mosquito studies (Díp­ tera, Culicidae). XII. A revision of the Neo­ tropical subgenus Howardina of Aedes. Amer. Entomol. Inst. Contrib. 4(2): 1-190. SCHICK, R. X. 1969. Mosquito studies (Diptera, Culicidae). XX. The Terrens Group of Aedes (Finlaya). Amer. Entomol. Inst. Contrib. 5(3): 1-158. ZAVORTINK, T. J. 1972. Mosquito studies (Dip­ tera, Culicidae). XXVIII. The New World species formerly placed in Aedes (Finlaya). Amer. Entomol. Inst. Contr. 8(3): 1-206.

Yellow Fever Mosquito Culicidae, Culicinae, A e d i n i , Aedes aegypli. Portuguese: P e r n i l o n g o rajado (Brazil). Tupi-Guaraní: C a r a p a n á p i n i m a (Brazil). Aedes aegypti ( C h r i s t o p h e r s 1960) is the only m e m b e r of the medically i m p o r t a n t , Old World m o s q u i t o s u b g e n u s Stegomyia in Latin America; it is a d v e n t i t i o u s in u r b a n situations over m u c h of the region a n d is easily r e c o g n i z e d by t h e u n i q u e , white, lyre-shaped m a r k i n g o n the back of a n otherwise black t h o r a x (fig. 11.3e). T h e larva is typical of most Aedes (fig. 11.3f). T h e species has long b e e n a b h o r e d as the most efficient vector of yellow fever a n d as an u r b a n vector species most difficult to eradicate (Groot 1980). Many times, o u t b r e a k s of this disease (known to t h e Spanish as "black vomit" a n d in English t r a d i t i o n as "yellow jack") may have c h a n g e d the c o u r s e of h u m a n lives a n d history. W h e n yellow fever b r o k e out a m o n g his t r o o p s , Sir G e o r g e Clifford, T h i r d Earl of C u m b e r l a n d , failed in his siege of San J u a n in 1598, a n d the colony r e m a i n e d u n d e r Spanish r u l e until the A m e r i c a n conquest. A similar fate befell

380

FLIES AND MIDGES

Napoleon's C a p t a i n - G e n e r a l LeClerc j n his a t t e m p t to take Haiti for France ¡ n 1802, a n d m u c h of America's future was probably t u r n e d from French domination ( C l o u d s l e y - T h o m p s o n 1976: 171). In 1741 A d m i r a l Vernon sailed from Britain with 27,000 m e n , with t h e intention of conquer­ ing Mexico a n d Peru. But h e lost 20,000 of his m e n to yellow fever, a n d his mission was a b o r t e d . A great stride in p r o v i n g the role of insects as h u m a n disease vectors a n d estab­ lishing the field of medical entomology resulted from the classical e x p e r i m e n t s of Walter Reed with soldier volunteers in H a v a n a following the Spanish-American w a r (Reed et al. 1900, 1901). T h e work confirmed the suspicions a n n o u n c e d al­ most twenty years earlier by the Cuban physician, Carlos Finlay (Sosa 1989).

References CHRISTOPHERS, S. R. 1960. Aedes aegypti (L.), the yellow fever mosquito: Its life history, bionom­ ics, and structure. Cambridge Univ. Press, Cambridge. CLOUDSLEY-THOMPSON, J. L. 1976. Insects and

history. St. Martin's, New York. GROOT, H. 1980. The reinvasion of Colombia by Aedes aegypti: Aspects to remember. Amer. J. Trop. Med. Hyg. 29: 330-338. REED, W., J. CARROL, AND A. AGRAMONTE.

190E

The etiology of yellow fever: An additional note. J. Amer. Med. Assoc. 36: 431-440. REED, W., J. CARROL, A. AGRAMONTE, ANDJ. W.

LAZEAR. 1900. The etiology of yellow fever: A preliminary note. Philadelphia Med. J. 6: 790-796. SOSA, JR., O. 1989. Carlos J. Finlay and yellow fever: A discovery. Entomol. Soc. Amer. Bull. 35(2): 23-25.

Blue Devils Culicidae, Culicinae, Aedini, Haemagogus. T h e d o r s u m of the t h o r a x of these small to medium-sized (BL 7 - 1 0 m m ) forest mos­ quitoes is completely covered with broad, flat, reflecting scales, giving t h e m a deep, steely blue or metallic greenish color (fig. 11.4a). T h e a n t e r i o r p r o n o t a l lobes are

«Mire 11.4 MOSQUITOES (CULICIDAE). (a) Blue devil (Haemagogus sp.). (b) Southern house mosquito (Culex quinquefasciatus). (c) Crabhole mosquito (Deinocerites cancer), (d) Sabethine mos­ quito (Sabethes sp.).

unusually large. T h e y a r e particularly per­ sistent biters a n d often noticed because of their d i u r n a l habits, r e s p l e n d e n t colors, and high-pitched w h i n e . Several species are i m p o r t a n t in arbovirus transmission and probably a r e responsible for maintain­ ing j u n g l e yellow fever in forest m a m m a l populations. Structurally, the larvae a r e like those of Aedes. T h e y often d e v e l o p in containers, bamboo s t u m p s , artificial vessels, a n d tree holes. This g e n u s of twenty-seven species is exclusive to t h e New World tropics a n d subtropics (Arnell 1973).

Reference ARNELL, J. H. 1973. Mosquito studies (Diptera, Culicidae). XXXII. A revision of the genus Haemagogus. Amer. Entomol. Inst. Contrib. 10(2): 1-174.

Gallinippers Culicidae, Culicinae, A e d i n i , Psorophora. Gallinippers a r e very large, fiercely biting mosquitoes in t h e g e n u s Psorophora, which contains over forty Neotropical species. Adults resemble Aedes b u t h a v e spiracular : bristles. T h e larvae, most of which d e v e l o p in ground pools, also r e s e m b l e those of Aedes but have a completely r i n g e d sclerotization a r o u n d the anal s e g m e n t , which is ¿.pierced along the m i d v e n t r a l line by tufted bristles. T h e y a r e poorly studied in Latin America but familiar because of their size.

Culex Mosquitoes Culicidae, Culicinae, Culicini, Culex. T h i s is by far the largest g e n u s of mosqui­ toes in the New World tropics. It contains over 300 species, m o r e t h a n half of these in one, characteristically Neotropical subge­ nus, Melanoconion. Adults a r e mostly d r a b gray with s u b d u e d thoracic coloration (ex­ cept the s u b g e n u s , which has m a n y o r n a ­ m e n t e d species); the spiracular a r e a is b a r e , a n d postspiracular bristles a r e ab­ sent; the tips of the tarsi possess pulvillar p a d s (lacking in o t h e r mosquitoes except Deinocerites). T h e larvae h a v e a n o r m a l b r e a t h i n g t u b e with lateral teeth at the base a n d several hair tufts along the u n d e r s i d e . Species in the g e n u s display varied h a b ­ its. Larval b r e e d i n g sites include all types, a n d adults of m a n y species a r e a n t h r o p o philic. T h e ubiquitous s o u t h e r n h o u s e mos­ quito (Culex quinquefasciatus; fig. 11.4b) is a p r i m e nuisance a n d is involved in the transmission of n u m e r o u s h u m a n a n d ani­ mal p a t h o g e n s . T h i s is a domestic species whose larvae d e v e l o p in all sorts of foul water in containers. Favored u r b a n b r e e d ­ i n g g r o u n d s a r e rain barrels, cisterns, flooded latrines, polluted p o n d s , r o a d ditches, gutters, a n d the like.

References BRAM, R. A. 1967. Classification of Culex subge­ nus Culex in the New World (Diptera: Culi­ cidae). U.S. Nati. Mus. Proc. 120: 1-120.

MOSQUITOES

381

FOOTE, R. H. 1952. The larval morphology and chaetotaxy of the Culex subgenus Melanoconion (Díptera, Culicidae). Entomol. Soc. Amer. Ann. 45: 445-472. FOOTE, R. H. 1954. The larvae and pupae of the mosquitoes belonging to the Culex subgenera Melanoconion and Mochlostyrax. U.S. Dept. Agrie. Tech. Bull. 1091: 1-126. SIRIVANAKARN, S. 1982. A review of the systematics and a proposed scheme of internal classification of the New World subgenus Melanoconion of Culex (Díptera, Culicidae). Mosq. Syst. 14: 265-333.

Galindomyia with a single species, G. leei from Pacific coastal Colombia, is structur­ ally similar to Deinocerites in the adult stage, but its larva a n d b r e e d i n g habits are unknown.

References ADAMES, A. J. 1971. Mosquito studies (Dipt e r a Culicidae). XXIV. A revision of the crabhole mosquitoes of the genus Deinocerites. Amer Entomol. Inst. Contrib. 7(2): 1-154. MCIVER, W., AND R. SIEMICKI. 1976. Fine struc­

Crabhole Mosquitoes Culicinae, Culicini, Deinocerites. T h i s is a n o t h e r exclusively Neotropical m o s q u i t o g r o u p , with e i g h t e e n species ( A d a m e s 1971). T h e m o n o c o l o r o u s graybrown adults possess t h e g e n e r a l charac­ teristics of Culex but h a v e sparsely h a i r e d a n t e n n a e , these o r g a n s usually m u c h l o n g e r t h a n t h e proboscis in both sexes, a n d the males often h a v e e n l a r g e d fore tarsal claws. T h e y use these claws a n d unusually long a n t e n n a e in a highly pecu­ liar m a t i n g strategy. T h e y r e m a i n in the flooded crab b u r r o w , skating on the water, while h o l d i n g the tips of the a n t e n n a e close to the surface (Mclver a n d Siemicki 1976). O n d e t e c t i n g a m a t u r e female p u p a fixed to the surface film by its b r e a t h i n g t r u m p e t s , they h o o k it with an e n l a r g e d fore tarsal claw (fig. 11.4c) a n d r e m a i n with it until t h e female e m e r g e s . C o p u l a t i o n immediately follows or even accompanies e m e r g e n c e , the m a l e some­ times g r a s p i n g the female's genitalia even before she has escaped h e r p u p a l skin (Provost a n d H a e g e r 1967). T h e larvae also r e s e m b l e those of Culex b u t possess a pair of c o n s p i c u o u s lateral p o u c h e s on the h e a d a n d m u c h r e d u c e d gills on the anal segment. B r e e d i n g a n d m u c h of a d u l t life is confined to the b u r r o w s of land crabs in the g e n e r a Cardisoma, Ucides, a n d Sesarma. T h e genus's distribution is entirely coastal.

382

FLIES AND MIDGES

ture of the antenna! tip of the crabhole mosquito, Deinocerites cancer Theobald (Dintera: Culicidae). Int. J. Ins. Morph. Embryol 5:319-334. PROVOST,

M.

W.,

AND F.

S.

HAEGER.

1957

Mating and pupal attendance in Deinocerites cancer and comparisons with Opifex fuscus (Diptera: Culicidae). Entomol. Soc. Amer. Ann. 60: 565-574.

Water Weed Mosquitoes Culicidae, Culicinae, Mansonini, Mansonia a n d Coquillettidia. T h e s e closely related g e n e r a (previously c o m b i n e d as Mansonia) have immatures that live a m o n g d e n s e beds of floating aquatic plants, c o m m o n l y water lettuce (Pistia stratiotes) a n d water hyacinth (Eichornia). Both the larvae a n d p u p a e fix them­ selves to the stems of these plants and extract oxygen from air vessels therein, the larvae by m e a n s of a u n i q u e , attenuated, h o o k e d siphonal t u b e with apical sawlike teeth a n d the p u p a e with scythelike respira­ tory h o r n s . T h e adults are easily distin­ guished from o t h e r medium-sized culicines by the very b r o a d , asymmetrical scales on the u p p e r wing surface. A m o n g the twentyfive Neotropical species, several a r e vectors of viral infections a n d filarial w o r m s (Ronderos a n d B a c h m a n n , 1963).

Reference RONDEROS, R. A., AND A. O. BACHMANN.

Sabethine Mosquitoes

Other Mosquitoes

Culicinae, Sabethini, Limatus, Sabethes, Wyeomyia, Phoniomyia, Tnchoprosopon, and relatives.

Culicinae, Culisetini, Culiseta: O r t h o p o d o m y i i n i , Orthopodomyia; Uranotaeniini, Uranotaenia.

These wholly Neotropical g e n e r a , contain­ ing together j u s t u n d e r 200 species, form a varied assemblage (Lane a n d C e r q u e i r a 1942, Zavortink 1979). All display a tuft of small hairs o n t h e p o s t n o t u m ( r o u n d e d area b e n e a t h t h e scutellum) a n d a trilobate scutellum; postspiracular bristles are ab­ sent. Many species are brilliantly metallic or iridescent, a n d o n e or m o r e tarsi often have conspicuous p a d d l e - s h a p e d areas of scales in the g e n u s Sabethes (fig. 11.4d). Most use various plant c o n t a i n e r s as larval and p u p a l habitats. Trichoprosopon digitatum commonly b r e e d s in the fetid water that accumulates in r o t t i n g cacao p o d s a n d is a major pest for plantation workers (Zavor­ tink et al. 1983); females have b e e n ob­ served b r o o d i n g t h e i r e g g rafts in t h e i r microhabitat (Lounibos a n d Machado-Allison 1983). T h e largest g e n u s is Wyeomyia with over 100 species, mostly distinguish­ able only by their highly c o m p l e x male genitalia. Highly specialized forest mosqui­ toes, they choose water-filled b a m b o o internodes, b r o m e l i a d leaf axils, inflores­ cences of Heliconia a n d Calathea, a n d palm spaths as b r e e d i n g sites.

Culiseta particeps is t h e only o n e truly Neo­ tropical m e m b e r of its g e n u s . It occurs at h i g h e r elevations in cold m o u n t a i n p o n d s . Orthopodomyia is an assemblage of seven regional species, all utilizing b a m b o o int e r n o d e s , bromeliad leaf axils, a n d tree holes for d e v e l o p m e n t . A b o u t thirty Urano­ taenia occur in the New World tropics; larvae are usually f o u n d in weedy g r o u n d pools (Zavortink 1968).

References LANE, J., AND N. L. CERQUEIRA. 1942.

LOUNIBOS, L. P., AND C. E. MACHADO-AI.LISON.

1983. Oviposition and egg brooding by the mosquito Trichoprosopon digitatum in cacao husks. Ecol. Entomol. 8: 475-478. ZAVORTINK, T. J. 1979. Mosquito studies (Dip­ tera, Culicidae). XXXV. T h e new sabethine genusjohnbelkinia and a preliminary reclassification of the composite genus Trichoprosopon. Amer. Entomol. Inst. Contrib. 17(1): 1-61. ZAVORTINK, T. J.,

1963.

Mansoniini Neotropicales. I (Diptera-Culicidae). Soc. Entomol. Argentina Rev. 26: 57-65.

Os sabe-

tínos da América (Diptera, Culicidae). Mus. Zool., Univ. Sao Paulo, Arq. Zool. 3: 473-849.

D.

R.

ROBERTS, AND A.

L.

HOCH. 1983. Trichoprosopon digitatum morphol­ ogy, biology and potential medical impor­ tance. Mosq. Syst. 15: 141-149.

Reference ZAVORTINK, T. J. 1968. Mosquito studies (Dip­ tera, Culicidae). VIII. A prodrome of the genus Orthopodomyia. Amer. Entomol. Inst. Contrib. 3(2): 1-221.

HORSEFLIES T a b a n i d a e . Spanish: T á b a n o s . TupiGuaraní ( a d a p t e d in Portuguese): Mutucas (Brazil). G r e e n h e a d s , m a n g o flies, deerflies, gadflies. T h e vicious bloodletting abilities of these flies are notorious. T h e y feed by l a p p i n g a n d s p o n g i n g u p the blood that oozes from w o u n d s they inflict by their sidewise slashing, bladelike m a n d i b l e s , a process distinctly m o r e painful to t h e host t h a n the delicate h y p o d e r m i c n e e d l i n g of o t h e r bloodsucking Diptera. Blood often flows from w o u n d s m u c h in excess of the a m o u n t ingested by the fly. Most females a r e h e m a t o p h a g o u s , b u t some, as well as all males, subsist only o n n e c t a r a n d p l a n t e x u d a t e s . Primarily m a m m a l s b u t also birds a n d reptiles, including crocodilians ( M e d e m 1981), are hosts. T h e proboscises of the subfamily P a n g o n i i n a e (Fidena, Scione) a r e very long a n d s l e n d e r a n d a r e used to extract nectar from flowers, n e v e r to take blood. S o m e species a n d strains

HORSEFLIES

383

probably can r e p r o d u c e w i t h o u t t h e fe­ males h a v i n g to feed o n blood (auto­ genous), like certain mosquitoes (Charlwood a n d Rafael 1980). Aside from t h e h e m a t o p h a g o u s habit of most of its m e m b e r s , this large family (approximately 1,000 species in t h e N e o tropics; Fairchild 1969) is recognized by their stout, n o n s p i n y bodies, large eyes ( m e e t i n g o n t o p of t h e h e a d in the males; narrowly s e p a r a t e d in t h e females), which often display iridescent r a i n b o w colors, large basal wing lobes, a n d a noticeable d i v e r g e n t fork in t h e wing vein (4th a n d 5th b r a n c h e s of t h e r a d i u s vein) that encloses the wing tip. T h e wings also a r e c o m m o n l y b a n d e d o r splotched with b r o w n p i g m e n t . T h e i r total body size r a n g e s from small to large (BL 5 - 2 5 m m ) . Females lay e g g masses on objects lo­ cated above t h e larval m e d i u m , usually s t a n d i n g water. T h i s explains the a b u n ­ d a n c e of t h e adults n e a r aquatic habitats. Yet they a r e s t r o n g fliers a n d may be f o u n d long distances from t h e s w a m p s , m a r s h e s , a n d weedy p o n d s that a r e their usual b r e e d i n g sites. A few a r e even terrestrial, while o t h e r s d e v e l o p in b r o m e l i a d tanks. T h e larvae (fig. 11.5c) a r e elongate, black or b r o w n i s h to g r e e n , often with a t r a n s l u c e n t cuticle that h a s linear p i g m e n t m a r k i n g s . T h e y have a well-developed head and a r e markedly segmented. T h e s e g m e n t s bear circlets of s h o r t lobes anteri­ orly, a n d t h e t e r m i n a l s e g m e n t has a short

telescoping respiratory tube. Most lay a m o n g dead leaves or o t h e r d e c o m p o s i n g plant m a t t e r on t h e bottom w h e r e they prey on o t h e r aquatic insects a n d invertebrates; a few a r e p h y t o p h a g o u s (Goodwin a n d Mur­ d o c h 1974). Because they a r e especially b o t h e r s o m e to h u m a n s a n d often colorfully marked m a n y species have attracted attention and are k n o w n by local n a m e s . Such are the widespread cabo v e r d e o r mosca congo (Lepiselaga crassipes; fig. 11.5a) (Fairchild 1940) in A m a z o n i a a n d C e n t r a l America a n d t h e colihuacho (Scaptia [ = Onca] lata) in Chile (Berg 1881). Both a r e very bother­ some biters. T h e f o r m e r is an insistent pest that is often very n u m e r o u s by rivers, c o m i n g a b o a r d boats to bite a r o u n d the ankles a n d legs. It is small (BL 8 - 9 mm) and all black except w h e r e t h e pigment abruptly t e r m i n a t e s beyond t h e middle of the wing, leaving its apical third transpar­ ent. Females favor the legs a n d feet of its hosts when bloodletting. D e v e l o p m e n t oc­ curs in m a t t e d floating vegetation, espe­ cially water lettuce (Pistia) in slow or stand­ ing water (Fairchild 1940). T h e second species is large (BL to 2 cm) a n d black with tufts of crimson hairs on the t h o r a x and a b d o m e n a n d can attack insidiously and in n u m b e r s by its river habitat (orig. obs.). These a n d most o t h e r tabanids are diur­ nal, a l t h o u g h s o m e a r e c r e p u s c u l a r o r even n o c t u r n a l in t h e tropics, such as the pale g r e e n Chlorolabanus. Major g e n e r a contain-

in2 biting f o r m s a r e Chrysops, Tabanus (fig. j 1.5b), a n d Dichaelacera. T h e significance of tabanids as disease vectors has h a r d l y b e e n studied in Latin America. C o n s i d e r i n g their considerable potential as such elsewhere (Krinsky 1976), they d e s e r v e a t t e n t i o n in this respect. T h e l i t e r a t u r e o n Neotropical tabanids is reviewed by C o s c a r o n (1977) a n d Fairchild (1981, 1982).

References BERG, F. W. K. 1881. Observaciones acerca de la Osea lata (Guér.) Lynch. Soc. Cient. Argentina An. (Bol.). Pp. 9 - 1 0 . CHARLWOOD, J. D., AND J. A. RAFAEL.

1980.

Autogeny in the River Negro Horse Ely, Lepiselaga crassipes, and an undescribed spe­ cies of Slenotabamis (Diptera: Tabanidae) from Amazonas, Brazil. J. Med. Entomol. 17: 519-521. COSCARON, S. 1977. Tabanidae. hi S. H. Hurlbert, ed., Biota acuática de sudamérica aus­ tral. San Diego State Univ., San Diego. Pp. 297-304. FAIRCHILD, G. B. 1940. A note on the early stages of Lepiselaga crassipes Fab. (Dipt., Taba­ nidae). Psyche 47: 8-13. FAIRCHILD, G. B. 1969. Notes on Neotropical Tabanidae. XII. Classification and distribu­ tion, with keys to genera and subgenera. Arq. Zool. (Sao Paulo) 17: 199-255. FAIRCHILD, G. B. 1981. Tabanidae. In S. H.

Hurlbert, G. Rodriguez, and N. Dias dos Santos, eds., Aquatic biota of tropical South America. Pt. 1. Arthropoda. San Diego State Univ., San Diego. Pp. 290-301. FAIRCHILD, G. B. 1982. Tabanidae. In S. H.

Hurlbert and A. Villalobos Figueroa, eds., Aquatic biota of Mexico, Central America and the West Indies. San Diego State Univ., San Diego. Pp. 452-460. GOODWIN, J. T , AND W. P. MURDOCH. 1974. A

Figure 11.5 FLIES, (a) Mosca Congo (Lepiselaga crassipes, Tabanidae). (b) Horse fly {Tabanus dorsiger, Tabanidae). (c) Horse 1ly, larva, (d) Giant mydas 1ly (Mydas sp., Mydidae). (e) Timber lly {Pantophthalmus sp., Pantophthalmidae). (f) Timber fly, larva.

384

FLIES AND MIDGES

study of some immature Neotropical Tabani­ dae (Diptera). Entomol. Soc. Amer. Ann. 67: 85-133. KRINSKY, W. L. 1976. Animal disease agents transmitted by horse flies and deer flies (Diptera: Tabanidae). J. Med. Entomol. 13: 225-275. MEDEM, F. 1981. Horse flies (Diptera: Tabani­ dae) as ectoparasites on caimans (Crocodylia: Alligatoridae) in eastern Colombia. Cespedesia 10: 123-191.

GIANT MYDAS FLIES Mydidae, Mydas. T h e giant mydas flies a r e very large d i p terans (BL 35—45 m m ) . Most species in t h e g e n u s a r e all black, save for t h e tips of its a n t e n n a e a n d its wings, which a r e b r i g h t o r a n g e - r e d ; o t h e r s have black wings as well (fig. 11.5d). Several a r e Batesian mimics of various large species of t a r a n t u l a hawk wasps (Pepsis, Pompilidae). T h e g e n u s ranges from n o r t h e r n Mexico to s o u t h e r n Brazil. Adults feed on flower nectar, a n d t h e larvae probably a r e p r e d a t o r s of o t h e r soildwelling insect larvae. Larvae of some Brazilian Mydas a r e k n o w n to live in leaf cutter a n t (Atta) nests, w h e r e they a r e p r e d a c e o u s on certain larval m y r m e c o philous Scarabaeidae (Zikán 1944).

Reference ZIKÁN, J. E 1944. Novas observacoes sobre a biología de Mydas (Dipt.) e sua relacáo com as formigueiros de saúva. Bol. Minis. Agrie. Rio de Janeiro 33: 43—55. [Not seen.]

TIMBER FLIES P a n t o p h t h a l m i d a e . Portuguese: (Brazil).

Moscardos

T i m b e r flies ( C a r r e r a a n d d ' A n d r e t t a 1957) a r e t h e real goliaths of t h e Diptera. A d u l t s reach body lengths of 40 to 50 millimeters a n d wingspans of 85 to 95 millimeters. T h e y generally r e s e m b l e horseflies but have small, n o n b i t i n g m o u t h p a r t s a n d m u c h r e d u c e d calypters. O t h e r family char­ acteristics a r e a flattened, squarish a b d o ­ m e n (female with a telescoping ovipositor) a n d n o o r very small tibial s p u r s . T h e r e a r e also five closed cells in the wing, as o p p o s e d to four in the horseflies. T h e wing m e m ­ b r a n e s of all species a r e p i g m e n t e d with large i r r e g u l a r blotches. Adults (fig. 11.5e) a r e usually seen rest­ ing on logs o r occasionally flying n e a r tree t r u n k s in forest habitats. They oviposit on

TIMBER FLIES

385

the crevices in t h e b a r k of trees o r fallen trees. After they hatch, t h e larvae b u r r o w into t h e s o u n d wood b e n e a t h t h e b a r k a n d s p e n d a p p r o x i m a t e l y two years w o r k i n g their way t h r o u g h t h e wood, f o r m i n g ex­ tensive t u b u l a r galleries. Living or d e a d h a r d w o o d trees of m a n y species m a y be attacked. It is u n c e r t a i n w h e t h e r or not t h e larvae feed o n t h e wood itself or o n fer­ m e n t e d sap that floods their galleries. T h e b u r r o w i n g is forceful a n d creates a r a s p i n g or g r a t i n g s o u n d that m a y be audible some m e t e r s away (orig. obs.). T h e larvae (fig. 11.5f) ( T h o r p e 1930) are cylindrical with a heavily sclerotized head a n d powerful m a n d i b l e s . T h e last s e g m e n t of t h e a b d o m e n is obliquely t r u n ­ cate with two series of s t r o n g spines, form­ ing a p u s h plate used t o eject t h e large quantities of sawdust p r o d u c e d by their b o r i n g activities. Ventrally, just a n t e r i o r to this s e g m e n t , is a u n i q u e o r g a n (Fiebrig's body), c o m p o s e d of m a n y a p p r e s s e d , fin­ gerlike processes; it a p p a r e n t l y functions as a gill that allows t h e larvae to r e s p i r e in their liquid microhabitat. T h e mobile p u p a e a r e strongly sclero­ tized a n d have a nearly spherical h e a d s t r u c t u r e b e a r i n g two powerful, r i d g e d , g r i n d i n g plates. T h e cylindrical body is abruptly t r u n c a t e posteriorly like t h e lar­ vae. T h e s e s t r u c t u r e s p e r m i t t h e p u p a e to plug t h e surface o p e n i n g of their galleries w h e n n e a r t h e log's e x t e r i o r j u s t p r i o r to adult e m e r g e n c e . T h i s is a small family with only two g e n e r a , Opetiops, with o n e species, a n d Pantophthalmus with n i n e t e e n . It is e n d e m i c to the New World tropics (Val 1976). P a n t o p h thalmids a r e most c o m m o n in t h e Amazo­ nian a n d C e n t r a l A m e r i c a n rain forests.

References CARRERA, M., AND M. A. V. D'ANDRETTA.

1957.

Sobre a familia Pantophthalmidae. Arq. Zool. (Sao Paulo) 10: 253-330. THORPE,

W. H.

1930. Observations on

the

structure, biology and systematic position of Pantophthalmus tabaninus Thunb. (Diptera:

386

FLIES AND MIDGES

Pantophthalmidae). Royal Entomol. Soc. Lon don Trans. 82: 5-22. VAL, F. C. 1976. Systematics and evolution of the Pantophthalmidae (Diptera, Brachycera) Arq. Zool. (Sao Paulo) 27: 51-164.

FLOWER FLIES Syrphidae. O v e r 1,600 species c o m p r i s e this g r o u p of c o m m o n , conspicuous, generally mediumsized (BL 4—15 m m ) flies with hovering flight a n d large eyes. Many a r e bright yellow or o r a n g e a n d b a n d e d , as in the g e n u s Metasyrphus, while o t h e r s a r e dull gray to black o r iridescent g r e e n o r blue, these p a t t e r n s a n d colors of m a n y giving t h e m a resemblance to stinging bees and wasps. T h e i r habit of visiting flowers makes t h e m beneficial as pollinators. T h e larvae of m a n y species a r e also valuable as carnivores o n pests such as a p h i d s a n d mealybugs (Goncalves a n d Goncalves 1976). A wide­ s p r e a d species is Metasyrphus americanus, which has a yellow-spotted a b d o m e n (fig. 11.6a). Syrphids a r e not well studied in Latin America. A n i m p o r t a n t t a x o n o m i c work is by T h o m p s o n (1972).

References GONCALVES, C. R., AND A. J. L. GONCALVES.

1976. Observances sobre moscas da familia Syrphidae predadoras de Homópteros. Soc. Entomol. Brasil. An. 5: 3-10. THOMPSON, F. C. 1972. A contribution to a

generic revision of the Neotropical Milesinae (Diptera: Syrphidae). Arq. Zool. (Sao Paulo) 23: 73-215.

Drone Fly Syrphidae, Milesiinae, Eristalini, Eristalis tenax. Spanish: Mosca abeja. T h i s flower fly (fig. 11.6b) is so similar in coloration, size ( B L 1 0 - 1 5 m m ) , a n d behav­ ior to t h e h o n e y b e e that it is avoided by h u m a n s a n d animals. It is also slightly pubescent, like t h e bee, a n d has a dark,

Figure 11-6 FLIES, (a) Flower fly (Metasyrphus americanus, Syrphidae). (b) Drone fly (Eristalis tenax, Syrphidae). (c) Green flower fly (Ornidia obesa, Syrphidae). (d) Drone fly, larva, (e) Wasp fly (Hermetia illuscens, Stratiomyidae), larva, (f) Wasp fly, adult. dull o r a n g e a n d black-banded a b d o m e n . Although variable (Heal 1979), t h e basal band is usually i n c o m p l e t e dorsally, b r o k e n by a m e d i a n d a r k bar, simulating t h e bee's narrow waist. F r e q u e n t i n g t h e same flow­ ers, d r o n e flies a n d honeybees have also evolved similar visual sensitivities (Bishop and C h u n g 1972). T h e larva ("rat-tailed maggot") (fig. 11.6d) possesses a long, telescoping, poste­ rior respiratory tube, consisting of t h r e e segments that can e x t e n d to several times the length of t h e r o b u s t , cylindrical body (BL 2 0 - 2 5 m m ) . T h e r e a r e eight pairs of stumpy false legs t i p p e d with fine spines developed o n t h e ventral side of t h e t r u n k . The larva is exceedingly h a r d y a n d lives in water c o n t a m i n a t e d by sewage, fresh excre­ ment, moist c a r r i o n , a n d o t h e r foul organic matter of liquid consistency. It is sometimes involved in cases of gastrointestinal myiasis. A cosmopolitan species, t h e d r o n e fly is found in most Latin A m e r i c a n countries, particularly in w a r m e r p a r t s , b u t is locally absent in m a n y areas. I n ancient times in Europe, its r e s e m b l a n c e to t h e h o n e y b e e and its habit of b r e e d i n g in decaying car­ casses led to a c u r i o u s p o p u l a r belief that ule bees w e r e s p o n t a n e o u s l y g e n e r a t e d from dead o x e n ("bugonia") (Atkins 1948).

BISHOP, L. G., AND D. W. CHUNG. 1972. Con­

vergence of visual sensory capabilities in a pair of Batesian mimics. J. Ins. Physiol. 18: 1501-1508. HEAL, J. 1979. Colour patterns of Syrphidae. I. Genetic variation in the dronefly Eristalis tenax. Heredity 42: 223-236.

Green Flower Fly Syrphidae, Milesiinae, Volucellini, Ornidia obesa. Portuguese: B e r n e i r a (Brazil). T h i s species (fig. 11.6c) r a n g e s t h r o u g h o u t the moist lowlands of most of t h e N e o t r o p i ­ cal Region w h e r e it is a familiar visitor on w a r m days to o u t h o u s e s , urinals, a n d rot­ ting fruit o r o t h e r foul materials. Its bril­ liant d e e p metallic color immediately iden­ tifies it even as it hovers in t h e air, its wings b l u r r e d by their rapid vibration. It is a b o u t the same size (BL 10 m m ) as similar blowflies a n d orchid bees a n d m a y mimic the latter. F u r t h e r identifying features a r e a d a r k spot midway on t h e leading e d g e of the wing, a m u c h smaller similar spot n e a r the wing tip, a n d a depression in t h e c e n t e r of t h e scutellum. T h e early stages a r e not well studied; t h e larva is said to be c o p r o p h a g o u s (Val 1972). Ornidia contains t h r e e o t h e r beautiful b u t less w i d e s p r e a d ( T h o m p s o n 1991).

equally species

References ATKINS, JR., E. L. 1948. Mimicry between the drone-fly, Eristalis tenex (L.), and the honeyDee, Apis mellifera L.: Its significance in anQent mythology and present day thought. Entomol. Soc. Amer. Ann. 41: 887-892.

References THOMPSON, F. C. 1991. The flower fly genus Ornidia (Diptera; Syrphidae). Proc. Entomol. Soc. Wash. 93: 248-261.

FLOWER FLIES

387

VAL, F. C. 1972. On the biometry and evolution of the genus Omidia (Diptera, Syrphidae). Univ. Sao Paulo Mus. Zool. Pap. Avul. Zool. 26:1-28.

WASP FLY Stratiomyidae, H e r m e t i i n a e , Hermetia illuscens. Spanish: G u a r e r o , b o r r a c h e r o (Central America). T h i s semidomestic species is found in all t e m p e r a t e a n d tropical p o r t i o n s of t h e world, owing to the b r o a d adaptive capaci­ ties of adults a n d t h e t r a n s p o r t a t i o n of larvae in c o n t a m i n a t e d food. Because all o t h e r m e m b e r s of its g e n u s a r e N e o t r o p i ­ cal, it probably o r i g i n a t e d in some w a r m a r e a of the New World. T h e larvae d e v e l o p in a wide variety of organic m e d i a , i n c l u d i n g decaying fruit a n d vegetables, g a r b a g e , compost, d u n g , carrion, a n d soil c o n t a m i n a t e d with these materials (Copello 1926). In C e n t r a l A m e r i ­ can b a n a n a p l a n t a t i o n s , t h e species d a m ­ ages fruit by ovipositing between t h e fin­ gers of y o u n g fruit a n d o n ripe b u n c h e s after they have b e e n picked. A s t r o n g attraction for yellow color by t h e female is t h o u g h t to account at least partly for t h e latter (Stephens 1975). W h e n they occupy t h e same food, wasp fly larvae m a y s u p p r e s s housefly p o p u l a ­ tions t h r o u g h some indirect competitive effect, since the f o r m e r d o n o t feed o n t h e latter ( F u r m a n et al. 1959). Larvae are also c o m m o n l y f o u n d in h o n e y b e e hives w h e r e they feed o n honey, wax, a n d waste materi­ als. T h e r e is o n e r e c o r d from the nest of a stingless bee ( B o r g m e i e r 1930) which may indicate a general habit in the wild for this a n d o t h e r m e m b e r s of t h e g e n u s . Larvae also a r e occasionally t h e cause of h u m a n intestinal myiasis. Identifying characteristics of t h e larval stage (BL 20 m m ) a r e a thick, leathery, d a r k b r o w n i n t e g u m e n t that is set with n u m e r o u s short bristles, a b r o a d , flat­

388

FLIES AND MIDGES

tened, ovate general shape, a n d a slende well-sclerotized head (fig. 11.6e). T h e moderately large (BL 1 5 - 2 0 mm\ adults (fig. 11.6f) m u c h resemble spider wasps in a n a t o m y a n d behavior. T h e y are elongate a n d d a r k bluish-black, with uni formly dusky wings a n d white tarsi (the h i n d tibia is also white); at t h e base of the a b d o m e n are paired t r a n s p a r e n t areas sep­ a r a t e d by a d a r k m e d i a n line, simulating a wasp's waist. T h e male a b d o m e n is bronzy in contrast to black in females. T h e fljej c o m p o r t themselves waspishly, r u n n i n g on the g r o u n d excitedly a n d flicking their wings. T h e y also buzz a n d feign a sting when h a n d l e d (Iide a n d Mileti 1976)

References BORGMEIER, T. 1930. Ueber das Vorkommen der Larven von Hermetia illuscens L. (Dipt. Stratiomyidae) in den Nestern von Meliponiden. Zool. Anz. 90: 225-235. COPELLO, A. 1926. Biología de Hermetia illuscens Latr. Rev. Entorno!. Argentina 1(2): 23-27. [Sic] Author of species is Linneaus. FURMAN, D. P., R. D. YOUNG, AND E. P. CATTS.

1959. Hermetia illuscens (Linnaeus) as a factor in the natural control of Musca domestica Linnaeus. J. Econ. Entomol. 52: 917-921. IIDE, P., AND D. I. C. MILF.TI. 1976. Estudos

morfológicos sobre Hermetia illuscens (Lin­ naeus, 1758) (Diptera, Stratiomyidae). Rev. Brasil. Biol. 36: 923-935. STEPHENS, C. S. 1975. Hermetia illuscens (Dip­ tera: Stratiomyidae) as a banana pest in Panama. Trop. Agrie. 52: 173-178.

POMACE FLIES Drosophilidae, Drosophilia. T h e s e gnatlike (BL 2 - 3 m m ) , yellowish to brownish flies (fig. 11.7a) a r e usually seen a r o u n d decaying vegetation a n d ripe fruit (Shorrocks 1980, Belo a n d d e Oliveira 1976). T h e y a r e best known, however, for the roles a few species have played in the a d v a n c e m e n t of t h e science of heredity. Early in this century, geneticist Thomas H u n t M o r g a n found that o n e species, Drosophilia melanogaster, r e p r o d u c e d witn

naure 11-7 FLIES, (a) Pomace fly {Drosophila melanogaster, Drosophilidae). (b) Pomace fly, larva, (c) Mediterranean fruit fly (Ceratitis capitata, Tephritidae). (d) Eye gnat (Liohippelates pusio, Chloropidae). (e) Stilt-legged fly (Taeniaptera sp., Micropezidae). great rapidity, was easily c u l t u r e d in t h e laboratory, r e q u i r e d b u t twelve days to develop from the e g g to maturity, a n d h a d larval cells c o n t a i n i n g only four, large, wellmarked c h r o m o s o m e s . In fact, t h e insect Has so well a d a p t e d for genetics that some­ one once q u i p p e d that it must have been created by the Almighty solely as an object of heredity r e s e a r c h . Since M o r g a n ' s days, many additional species h a v e b e e n studied in great detail a n d c o n t r i b u t e d m o r e to o u r understanding of evolutionary mecha"!~ms, p o p u l a t i o n dynamics, phylogeny, cylogy, a n d so o n , t h a n a n y o t h e r single 3up of h i g h e r o r g a n i s m s ( A s h b u r n e r J 9 7 6 - I 9 8 1 , R u b i n 1988, T h r o c k m o r t o n 975). As a result, a t r e m e n d o u s body of erature now exists c o n c e r n i n g their tax~omy a n d biology. (Although also comnly called "fruit flies," they are not to be nfused with the T e p h r i t i d a e . ) T h e family is large in t h e Neotropics, ntaining almost 700 species, the majority which belong to the g e n u s Drosophila (Val al. 1981). T h e vast majority of species a r e nizens of w o o d l a n d s ; a few a r e desert «fellers. A d u l t s are d i u r n a l a n d active for ef periods d u r i n g t h e day. T h e larvae "■ 11.7b) of these mostly feed o n microorisms, especially yeasts (da C u n h a et al. 57), associated with spoiled fruit, slime p) fluxes o n tree t r u n k s a n d roots, in : ngcacti ( B e n a d o e t a l . 1984), a n d fungi similar f e r m e n t i n g vegetable m a t t e r ¡eth a n d H e e d 1972). S o m e species and feed as adults in living flowers

(often Calathea a n d Helwonia) (Pipkin et al. 1966). Two r e m a r k a b l e species, D. carcinophila a n d D. endobmnchia, in t h e C a r i b b e a n live o n the bodies of land crabs, probably as commensals (Carson 1967, C a r s o n a n d W h e e l e r 1968).

References ASHBURNER, M., ed. 1976-1981. T h e genetics and biology of Drosophila. 3 vols. Pergamon, Oxford. BELO, M., AND I. J.

J. DE OLIVEIRA.

1976.

Especies domésticas de Drosophila. V. Influen­ cias de factores ambientáis no número de individuos capturados. Rev. Brasil. Biol. 36: 903-909. BENADO, M., A. FONTDEVII.A, H. G. CERCA, G. GARCIA, A. Ruiz, AND C. MONTERO. 1984. On

the distribution and the cactiphilrc niche of Drosophila martens is in Venezuela. Biotropica 16: 120-124. CARSON, H. L. 1967. T h e association between Drosophila carcinophila Wheeler and its host, the land crab Gecarcinus niñeóla (L.). Amer. Midi. Nat. 78: 324-343. CARSON,

H. L., AND M. R. WHEELER.

1968.

Drosophila endobmnchia, a new drosophilid associated with land crabs in the West Indies. Entomol. Soc. Amer. Ann. 61: 675—678. DA CUNHA, A. B., A. M. EL-TABAY SHEHATE,

AND W. DE OLIVEIRA. 1957. A study of the

diets and nutritional preferences of tropical species oí Drosophila. Ecology 38: 98—106. PIPKIN, S. B., R. L. RODRÍGUEZ, AND J. LEÓN.

1966. Plant host specificity among flowerfeeding Neotropical Drosophila (Diptera: Dro­ sophilidae). Amer. Nat. 100: 135-156. RUBIN, G. M. 1988. Drosophila melanogaster as an experimental organism. Science 240: 1453-1459. SHORROCKS, B. 1980. Drosophila. Pergamon, Oxford.

POMACE FLIES

389

VAL, F. C. 1972. On the biometry and evolution of the genus Ornidia (Diptera, Syrphidae). Univ. Sao Paulo Mus. Zool. Pap. Avul. Zool. 26:1-28.

WASP FLY Stratiomyidae, H e r m e t i i n a e , Hermetia illuscens. Spanish: G u a r e r o , b o r r a c h e r o (Central America). T h i s semidomestic species is f o u n d in all t e m p e r a t e a n d tropical p o r t i o n s of t h e world, owing to the b r o a d adaptive capaci­ ties of adults a n d t h e t r a n s p o r t a t i o n of larvae in c o n t a m i n a t e d food. Because all o t h e r m e m b e r s of its g e n u s a r e N e o t r o p i ­ cal, it probably o r i g i n a t e d in some w a r m area of the New World. T h e larvae d e v e l o p in a wide variety of organic media, i n c l u d i n g decaying fruit a n d vegetables, g a r b a g e , compost, d u n g , carrion, a n d soil c o n t a m i n a t e d with these materials (Copello 1926). In C e n t r a l A m e r i ­ can b a n a n a plantations, t h e species d a m ­ ages fruit by ovipositing between t h e fin­ gers of y o u n g fruit a n d o n ripe b u n c h e s after they have b e e n picked. A s t r o n g attraction for yellow color by t h e female is t h o u g h t to account at least partly for t h e latter (Stephens 1975). W h e n they occupy t h e same food, wasp fly larvae m a y s u p p r e s s housefly p o p u l a ­ tions t h r o u g h some indirect competitive effect, since the f o r m e r d o not feed o n t h e latter ( F u r m a n et al. 1959). Larvae a r e also c o m m o n l y f o u n d in h o n e y b e e hives w h e r e they feed o n honey, wax, a n d waste materi­ als. T h e r e is o n e r e c o r d from t h e nest of a stingless b e e ( B o r g m e i e r 1930) which may indicate a g e n e r a l habit in the wild for this a n d o t h e r m e m b e r s of t h e g e n u s . Larvae also a r e occasionally t h e cause of h u m a n intestinal myiasis. Identifying characteristics of t h e larval stage (BL 20 m m ) a r e a thick, leathery, d a r k b r o w n i n t e g u m e n t that is set with n u m e r o u s short bristles, a b r o a d , flat­

388

FLIES AND MIDGES

t e n e d , ovate general s h a p e , a n d a slender well-sclerotized head (fig. 1 1.6e). T h e moderately large (BL 1 5 - 2 0 xx\vc\\ adults (fig. 11.6f) m u c h resemble spider wasps in a n a t o m y a n d behavior. T h e y are elongate a n d d a r k bluish-black, with uni­ formly dusky wings a n d white tarsi (the h i n d tibia is also white); at t h e base of the a b d o m e n are paired t r a n s p a r e n t areas sep­ arated by a d a r k m e d i a n line, simulating a wasp's waist. T h e male a b d o m e n is bronzy in contrast to black in females. T h e flies c o m p o r t themselves waspishly, r u n n i n g on the g r o u n d excitedly a n d flicking their wings. T h e y also buzz a n d feign a sting when h a n d l e d (Iide a n d Mileti 1976)

References BORGMEIER, T. 1930. Ueber das Vorkommen der Larven von Hermetia illuscens L. (Dipt. Stratiomyidae) in den Nestern von Meliponiden. Zool. Anz. 90: 225-235. COPELLO, A. 1926. Biología de Hermetia illuscens Latr. Rev. Entomol. Argentina 1(2): 23-27. [Sic] Author of species is Linneaus. FURMAN, D. P., R. D. YOUNG, AND E. P. CATTS.

1959. Hermetia illuscens (Linnaeus) as a factor in the natural control of Musca domestica Linnaeus. J. Econ. Entomol. 52: 917-921. IIDE, P., AND D. I. C. MILETI. 1976. Estudos

morfológicos sobre Hermetia illuscens (Lin­ naeus, 1758) (Diptera, Stratiomyidae). Rev. Brasil. Biol. 36: 923-935. STEPHENS, C. S. 1975. Hermetia illuscens (Dip­ tera: Stratiomyidae) as a banana pest in Panama. Trop. Agrie. 52: 173-178.

POMACE FLIES Drosophilidae, Drosophilia. T h e s e gnatlike (BL 2 - 3 m m ) , yellowish to brownish flies (fig. 11.7a) a r e usually seen a r o u n d decaying vegetation a n d ripe fruit (Shorrocks 1980, Belo a n d d e Oliveira 1976). T h e y a r e best known, however, for the roles a few species have played in the a d v a n c e m e n t of t h e science of heredity. Early in this century, geneticist Thomas H u n t M o r g a n f o u n d that o n e species, Drosophilia melanogaster, r e p r o d u c e d with

Figure 11.7 FLIES, (a) Pomace fly (Drosophila melanogaster, Drosophilidae). (b) Pomace fly, larva, (c) Mediterranean fruit fly (Ceratitis capitata, Tephritidae). (d) Eye gnat (Liohippelates pusio, Chloropidae). (e) Stilt-legged fly (Taeniaptera sp., Micropezidae). great rapidity, was easily c u l t u r e d in t h e laboratory, r e q u i r e d b u t twelve days to develop from t h e e g g to maturity, a n d h a d larval cells c o n t a i n i n g only four, large, wellmarked c h r o m o s o m e s . I n fact, t h e insect was so well a d a p t e d for genetics that some­ one once q u i p p e d that it must have b e e n created by the Almighty solely as a n object of heredity r e s e a r c h . Since M o r g a n ' s days, many additional species have b e e n studied in great detail a n d c o n t r i b u t e d m o r e to o u r understanding of evolutionary mecha­ nisms, p o p u l a t i o n dynamics, phylogeny, cy­ tology, a n d so o n , t h a n a n y o t h e r single group of h i g h e r o r g a n i s m s ( A s h b u r n e r 1976-1981, R u b i n 1988, T h r o c k m o r t o n 1975). As a result, a t r e m e n d o u s body of literature now exists c o n c e r n i n g their tax­ onomy a n d biology. ( A l t h o u g h also com­ monly called "fruit flies," they are not to b e confused with t h e T e p h r i t i d a e . ) T h e family is large in t h e Neotropics, containing almost 700 species, the majority of which b e l o n g to the g e n u s Drosophila (Val etal. 1981). T h e vast majority of species are denizens of w o o d l a n d s ; a few a r e d e s e r t dwellers. Adults a r e d i u r n a l a n d active for ¿brief periods d u r i n g t h e day. T h e larvae :(fig. 11.7b) of these mostly feed on microor­ ganisms, especially yeasts (da C u n h a et al. 1957), associated with spoiled fruit, slime (*ap) fluxes o n tree t r u n k s a n d roots, in lotting cacti ( B e n a d o et al. 1984), a n d fungi *nd similar f e r m e n t i n g vegetable m a t t e r pieth a n d H e e d 1972). S o m e species d a n d feed as adults in living flowers

(often Calalhea a n d Heliconia) (Pipkin et al. 1966). Two r e m a r k a b l e species, D. carcinophila a n d D. endobranchia, in the C a r i b b e a n live on the bodies of land crabs, probably as c o m m e n s a l s (Carson 1967, C a r s o n a n d W h e e l e r 1968).

References ASHBURNER, M., ed. 1976-1981. T h e genetics and biology of Drosophila. 3 vols. Pergamon, Oxford. BELO, M., AND 1. J. J. DE OLIVEIRA.

1976.

Especies domésticas de Drosophila. V. Influen­ cias de factores ambientáis no número de individuos capturados. Rev. Brasil. Biol. 36: 903-909. BENADO, M., A. FONTDEVILA, H. G. CERCA, G. GARCIA, A. Ruiz, AND C. MONTERO. 1984. On

the distribution and the cactiphilic niche of Drosophila marlensis in Venezuela. Biotropica 16: 120-124. CARSON, H. L. 1967. The association between Drosophila carcinophila Wheeler and its host, the land crab Gecarcinus ruricola (L.). Amer. Midi. Nat. 78: 324-343. CARSON, H. L., AND M. R. WHEELER.

1968.

Drosophila endobranchia, a new drosophilid associated with land crabs in the West Indies. Entomol. Soc. Amer. Ann. 61: 675-678. DA CUNHA, A. B., A. M. EL-TABAY SHEHATE,

AND VV. DE OLIVEIRA. 1957. A study of the

diets and nutritional preferences of tropical species of Drosophila. Ecology 38: 98-106. PIPKIN, S. B., R. L. RODRÍGUEZ, AND J. LEÓN.

1966. Plant host specificity among flowerfeeding Neotropical Drosophila (Diptera: Dro­ sophilidae). Amer. Nat. 100: 135-156. RUBIN, G. M. 1988. Drosophila melanogaster as an experimental organism. Science 240: 1453-1459. SHORROCKS, B. 1980. Drosophila. Pergamon, Oxford.

POMACE FLIES

389

SPIETH, H. T , AND W. B. HEED. 1972. Experi­

mental systematics and ecology of Drosophila. Ann. Rev. Ecol. Syst. 3: 269-288. THROCKMORTON, L. H. 1975. In R. C. King,

ed., Handbook of genetics. 3. Invertebrates of genetic interest. Plenum, New York. Pp. 421-469. VAL, F. C , C. R. VILELA, AND M. D. MARQUES.

1981. Drosophilidae of the Neotropical Re­ gion. In M. Ashburner, ed., The genetics and biology of Drosophila. 3a: 123—168. Pergamon, Oxford.

FRUIT FLIES T e p h r i t i d a e . Spanish: Moscas d e la fruta. Portuguese: Moscas d a fruta. T h e s e a r e small t o m e d i u m - s i z e d flies (BL 3—7 m m ) with s p o t t e d o r b a n d e d wings in colors of b r o w n a n d white, often f o r m i n g complicated o r attractive p a t t e r n s . Viewed from b e h i n d , living flies may mimic j u m p ­ ing spiders: t h e wing bars simulate legs, a n d d a r k spots o n t h e a p e x of t h e a b d o ­ m e n look like t h e spider's eyes (Eisner 1984). T h e a p e x of t h e subcostal vein angles sharply forward to distinguish t h e m from o t h e r p i c t u r e d - w i n g e d flies. Adults are c o m m o n l y observed feeding o n nectar from flowers o r ovipositing o n fruit. W h e n at rest o r walking, s o m e have t h e o d d behavior of slowly m o v i n g their wings u p a n d d o w n as if displaying t h e m . T h e fe­ males of most species insert their eggs in living, healthy plant tissue. T h e larvae, which a r e typical maggots, feed in stalks, leaves, flowers, a n d often in fruits, a n d thus, m a n y species have b e c o m e major agricultural pests ( B a t e m a n 1976, Cavalloro 1983). T h e most serious a n d w i d e s p r e a d is t h e M e d i t e r r a n e a n fruit fly or so-called Medfly (Ceratitis capitata; fig. 11.7c), which attacks citrus, papaya, m a n g o , p i n e a p p l e , a n d as m a n y as 260 o t h e r hosts from C e n t r a l Mexico south t h r o u g h most of South A m e r i c a (not in t h e Antilles) (Weems 1981). Many c o n s p i c u o u s , economically im­ p o r t a n t species a r e f o u n d also in t h e g e n u s Anastrepha, such as t h e S o u t h A m e r i c a n (A.

390

FLIES AND MIDGES

fraterculus), Mexican (A. ludens), a n d Carib­ bean fruit fly (A. suspensa). Larvae of the Medfly a r e ubiquitous in guayaba fruits and i n c o r p o r a t e d unavoidably into jelly and o t h e r p r o d u c t s m a d e from t h e m . T h e pres­ ence of larvae of these species in fruit c o n s u m e d by h u m a n s sometimes causes an intestinal form of myiasis, t h e principal s y m p t o m of which is d i a r r h e a (Jirón and Zeledón 1979).

BATEMAN, M. A. 1972. T h e ecology of fruit flies. Ann. Rev. Entomol. 17: 493-518. BATEMAN, M. A. 1976. Fruit flies. In V. L. Delucchi, ed.. Studies in biological control. Int. Biol. Prog. 9: 11-49. Cambridge Univ. Press, Cambridge. ROLLER, E. F., AND R. J. PROKOPY. 1976. Bionom­

ics and management of Rhagoletis. Ann. Rev. Entomol. 21: 223-246. CAVALLORO, R., ed. 1983. Fruit flies of economic importance. Proc. CEC/IOBC Int. Symp. 1982. Balkema, Rotterdam.

Some m e m b e r s of t h e mainly temper­ ate genus Rhagoletis a r e also of major i m p o r t a n c e (Boiler a n d Prokopy 1976). Unlike t h e others m e n t i o n e d , t h e species have relatively n a r r o w food preferences for e x a m p l e , R. lycopersella, a species in­ d i g e n o u s t o cultivated valleys in t h e dry coastal plains of western Peru which at­ tacks only tomatoes (Smyth 1960) and Toxotripana curvicauda, which attacks only papaya (Adarve 1979).

CHRISTENSON, L. O., AND R. H. FOOTE. 1960.

Tropical fruit flies a r e reproductively active t h r o u g h o u t t h e year. In temperate areas, t h e r e a r e species with only o n e gen­ eration p e r year, t h e adults having a winter h i b e r n a t i o n period. Both types tend to form local transient p o p u l a t i o n s through dispersal of t h e s t r o n g flying females. Except for a few economically impor­ tant species (Prokopy a n d Roitberg 1984), the Neotropical fauna is generally poorly studied (Christenson a n d Foote 1960, Bate­ m a n 1972). T h e family is a fairly diverse o n e in t h e region, with m o r e than 680 species a n d 88 g e n e r a (Foote 1980). Adults feed o n a variety of natural soupy liquids such as t h e juices oozing from d a m a g e d o r decaying fruit, plant sap, flower nectar, a n d bird feces. Adults and larvae of many species h a r b o r symbiotic microorganisms in their intestines, which most likely provide specific nutritive sub­ stances from vegetable tissues.

PROKOPY, R. J., AND B. D. ROITBERG.

References ADARVE, R. 1979. Observaciones sobre los há­ bitos del Toxotripana curvicauda Gerst (Tephri­ tidae) que ataca al Carica papaya. Ceiba 23: 63-75.

Biology of fruit flies. Ann. Rev. Entomol. 5: 171-192. EISNER, T. 1984. Consumer fraud. Nat. Hist. 93: 112. FOOTE, R- H. 1980. Fruit fly genera south of the United States. U.S. Dept. Agrie. Tech. Bull. 1600: 1-79. [IRON, L. F., AND R. ZELEDÓN. 1979. El género

Anastrepha (Diptera: Tephritidae) en las prin­ cipales frutas de Costa Rica y su relación con pseudomiasis humana. Rev. Biol. Trop. 27: 155-161.

Neotropical species ( L e g n e r a n d Bay 1965, Sabrosky 1984). All a r e small (BL 1.5-2.5 m m ) a n d generally shiny black with clear wings; most were formerly included in t h e g e n u s Hippelates. O t h e r small gnatlike flies have similar noxious habits. Paraleucopis mexicana (Chamaemyiidae), called "bobos," walk o n ex­ posed skin a n d hair. T h e y d o not bite, b u t large n u m b e r s swarm a b o u t t h e face a n d h e a d , m a k i n g it very u n p l e a s a n t to be n e a r their seaside habitat. T h e y also cluster a b o u t w o u n d s . T h e s e flies c o m e every year in March to t h e u p p e r Gulf of California (Smith 1981) w h e r e they m a k e t h e local residents miserable with their persistent aggravations. T h e i r n o r m a l feeding seems to b e o n secretions e m a n a t i n g from t h e eyes of lizards a n d m a r i n e birds. T h e y have not b e e n incriminated in t h e s p r e a d of any diseases.

1984.

Foraging behavior of true fruit flies. Amer. Sci. 7 2 : 4 1 - 4 9 . SMYTH, E. G. 1960. A new tephritid fly injuri­ ous to tomatoes in Peru. Dept. Agrie. State Calif Bull. 49(1): 16-22. WEEMS, H. V. 1981. Mediterranean fruit fly, Ceratitis capitata (Wiedmann) (Diptera: Tephri­ tidae). Fia. Dept. Agrie. Cons. Ser., Div. Plant Indus., Entomol. Circ. 230: 1-4 (unnum­ bered) + 1-8.

EYE GNATS Chloropidae, Liohippelates pusio c o m p l e x . Ulcer flies (West Indies). Although they d o n o t bite, eye gnats a r e a major source of t o r m e n t because of their persistent habit of e n t e r i n g t h e eyes a n d clustering a b o u t w o u n d s a n d t h e e x p o s e d genitals of m a m m a l s ("dog pecker gnats"). They were implicated in t h e mechanical transmission of yaws in t h e West Indies when t h e disease was p r e v a l e n t (Nicholls 1936). T h e c o m m o n species h e r e a r e f o u n d •n the L. pusio c o m p l e x (fig. 11.7d) in a Widely distributed g e n u s of twenty-nine

References LEGNER, E. F , AND E. C. BAY. 1965. Culture of

Hippelates pusio (Diptera: Chloropidae) in the West Indies for natural enemy exploration and some notes on behavior and distribution. Entomol. Soc. Amer. Ann. 58: 436-440. NICHOLLS, L. 1936. Framboesia trópica: A short review of a colonial report concerning statis­ tics and Hippelates flavipes. Amer. J. Trop. Med. Paras. 30: 331-335. SABROSKY, C. W. 1984. Family Chloropidae. In N. Papavero, ed., A catalogue of the Diptera of the Americas south of the United States. Mus. Zool., Univ. Sao Paulo, Sao Paulo. Pp. 81.1-81.63. SMITH, R. L. 1981. T h e trouble with "bobos," Paraleucopis mexicana Steyskal, at Kino Bay, Sonora, Mexico (Diptera: Chamaemyiidae). Entomol. Soc. Wash. Proc. 83: 406-412.

STILT-LEGGED FLIES Micropezidae ( = Tylidae). T h e s e a r e slender, medium-sized (BL 10— 15 m m ) flies with very long mid a n d h i n d legs, c o m p a r e d to t h e forelegs, which a r e a b o u t half their length. T h e tip of t h e

STILT-LEGGED FLIES

391

a b d o m e n is usually r e c u r v e d ventrally. Many a r e d a r k colored with b a n d e d wings (such as m e m b e r s of the g e n u s Taeniaptera; fig. 11.7e) a n d c o n t r a s t i n g white-tipped tarsal s e g m e n t s ; the h e a d is also often light h u e d in shades of red or o r a n g e . Adults are d i u r n a l , slow to fly, a n d usually seen walking nervously a b o u t o n the u p p e r surfaces of leaves with the wings closed over the d o r s u m a n d the forelegs held erect a n d waving in front of the h e a d . T h e i r f o r m , coloration, a n d this behavior gives t h e m a s t r o n g r e s e m b l a n c e to ants. T h e mimicry is e n h a n c e d by the conspicu­ ous white-tipped forelegs, stimulating an­ t e n n a e , a n d suitably placed d a r k wing b a n d s , which give t h e impression of a n a r r o w e d , antlike waist. T h i s is mainly a tropical g r o u p of a b o u t 280 species (Steyskal 1968), the adults fre­ q u e n t i n g shady, moist forest habitats. T h e i m m a t u r e s a r e poorly k n o w n . T h e maggots of s o m e species have b e e n f o u n d living in feces, r o t t i n g fruit, a n d d e c o m p o s i n g plant parts such as the p s e u d o - s t e m s of b a n a n a s (Fischer 1932). O t h e r s may b o r e into living plant tissue, a n d at least o n e species is a m i n o r pest by i n v a d i n g g i n g e r roots in the Old World (Steyskal 1964). T h e s e flies are also k n o w n for their bizarre c o u r t s h i p habits. T h e antics of Plocoscelus arthriticus ( = Cardiacephala myrrnex) in P a n a m a w e r e r e c o r d e d by W h e e l e r (1924). Males a p p r o a c h willing females h e a d o n a n d p e r f o r m a peculiar d a n c e , s t e p p i n g first to o n e side a n d t h e n to the other, swaying t h e a b d o m e n . After witness­ ing this d a n c e , the female b e n d s h e r body in an arc by t h r o w i n g back t h e h e a d a n d t u r n i n g u p the tip of t h e a b d o m e n . T h e male t h e n m o u n t s h e r a n d brings his p r o ­ boscis in contact with h e r s , a d r o p of food o n it, a n d simultaneously inserts his introm i t t e n t o r g a n . Occasionally, h e reaches for­ ward with a foreleg a n d scratches the fe­ male's eye a n d places m o r e food o n the c o r n e r of it. T h e s e acts, with o t h e r detailed m o v e m e n t s , may be r e p e a t e d several times.

392

FLIES AND MIDGES

The female finally kicks the male free, and copulation is c o m p l e t e .

References FISCHER, C. R. 1932. Contribuicáo para o conhecimento da metamorphose e posicáo systematica da familia Tylidae (Micropezidae Dipt.). Rev. Entomol. 2: 15-24. STEYSKAL, G. C. 1964. Larvae of Micropezidae (Diptera), including two species that bore in ginger roots. Entomol. Soc. Amer. Ann 57292-296. STEYSKAL, G. G. 1968. Family Micropezidae. In N. Papavero, A catalogue of the Diptera of the Americas south of the United States Dept. Zool., Sec. Agrie, Sao Paulo, no 48 48.1-48.33. ' ' ' WHEELER, W. M. 1924. Courtship of Calobatos the kelep ant and the courtship of its mimic Cardiacephala myntiex. J. Heredity 15: 487494.

Figure 11.8 FLIES, (a) Kelp fly {Fucellia sp., Anthomyiidae). (b) Stable fly (Stomoxys calcitrans, Muscidae). (c) House fly (Musca domestica, Muscidae). (d) House fly, puparium. (e) House fly, larva. (f) Lesser house fly (Fannia canicularis, Muscidae). (g) Lesser house fly, larva. widely spaced, coarse setae on the lower surface of the a n t e r i o r m o s t wing vein. A d d i ­ tionally, the males have a heavily spined protuberance at t h e base of t h e h i n d f e m u r on the i n n e r side. Very little has b e e n written on the Neo­ tropical Fucellia. T h e life history of the European F. marilima may serve as a m o d e l for the g e n u s (Egglishaw 1960).

KELP FLIES A n t h o m y i i d a e , Fucellia. Larval kelp flies live in old piles of the b r o w n seaweeds (kelp, Fucus, Macrocystis, etc.) cast u p by the waves along sea beaches. T h e y feed o n the d e a d plant and are i m p o r t a n t in its decomposition a n d nutri­ ent cycling in the coastal habitat. T h e adults can be seen mostly d u r i n g the warmer m o n t h s of the year on h e a p s of such mate­ rial (wrack beds) a n d may develop enor­ m o u s populations. Sometimes d u r i n g the winter or on cold days, d e n s e masses of flies are seen packed into crevices on rocky cliffs above the s h o r e , a p p a r e n t l y overwintering. Species are f o u n d t h r o u g h o u t the tropi­ cal A m e r i c a n region on b o t h the Atlantic a n d Pacific seaboards, including offshore islands a n d the Caribbean a n d J u a n Fernan­ dez islands. T h e y are medium-sized ( B L 4 5 m m ) a n d gray-brown, somewhat resem­ bling the h o u s e fly, but with fine hairs b e n e a t h the scutellum, small, bulging r o u n d eyes, a n d with a row of 3 to 4 stout, erect bristles o n the u p p e r side of the hind tibia (fig. 11.8a) (Aldrich 1918). These are

References ALDRICH, J. M. 1918. The kelp-flies of North America (Genus Fucellia, Family Anthomyidae). Calif. Acad. Sci. Proc. (4 ser.) 8: 157179. EGGLISHAW, H. J.

1960.

The

life-history

of

Fucellia marilima (Haliday) (Diptera, Musci­ dae). Entomologist 93: 225-231.

MUSCID FLIES Muscidae. This is the largest family of muscoid Dip­ tera, with 830 species in the Neotropics (Font 1972) h a v i n g varied forms a n d bio­ logies (Skidmore 1985). Adults are gener­ ally small to m e d i u m - s i z e d , dull gray to brown flies (a few, such as Muscina, are metallic g r e e n or blue like calliphorids). are m o d e r a t e l y hairy but a r e u n i q u e ong Calypterates (a feature s h a r e d only the closely related A n t h o m y i i d a e ) in king bristles o n the thoracic sclerite mediately d o r s a l to the base of the h i n d (= h y p o p l e u r o n ) . Larvae are mostly typical maggots, al­

t h o u g h some are flattened a n d exhibit projections (Fannia) or o t h e r peculiar m o d i ­ fications of the cuticle. T h e y a r e e x t r e m e l y varied in habits, scavenging animal feces a n d feeding o n c a r r i o n a n d o t h e r insects, in m a n y habitats, i n c l u d i n g r u n n i n g a n d s t a n d i n g water (Limnophora). T h e larvae of the m o r e t h a n thirty species of Philornis ( = Neomusca) are found in birds' nests w h e r e they feed on feces a n d o t h e r o r g a n i c mat­ ter, occasionally attacking the nestlings a n d causing s u b c u t a n e o u s myiasis ( G u i m a r a e s et al. 1983). T h e conspicuous, noisy adults place their eggs or larvae o n t h e chicks of o r e p e n d o l a s a n d caciques. T h e larvae b u r ­ row into the chick's body, often killing it. Curiously, nests of these birds parasitized by the giant cowbird in P a n a m a suffer less h a r m from Philornis because of the p r o p e n ­ sity for p r e e n i n g exhibited by the alien chick (Smith 1968). T h e reverse situation is described by Fraga (1984) for cowbirds. Several Muscidae a r e cosmopolitan asso­ ciates of civilization a n d a r e i m p o r t a n t as food c o n t a m i n a t o r s a n d h o u s e h o l d invad­ ers. O t h e r s (subfamily S t o m o x y i n a e : Sto­ moxys, Haematobia, a n d Neivamyia) bite or are e x t r e m e nuisances to h u m a n s a n d domestic animals (Pinto a n d d e Souza Lopes 1933).

References FRAGA, R. M. 1984. Bay-winged cowbirds (Molothrus badius) remove ectoparasites from their brood parasites, the screaming cowbirds (M. rufoaxillaris). Biotropica 16: 223-226.

MUSCID FLIES

393

GUIMARÁES, J. H., N. PAPAVERO, AND A. P. Do

PRADO. 1983. As miíases na regiáo Neotropi­ cal (Identificacáo, biología, bibliografía). Rev. Brasil. Zool. 1: 239-416. PINTO,

C,

AND H.

DE SOUZA

LOPES.

1933.

Anatomía, biología e papel patogénico da Neivamyia lutzi, mosca sugadora de sangue dos equídeos do Brasil. Escol. Sup. Agrie. Med. Vet. Arch. 10: 77-88. PONT, A. C. 1972. Family Muscidae. in N. Papavero, ed., A catalogue of the Díptera of the Americas south of the United States. Mus. Zool., Univ. Sao Paulo, Sao Paulo. Pp. 97.1-97.111. SKIDMORE, P. 1985. T h e biology of the Muscidae of the World. Junk (Series Entomológica), The Hague. SMITH, N. G. 1968. The advantage of being parasitized. Nature 219: 690-694. ZUMPT, F. 1973. T h e Stomoxyine biting flies of the world. Diptera: Muscidae. Taxonomy, biology, economic importance and control measures. Fischer, Stuttgart.

House Fly Muscidae, Muscinae, Musca domestica. Spanish: Mosca casera. Portuguese: Mosca doméstica, mosca c o m ú n d a s casas. T y p h o i d fly. T h i s species is k n o w n by its ubiquity a n d close association with m a n k i n d . I n d e e d , its n u m b e r s in slums, u n k e m p t corrals of domestic animals, d u m p s , a n d o t h e r sites of h u m a n neglect ( B a u m g a r t n e r 1988) may be astronomical a n d t h r e a t e n public health t h r o u g h m e c h a n i c a l transmission of dysentery-causing bacteria a n d protozoa ( C o u t i n h o et al. 1957). T h e species was not always so a b u n d a n t in t h e A m e r i c a n t r o p ­ ics; not long after its colonization by Eu­ r o p e , t h e New World is t h o u g h t to have received Musca domestica with h u m a n traf­ fic from t h e O l d World. Only in undis­ t u r b e d forests a n d t h e most r e m o t e u n i n ­ habited regions is it still absent. T h e a d u l t is small (BL 6—9 m m ) , g e n e r ­ ally gray, with four d a r k lines r u n n i n g longitudinally o n t h e back of t h e t h o r a x ; the lateral p o r t i o n s of t h e a b d o m e n a r e translucent b r o w n (fig. 11.8c). T h e larva (fig. 11.8e) is a typical, elongate m a g g o t

394

FLIES AND MIDGES

(BL to 12 m m w h e n m a t u r e ) , creamy white, a n d may be distinguished from that of o t h e r similar domestic fly larvae by the s h a p e of t h e posterior spiracles, t h r e e ser­ p e n t i n e slits s u r r o u n d e d by a complete sclerotized ring. T h e p u p a r i u m (fig. 11.8d) is outwardly indistinguishable from that of o t h e r similar-sized muscoid flies. T h e species b r e e d s in a wide r a n g e of d e c o m p o s i n g organic m a t t e r b u t finds horse m a n u r e an especially favored food m e d i u m . O t h e r materials c o m m o n l y har­ b o r i n g larvae a r e cow d u n g , h u m a n feces a n d refuse h e a p s of vegetables a n d fruit, all as long as they a r e fairly moist a n d warm. T h e larvae a r e not normally found in carcasses o r c o m m o n l y involved in myiasis. Details of t h e life cycles a r e available in n u m e r o u s publications (Milani 1975, West 1951, West a n d Peters 1973). T h e house fly is highly prolific, with a fantastic potential for multiplication, which, fortunately, is never realized d u e to a substantial mortal­ ity rate from p r e d a t i o n , parasitism, and food a n d e n v i r o n m e n t a l limitations. How­ ever, it is difficult to control artificially (Schenone 1962).

Stable Fly Muscidae, S t o m o x y i n a e , Stomoxys calcitrans. Spanish: Mosca d e los establos. Portuguese: Mosca d o bagado. Dog flyThis m e d i u m / s m a l l (BL 5—6 m m ) muscoid fly also superficially resembles t h e com­ mon house fly with its mostly gray, blackstriped t h o r a x (fig. 11.8b). It differs signifi­ cantly, however, in possessing a spotted abdomen a n d a rigid, elongate sucking beak with m i n u t e biting teeth at its a p e x . This o r g a n is used to p u n c t u r e t h e skin of mammals a n d w i t h d r a w blood. T h e stable fly is a major pest of horses, cattle, a n d humans a n d o t h e r large domestic animals. Both sexes suck blood. T h e typical maggotlike larvae b r e e d in a variety of r o t t i n g plant materials, com­ monly straw in stables, weeds washed u p on lakeshores, a n d seacoast wrack (Kunz et al. 1977). I m m e n s e p o p u l a t i o n s may d e ­ velop n e a r such larval food sources a n d constitute a major nuisance to equestrians and beach b a t h e r s .

Reference KUNZ, S. E., L. L. BERRY, AND K. W. FOF.RSTER

References BAUMGARTNER, D. L. 1988. The housefly, Musca domestica (Diptera, Muscidae), in central Peru: Ecological studies of medical importance. Rev. Brasil. Entomol. 32: 455-463. COUTINHO, J. O., A. DE E. TAUNAY, AND L. P. DE

CARVALHO LIMA. 1957. Importancia da Musca

domestica como vector de agentes patogénicos para o homen. Inst. Adolfo Lutz Rev. 17: 5 23. MILANI, R. 1975. T h e house fly, Musca domestica. in R. C. King, ed., Handbook of genetics. Vol. 3. Invertebrates of genetic interest. Plenum, New York. Pp. 377-399. SCHENONE, H. 1962. Medidas de profilaxis de las moscas. Bol. Chilena Parasit. 17: 23-25. WEST, L. S. 1951. T h e housefly. Comstock, Ithaca. WEST,

L.

S.,

AND O.

B.

PETERS.

1973.

An

annotated bibliography of Musca domestica Linnaeus. Dawsons of Pall Mall, Folkestone and London.

1977. T h e development of the immature forms of Stomoxys calcitrans. Entomol. Soc. Amer. Ann. 70: 169-172.

Lesser House Fly Muscidae, F a n n i i d a e , Fannia

caniculans.

This species r e s e m b l e s t h e h o u s e fly b u t is somewhat smaller (BL 6 - 7 m m ) a n d has a more elongate body s h a p e a n d yellow spots at the base of t h e a b d o m e n (fig. 11.81'). It is most easily recognized by its habit of hover­ ing in small g r o u p s in shady places o n hot days, individuals lazily zigzagging in t h e air, usually j u s t a few feet off t h e g r o u n d , never landing. It is a n a b u n d a n t fly in a n d near h u m a n habitations a n d is f r e q u e n t in outdoor toilets, privies, stables, pigsties, and other places w h e r e e x c r e m e n t accumu­ lates. T h e darkly p i g m e n t e d larvae have a

flattened, fusiform body with series of elongate, basally fringed projections ex­ t e n d i n g from t h e sides a n d d o r s u m (fig. 11.8g). T h e y tolerate very moist substrata a n d b r e e d in t h e fresh liquid feces of most animals a n d poultry.

Green House Fly Muscidae, Muscinae, Muscina False stable fly.

stahulans.

T h i s muscoid is u n u s u a l in being metallic blue-green in color, a p p e a r i n g very similar to many blowflies b u t generally smaller (BL 8 m m ) . Adults a r e invariably found in the n e a r vicinity of h u m a n habitations in r u r a l situations; they a r e scarce in towns. T h e y a r e filth feeders b u t also take honeydew a n d tree sap. T h e larvae feed in h u m a n a n d o t h e r animal feces a n d a wide variety of r o t t i n g m a t t e r (rarely c a r r i o n ; m o r e likely decaying fruit, bird's nests, d e a d insects, etc.).

Horn Fly Muscidae, Stomoxyinae, Haematohia irritans. Spanish: Mosca (mosquilla) del g a n a d o (General). T h e h o r n fly is primarily a p a s t u r e pest of cattle (McLintock a n d D e p n e r 1954, Vo­ gelsang a n d d e A r m a s 1940). It is fairly w i d e s p r e a d b u t has been found in Brazil only relatively recently (Valerio a n d Gui­ m a r á e s 1983). It b r e e d s exclusively in fresh cattle d u n g , a n d t h e small (BL 4 m m ) adults of both sexes c o n g r e g a t e in masses at t h e base of t h e h o r n s a n d suck blood. W h e n very a b u n d a n t , they a r e excessively b o t h e r s o m e to t h e host a n d c o n t r i b u t e significantly to weight a n d milk losses. T h e y a r e attracted to their host by olfac­ tory, heat, a n d visual stimuli, t h e last seem­ ing to be t h e most i m p o r t a n t ( H a r g e t t a n d G o u l d i n g 1962). T h i s species was formerly placed in ei­ t h e r t h e g e n e r a Lyperosia or Siphona. T h e adult superficially resembles t h e lesser

MUSCID FLIES

395

Figure 11.9 DOMESTIC FLIES, (a) Horn fly (Haematobia irritans, Muscidae). (b) Black garbage flv (Ophyra aenescens, Muscidae). (c) Flesh fly (Bercaea haemorrhoidalis, Sarcophagidae), adult (d) Flesh fly, larva, (e) Green blowfly (Phaenicia sericata, Calliphoridae). (f) Green blowfly, larva h o u s e fly (BL a b o u t 3 m m ) b u t has a short rigid beak like t h e stable fly (fig. 11.9a).

References 1962.

Studies on the behavior of the horn fly [Haematobia irritans (Linn.)]. Agrie. Exper. Sta., Ore. State Univ., Tech. Bull. 61: 1-27. MCLINTOCK, J., AND K. R.

DEPNER.

1954.

AND J.

H.

GUIMARÁES.

1983.

Sobre o ocorréncia de urna nova praga, Haematobia irritans (L.) (Diptera, Muscidae), no Brasil. Rev. Brasil. Zool. 1: 417-418. VOGELSANG, E. G., AND J. C. DE ARMAS. 1940. La

mosquilla del ganado, Lyperosia irritans (L. 1761) en Venezuela. Rev. Med. Vet. Parasit. (Min. Agrie. Cria, Venezuela) 2: 95-98.

Black Garbage Fly Muscidae, Muscinae, H y d r o t a e i n i , Ophyra aenescens. Adults of this small (BL 6 m m ) , shiny black species (fig. 11.9b) a r e a t t r a c t e d to h u m a n a n d animal e x c r e m e n t . T h e hairlike bristle of t h e a n t e n n a is very long a n d slightly p u b e s c e n t . T h e m a g g o t s d e v e l o p in t h e a f o r e m e n t i o n e d materials a n d often p r o ­ d u c e heavy p o p u l a t i o n s t h a t become a nui­ sance in u r b a n a n d s u b u r b a n settlements. T h e y show a fair d e g r e e of tolerance for high salinity in their food a n d have b e e n f o u n d c o m m o n also in c a r r i o n of m a r i n e origin a n d even in salted m e a t s ( J o h n s o n a n d V e n a r d 1957, d e Oliveira 1941).

FLIES AND MIDGES

W.

T,

AND C.

E.

VENARD.

1957

Observations on the biology and morphology of Ophyra aenescens (Diptera: Muscidae) Ohio J. Sci. 57: 21-26.

A

review of the life-history and habits of the horn fly, Siphona irritans (L.) (Diptera: Musci­ dae). Gan. Entomol. 86: 2 0 - 3 3 .

396

DE OLIVEIRA, S. J. 1941. Sobre Ophyra aenescens (Wiedmann, 1830) (Diptera: Anthomyidae) Arq. Zool. (Sao Paulo) 2: 341-355. JOHNSON,

HARGETT, L. T., AND R. L. GOULDING.

VALERIO, J. R.,

References

FLESH FLIES S a r c o p h a g i d a e . Spanish: Moscas d e la c a r n e (Argentina). A d u l t flesh flies of most of t h e common species a r e similar, medium-sized muscoid types, gray in general color a n d frequently m a r k e d with longitudinal black bands on the back of t h e t h o r a x a n d checkerboard p a t t e r n s o n t h e a b d o m e n that c h a n g e with varied light incidence. T h e eyes a r e widely s e p a r a t e d in both sexes a n d often bright g r e e n or brick r e d ; t h e genitalia may also be glaring r e d . T h e body is strongly bristled. Larvae a r e typical pale muscoid mag­ gots, often fairly large (BL when mature 17 m m ) a n d with t r u n c a t e posteriors. T h e latter has a d e e p concavity rimed with p r o m i n e n t fleshy tubercles, in the bottom of which a r e located t h e paired r e a r spira­ cles with t h r e e straight slits each. T h e family displays a great variety of biodc types (Jirón a n d Marín 1982) and a wider s p e c t r u m of hosts t h a n any other, a l t h o u g h , as a rule, t h e larvae a r e pred a c e o u s o r e n d o p a r a s i t o i d s o n o t h e r ani­ mals (Souza Lopes 1973). Most attack other

invertebrates, such as snails, free-living (rrasshoppers (Doringia acridiorum para­ sitizes t h e S o u t h A m e r i c a n locusts), a n d tatydids, b u t s o m e scavenge d e a d insects, even including provisions in wasp's nests. A minority b r e e d in e x c r e m e n t a n d d e c o m ­ posing o r g a n i c m a t t e r o r a r e involved in various m o d e s of myiasis in m a m m a l s a n d humans; they rarely d e v e l o p in carrion. These mostly belong to various species formerly placed in Sarcophaga, an O l d World g e n u s , b u t now a r e dispersed a m o n g several g e n e r a , primarily Peckia. T h e g e n u s Dexosarcophaga lives in t e r m i t e a n d a n t nests o r in t h e galleries of some w o o d - b o r i n g insects. T h e adults of m a n y species associate with h u m a n s a n d may be a factor in mechanical disease transmission (Gregor 1972). Bercaea haemorrhoidalis (fig. 11.9c), widely cited in t h e g e n u s Sarcophaga, is a cosmopolitan species often associated with man. Its larvae (fig. 11.9d) feed on carrion, excrement, a n d e x p o s e d meats a n d some­ times cause myiasis. T h e family is large, with over 600 spe­ cies in t h e N e o t r o p i c a l Region.

References GREGOR, F. 1972. Synanthropy of Sarcophaginae (Diptera) from Guba. Fol. Parasit. 19: 155-163. JIRÓN, L. E, AND R. E. MARÍN. 1982.

Moscas

sarcofágidas de Costa Rica (Diptera; Cyclorrhapha). Rev. Biol. Trop. 30: 105-106. ' SOUZA LOPES, H. 1973. Collecting and rearing

sarcophagid flies (Diptera) in Brazil during forty years. Acad. Brasil. Cien. An. 45: 2 7 9 291.

CARRION FLIES Calliphoridae. Spanish: Moscas verdes (azules) d e la c a r n e (Argentina). Quechua: Shinguitos (Peru, larvae). 5. Blowflies, bluebottle flies. ¡embers of this family (Hall 1948, Norris 1965) a r e t h e familiar, m e d i u m to small

(BL 5—10 m m ) , metallic g r e e n a n d blue muscoid flies that buzz a b o u t fresh c a r r i o n (in t h e "blown" stage, h e n c e t h e c o m m o n n a m e ) . T h e i r maggots (fig. 11.9f) thrive in d e a d animal tissue, a n d their f e e d i n g con­ stitutes a d o m i n a n t factor in t h e p r i m a r y r e d u c t i o n of most u n t e n d e d v e r t e b r a t e cadavers a n d thus is of considerable i m p o r ­ tance in ecological hygiene a n d n u t r i e n t cycling. T h o s e of a few species a r e obliga­ tory parasites, attacking healthy flesh, b u t most may assume facultative roles in my­ iasis a n d as veterinary a n d medical pests. Adults may also transmit p a t h o g e n s m e ­ chanically from putrefactive or fecal m a t e ­ rial to h u m a n food. Because c a r r i o n flies b r e e d in cadavers, their p r e s e n c e a n d d e ­ velopmental rates a r e frequently used in forensic medicine to d e t e r m i n e t h e time of d e a t h of cast-off h u m a n bodies (Easton a n d Smith 1970). Notable a m o n g t h e a p p r o x i m a t e l y o n e h u n d r e d native A m e r i c a n tropical species (Dale 1987, J a m e s 1970) a r e t h e black blowfly (Phormia regina), a variable-sized, blackish-green species, a n d t h e g r e e n b o t t l e fly (Lucilia illustris), a small, b l u i s h - g r e e n fly. Both of these occur n o f a r t h e r s o u t h t h a n central Mexico. T h e g r e e n blowflies (sheep blowflies, fleeceworms, Phaenicia sericata [fig. 11.9e], P. cuprina, a n d P. eximia), bright, metallic green to b r o n z e , a r e t h e usual types found in h u m a n refuse ( Q u a t t r o a n d Wasti 1978). T h e bluebottles (Calliphora species), with shiny d a r k blue bodies, a r e normally associated with carcasses. All t h e p r e c e d i n g flies a r e w i d e - r a n g i n g a n d e x t r e m e l y com­ m o n in u r b a n as well as in r u r a l situations. (See G r e e n b e r g a n d Szyska 1984 a n d J i r ó n a n d Marín 1984, for information o n these a n d additional species.) In recent years, four species of t h e economically i m p o r t a n t O l d world g e n u s Chrysomya have b e c o m e established a n d a r e s p r e a d i n g in Latin A m e r i c a ( B a u m g a r t n e r a n d G r e e n b e r g 1984, G u i m a r á e s et al. 1979, J i r ó n 1979). T h e s e have habits simi­ lar to those of t h e screwworm flies a n d a r e

CARRION FLIES

397

displacing native calliphorid species. T h e y all associate with h u m a n s a n d live n e a r dwellings to a g r e a t e r o r lesser d e g r e e a n d a r e potential public h e a l t h a n d veterinary nuisances ( B a u m g a r t n e r a n d G r e e n b e r g 1984). T h e most serious m y i a s i s - p r o d u c i n g spe­ cies, t h e screwworms, a r e discussed below.

References

JIRÓN,

L. E, AND F. J. MARÍN.

1984. Notas

complementarias sobre moscas califóridas de Costa Rica (Diptera: Calliphoridae). Brenesia a 22: 65-68. NORRIS, K. R. 1965. T h e bionomics of blow flies. Ann. Rev. Entomol. 10: 47-68. QUATTRO, M. J., AND S. S. WASTI. 1978. Olf ac .

tory and oviposition responses of the green bottle fly, Phoenicia sericata (Meig.), to a vari­ ety of natural baits (Diptera: Calliphoridae) Rev. Brasil. Biol. 38: 115-119.

BAUMGARTNER, D. L., AND B. GREENBERG. 1984.

The genus Chrysomyia (Díptera: Calliphoridae) in the New World. J. Med. Entomol. 21: 105-113. DALE, W. E. 1987. Identidad de las moscas Calliphoridae en la costa central del Perú. Rev. Peruana Entomol. 28: 6 3 - 7 0 .

Screwworms

GuiMARÁES, J . H . , A . P. DO PRADO, AND G . M .

Calliphoridae, Cochliomyia hominovorax (= americana). Spanish: L a r v a e — g u s a n o s tornillos (General), gusanos d e las h e r i d a s ( P u e r t o Rico), g u s a n o s b a r r e n a d o r e s (Mexico). Adults—mosca a m e r i c a n a d e larva-tornillo, moscas de la q u e r e s a (Panama). Portuguese: Larvae—vermes torneiros, mororó, coró. A d u l t s — m o s c a s varejeiras, b e r o n h a s (Brazil). Wounds—bicheiras. Tupi-Guaraní: L a r v a e — t a p u r u (Brazil).

BURALLI. 1979. Dispersal and distribution of three newly introduced species of Chrysomya Robineau-Desvoidy in Brazil (Diptera, Calli­ phoridae). Rev. Brasil. Entomol. 23: 245-255. HALL, D. G. 1948. T h e blowflies of North America. Entomol. Soc. Amer., Washington, D.C. JAMES, M. T 1970. Family Calliphoridae. In N. Papavero, ed., A catalogue of the Diptera of the Americas south of the United States. Dept. Zoof, Sec. Agrie, Sao Paulo, no. 102. JIRÓN, L. F. 1979. Sobre moscas califóridas de Costa Rica (Diptera: Cyclorrhapha). Brenesia 16:221-223.

T h i s is o n e of t h e most serious pests of livestock in subtropical a n d tropical Amer­ ica, n a m e d t h e screwworm from t h e wrig­ gling a n d twisting a p p e a r a n c e of the larva (fig. 11.10b) w h e n b u r r o w i n g into flesh. Adults oviposit in a n y o p e n w o u n d , the umbilicus of n e w b o r n s , even m i n o r cuts. T h e eggs hatch into white, slender mag­ gots that feed on healthy tissue within the w o u n d . Infested animals b e c o m e nervous a n d m a k e frantic a t t e m p t s to scratch and

EASTON, A. M., AND K. G. V. SMITH. 1970. T h e

entomology of the cadaver. Med. Sci. Law 10: 208-215. GREENBERG, B., AND M. L. SZYSKA. 1984. Imma­

ture stages and biology of fifteen species of Peruvian Calliphoridae (Diptera). Entomol. Soc. Amer. Ann. 77: 488-517.

Figure 11.10 PARASITIC FLIES, (a) Screwworm (Cochliomyia hominovorax, Calliphoridae). (b) Screwworm, larva, (c) Sheep botfly (Oestris ovis, Oestridae). (d) Common cattle grub (Hypoderma tineatum, Hypodermatidae), larva, (e) Cattle grub, adult, (f) Horse botfly (Gasterophilus intestinalis, Gasterophilidae). (g) Horse botfly, larva.

398

FLIES AND MIDGES

lick t h e afflicted area; t h e majority of untreated cases e n d i n t h e d e a t h of t h e animal, which m a y be any species of d o m e s He animal, a l t h o u g h cattle, goats, s h e e p , and hogs a r e t h e usual hosts; h u m a n s may also suffer infections with d i r e results (Aubertin a n d B u x t o n 1934, Mazza a n d Jórg 1939). Xhe adult s c r e w w o r m fly is m e d i u m sized (BL 8 - 1 0 m m ) a n d is generally metallic blue t o bluish-green (fig. 11.10a). There a r e t h r e e indistinct d a r k longitudi­ nal bands o n t h e back of t h e t h o r a x , a n d the head is c o n t r a s t i n g r e d d i s h - o r a n g e . Por many years, it was confused with t h e secondary s c r e w w o r m (C. macellaria) b u t found ultimately to be distinct (Laake et al. 1936). T h e species occurs widely in the A m e r i ­ cas and r e m a i n s a major p r o b l e m in spite of the availability of good control tech­ niques. I t has b e e n e r a d i c a t e d for the time being from t h e s o u t h e r n parts of t h e United States t h r o u g h t h e use of the sterile male p r o g r a m . Entomologists first estab­ lished a screwworm b a r r i e r zone along the 2,000-mile b o r d e r between t h e United States a n d Mexico to r e d u c e the c h a n c e of migration n o r t h w a r d ; in 1972, an agree­ ment was signed between the two countries to move t h e b a r r i e r zone to t h e narrowest part of Mexico, t h e T e h u a n t e p e c I s t h m u s (Spencer e t a l . 1981). A t t e m p t s w e r e u n d e r ­ taken to apply this t e c h n i q u e in Mexico (Brenner 1984) a n d C e n t r a l America (Snow et al. 1985) with limited success. As yet, no serious efforts have been m a d e to apply this w i d e - r a n g i n g s u p p r e s s i o n mea­ sure in o t h e r p a r t s of Latin America, partly because of its rising ineffectiveness in parts of the United States a n d Mexico, probably due to the genetic diversity of wild p o p u l a ­ tions (Richardson et al. 1982). However, tome feel that it is feasible to d e v e l o p plans for eradication at least t h r o u g h C e n t r a l America (Snow et al. 1985). It is of histori­ cal interest to note, however, that t h e first Complete eradication of t h e species was

accomplished experimentally o n the island of C u r a c a o in 1954 ( B u s h l a n d et al. 1955), yet t h e fly d e m o n s t r a t e d its tenacity by reestablishing itself o n t h e island in 1975 (Snow e t a l . 1978). Victims of screwworms a r e t r e a t e d by various folk r e m e d i e s , usually consisting of poultices m a d e from t h e leaves of medici­ nal plants. I n Brazil, magical m e a n s involv­ ing p r a y e r s a n d incantations believed to force t h e exit of t h e larvae from w o u n d s have also b e e n applied widely. O n e such is p e r f o r m e d by a n exorcist, w h o first ties t o g e t h e r t h e e n d s of a flexible stick t o m a k e a circle a b o u t t h e d i a m e t e r of t h e w o u n d . H o l d i n g this c h a r m over t h e af­ flicted p a r t of t h e animal, h e u t t e r s t h e supplication: "Foge doenca, De bicho m a u , Da santa presenca, De Sao Nicolau, V e r m e d a terra, Na t e r r a d u r a , Sao Nicolau fez tua s i p u r t u r a " (Go away sickness of t h e b a d g r u b , from the p r e s e n c e of Saint Nicholas. G r u b of t h e e a r t h in t h e e a r t h will Saint Nicholas m a k e your grave) (Lenko a n d P a p a v e r o 1979: 423). Two figures in t h e Mayan T i r o - C o r t e sianus C o d e x a p p e a r to depict larvae of these flies attacking gods. I n each case, o n e of the maggots is p r o x i m a t e to t h e nose, a c o m m o n focus of screwworms. T h e hosts may have been victims of h u m a n sacrifice (Tozzer a n d Allen 1910).

References AUBERTIN, D., AND P. A. BUXTON. 1934. Coch­

liomyia and myiasis in tropical America. Ann. Trop. Med. Parasit. 28: 245-254. BRENNER, R. J. 1984. Dispersal, mating, and oviposition oí the screwworm (Diptera: Calli­ phoridae) in southern Mexico. Entomol. Soc. Amer. Ann. 77: 779-788. BUSHLAND, R. C , A. W. LINDQUIST, AND E. E

KNIPLING. 1955. Eradication of screw-worms through release of sterilized males. Science 122:287-288. LAAKE, E. W., E. C. CUSHING, AND H. E. PARISH.

1936. Biology of the primary screw worm fly, Cochliomyia americana, and a comparison of its stages with those of C. macellaria. U.S. Dept. Agrie. Tech. Bull. 500: 1-24.

CARRION FLIES

399

LENKO, K., AND N. PAPAVERO. 1979. lnsetos no

folcJore. Sec. Cult. Cien. Tech., Sao Paulo. MAZZA, S., AND M. E. J Ó R G . 1939. Cochliomyia

hominivorax americana C. y P., estudio de sus larvas y consideraciones sobre miasis. Univ. Buenos Aires Mis. Estud. Pat. Reg. Argentina Publ. 41: 3-46. RICHARDSON, R. H., J. R. ELLISON, AND W. W.

AVERHOFF. 1982. Autocidal control of screwworms in North America. Science 215: 361 — 370. SNOW, J. W., J. R. COPPEDGE, AND A. H. BAUM-

HOVER. 1978. T h e screwworm Cochliomyia hominivorax (Díptera: Calliphoridae) reinfests the island of Curacao, Netherlands Antilles. J. Med. Entomol. 14: 592-593. SNOW, J. W., C. J. WHITTEN, A. SALINAS, J. FERRER, AND W. H. SUDLOW. 1985. T h e

screwworm, Cochliomyia hominivorax (Díptera: Calliphoridae), in Central America and pro­ posed plans for its eradication south to the Darien Gap in Panama. J. Med. Entomol. 22: 353-360. SPENCER, J. P., J. W. SNOW, J. R. COPPEDGE, AND

C. J. WHITTEN. 1981. Seasonal occurrence of the primary and secondary screwworm (Diptera: Calliphoridae) in the Pacific coastal area of Chiapas, Mexico, during 1978-1979. J. Med. Entomol. 18: 240-243. TOZZER, A. M., AND G. M. ALLEN. 1910. Animal

figures in the Mayan codices. Harvard Univ., Peabody Mus. Pap. Araer. Archaeol. Ethnol. 4: 273-372.

BOTFLIES Cuterebridae, Gasterophilidae, Hypodermatidae, a n d Oestridae. Warble flies, b r e e z e flies, clegs, heel flies, gadflies (adults), g r u b s (larvae). Botflies a n d warble flies all a r e s o m e w h a t large (BL 1 2 - 2 0 m m ) , r o b u s t , a n d usually hairy, m u s c o i d flies, which d o n o t feed, possessing only vestigial m o u t h p a r t s . T h e e n d o p a r a s i t i c larvae a r e likewise large, thick s k i n n e d a n d heavy bodied, always with r e a r w a r d d i r e c t e d spines o r denticles projecting from t h e cuticle, which p r e v e n t t h e m from b e i n g easily dislodged from t h e host. T h e p u p a l stage is passed in a very h a r d , heavily p i g m e n t e d p u p a r i u m t h a t

400

FLIES AND MIDGES

develops o n the g r o u n d , in surface debris o r shallowly b u r i e d in t h e soil. T h e r e a r e various types, t h e larvae of a | | living obligatorily in m a m m a l s (Guimaráes et al. 1983). Some types a r e n o t native to the New World, having been introduced with infected livestock into a few areas from elsewhere. T h e s e i n c l u d e t h e sheen warbles (Oestrus ovis; fig. 11.10c), which infest t h e sinuses a n d o t h e r cranial tissues of s h e e p (estro del borrego, mosca de la nariz in Peru) (Rogers a n d K n a p p 1973) a n d occa­ sionally e n t e r t h e conjunctiva of t h e hu­ m a n eye as first instars (Adas et al. I960) a n d two species of cattle g r u b s . T h e latter Hypoderma lineatum (fig. l l . l O d ) a n d / / bovis, pass p a r t of t h e i r life in t h e internal o r g a n s of cattle a n d o x e n b u t later lodge in o p e n boils b e n e a t h t h e skin o n t h e back from which they e m e r g e later to pupate, d o i n g d a m a g e to t h e h i d e . T h e adults resemble b u m b l e b e e s (fig. 1 l.lOe) and are m u c h feared by livestock. T h e species have b e c o m e pests in certain cattle areas of Latin America b u t a r e not well established. T h e p r e c e d i n g species all belong to the families H y p o d e r m a t i d a e a n d Oestridae. T h e family G a s t e r o p h i l i d a e contains three similar parasites in the g e n u s Gasterophilus, G. intestinalis, G. haemorrhoidalis, a n d G. nasalis, all occupying t h e intestinal tract of horses. T h e larvae (fig. 11.1 Og) most com­ m o n l y attach to t h e internal stomach wall a n d cause considerable irritation to the host. Like t h e f o r e g o i n g species, horse bots (fig. 11.1 Of) a r e of local o c c u r r e n c e in the c o u n t r i e s of t h e New World tropics, only w h e r e they h a v e been carried in with i m p o r t e d animals. A m o n g t h e bots, only t h e family Cutere­ b r i d a e , the r o d e n t botflies, is indigenous to the Western H e m i s p h e r e . T h e larvae of most of the eighty-three species ol Cuterebra a n d five related g e n e r a parasitize a wide variety of r o d e n t s (Sciurus, Thomomys, Neotoma, Oryzomys, Mus, etc.) a n d rabbits (Lepus, Sylvilagus) (Guimaráes 1971) as well as marsupials a n d carnivores (Séguy 1948).

T h e life cycle of most species is similar. The female oviposits in e n v i r o n s fre­ quented by t h e host, a n d t h e eggs hatch in response to t h e h e a t a n d m o v e m e n t of its uody nearby. T h e newly h a t c h e d larvae a r e w e t a n d sticky from e g g fluids a n d readily adhere to t h e host as it contacts t h e m . Entry to the host's b o d y is via moist b o d y o p e n i n g s (mouth, nostrils, eyes, etc.) o r skin lacera­ tions. Following entry, t h e larvae m i g r a t e internally to s u b c u t a n e o u s sites w h e r e they settle, molt, a n d form feeding pockets with external b r e a t h i n g p o r e s . M a t u r e larvae drop from t h e p o r e a n d b u r r o w into sur­ face soil o r d e b r i s for p u p a t i o n (Catts 1982). Alouattamyia contains a single species, the monkey botfly, d e v e l o p i n g s u b c u t a n e ously primarily in t h e t h r o a t s of howler monkeys (Zeledón et al. 1957). Little is known of the biology of the four r e m a i n ­ ing genera, save Dermatobia, which con­ tains a single, most c u r i o u s species, b e ­ cause of its u n i q u e m o d e of oviposition and frequent u s e of h u m a n s as hosts (see below).

References ATLAS, A., R. DONCKASTER, H. SCHENONE, AND

M. OLIVARES. 1960. Myiasis ocular producida por larvas de Oestrus ovis. Bol. Chilena Parasit. 15: 37-38. TTS, E. P. 1982. Biology of New World bot flies: Cuterebridae. Ann. Rev. Entomol. 27: 513-338. "IMARÁES, J. H. 1971. Notes on the hosts of Neotropical Cuterebrini (Díptera, Cuterebri­ dae), with new records from Brazil. Univ. Sao '' Paulo, Mus. Zool., Pap. Avul. Zool. 25: 89-94. ""IMARAES, J. H., N. PAPAVERO, AND A. P. DO

i PRADO. 1983. As miíases na regiáo Neotropi­ cal (Identificafáo, biología, bibliografía). Rev. Brasil. Zool. 1: 239-416. IERS, C. E., AND F. W. KNAPP. 1973.

Bionomics of the sheep botfly,Oestris ovis. Environ. Entomol. 2: 11-23. -UY, E. 1948. Introduction á l'étude des myiases. Rev. Brasil. Biol. 8: 9 3 - 1 1 1 . SDÓN, R., O. JIMÉNEZ, AND R. R. BRENES.

1957. Cuterebra baeri Shannon y Greene, 1926 en el mono aullador de Costa Rica. Rev. Biol. TVop. 5: 129-134.

Human Botfly C u t e r e b r i d a e , Dermatobia hominis. Spanish: T ó r s a l o , torcel (Central America); tornillo (Peru, A r g e n t i n a ) ; g u s a n o d e m o n t e , n u c h e (central a n d n o r t h e r n South America); b a r r o (Bolivia); g u s a n o p e l u d o (Colombia). Portuguese: B e r n e (Brazil). Náhuatl: Colmoyotl, moyocuil (Mexico, G u a t e m a l a ) . TupiGuaraní: Ura. Kaingang Indian: B i k u r u . T h e m e t h o d used by t h e female of this botfly to infect its host is o n e of t h e most devious a n d a m a z i n g e m p l o y e d by verte­ b r a t e endoparasites (Catts 1982). T h e fe­ male does n o t lay its eggs directly o n its wary hosts, large m a m m a l s (cattle, horses, dogs, pigs, tapirs, deer, etc.) a n d s o m e birds, b u t c o m m a n d e e r s o t h e r insects to carry them, always choosing a bloodsuck­ ing o r zoophilous type (mosquito—viróle zancudo—deerfly, stable fly) that will surely be attracted to a n animal in search of a meal. It attaches its eggs to t h e body of t h e vector, which t h e n t r a n s p o r t s t h e m to t h e host's skin. Stimulated by b o d y h e a t a n d agitation, t h e completely m a t u r e larva with­ in t h e e g g capsule hatches instantly w h e n the vector touches t h e animal a n d i m m e d i ­ ately b u r r o w s into its skin. It is interesting that the A m a z o n i a n I n d i a n a p p a r e n t l y h a d knowledge of this u n i q u e biological p h e ­ n o m e n o n , as evidenced by t h e existence of a w o r d in t h e T u p i - G u a r a n í l a n g u a g e for c r a n e flies, t h o u g h t t o b e carriers, campana ura. T h e use of t h e w o r d rnoyotl (mosquito) in combinations of w o r d s for this insect by the Aztecs suggests a similiar k n o w l e d g e by these people (orig. obs.). After feeding a week a n d a half subcutaneously, t h e larvae m a k e a b r e a t h i n g a p e r ­ t u r e to t h e outside to a c c o m m o d a t e their increased oxygen n e e d s , having molted t o t h e second stage. First-stage larvae a r e small (BL 6 m m ) , fusiform, a n d with n u m e r o u s fine to coarse surface denticles. T h e second stage is l a r g e r (BL 10—15 m m ) a n d strangely s h a p e d with a s p h e r i -

BOTFLIES

401

Figure 11.11 PARASITIC FLIES, (a) Human botfly (Dermatobia hominis, Cuterebridae), larva (b) Human botfly, adult, (c) Bat tick fly (Basilia ferrisi, Nycteribiidae). (d) Bat fly (Trichobius dugesii Streblidae). (e) Louse fly {Olfersia fassulata, Hippoboscidae).

dary forest. I n p a r t s of its r a n g e , cattle a r e severely affected, a n d infestations account for significant losses in meat, milk, a n d hide value. Insecticides a r e used against larvae in livestock, b u t possible control through t h e use of chemosterilants is u n ­ der study. A bibliography on the species, c o m p l e t e to 1966, was p u b l i s h e d by G u i m a r á e s a n d Papa vero (1966).

References cal a n t e r i o r p o r t i o n with coarse denticles a n d elongate, s m o o t h taillike extension. T h e t h i r d instar (fig. 11.11a) is large (BL to 2 cm) a n d grublike, with a slightly narrow, u n a r m e d p o s t e r i o r section a n d n u m e r o u s small r e a r w a r d - p o i n t i n g denti­ cles a r r a n g e d in t r a n s v e r s e rows o n t h e a n t e r i o r s e g m e n t s . All stages r e m a i n in a feeding pocket b e n e a t h t h e b r e a t h i n g a p ­ e r t u r e ; o n fully m a t u r i n g , they squeeze out of the pocket a n d d r o p to the g r o u n d to p u p a t e . T h e whole d e v e l o p m e n t a l pe­ riod, from e g g to adult, r e q u i r e s a little over two m o n t h s ( J o b s e n a n d M o u r i e r 1972). T h i s life cycle may take place in h u m a n s (mirunta, Peru) as well as domestic a n d wild m a m m a l s , a n d the incidence of infestation in some parts of Latin America, particu­ larly in cattle areas, is irritatingly high a n d of l o n g s t a n d i n g ( B l a n c h a r d 1892, T h o m a s 1988). T h e larvae a r e s e l d o m able to com­ plete their d e v e l o p m e n t , however, because they a r e not l o n g tolerated a n d a r e re­ m o v e d . T h e y frequently twist o n their axis, causing pain, a n d occasionally cut t h r o u g h blood vessels while feeding, releasing copi­ ous flows of blood. Removal of larvae is accomplished in m a n y ways by the locals, d e p e n d i n g o n tradition. Because they a r e virtually impossi­ ble to r e m o v e w h e n healthy a n d vigorous, the m a g g o t s ' body b e i n g covered with re­ c u r v e d denticles that p r e v e n t its with­ drawal, they a r e i n d u c e d to relax their g r i p in the feeding pocket by an o v e r n i g h t

402

FLIES AND MIDGES

application of a kind of poultice, often a piece of raw bacon a n d / o r tobacco (ampiri Peru). T h i s also suffocates the larvae, which in a limp state can t h e n be extracted with forceps or squeezed out. T h e m o d e r n ver­ sion of this trick is the application of a tight covering of t r a n s p a r e n t adhesive tape. Usu­ ally, the larvae occur singly, but multiple infections of two, t h r e e , or m o r e at a single site a r e not u n c o m m o n . A l t h o u g h h a r b o r i n g larvae is very pain­ ful a n d u n p l e a s a n t , the w o u n d bleeding often a n d oozing noxious liquids, at least a few intrepid entomologists have allowed t h e m to c o m p l e t e d e v e l o p m e n t in their own bodies a n d r e c o r d e d the e x p e r i e n c e (Dunn 1930, Busck 1913). Usual sites of infection are the f o r e a r m s , s h o u l d e r s , a n d scalp. In infants, the last site is potentially dangerous because larvae have b e e n k n o w n to bore t h r o u g h the soft fontanel a n d lodge in the brain, causing d e a t h (Rossi a n d Zucoloto 1973). Adults of both sexes a r e heavy bodied a n d large (BL 1 2 - 1 5 m m ; fig. 11.11b). (For a detailed study of the male genitalia, see Leite 1990.) T h e h e a d is mainly yellow, the t h o r a x d a r k bluish-grey, a n d the abdo­ m e n a brilliant, shiny d a r k blue. They are rarely seen in n a t u r e but a r e occasionally observed d u r i n g the daytime in the forest n e a r stagnant water w h e r e their most com­ m o n mosquito egg vectors a r e emerging. Dermatobia hominis has an equatorial dis­ tribution from 25 deg. N to 18 deg. S, in which it favors moist tropical, hilly, secon-

BLANCHARD, R. 1892. Sur les Oestrides amé-

ricains dont la larve vit dans la peau de I'homme. Soc. Entomol. France Ann. 61: 109-154. BUSCK, A. 1913. On the rearing of a Dermatobia hominis Linnaeus. Entomol. Soc. Wash. Proc. 14:9-11. CATTS, E. P. 1982. Biology of New World bot flies: Cuterebridae. Ann. Rev. Entomol. 27: 313-338. DUNN, L. H. 1930. Rearing the larvae of Derma­ tobia hominis Linn., in man. Psyche 37: 3 2 7 342. GUIMARÁES, J. H.,

AND N. PAPAVERO. 1966.

A

tentative annotated bibliography of Derma­ tobia hominis (Linnaeus 1781) (Diptera, Cutere­ bridae). Arq. Zool. (Sao Paulo) 14: 223-294. JOBSEN, J.

A.,

AND H.

MOURIER.

1972.

The

morphology of the larval instars and pupa of Dermatobia hominis L. Jr. (Diptera: Cuterebri­ dae). Entomol. Berichi. 32: 218-224. LEITE, A. C. R. 1990. Scanning electron micros­ copy of male genitalia of Dermatobia hominis (Diptera: Cuterebridae). J. Med. Entomol. 27: 706-708. ROSSI, M.

A.,

AND S. ZUCOLOTO. 1973.

Fatal

cerebral myiasis caused by the tropical warble fly, Dermatobia hominis. Amer. J. Trop. Med. Hyg. 22: 267-269. THOMAS, JR., D. B. 1988. The pattern of Derma­ tobia (Diptera: Cuterebridae) myiasis in cattle in tropical Mexico. J. Med. Entomol. 25: 1 3 1 135.

ECTOPARASn IC FLIES Streblidae, bat flies. Nycteribiidae, bat tick flies. H i p p o b o s c i d a e , louse flies. Three families of D i p t e r a have evolved a Parasitic way of life a m o n g the hairs of bats

a n d feathers of birds. T h e i r a d a p t a t i o n s include flattening of the body, develop­ m e n t of combs of flat bristles for scuttling a m o n g the host's pelage, a n d e n l a r g e d tarsal claws for grasping. T h e blind, straw-colored or yellowish bat flies (Streblidae, fig. 11.1 Id) a n d bat tick flies (Nycteribiidae, fig. 11.1 le) ( G u i m a r á e s a n d d ' A n d r e t t a 1956) a r e small (BL 2 - 4 m m ) a n d live p e r m a n e n t l y a n d solely on bats. T h e s e d i p t e r a n s a r e well studied only taxonomically a n d from few areas of Latin America, primarily P a n a m a ( G u i m a r á e s 1966, Wenzel et al. 1966) a n d Venezuela (Wenzel 1976), as a result of special surveys t h e r e . A bibliography to 1971 is p r o v i d e d by M a a ( 1 9 7 1 ) . T h e streblids (Wenzel 1976), with 94 neotropical species in 23 g e n e r a , a r e cylin­ drical, have a m o r e or less n o r m a l a p p e a r i n g h e a d a n d relatively short legs, a n d are usually winged (some have r e ­ d u c e d or n o wings); nycteribiids a r e flat a n d spiderlike, with long legs, without wings, a n d the h e a d is uniquely folded back into a groove o n t h e back of t h e t h o r a x . T h e r e are 37 species in two g e n e r a (almost all in Basilia, o n vespertilionid bats). Louse flies ( H i p p o b o s c i d a e , fig. 11.1 le) a r e larger (BL 3—11 m m ) a n d h a v e welldeveloped eyes (Bequaert 1953—1957, Maa 1963). T h e i r i n t e g u m e n t is d a r k a n d leath­ ery, they a r e usually completely winged, a n d they lack bristle combs; the tarsal claws a r e hooked a n d heavy for g r a s p i n g . Most species live on birds; the d a r k - c o l o r e d adults are often seen r u n n i n g conspicu­ ously over the white p l u m a g e of seabirds ( i m p o r t a n t pests, moscas de gallinazos, of g u a n o birds in Peru b e l o n g to the g e n u s Olfersia; Dale 1969). Altogether, t h e r e a r e 43 regional species (Maa 1969). T h e r e a r e also many kinds infesting m a m m a l s such as d e e r (Lipoptena mazamae) a n d domestic q u a d r u p e d s (e.g., the h o r s e louse fly, Hippobosca equina, a n d s h e e p k e d , Melophagus ovinas; they have never b e e n f o u n d

ECTOPARASIT1C FLIES

403

on bats. T h e last two are cosmopolitan pests, s p r e a d by m a n with their hosts, as is the u b i q u i t o u s pigeon louse fly {Pseudolynchia canariensis). In all hippoboscids, a single larva at a time is r e t a i n e d within t h e female's abdo­ m e n w h e r e it is n o u r i s h e d to m a t u r i t y by special "milk" glands. It is t h e n e x t r u d e d a n d falls to the g r o u n d w h e r e it p u p a t e s . T h e blood-feeding habits of these Díp­ tera would m a k e t h e m potential s p r e a d e r s of disease, b u t they so far have n o t b e e n i n c r i m i n a t e d as p r i m a r y vectors for any p a t h o g e n s save s o m e b i r d blood p r o t o ­ zoans t r a n s m i t t e d by h i p p o b o s c i d s .

References BEQUAERT, J. 1953-1957. The Hippoboscidae or louse-flies (Diptera) of mammals and birds. Pt. I. Structure, physiology and natu­ ral history. Entomol. Americana (n.s.) 33: 1 — 209, 211-442, figs. 1-21; 34: 1-232, figs. 22-44; 35: 233-416, figs. 4 5 - 8 2 ; 36: 4 1 7 611, figs. 83-104.

404

FLIES AND MIDGES

DALE, W. E. A. 1969. Hippoboscidae (Díptera) del Peru. I. Nuevas identificaciones: R e . gistros hallados en la literatura Peruana Biota 8: 41-52. GUIMARAES, L. R. 1966. Nycteribiid batflies from Panama. In R. L. Wenzel and V. I. Tipton, eds., Ectoparasites of Panama. Field Mus. Nat. Hist., Chicago. Pp. 393-404.

12

SAWFLIES, WASPS, ANTS, AND BEES

GUIMARAES, L. R., AND M. A. V. D'ANDRETTA.

1956. Sinopse dos Nycteribiidae (Diptera) do Novo Mundo. Arq. Zool. (Sao Paulo) 9: 1184. MAA, T. C. 1963. Genera and species of Hippo­ boscidae (Diptera): Types, synonymy, habitats and natural groupings. Pacific Ins. Monogr. 6: 1-186. MAA, T. C. 1969. A revised checklist and concise host index of Hippoboscidae (Diptera). Pa­ cific Ins. Monogr. 20: 262-299. MAA, T. C. 1971. An annotated bibliography of batflies. Pacific Ins. Monogr. 28: 119-211. WENZEL, R. L. 1976. The streblid batflies of Venezuela (Diptera: Streblidae). Brigham Young Univ. Sci. Bull. (Biol Ser.) 10(4): 1-177. WENZEL, R. L., V. ). TIPTON, AND A. KIF.WLICZ.

1966. The streblid batflies of Panama (Dip­ tera: Streblidae). In. R. L. Wenzel and V. J. Tipton, eds., Ectoparasites of Panama. Field Mus. Nat. Hist., Chicago. Pp. 405-675.

Hymenoptera. Although t h e primitive sawflies a r e freeliving, p h y t o p h a g o u s , a n d u n a r m e d , out­ standing characteristics of this familiar or­ der are their social or subsocial living habits, p r e d a c e o u s n e s s , a n d ability to sting. A m o n g the H y m e n o p t e r a , all the ants and t h o u s a n d s of species of bees a n d wasps (Snelling 1981) exhibit some level of so­ ciality, from simple m a t e r n a l care to the complex c o m m u n i t y organization of the honeybee hive. A progression of stages exists, r e c a p i t u l a t i n g the evolution of so­ ciality in this o r d e r (see social insects, c h a p . 2). T h e first stage is simple mass provi­ sioning for larvae in a n a t u r a l b u r r o w such as found in certain spider wasps, followed by forms, such as the sand wasps, that construct nests that the p a r e n t provisions continually to e n s u r e a supply of fresh food for the d e v e l o p i n g larvae. A f u r t h e r step beyond this is increased longevity of the female p a r e n t a n d a t e n d e n c y for offspring to r e m a i n with h e r a n d assist in the care of s u b s e q u e n t b r o o d s , the p a t t e r n in polistine p a p e r wasps. T h e final evolu­ tionary step is a division of labor a n d a correlated f o r m a t i o n of different body types, or "castes," a n d e n o r m o u s n u m b e r s of colony m e m b e r s living in elaborate nests such as the polybiine p a p e r wasps, leaf cutter ants, a n d social bees. Nest building and larval care a r e major c o n c e r n s of the females; the males' adult lives a r e mainly spent in p u r s u i t of females, often in com­ plex ways (Alcock et al. 1978). To u n d e r ­

stand h y m e n o p t e r a n social s t r u c t u r e , it is i m p o r t a n t to recall that the m e m b e r s of the colony are all closely related family m e m ­ bers. All the so-called workers or sterile females a r e d a u g h t e r s of a single f o u n d i n g mother. ( T h e latter is the n o r m a l case a n d is r e f e r r e d to as monogyny. In some spe­ cies a n d g r o u p s , t h e r e are multiple q u e e n s at the head of a colony, r e f e r r e d to as polygyny, but this is unusual.) T h e q u e e n (or queens) mates a n d carries the s p e r m in storage pouches off the r e p r o d u c t i v e tract. T h e fathers of colonies die soon after inseminating the q u e e n . T h e m a n y kinds of H y m e n o p t e r a capa­ ble of stinging d o so with a specially modified ovipositor, the egg-laying device of the female, which is located in the tip of the a b d o m e n ( H e r m a n n a n d Blum 1981). Males, lacking this o r g a n , are incapable of stinging. A sting is an effective way to discourage or p u n i s h e n e m i e s but also is used to s u b d u e prey. T h e pain, swelling, a n d o t h e r adverse s y m p t o m s caused in h u m a n s is s h a r e d by o t h e r v e r t e b r a t e ani­ mals, a l t h o u g h m a n y have a n a t u r a l i m m u ­ nity to stings. T h e s e h a r m f u l effects a r e d u e to proteins a n d enzymes in the v e n o m which are injected into the w o u n d a n d are foreign to the physiology of the recipient. Direct toxicity a n d allergic reactions result, a n d either may be sufficiently severe to cause d e a t h in some cases. S o m e birds take a d v a n t a g e of the aggressiveness a n d sting­ ing habits of ants, bees, a n d wasps by situating their own nests in close proximity (Myers 1935, Smith 1968).

405

T h e r e a r e s o m e 105,000 k n o w n species of H y m e n o p t e r a in t h e world, a large p e r c e n t a g e of these in Latin America (Willink 1982). Previously u n k n o w n to sci­ ence, m a n y a r e being discovered every year, especially a m o n g t h e small, parasitic g r o u p s . Because of t h e diversity of t h e order, it c a n n o t b e easily characterized structurally. I n g e n e r a l , these insects have two pairs of m e m b r a n o u s wings, t h e fore larger t h a n t h e h i n d , with m o d e r a t e l y c o m p l e x venation, a l t h o u g h m a n y a r e wingless (e.g., w o r k e r ants), a n d o t h e r s exhibit simplified v e n a t i o n . T h e m o u t h parts a r e a d a p t e d basically for biting, b u t bees h a v e e l o n g a t e d maxillae a n d labia for feeding o n liquids. T h e r e a r e two s u b o r d e r s , t h e Symphyta (Chalastrogastra) a n d A p o c r i t a (Clistogastra, Petiolata). T h e f o r m e r is defined by the b r o a d a t t a c h m e n t of t h e a b d o m e n to the t h o r a x , t w o - s e g m e n t e d t r o c h a n t e r s in the legs, at least t h r e e closed cells in t h e h i n d wing venation, a n d p h y t o p h a g o u s larval habits. M e m b e r s of t h e g r o u p also possess a well-developed c u t t i n g ovipositor that is s o m e w h a t sawlike in s o m e families ("sawflies") o r e l o n g a t e for drilling into wood in o t h e r s ("horntails"). T h e larvae of those f e e d i n g externally o n plants a r e often caterpillarlike, with walking legs a n d often cryptic colors; t h e wood borers a r e wormlike, e l o n g a t e , pale, a n d legless. O t h ­ ers form galls o r b o r e in wood a n d a r e grublike. T h e O r u s s i d a e a r e parasitic o n w o o d - b o r i n g insects. I n t h e Apocrita, t h e basal s e g m e n t of the a b d o m e n is fused t o t h e t h o r a x , a n d the next s e g m e n t forms a stalk o r n a r r o w waist. T h e t r o c h a n t e r s a r e o n e o r two s e g m e n t e d , a n d t h e h i n d wing venation never h a s m o r e t h a n two closed cells. T h e grublike larvae of most a r e either internal or e x t e r n a l parasitoids o f o t h e r a r t h r o p o d s (Parasitica) o r a r e h o u s e d in nests a n d fed a diet of processed p l a n t o r animal tissue, honey, o r pollen by t h e adults (Aculeata).

406

SAWFLIES, WASPS, ANTS, AND BEES

Some specialized types a r e leaf o r fruit m i n e r s o r gall m a k e r s ( A g a o n i d a e , CynipJ. dae, etc.). H y m e n o p t e r a a r e t r e m e n d o u s l y impor­ tant a n d interesting ecologically (Rau 1933), bees especially, for t h e ecological roles they play as pollinators of flowering plants, many of which a r e h u m a n food staples. I n fact, t h e d e p e n d e n c e of certain plants o n these insects is so g r e a t that they possess highly specialized flower structures a n d colors to e n s u r e successful fertilization by specific h y m e n o p t e r a n s . A l t h o u g h a few p h y t o p h a g o u s species a r e pests of tropical crops, t h e large n u m b e r of "para­ sites" (actually parasitoids) far offset their injurious effects by k e e p i n g o t h e r noxious insects at bay. T h e s e insects a r e familiar to everyone a n d have a particular cultural importance to m a n y A m e r i n d g r o u p s . Por instance, it is believed by t h e Kayapó I n d i a n s of Brazil that they l e a r n e d h o w to live as social beings from a n ancestral wise m a n (wayanga) who gained this knowledge from t h e study of ant, bee, a n d wasp behavior (Posey 1982).

References ALCOCK, J., E. M. BURROWS, G. GORDH, L. J. HUBBARD, L . KlRKENDALL, D . W . P Y L E , T . L. PONDER, AND F. G. ZALOM. 1978. The ecology

Hymenoptera. In H. R. Hermann, ed.. Social insects. 2: 369-453. Academic, New York. WILLINK, A. 1982. Himenópteros Neotropicales, su origen, ecología, comportamiento y distribución. In P. J. Salinas, ed., Zoología Neotropical. 1: 71-90. 8th Cong. Latino­ americano Zool. (Mérida) Actas.

SAWFLIES AND HORNTAILS Symphyta. These two g r o u p s a r e very poorly studied in Latin A m e r i c a . T h e y a r e less c o m m o n l y collected in t h e region t h a n in n o r t h e r n latitudes a n d a r e n o t found in n u m b e r s as they a r e in subarctic a n d t e m p e r a t e N o r t h America (Smith 1988). Most Neotropical sawflies belong t o t h e families P e r g i d a e , A r g i d a e , a n d T e n t h r e d i nidae, whose larvae feed o n a large variety of plants, i n c l u d i n g ferns. Hosts include forest a n d o r n a m e n t a l trees, s h r u b s , a n d agricultural c r o p s . M e m b e r s of t h e native genus Acordulecera (Pergidae) a r e pests of potatoes in Peru a n d Bolivia (Smith 1980). Among t h e t e n t h r e d i n i d pests a r e m a n y species of t h e g e n u s Waldheimia (fig. 12.1a); Syzygonia cyanocephala attacks t h e Q u a r e s meira {Tibouchina, Melanostomaceae) in Brazil (Marques 1933).

the coniferous forests into C e n t r a l A m e r ­ ica a n d a r e mostly in t h e g e n e r a Sirex a n d Urocerus (fig. 12.1b). O n e palearctic h o r n tail, Urocerus gigas gigas, has been i n t r o ­ d u c e d into Chile a n d h a s b e c o m e estab­ lished t h e r e (Smith 1988). A n o t h e r indige­ n o u s species in t h e same g e n u s (U. patagonicus) is k n o w n as a Paleocene fossil from A r g e n t i n a (Fidalgo a n d Smith 1987). In Mexico, horntail larvae m a y a s s u m e i m p o r t a n c e as pests w h e n b u r r o w i n g in pine a n d fir wood used for building con­ struction o r f u r n i t u r e . Adults of both g r o u p s a r e small to large, wasplike insects with well-veined wings a n d m a n y - s e g m e n t e d a n t e n n a e . T h e latter a r e sometimes highly modified, with t e r m i n a l clubs o r p l u m o s e b r a n c h e s .

References FIDALGO, P., AND D. R. SMITH. 1987. A fossil

Siricidae (Hymenoptera) from Argentina. Entomol. News 98: 63-66. MARQUES, L. A. 1933. Tenthredinidae conhecida por "Mosca de Serra," cuja larva ou "falsa lagarta" é novica a varias especies do género Tibouchina. Inst. Biol. Dept. Agrie. Rio de Janeiro 1933: 1-11. SMITH,

D. R.

1980. Identification

of the

Acordulecera "potato" sawflies of Peru and Bolivia, with descriptions of these and related species from South America (Hymenoptera: Pergidae). J. Wash. Acad. Sci. 70: 89-103. SMITH, D. R. 1988. A synopsis of the sawflies (Hymenoptera: Symphyta) of America south of the United States: Introduction, Xyelidae,

HERMANN, H. R., AND M. S. BLUM. 1981. Defen­

Horntails a r e placed in t h e family Siricidae, a primarily n o r t h e r n g r o u p asso­ ciated with conifers a n d a r b o r e a l angiosperms. T h e y occur only as far south as

sive mechanisms in the social Hymenoptera. In H. R. Hermann, ed., Social insects. 2: 77197. Academic, New York. MYERS, J. G. 1935. Nesting associations of birds with social insects. Royal Entomol. Soc. Lon­ don Trans. 83: 11-22. POSEY, D. A. 1982. The importance of bees to the Kayapó Indians of the Brazilian Amazon. Fia. Entomol. 65: 452-458. RAU, P. 1933. The jungle bees and wasps of Barro Colorado Island (with notes on other insects). Publ. by author, Kirkwood, Mo. SMITH, N. G. 1968. T h e advantage of being parasitized. Nature 219: 690-694. SNELLING, R. R. 1981. Systematic^ of social

R a j " * 12.1 WASPS, (a) Sawfly (Waldheimia ochra, Tenthredinidae). (b) Horntail (Urocerus ca■wi/ci/s, Siricidae). (c) Braconid wasp (Apanteles congregatus, Braconidae). (d) Ichneumon wasp [Thyreodon sp., Ichneumonidae).

and evolution oí male reproductive behavior in the bees and wasps. Zool. J. Linnean Soc. 64: 293-326.

SAWFLIES AND HORNTAILS

407

Pamphilidae,Cimbicidae,Diprionidae,Xiphydriidae, Siricidae, Orussidae, Cephidae. Syst. Entomol. 13: 205-261.

PARASITOID WASPS Braconid and Ichneumon Wasps Braconidae and Ichneumonidae.

WASPS H y m e n o p t e r a . Spanish: Avispas. Portuguese: Vespas, m a r i m b o n d o s (larger, stinging types, Brazil), cabas (Brazil). T h e t e r m "wasp" is loosely applied to all winged adult H y m e n o p t e r a , except the bees a n d ants. T h u s , a vast assemblage of diverse, often unrelated, forms are l u m p e d t o g e t h e r into o n e b r o a d category (Evans a n d W e s t - E b e r h a r d 1970, S p r a d bery 1973). Within the wasps, however, it is useful to recognize several basic biologi­ cal types: the "parasitic" (actually parasi­ toid) wasps, gall wasps, solitary wasps, a n d social wasps. Parasitoid wasps a r e c o n s i d e r e d by some a u t h o r s to be less n u m e r o u s in species in the tropics t h a n in the t e m p e r a t e latitudes. Probably, because they t e n d to be niche specific r a t h e r t h a n host specific, this may be t r u e for I c h n e u m o n i d a e ( J a n z e n 1981) but may be only an artifact of collecting for chalicidoids ( H e s p e n h e i d e 1979). Also, ichneumons need a humid environment a n d k e e p mainly to forests; they are com­ paratively r a r e in deserts a n d high m o u n ­ tains, while chalcidoids as a g r o u p a r e m o r e widely tolerant of the e n v i r o n m e n t .

References EVANS, H. E., AND M. J. WEST-EBERHARD.

1970.

The wasps. Univ. Michigan Press, Ann Arbor. HESPENHEIDE, H. A. 1979. Are there fewer parasitoids in the tropics? Amer. Nat. 113: 766-769. JANZEN, D. H. 1981. The peak in North Ameri­ can ichneumonid species richness between 30° and 42°N. Ecology 62: 532-537. SPRADBERY, J. P. 1973. Wasps. Univ. Washington Press, Seattle.

408

SAWFLIES, WASPS, ANTS, AND BEES

A m o n g the Neotropical parasitic Hy­ m e n o p t e r a , two very large families are d o m i n a n t , the braconid wasps (fig. 12.1c) (Matthews 1974) a n d the i c h n e u m o n wasps (fig. 12.Id) (Porter 1980). T h e y are gener­ ally similar, small to medium-sized (BL 3 25 m m ) , somewhat frail-bodied wasps with m a n y - s e g m e n t e d a n t e n n a e (16 or more) a n d with parallel e n t o m o p h a g o u s parasi­ toid habits, utilizing the larvae a n d pupae of almost any h o l o m e t a b o l o u s insect (but especially L e p i d o p t e r a ) as hosts (Gauld 1988). O n e to m a n y larvae develop inside the host, feeding on various tissues and eventually killing the insect. Ichneumons generally d o not spin cocoons outside the host; braconids attach individual white silken cocoons on the host's exterior, and parasitized caterpillars are often seen a d o r n e d with masses of such cocoons cling­ ing to the surface of the skin. Sometimes, the cocoons are located a p a r t from the host either singly or in a mass on vegetation or other substratum. T h e two families are distinguished from o t h e r parasitoid wasps by the ab­ sence of a costal cell in the fore wing venation, the leading m a r g i n of the fore wing being a single, heavy vein. Braconids are usually smaller (BL 14 m m maximum) a n d have only o n e r e c u r r e n t vein in the fore wing (i.e., n o closed cell in the outer, posterior p a r t of the wing); the females also usually have abbreviated abdominal petioles (waists) a n d short ovipositors, al­ t h o u g h in Iphiaulax, the latter is fourteen times the length of the body. Ichneumons a r e all sizes but are often large (BL up to 2 cm or m o r e ) a n d with two recurrent veins in the fore wing (a closed cell is p r e s e n t in the o u t e r posterior part of the wing); the females of m a n y have ex-

tremely long, slender ovipositors a n d long waists c o n n e c t i n g the a b d o m e n to the thorax. T h e ovipositor is inserted directly into the host or used to drill t h r o u g h wood, leaf tissue, cocoons, soil, a n d so on, to oviposit in h i d d e n insects. T h e a b d o ­ men is often m u c h c o m p r e s s e d . An addi­ tional, useful identifying characteristic is the fusion of t h e second a n d third a b d o m i ­ nal tergites in most braconids; nearly all ichneumons h a v e a freely movable articu­ lation b e t w e e n these two s e g m e n t s . Because they a r e parasitoids, these n u ­ merous (just u n d e r 20,000 described Neotropical I c h n e u m o n i d a e a n d p e r h a p s several t h o u s a n d B r a c o n i d a e ; Townes a n d Townes 1966) wasps a r e of g r e a t value in controlling insect p o p u l a t i o n s , including those of m a n y pests, naturally or t h r o u g h human i n t r o d u c t i o n . Hosts consist mainly of caterpillars, b u t a p h i d s , beetle larvae, and o t h e r insects a r e used also. T h e y subdue their p r e y with paralyzing v e n o m s (Beard 1978). A few i c h n e u m o n i d s have a Stinging a p p a r a t u s sufficiently powerful to penetrate h u m a n skin. T h e potency of the Sting of certain Tetragonochora is a p p a r e n t l y sufficient to qualify t h e m as m o d e l s for Batesian mimics in the katydid g e n u s Aganacris, whose n y m p h s r e s e m b l e t h e m remarkably ( d a r k o r a n g e a b d o m e n , white maculae on sides of black t h o r a x , a n d black antenna with m e d i a n white b a n d ) . S o m e large species of i c h n e u m o n i d s in the sub­ family O p h i o n i n a e , for e x a m p l e , Rhynchophion, resemble t a r a n t u l a hawks (Pepsis, Fbmpilidae) a n d also u n d o u b t e d l y e n t e r into mimicry c o m p l e x e s with t h e m .

References BEARD, R. L. 1978. Venoms of Braconidae. In S. Bettini, ed., Arthropod venoms. Springer, Berlin. Pp. 773-800. GAULD, I. D. 1988. Evolutionary pattern oí host Utilization by ichneumonoid parasitoids (Hy. menoptera: Ichneumonidae and Braconi­ dae). Bid. J. Linnean Soc. 35: 351-377.

MATTHEWS, R. W. 1974. Biology oí Braconidae. Ann. Rev. Entomol. 19: 15-32. PORTER, C. 1980. Zoogeografía de las Ichneu­ monidae Latino-Americanas (Hymenoptera). Acta Zool. Lilloana 36: 1-52. TOWNES, H.,

AND M.

TOWNES,

1966.

A cata­

logue and reclassification of the Neotropic Ichneumonidae. Amer. Entomol. Inst. Mem. 8: 1-367.

Chalcidoid Wasps Chalcidoidea. T h i s is an e n o r m o u s assemblage of mostly parasitoid wasps, which occur worldwide a n d occupy almost every habitat. Despite their o m n i p r e s e n c e , they a r e seldom n o ­ ticed because of their small to m i n u t e size. Most have body lengths within the limits of 0.5 to 3 or 4 millimeters; in fact, the smallest insect k n o w n belongs to this g r o u p a n d is only 0.2 millimeters long (Alaptus, Mymaridae). The adults are recognized not only by their m i n u t e n e s s but by the possession of elbowed a n t e n n a e , a p r o n o t u m not e x t e n d ­ ing to the small caps covering the anterior, o u t e r c o r n e r s of the t h o r a x (tegulae), a n d wings almost without venation. Usually only the anterior, heavier wing veins are present, a n d sometimes all traces of n e r v a t u r e are absent. T h e wings a r e clear a n d at rest are held flat over the body. T h e bodies of the majority a r e black or d a r k , a l t h o u g h m a n y a r e d e e p metallic g r e e n or blue; some may have c o n t r a s t i n g yellow or white p a t t e r n s . T h i s category contains some twenty fami­ lies a n d exhibits a wide variety of a d a p t a ­ tions a n d habits. Most are e n d o p a r a s i t o i d s , attacking m o r e host types, in different major taxonomic categories, than any o t h e r g r o u p of parasitoid insects. T h e s p e c t r u m includes spider eggs, ticks, aquatic beetles, ants, aculeate wasps, scale insects, l e p i d o p terous eggs a n d larvae, a n d flies. Because of this characteristic, they a r e a m o n g the most useful biological control agents, a n d m u c h

PARASITOID WASPS

409

of what has b e e n l e a r n e d about chalcidoid biology has c o m e from studies relating to their use in this r e g a r d . T h e wasps' larva develops internally in t h e host, nearly al­ ways killing it. H y p e r p a r a s i t o i d i s m (parasitoidism of a parasitoid), s u p e r p a r a s i t o i d i s m (host attacked by multiple parasitoids), a n d p o l y e m b r y o n y (multiplication of parasitoid's eggs in host) a r e c o m m o n p h e n o m ­ e n a exhibited by different parasitoid spe­ cies. A considerable n u m b e r of chalcidoids are p h y t o p h a g o u s , i n c l u d i n g o n e family, the A g a o n i d a e , that develops in the fruit of Ficus trees a n d has evolved an intimate association with these plants, including an essential role in their pollination (see fig wasps, below). A few types p r o d u c e galls o r live in t h e m as inquilines or guests of the t r u e gall m a k e r s . T h e s e insects a r e diverse (de Santis 1967) a n d difficult to identify, a n d very little is k n o w n of t h e m in the Neotropics. De Santis (1971) r e c o r d e d 1,581 species in 421 g e n e r a in c o n t i n e n t a l S o u t h America. T h i s certainly r e p r e s e n t s only a small frac­ tion of the total fauna, literally t h o u s a n d s of species r e m a i n i n g to be discovered (some of which a p p e a r already in d e Santis's later works [1979, 1981]). S o m e a r e of e c o n o m i c i m p o r t a n c e , ei­ t h e r as pests, for e x a m p l e , the e u r y t o m i d Bruchophagus platyptera, which destroys seeds of alfalfa a n d clover, or as beneficial, biological control a g e n t s . T h e n u m b e r of

the latter currently used within Latin America is only a b o u t thirty, all introduced from elsewhere, such as Tnchogramma minutum (fig. 12.2d) to infect the egg stage of l e p i d o p t e r o u s pests a n d Aphelinus mali (fig. 12.2c), which kills a p h i d s . A great potential probably exists a m o n g the un­ k n o w n forms in these applications. As for the world generally, the largest families in the A m e r i c a n tropics are Trichogrammatidae,Eupelmidae,Pteromalidae, a n d Chalcididae.

References nr. SANTIS, L. 1967. Catálogo de los himenópteros Argentinos de la Serie Parasitica incluyendo Bethyloidea. Prov. Buenos Aires Comisión [nvestig. Cien., La Plata, Publ. Especial. DE SANTIS, L. 1971. La fauna de chalcidoideos de América del Sur. Soc. Entomol. Peru Bol 6: 57-63. DE SANTIS, L. 1979. Catálogo de los himenópteros chalcidoideos de América al sur de los Estados Unidos. Prov. Buenos Aires, Comi­ sión lnvestig. Cien., La Plata, Publ. Especial. Exeludes Argentina and Brazil. DE SANTIS, L. 1981. Catálogo de los himenópteros chalcidoideos de América al sur de los Estados Unidos. Primer supplemento. Rev. Peruana Entomol. 24: 1—38. Fig Wasps Chalcidoidea, A g a o n i d a e . In the New World figs (Ficus), the female flowers are i n t e r m i x e d with the male flow­ ers a n d are scattered over the internal

Figure 12.2 CHALCIDOID WASPS, (a) Fig wasp {Blastophaga dugesi, Agaonidae), female. (b) Fig wasp, male, (c) Parasitic chalcidoid (Aphelinus mali, Eulophidae). (d) Minute Egg Parasite (Tnchogramma minutum, Trichogrammatidae). (e) Gall wasp (Atrusca spinuli, Cynipidae).

410

SAWFLIES, WASPS, ANTS, AND BEES

surface of the swollen, invaginated recepta­ cle, called a s y n c o n i u m or fig fruit. T h e female flowers are of two types, "gall flowers" with s h o r t styles a n d " r e p r o d u c ­ tive flowers" with long styles. T h e a r r a n g e ­ ment provides for a u n i q u e symbiotic rela­ tionship with m a n y species of fig wasps, in the New World mostly with the g e n e r a Blastophaga a n d Tetrapus (Wiebes 1966), which are responsible for pollinating the plant as p a r t of the process of r e a r i n g their young in the s y n c o n i u m . Wasps of the first genus carry t h e pollen on special setose areas (corbiculae, R a m i r e z 1978) on the body a n d legs; those of the second g e n u s , in the digestive tract. Fully w i n g e d female wasps (fig. 12.2a), after e m e r g i n g from ripe figs, fly in search of new, y o u n g figs at the right stage for pollination. T h e y e n t e r the synconium through the a p e r t u r e at its a p e x , forcing their way t h r o u g h the tight o p e n i n g a n d in doing so lose their wings a n d (Blastophaga only) the apical six s e g m e n t s of their a n t e n ­ nae. O n c e inside, the females use their long ovipositors to b o r e d o w n t h r o u g h the styles of female flowers a n d deposit eggs among the i m m a t u r e ovules. As they d o so, they inject a g a l l - p r o d u c i n g substance as well. T h e ovipositor is long e n o u g h , how­ ever, only to r e a c h the ovules of the shortStyled flowers; long-styled flowers are merely p r o b e d . At the same time, both types of flowers a r e fertilized by the fe­ males with pollen from their bodies (Ramirez 1978), but only the long-styled flowers d e v e l o p seeds. T h e short-styled flowers form galls in which the wasp larvae feed and m a t u r e . Wingless m a l e wasps (fig. 12.2b) de­ velop first, in figs that a r e still u n r i p e , thus avoiding b e i n g d e s t r o y e d by fig-eating ani­ mals. T h e y seek o u t m a t u r e female p u p a e and fertilize the females t h e r e i n . T h e n they make t u n n e l s t h r o u g h the wall of the synconium a n d escape. T h e s e events are followed by the e m e r g e n c e of i m p r e g n a t e d females, which, also in leaving the fruit,

crawl over a n d pick u p pollen from the male flowers that have now o p e n e d . T h e females escape the synconium t h r o u g h its n a t u r a l o p e n i n g with the aid of the males, which h e l p bite t h r o u g h any tissues inhibit­ ing their progress. T h e males t h e n die, a n d the females go in search of new trees with receptive fruit to begin a new cycle. Finally, the fig r i p e n s , a n d its seeds are dispersed, usually by h e r b i v o r o u s animals. T h u s , while accomplishing its own r e p r o ­ duction, the wasp e n s u r e s the life of the fig, which sacrifices only a portion of g e r m i n a l tissue as food for the carriers of its pollen. In most Ficus, virtually every s y n c o n i u m of an individual tree is pollinated the same day, or at least over a short p e r i o d of u p to t h r e e days. If the y o u n g figs are not pollinated d u r i n g this p e r i o d , even if wasps enter, they stop growing, shrink, a n d d r o p from the tree. T h e time of develop­ m e n t for each species of wasp is c o r r e l a t e d with the r i p e n i n g time of the fruit, a r o u n d a m o n t h . T h e r e is also considerable speci­ ficity between the species of wasp a n d fig (Ramirez 1970a, Wiebes 1979). Many m o r e details of these relationships a r e available (Ramirez 19706, 1976). O t h e r nonpollinating chalcidoids also inhabit a n d develop in figs, for e x a m p l e , the genus Idarnes (Torymidae) a n d various E u r y t o m i d a e . T h e i r relations to agaonids a n d the figs have not b e e n established ( G o r d h 1975). Blastophaga species of the New World are all placed in the s u b g e n u s Pegoscapus, which are characterized by hav­ ing the pollen-carrying o r g a n s located o n the front coxae.

References GORDH, G. 1975. The comparative external morphology and systematic^ of the Neo­ tropical parasitic fig wasp genus Idarnes (Hymenoptera: Torymidae). Univ. Kansas Sci. Bull. 50: 389-455. RAMÍREZ, W. 1970a. Hosi specificity of fig wasps (Agaonidae). Evolution 24: 680-691. RAMÍREZ, W. 1970/;. Taxonomic and biological

PARASITOID WASPS

411

studies of Neotropical fig wasps (Hymenop­ tera: Agaonidae). Univ. Kansas Sci. Bull. 49: 1-44. RAMÍREZ, W. 1976. Evolution of blastophagy. Brenesia9: 1-13. RAMÍREZ, W. 1978. Evolution of mechanisms to carry pollen in Agaonidae (Hymenoptera: Chalcidoidea). Tijd. Entomol. 121: 279-293. WiF.BF.s, J. 1". 1966. Provisional check list of fig wasps (Hymenoptera, Chalcidoidea). Zool. Verh. 83: 1-44. WIEBES, J. T. 1979. Co-evolution of figs and their insect pollinators. Ann. Rev. Ecol. Syst. 10: 1-12.

GALL WASPS Cynipidae, C y n i p i n a e . Spanish: galígenas (General).

Avispas

It is t h e galls, i n d u c e d o n plants by these wasps, that a r e usually noticed r a t h e r t h a n the wasps themselves (Mani 1964). Each species of wasp causes a particular part of a particular plant to f o r m characteristic galls. T h e latter a r e swollen masses of tissue in which the wasp's larvae feed a n d develop. T h e m e c h a n i s m of gall f o r m a t i o n is poorly u n d e r s t o o d b u t is t h o u g h t mainly to involve r e d i r e c t e d g r o w t h of u n d i f f e r e n tiated plant tissues by substances in the larval saliva (Cornell 1983). Galls are of all shapes, sizes, a n d loca­ tions on the host plant. Many are smooth, spherical, a n d singular, while o t h e r s are i r r e g u l a r masses or swellings often with a coarse or even hairy surface. T h e y usually grow from twigs o r leaves, often along the m i d r i b of the latter. T h e largest n u m b e r of gall wasps use oaks as hosts a n d are t h e r e ­ fore most n u m e r o u s in the n o r t h e r n half of Latin A m e r i c a w h e r e these trees occur. Typical spherical galls ( 1 . 5 - 2 cm d i a m e t e r ) are the p r o d u c t of m e m b e r s of the wide­ s p r e a d g e n u s Atrusca (fig. 12.2e). Not all gall wasps are p h y t o p h a g o u s a n d p r o d u c e their own galls; s o m e live as inquilines in t h e galls m a d e by o t h e r wasps,

412

SAWFL1ES, WASPS, ANTS, AND BEES

a n d some are parasitic. L a r g e r galls often house a varied assemblage of o t h e r insects besides inquiline gall wasps, for example chalcidoid wasps a n d parasitic Diptera (Shorthouse 1973). P h y t o p h a g o u s gall wasps (Weld 1952) are small to m i n u t e (BL 1-4 m m ) and black to pale r e d d i s h - b r o w n a n d have filiform a n t e n n a e , a p r o n o t u m with exten­ sions laterally to touch the tegulae, and a shiny, large, oval, c o m p r e s s e d abdomen whose second dorsal sclerite is greatly en­ larged a n d covers over half of this body region. Many gall wasps have a complex life cycle, with two very different generations in vastly different galls a p p e a r i n g at differ­ ent times of the year. O n e g e n e r a t i o n may be p a r t h e n o g e n e t i c . T h e study of galls a n d their m a k e r s is in a b e g i n n i n g phase in Latin A m e r i c a (Occhioni 1979). Few life cycles of indigenous species are known, a n d their unraveling will r e m a i n a fertile field for investigation for a long time to come. S o m e landmark studies were accomplished on the northern fauna by A. C. Kinsey (1930, 1936).

References CORNELL, B. 1983. Why and how wasps form galls: Cynipids as genetic engineers? Antenna 7(2): 53-58. KINSEY, A. C. 1930 [1929]. The gall wasp genus Cynips: A study in the origin of species. Indiana Univ. Stud. 16: 1-577. KINSEY, A. C. 1936. The origin of higher categories in Cynips. Indiana Univ. Publ. Sci. Ser. 4: 1-334. ' MANÍ, M. S. 1964. Ecology of plant galls. Junk, The Hague. OCCHIONI,' P. 1979. "Galhas," "cecideas" ou "tumores vegetáis" em plantas nativas da flora do Brasil. Leandra 8-9: 5-35. SHORTHOUSE, J. D. 1973. The insect community associated with rose galls of Diplolepid poliia (Cynipidae, Hymenoptera). Quaest. Ento­ mol. 9: 55-98. WELD, L. H. 1952. Cynipoidea (Hym.) 19051950. L. Weld, Ann Arbor.

WASP NESTS Only certain solitary a n d social wasps con­ struct t r u e nests. O t h e r wasps r e a r their young in a variety of plant or animal hosts. Several a u t h o r s (Richards 1978: 1 9 - 2 1 , leanne 1975, Wilson 1971) have a t t e m p t e d to classify t h e different types of nests. Perhaps the most useful system is the following (simplified from Evans a n d West-Eberhard 1970): I. A p r e e x i s t i n g cavity, modified to suit the n e e d s of the species (e.g., Pepsis, Pompilidae). II. An e l o n g a t e b u r r o w d u g in the g r o u n d , r o t t e n wood, or pith (Bembix, Sphecidae). III. Fabricated of foreign material a n d usually placed a b o v e g r o u n d . A. C o n s t r u c t i o n material primarily m u d (some Polybia, Vespidae). B. C o n s t r u c t i o n material wood p u l p or o t h e r vegetable substance. 1. Spherical cells in i r r e g u l a r clus­ ters inside a ball of plant wool s u s p e n d e d by a filament (Microstigmus, Sphecidae). 2. T u b u l a r cells in a cluster o r series on plant stems (Parischnigaster, Vespidae). 3. N a k e d p a p e r c o m b or combs s u s p e n d e d by a pedicel (Polistes, Vespidae). 4. P a p e r combs s u r r o u n d e d by an e n v e l o p e (most Polybia, Vespidae). T h e large size, internal s t r u c t u r e a n d shape of m a n y social wasp nests m a k e them true w o n d e r s of n a t u r e (Rau 1943). Some of the so-called ceramic types, com­ posed of smoothly polished, colored m u d (e.g., by Polybia singularis), are c o n s i d e r e d Works of art a n d sold as such in curio shops in South America. In the Neotropics, t h e greatest develop­

m e n t of nest building is found in the vespid subfamily Polistinae. Generally, these wasps build rows of horizontal combs that they attach directly to a flat surface or s u s p e n d from o v e r h a n g i n g objects with a central or lateral filament. T h e y m a k e the combs of relatively fragile, p a p e r l i k e material, which they p r o d u c e by chewing wood fibers a n d mixing t h e m with salivary secretions ("wasp p a p e r " ) . T h e combs a r e usually protected by an o u t e r baglike e n v e l o p e of m u c h s t r o n g e r material, either m u d or a m u c h heavier form of wasp paper. T h e latter may exceed the thickness a n d tenacity of highg r a d e , commercial c a r d b o a r d in some nests, such as those of Chartergus chartarius. Both the i n n e r combs a n d o u t e r wall a r e m a d e simultaneously so that the entire s t r u c t u r e is finished at the same time. S o m e wasps may "add o n " to a c o m p l e t e nest from time to time, a typical practice of Synoeca a n d Polybia rejecta. T h e same nest may be occupied year after year (perennial) or a b a n d o n e d after a single r e p r o d u c t i v e sea­ son (annual). E x a m p l e s of the f o r m e r can persist as long as twenty-five years. T h e wasps carefully form single or multi­ ple o p e n i n g s in the o u t e r e n v e l o p e for passage in a n d o u t of the nest. Usually, t h e r e is only o n e such doorway at the bottom, but slitlike or d o u b l e o p e n i n g s may be placed at the sides or o u t e r c o r n e r s by some species. T h e combs also a r e p e n e ­ trated by passageways to allow access to all parts of the nest interior. T h e n u m b e r of individuals p e r colony varies considerably, from a few to h u n d r e d s . It is not u n u s u a l for large nests of several years age to be h o m e for several t h o u s a n d adult wasps at a time. In general, the b r o o d cells a r e r e u s e d for the d e v e l o p m e n t of several g e n e r a t i o n s a n d must be cleaned of debris a n d excre­ tions of prior larvae before receiving their new occupants ( J e a n n e 1980). Most wasp nests are h u n g from b r a n c h e s high in trees to e n s u r e protection from climbing m a m m a l i a n o r reptilian p r e d a -

WASP NESTS

413

tors. T h e globular d a r k forms of these nidos de avispas are familiar sights to c o u n t r y folk. To y o u n g s t e r s in search of a d v e n t u r e , nests serve as attractive targets for rock t h r o w i n g a n d o t h e r forms of g e n e r a l mischief. T h e d a n g e r involved in molesting a large colony is quickly a p p r e c i a t e d as scores of stinging wasps soon d e s c e n d o n any would-be at­ tacker a n d can inflict e x t r e m e l y painful a n d occasionally fatal r e t r i b u t i o n . C e r t a i n A m a z o n i a n I n d i a n s actually in­ vite attacks from wasps as p a r t of rituals. A m o n g t h e K a y a p ó of c e n t r a l Brazil, ag­ gressive Polybia species a r e sought o n r e g u ­ lar occasions to take part in the reenactm e n t of a n ancient m y t h describing their fight with t h e giant r h i n o c e r o s beetle god. A scaffold is c o n s t r u c t e d which t h e warrior uses to reach t h e nests. With b a r e h a n d s , the I n d i a n s strike the nests a n d receive the stings of wasps until they b e c o m e u n c o n ­ scious from t h e pain a n d v e n o m . T h e c e r e m o n y is i m p o r t a n t to the Kayapó as a s t a t e m e n t of their place in the universe (Posey 1981). Several kinds of Neotropical birds take a d v a n t a g e of the aggressiveness of wasps by placing their own nests in proximity to those of the insects (see above).

References EVANS, H. E., AND M. J. WEST-EBERHARD.

Solitary wasps are defined as species in which t h e r e is n o c o o p e r a t i o n involving division of labor between m o t h e r and d a u g h t e r s or between females of the same g e n e r a t i o n . Many a r e subsocial, living ¡ n nesting aggregations a n d building free nests of m u d a n d plant materials. One genus of the otherwise totally solitary fam­ ily Sphecidae (Microstigmus) is social. Females of most form a b u r r o w in the g r o u n d in which to r e a r their young usually fed o n insect prey paralyzed by her sting. Provisioning may be progressive fresh food being b r o u g h t to the develop­ ing larvae until they m a t u r e , or prey is p r o v i d e d in a single mass for the larvae to d e v o u r without f u r t h e r attention from the adults. T h i s assemblage is s e p a r a t e d from the "parasitoid wasps" on the basis of their well-developed nesting habits a n d larvae that feed externally o n the host (Evans 1966). It is comprised of generally larger wasps that are agile fliers a n d capable of stinging painfully. T h e sting is used to s u b d u e a n d paralyze prey, in constrast to its purely defensive use in the social wasps. G r o u p s of adults sleeping o n plants are sometimes observed.

1970.

The wasps. Univ. Michigan Press, Ann Arbor. JEANNE, R. L. 1975. The adaptiveness of social wasp nest architecture. Quart. Rev. Biol. 50: 267-287. JEANNE, R. L. 1980. Observacóes sobre limpeza e reutilizacáo de células em ninhos de vespas sociais (Hymenoptera: Vespidae). Mus. Paraense Emilio Goeldi Bol. Zool. 101: 1-7. POSEY, D. A. 1981. Wasps, warriors and fearless men: Ethnoentomology of the Kayapó Indi­ ans of central Brazil. J. Ethnobiol. I: 165-174. RAU, P. 1943. The nesting habits of Mexican social and solitary Vespidae. Entomol. Soc. Amer. Ann. 36: 515-536. RICHARDS, O. W. 1978. The social wasps of the Americas, excluding the Vespinae. Brit. Mus. Nat. Hist., London. WILSON, E. O. 1971. T h e insect societies. Belknap Press, Harvard Univ., Cambridge.

414

SOLITARY WASPS

SAWFLIES, WASPS, ANTS, AND BEES

Reference EVANS, H. 1966. The behavior patterns of soli­ tary wasps. Ann. Rev. Entomol. 11: 123-154.

Figure 12.3 SOLITARY WASPS, (a) Cuckoo wasp (Neochrysis carina, Chrysididae). (b) Velvet ant (Traumatomutilla indica, Mutillidae). (c) Mammoth wasp (Campsomeris ephippium, Scoliidae). (d) Taruntula hawk (Pepsis sp., Pompilidae).

food left in t h e cell for the host's y o u n g , killing a n d e a t i n g the latter in the process. These are a m o n g the most beautiful of insects because of their brilliantly colored bodies. T h e n u m e r o u s species a r e bright, metallic p u r p l e , blue, or g r e e n a n d a r e sometimes m i x t u r e s of o n e or m o r e of these colors in r e s p l e n d e n t combinations, appreciated only with the aid of a s t r o n g magnifying lens. Most are small (BL rarely over 12 m m ) a n d also recognized by a coarse body s c u l p t u r e a n d ventrally con­ cave a b d o m e n , t h e latter consisting of only three or four visible s e g m e n t s . W h e n the wasp is d i s t u r b e d , it curls u p into a ball, the head a n d t h o r a x nesting snugly in the hollow of the a b d o m e n , a n d r e m a i n s i m m o ­ bile until the d a n g e r has passed. T h e two largest g e n e r a in t h e tropics are Trichrysis a n d Neochrysis 12.3a). T h e family is not large h e r e , 111 species in 18 g e n e r a (Kimsey Bohart 1980).

Neo(fig. only and

Cuckoo Wasps Chrysididae. T h e c o m m o n n a m e of this family comes from the habit m a n y species have of e n t e r i n g the nests of their hosts, which are most often solitary wasps a n d bees, as they are being provisioned. (One anomalous subfamily—Amiseginae—of these wasps specializes on walkingsticks as prey.) In a m a n n e r analogous to that of their avian namesakes, the wasp larvae devour the

Reference KIMSEY, L.

S.,

AND R.

M.

BOHART.

1980.

A

synopsis of the chrysidid genera of Neo­ tropical America (Chrysidoidea, Hymenop­ tera). Psyche 87: 7 5 - 9 1 .

Velvet Ants Mutillidae. Spanish: Perritos d e Dios, hormigas terciopelas, a r a ñ a s p u s - p u s (Argentina). Portuguese: Formigas feíticeiras, c a c h o r r i n h o s d e Nossa

S e n h o r a , formigas d e onca, o n c i n h a s (Brazil). Quechua: Sisi h u a k a n ñ a h u i (ant that makes you cry; Peru). T h e females of these wasps a r e usually seen walking agilely on the g r o u n d or o n logs a n d s t u m p s , their brightly m a r k e d , hairy bodies attracting attention. T h e p u ­ bescence of most is d a r k velvety blue or black with contrasting spots of brilliant red, white, or yellow. Most a r e relatively small (BL 3—5 m m ) , b u t s o m e Neotropical representatives, Hoplomutilla, Leucospilomutilla, a n d Traumatomutilla (fig. 12.3b), r e a c h fairly great size, lacking wings a n d stinging painfully; they are sometimes mistakenly identified as large ants (e.g., H. xanthocerata = folofilla for Paraponera clavata = folofa in P a n a m a ; M é n d e z 1987). Species of Hoplocrates are also large a n d have outsized heads. Males are winged b u t less often observed. Both sexes a r e capable of p r o ­ d u c i n g a squeaking noise by m o v i n g t h e third a b d o m i n a l s e g m e n t in a n d out of the second, thus b r i n g i n g stridulatory surfaces on each into contact. All are solitary a n d d e v e l o p as e x t e r n a l parasitoids o n the i m m a t u r e s of various wasps, bees, beetles, a n d flies. T h e exces­ sively long stinger of the females enables t h e m to pierce the nest cells of their hosts, into which they inject their v e n o m a n d o n which they place their eggs. In some cultures, o d d beliefs a n d prac­ tices have arisen s u r r o u n d i n g these wasps.

SOLITARY WASPS

415

Some p e o p l e in Brazil e m p l o y t h e m in love magic. A m a n , desiring the attentions of a w o m a n , obtains t h r e e of h e r hairs, coats t h e m with sweet s y r u p , a n d p u t s t h e m in a box with a velvet ant. As the insect eats the hairs, the w o m a n gradually falls in love with the p e r p e t r a t o r . O t h e r powers of sorcery a t t r i b u t e d to mutillids, c o m m o n especially a m o n g the caboclos a n d I n d i a n s of Brazil, include the ability to c u r e a variety of illnesses a n d to e n h a n c e talents. For e x a m p l e , playing t h e violin can be i m p r o v e d by crossing the palm of the right h a n d with the insect o n Friday ( L e n k o a n d P a p a v e r o 1979: 218). T h e Neotropics is rich in species: j u s t u n d e r 1,100 a r e now k n o w n (Fritz p e r s . comm.), a n d m a n y m o r e a r e certain to be discovered ( B r o t h e r s 1975: 5 8 9 - 6 3 8 , Schu­ ster 1949). T h e y are widely distributed as a g r o u p but especially a b u n d a n t in w a r m e r climates (Mickel 1952).

References BROTHERS, D. M. ] 975. Phylogeny and classifica­ tion of the Aculeate Hymenoptera, with spe­ cial reference to Mutillidae. Univ. Kansas Sci. Bull. 50: 483-648. LENKO, K., AND N. PAPAVERO. 1979.

Insetos no

folclor. Cons. Estad. Artes Cien. Hum., Sao Paulo. MÉNDEZ, E. 1987. Elementos de la fauna panameña. Priv. publ., Panama. MICKEL, C. E. 1952. The Mutillidae (wasps) of British Guiana. Zoológica 37: 105—150. SCHUSTER, R. M. 1949. Contributions toward a monograph of the Mutillidae of the Neo­ tropical Region. III. A key to the subfamilies represented and descriptions of several new genera (Hymenoptera). Entomol. Americana 29: 59-140.

Mammoth Wasps Scoliidae, C a m p s o m e r i n a e , C a m p s o m e r i n i , Campsomeris. T h e s e gigantic (BL males to 20 m m , fe­ males to 35 m m ) , d a r k wasps with black, hairy, r o b u s t bodies a n d shiny blue-black wings usually sport c o n s p i c u o u s , d e e p o r a n g e , transverse b a n d s or spots at the

416

SAWFLIES, WASPS, ANTS, AND BEES

base of the a b d o m e n dorsally (fig. 12.3 C \ (Bradley 1945). T h e smaller males differ somewhat from the females also in having longer, straighter a n t e n n a e a n d finer body hairs. Also, the legs of the female are s h o r t e r a n d a d a p t e d for digging, unlike the m o r e delicate slender a p p e n d a g e s of the male. A l t h o u g h the several species (Bradley 1957) are fairly c o m m o n over almost all of the A m e r i c a n tropics, very little is known of their habits. T h e r e are several similar species in the g e n u s , all of which are parasitoids of scarab beetle larvae. The latter may be so-called white grubs (Phyllophaga) or o t h e r large types that burrow in the g r o u n d (Strategus, Oryctes). T h e female wasp's powerful fossorial legs a d a p t it par­ ticularly well for digging into the ground after these hosts.

References BRADLEY, J. C. 1945. The Scoliidae (Hymenop­ tera) of northern South America, with especial reference to Venezuela. 1. The genus Camp­ someris. Bol. Entomol. Venezolana 4: 1-36. BRADLEY, J. C. 1957. The taxa of Campsomeris. (Hymenoptera: Scoliidae) occurring in the New World. Amer. Entomol. Soc. Trans. 83: 65-77.

Spider Wasps Pompilidae. T h i s is a large a n d diverse family, mostly c o m p o s e d of medium-sized to very large (BL 1.5—5 cm) wasps with slender bodies a n d long, spiny legs (Evans 1953). Most are d a r k with p i g m e n t e d wings, which they often flick nervously w h e n walking on the ground. Females seek spiders, which they para­ lyze with a sting a n d pack into subterra­ nean cells or in existing cavities in wood to provide food for the developing young. S o m e construct elevated m u d nests. A single prey is p r o v i d e d for each cell. T h e female uses the tip of the a b d o m e n to tamp

¿own the e a r t h when closing the cell or to mold m u d .

Reference EVANS, H. E. 1953. Comparative ethology and the systematics of spider wasps. Syst. Zool. 2: 155-172. Tarantula Hawks Pompilidae, Pepsis. Spanish: San J o r g e s , avispones, matacaballos (Argentina). It is curious that these conspicuous spider wasps d o not carry m o r e vernacular n a m e s among the p e o p l e of the region. T h e i r gigantic size, impressive steely blue bodies and the b r i g h t o r a n g e wings of m a n y species often attract attention a n d i m m e d i ­ ately identify t h e m . T h e y are c o m m o n l y seen taking nectar from flowers (often milkweed) a n d can be h e a r d m a k i n g a loud buzzing s o u n d while in flight. Females of t h e largest species have body lengths of over 4.5 c e n t i m e t e r s ; males are a little smaller, 2.5 to 3.5 centimeters (fig. 12.3d). Species have o n e of two wing colors: bright or b u r n t o r a n g e or a d a r k , smoky h u e . T h e large size of these c r e a t u r e s a d a p t s them to their prey, the g r e a t m y g a l o m o r p h spiders, or t a r a n t u l a s . Females h u n t these spiders a n d e n g a g e t h e m in battles that have been witnessed by naturalists a n d described in m a n y p h o t o g r a p h i c essays (Petrunkevitch 1926). T h e y a p p r o a c h their arachnid adversary with wings raised, then grasp it by a leg with the mandibles. T h e Spider retaliates, a t t e m p t i n g to bite a n d kill the wasp, in which it is sometimes success­ ful. But almost always, the wasp wins the foray by stinging the hapless t a r a n t u l a on the u n d e r s i d e . She t h e n d r a g s its paralyzed body to a b u r r o w , often the spider's own, and buries it t h e r e after d e p o s i t i n g a single ^ggon it. T h e larva feeds externally on the interred carcass, finally t r a n s f o r m i n g into *n adult within a silken cocoon s p u n nearby in t h e b u r r o w . Wasps excited by

battle or threat emit a p u n g e n t o d o r whose function is u n k n o w n . A l t h o u g h not well known, Pepsis a p p e a r to be very specific with r e g a r d to prey species a n d a r e a d e p t at discriminating t h e m , a p p a r e n t l y by o d o r clues picked u p as the wasp taps the s p i d e r with its a n t e n n a e . M e m b e r s of this g e n u s are restricted to the New World, in the tropical p o r t i o n s of which t h e r e are several h u n d r e d species ( H u r d 1952). T h e g r o u p seems c e n t e r e d in Amazonia w h e r e most of the 300 species of n o r t h e r n South America are f o u n d . Many species also live in Middle A m e r i c a , includ­ ing the West Indies (Alayo 1954), a n d occupy all habitats from sea level to 4,000 m e t e r s in the A n d e s . Several d i s p a r a t e insect types have evolved color p a t t e r n s a n d behaviors m i m ­ icking t a r a n t u l a hawks. T h e resemblances are very convincing a n d incredible in the way unwasplike portions of the mimic's body are m o l d e d a n d colored to r e s e m b l e the wasp's. Such mimics a r e f o u n d a m o n g certain reduviid bugs (Spiniger ater), katy­ dids (Scaphura, Aganacris), m y d i d flies (Mydas rubidapex), a n d arctiid moths (Macrocneme).

References ALAYO, P. 1954. El género Pepsis Fabr. en Cuba (Hymenoptera-Pompilidae). Univ. Oriente Cuadernos 37: 1-25. HURD, P. 1952. Revision of the nearctic species of the pompilid genus Pepsis (Hymenoptera, Pompilidae). Amer. Mus. Nal. Hist. Bull. 98; 257-334. PETRUNKEVITCH, A. 1926. Tarantula versus tarantula-hawk: A study in instinct. J. Exper. Zool. 45: 367-397.

Digger Wasps Sphecidae. Digger wasps c o m p r i s e a large family (Bohart a n d M e n k e 1976) with varied form a n d habits. T h e y are distinguished by the s h a p e of the lateral p o r t i o n of t h e p r o n o t u m (dorsal sclerite of p r o t h o r a x ) , which is f o r m e d into a r o u n d e d lobe, well

SOLITARY WASPS

417

A lohnston 1978); its place is a s s u m e d by asiaticum a n d S. fistularium in South erica. Species occupy islands, including in t h e r e m o t e Pacific (Coco Island), me here they h a v e p r e s u m a b l y b e e n intro­ duced inadvertently by h u m a n action.

Figure 12.4 DIGGER WASPS (SPHECIDAE). (a) Digger wasp (Trypoxylon albitarsi) (b) M i * dauber {Sceliphron assimile). (c) Sand wasp {Bembix citripes). (d) Social sphecid (Microstiami comes)

s e p a r a t e d from t h e base of t h e wing. T h e wings a r e n e v e r folded longitudinally when at rest. All a r e solitary, with t h e exception of t h e Microstigmus (see social sphecids, below), m a k i n g their nest in many situations a n d provisioning t h e m with various kinds of insects, such as plant bugs, spiders, g r a s s h o p p e r s , a n d caterpil­ lars (Fritz a n d Genise 1980). A m o n g t h e g r o u n d a n d m u d nest m a k e r s , females apply their h e a d to s h a p i n g a n d m a n i p u l a t ­ ing e a r t h . T h e sting is used to s u b d u e prey, not j u s t for defense, as in t h e social wasps. I n Brazil, t h e m u d nest m a k i n g species of Trypoxylon (fig. 12.4a) a r e r e g a r d e d with superstition by t h e caboclos a n d a r e k n o w n by m a n y local n a m e s : minguita, nhá fina, mariambola. T h e i r nest material is applied widely in folk t h e r a p e u t i c s , uses r a n g i n g from aphrodisiacs t o c u r e s for constipa­ tion, s p i d e r bites, a n d b u r n s . It is a very large g e n u s (Richards 1934). A n interest­ ing feature of m a n y species is t h e peculiar n e s t - g u a r d i n g b e h a v i o r of males in t h e absense of t h e females (Coville a n d Griswold 1984). T h e m a l e sits j u s t inside t h e nest with its h e a d p r o t r u d i n g from t h e entrance.

References BOHART, R. M., AND A. S. MENKE. 1976. Sphecid

wasps of the world. Univ. California Press, Berkeley and Los Angeles. COVILLE, R. E., AND C. GRISWOLD. 1984. Biology

of Trypoxylon (Trypargilum) superbum (Hymenoptera: Sphecidae), a spider-hunting

418

SAWFLIES, WASPS, ANTS, AND BEES

References PBEEMAN, B. E. 1973. Preliminary studies on the nopulation dynamics of Sceliphron assimile Dahlbom (Hymenoptera: Sphecidae) in Ja­ maica-J- Anim. Ecol. 42: 173-182. ftoEEMAN, B. E., AND B. JOHNSTON. 1978. The

wasp with extended guarding of the brood bv males. Kans. Entomol. Soc. J. 57: 365-376 FRITZ, M., AND J. GENISE. 1980. Nido de barro

Sphecidae, constructores, inquilinos, parasitoides, cleptoparásitos, detritívoros. Soc. Ento­ mol. Argentina Rev. 39: 6 7 - 8 1 . RICHARDS, O. W. 1934. The American species of the genus Trypoxylon (Hymenopt., Sphecoidea). Royal Entomol. Soc. London Trans 123: 173-362. Mud Daubers Sphecidae, Sphecinae, Sceliphronini, Sceliphron. Spanish: Celifrónes (General). Just six of t h e thirty worldwide species in this g e n u s live in t h e New World tropics (van d e r Vecht a n d van Breugel 1968), but these a r e conspicuous both for their com­ m o n n e s s a n d for t h e nests they often build on h u m a n habitations (Shafer 1949). T h e adults a r e readily recognized by their extra-long waist a n d color pattern of sharply contrasting designs of black and yellow. T h e sides of t h e r e a r of the thorax ( p r o p o d e u m ) a r e built u p also as ridges to form a U-shaped enclosure. Nests a r e constructed of m u d a n d con­ sist of a series of parallel tubes or elon­ gated cells. Each cell is mass provisioned with spiders. O n e species, Sceliphron assi­ mile (fig. 12.4b), thrives o n civilization and often builds its nest o n houses or other m a n - m a d e structures. T h i s species occurs t h r o u g h t h e West Indies, Mexico, and Central America ( F r e e m a n 1973, Freeman

biology in Jamaica of the adults of the sphecid wasp Sceliphron assimile Dahlbom. Ecol. Ento­ mol. 3: 39-52. SHAFER, G. 1949. T h e ways of a mud dauber. Stanford Univ., Stanford. VAN DER VECHT, J., AND F. M. A. VAN BREUGEL.

1968. Revision of the nominate subgenus Sceliphron Latreille (Hymenoptera, Spheci­ dae) (Studies on the Sceliphronini, Part 1). Tijd. Entomol. I l l : 185-255.

provision t h e nests progressively with vari­ ous insects, most often noxious flies, includ­ ing horseflies (frequently p l u c k i n g t h e m r i g h t off horses a n d cattle), S y r p h i d a e , Muscidae, a n d o t h e r large species. Al­ t h o u g h they s u b d u e their prey with a p o t e n t stinger, they a r e n o t p r o n e to sting h u m a n s (Cane a n d Miyamoto 1979). T h e s e a r e truly solitary wasps, b u t they occasionally e n g a g e in mass attacks against i n t r u d e r s of "colonies" (where m a n y indi­ viduals a r e nesting in proximity). Males also take p a r t in "sun d a n c e s , " o r flight rituals wherein females a r e m e t in t h e air a n d aggressively b r o u g h t to bay for copulation. Only a couple of dozen species a r e found in t h e Neotropics, either in inland areas of sandy soil o r m o r e c o m m o n l y o n coastal beaches. T h e most typical a n d wide­ s p r e a d a r e Bembix americana in t h e Carib­ bean a n d B. citripes (fig. 12.4c) in South America.

Sand Wasps Sphecidae, Nyssoninae, Bembicini, Bembix. Spanish: Insectos policias (General). Cowfly tigers, h o r s e g u a r d s .

CANE, J. H., AND M. M. MIYAMOTO. 1979. Nest

The sand wasps a r e a familiar sight o n beaches o r sand d u n e s cruising a n d dart­ ing about in search of prey o r busily excavating nests (Evans 1957). T h e y a r e medium to l a r g e (BL 1 2 - 1 7 m m ) , stoutbodied wasps, basically black b u t usually elaborately m a r k e d with u n d u l a t i n g white or yellow b a n d s o n t h e a b d o m e n . T h e y a r e also recognized by t h e r e d u c e d simple eyes (ocelli), t h e a n t e r i o r m o s t of which m a y b e vestigial, a n d e l o n g a t e m o u t h p a r t s . T h e anterior legs normally a r e fringed with long hairs useful to t h e wasp in d i g g i n g in soft sand.

Social Sphecids Sphecidae, P e m p h r e d o n i n a e , Microstigmus.

Nests a r e sloping t u n n e l s , a few to several c e n t i m e t e r s in d e p t h , in sandy soil. At the t e r m i n u s of t h e e n t r y t u n n e l is a horizontal b r a n c h in which t h e y o u n g a r e Wared a n d a final vertical s p u r w h e r e t h e females may rest o r take refuge. T h e y

References defense and foraging ethology of a Neo­ tropical sand wasp, Bembix multipicta (Hy­ menoptera: Sphecidae). Kans. Entomol. Soc. J. 52: 667-672. EVANS, H. E. 1957. Studies on the comparative ethology of digger wasps of the genus Bembix. Comstock, Ithaca.

T h i s g e n u s of nearly fifty species (WestE b e r h a r d 1977), k n o w n from Costa Rica to Paraguay, is considered to be t h e only g e n u s of t h e family with s o m e species having t r u e social habits. Sociality was discovered r e ­ cently in o n e species (Microstigmus comes, fig. 12.4d; Matthews 1968a, 19686) a n d subse­ quently verified in m a n y o t h e r s in t h e g e n u s . O t h e r species a r e solitary a n d prac­ tice progressive provisioning. T h e small ( 0 . 5 - 1 . 5 c m ) , u n i q u e nests

SOLITARY WASPS

419

(pi. 4b) are baglike a n d c o n s t r u c t e d of the waxy material coating the u n d e r s i d e s of fan-palm leaves (in M. comes, see Matthews a n d Starr 1984). T h i s material, or wood flecks, moss, a n d lichens in o t h e r species, is b o u n d t o g e t h e r by c o o p e r a t i n g females with silk p r o d u c e d from a b d o m i n a l glands. Nests are usually s u s p e n d e d from the u n ­ dersides of leaves b u t sometimes h a n g from o t h e r inclined objects (rocks, logs, eaves of dwellings) by a slender coiled or straight pedicel. T h e u p p e r p o r t i o n of the nest is hollow a n d acts as a c h a m b e r for the adult male a n d female wasp o c c u p a n t s . T h e lower p o r t i o n consists of a few to several cells in which the b r o o d is r e a r e d . T h e s e cells a r e mass p r o v i s i o n e d with very small insects, n o r m a l l y springtails, thrips, or l e a f h o p p e r s (Cicadellidae) by females w o r k i n g in c o o p e r a t i o n . T h e prey type a n d nest s h a p e are species specific. O n e female may show r e p r o d u c t i v e d o m i n a n c e a n d , a l t h o u g h structurally indistinct from the others, m a y be c o n s i d e r e d a kind of " q u e e n . " Foraging a n d o t h e r activities are diurnal. T h e wasps themselves a r e m i n u t e (BL 2 - 5 m m ) a n d usually r e d d i s h or pale yellowish-brown a n d n a r r o w waisted. In life, the yellow species have b r i g h t g r e e n eyes. T h e lower tooth of the m a n d i b l e is often very long.

References MATTHEWS, R. W. 1968a. Microstigrnus comes: Sociality in a sphecid wasp. Science 160: 787-788. MATTHEWS, R. W. 19686. Nesting biology of the social wasp Microstigrnus comes (Hymenoptera: Sphecidae, Pemphredoninae). Psvche 75: 23-45. MATTHEWS, R.

W.,

AND C.

K.

STARR.

1984.

Microstigrnus comes wasps have a method of nest construction unique among social in­ sects. Biotropica 16: 5 5 - 5 8 . WEST-EBERHARD, M. J. 1977. Morphology and

behavior in the taxonomy of Microstigrnus wasps. 8th Int. Cong. Int. Union Stud. Soc. Ins. Proc. Wageningen, the Netherlands. Pp. 123-125.

420

SAWFLIES, WASPS, ANTS, AND BEES

Caterpillar Hunters Eumenidae. T h e s e a r e builders of a g r e a t e r variety 0 r nests than any o t h e r g r o u p of solitary wasps, but they almost always e m p | o v m u d in some way. Females may choose a simple hollowed twig or b u r r o w into clav or compact soil banks. M o r e familiar are the so-called m a s o n wasps, which con­ struct free m u d nests. T h o s e of the "pot­ ter wasps" may take the s h a p e of a wellf o r m e d j u g (spherical with flared spout) placed a t o p a twig. A few actually camou­ flage their m u d cells with chewed wood fibers o r build with m a c e r a t e d leaves plant fibers, a n d resins, a practice thought to r e p r e s e n t a step in the direction of p a p e r nest building, as p e r f o r m e d by their h i g h e r social relatives. T h e nests are usually supplied with caterpillars, a l t h o u g h beetle larvae are sometimes used. T h e female may provi­ sion singly or r e t u r n r e p e a t e d l y to provide h e r d e v e l o p i n g larvae with a continuous supply of food (progressive provisioning). Caterpillar h u n t e r s are like their close relatives, the social wasps, in general ap­ p e a r a n c e , all having the wings folded longi­ tudinally at rest a n d some even with nar­ r o w e d waists. T h e y differ, however, in possessing long mandibles, which often cross when closed, o n e apical s p u r on the m i d d l e tibia, bifurcate tarsal claws, and, of course, lacking the social habit. Potter Wasps E u m e n i d a e , E u m e n i n a e , Eumenes. Portuguese: Mariambolas (Brazil). T h e q u a i n t little ( 1 - 2 cm diameter) mud j u g nests of these wasps are unique. They a r e spherical, with the edges of the opening (in the c e n t e r or at o n e e n d ) on a slight projection a n d flared like the spout of an old-fashioned, clay water pot. They are constructed by all the h u n d r e d or so species in this g e n u s , the most c o m m o n and wide­ s p r e a d b e i n g E. consobnnus (fig- 12.5a).

piaure 12.5 CATERPILLAR HUNTING WASPS (EUMENIDAE). (a) Potter wasp (Eumenes con¡obrinus). (b) Montezuma wasp (Montezumia azurescens). (c) Wanderer (Pachodynerus nasidens). U¡) Zethus wasp [Zethus matzicatzin).

Their t a x o n o m y is c o m p l e x a n d unsettled (Soika 1978). These are medium-sized (BL 10—15 mm), solitary wasps with long mandibles and elongate waists like zethus wasps (see below). T h e y a r e distinguished from the latter, however, by the strongly convex thorax a n d by the swollen p a r t of the waist segment n e a r the j o i n t with the a b d o m e n (rather t h a n n e a r t h e middle). Neither is the basal p o r t i o n of the a b d o m e n e x t e n d e d as it is in their n e a r relatives.

c o m m o n Polistes p a p e r wasps in their black a n d yellow a n d o t h e r maculate color pat­ terns a n d fairly large size (BL 2 cm). T h e y are distinguished by family characteristics a n d by the combination of a completely sessile a b d o m e n , a t h i c k e n i n g in the veins immediately before the spot (pterostigma) o n the o u t e r leading e d g e of the fore wing, a n d the peculiar s h a p e of the small cap (tegula) overlying the base of the wing, this having a posterior lobe or extension that curves inward.

T h e nests are placed, often evenly ced in linear series, sometimes in clus­ ters, along t h e u p p e r sides of slender T h e y a r e provisioned with caterpilS, and each is the n u r s e r y for a single va. T h e latter hatches from an egg spended from the roof of the nest on a readlike stalk.

Not m u c h is k n o w n of their habits, save that nests seem to be m a d e both in the g r o u n d a n d banks of soil as well as in the form of free m u d structures affixed to h a r d substrates, d e p e n d i n g o n the species. Free nests contain only a few cells, generally a r r a n g e d in parallel fashion in a single layer, m u c h as in the nests of m u d d a u b e r s but m u c h m o r e irregularly a n d roughly s h a p e d . Females a p p a r e n t l y provision these cells progressively (Evans 1973).

rence IKA, A. G. 1978. Revisione degli Eumenidi Neotropicali appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pamraphidoglossa Schulth. et affini. Mus. Civ. Storia Nat. Venezia Bol. 29: 1-420. (The several new genera proposed are considered synony­ mous with Eumenes bv most other authori. ties.) Montezuma Wasps enidae, E u m e n i n a e ,

Montezumia.

bers of this g e n u s (fig. 12.5b) of solimason wasps superficially resemble

T h e g e n u s is entirely A m e r i c a n a n d contains fifty-two species mostly living in w a r m lowland climes (Willink 1982).

References EVANS, H. E. 1973. Notes on the nests of Montezumia (Hymenoptera, Eumenidae). Entomol. News 84: 285-290. WILLINK, A. 1982. Revisión de los géneros Montezumia Saussure y Monobiu Saussure (Hy­ menoptera: Eumenidae). Acad. Nac. Cien. (Argentina) Bol. 55: 1-321.

SOLITARY WASPS

421

Wanderer Eumenidae, Eumeninae, nasidens.

Pachodynerus

egg is s u s p e n d e d from the food or side of the cell by a fine t h r e a d .

References T h i s m a s o n wasp is n a m e d for its p e n c h a n t for t u r n i n g u p in out-of-the-way places. It is almost c o n t i n u o u s l y distributed over Mexico, C e n t r a l A m e r i c a , a n d northwest­ ern South A m e r i c a b u t also is f o u n d o n the Antilles, o n Coco Island, a n d across the Pacific in Hawaii a n d Micronesia. T h e very similar Pachyodynerus galapagensis occurs in the Galápagos A r c h i p e l a g o . T h e w a n d e r e r so closely resembles the h o n e y wasp (Brachygastra lecheguana) that they a r e be­ lieved to be Müllerian mimics (Snelling pers. c o m m . ) . It is a small species (BL 8—14 m m ) a n d recognizable by its a b d o m i n a l color pat­ t e r n , black with yellow b a n d s completely r i n g i n g the h i n d b o r d e r s of each s e g m e n t , except t h e first (fig. 12.5c). T h e t h o r a x is black with a thick, rear, m a r g i n a l yellow b o r d e r a n d line of yellow a r c h i n g across the front. T h e t h o r a x also bears a d e n s e vestiture of short g o l d e n hairs. A u n i q u e trait identifying t h e males in t h e g e n u s to which it belongs is t h e greatly r e d u c e d t e r m i n a l a n t e n n a l s e g m e n t s , which are tiny lobes residing in a pit o n the tip of the organ. S o m e has b e e n written a b o u t the species' biology ( F r e e m a n a n d Jayasingh 1975, Jayasingh a n d Taffe 1982). Its m u d nests of a few cells a r e mostly f o u n d o n m a n m a d e objects, cracks in windows, keyholes, electrical sockets, wall recesses, a n d the like. T h e wasps m u s t seek n a t u r a l holes a n d cavities such as hollow twigs in the wild, b u t such nests h a v e not yet been observed. T h e only well-known n a t u r a l sites are cells in a b a n d o n e d m u d d a u b e r (Sceliphron) a n d m a s o n wasp (Zeta) nests. T h e female wasps stock their b r o o d cells with caterpillars of various species, show­ ing little specificity as to type. T h e y lay their eggs in t h e cells b e f o r e provisioning t h e m with four to eighty prey pieces. Each

422

SAWFLIES, WASPS, ANTS, AND BEES

FREEMAN, B. E., AND D. B. JAYASINGH.

1975

Population dynamics oí Pachodynerus nasidens (Hymenoptera) in Jamaica. Oikos 26: 86-91 JAYASINGH, D. B., AND C. A. TAFFE. 1982.

longer t h a n those of their vespid relatives. Color p a t t e r n s a r e typically vespoid, black and yellow or o t h e r light-colored b a n d s a n d thoracic patches o n a d a r k b a c k g r o u n d .

The

biology of the eumenid mud-wasp Pachody. nerus nasidens in trap nests. Ecol. Entomol 7283-289. Zethus Wasps E u m e n i d a e , Discoeliinae, Zethus. T h e 195 species of solitary wasps in this g e n u s (fig. 12.5d) a r e typically Neotropical, o c c u r r i n g t h r o u g h o u t almost all of the region with the exception of the Lesser Antilles, western A n d e s , a n d coastal des­ erts (one species is k n o w n from the Chil­ ean desert) (Bohart a n d Stange 1965). T h e g e n u s is of great interest ecologi­ cally, as it a p p e a r s to b r i d g e the gap be­ tween social a n d solitary wasps. Two dis­ tinct types of nesting behavior a r e found: utilization of old insect b u r r o w s in twigs, wood, o r the g r o u n d by t h e less specialized species, a n d construction of elevated nests from masticated a n d salivated plant mate­ rial by the m o r e a d v a n c e d species. Females practice both progressive a n d mass provi­ sioning, with caterpillars always forming the p r e f e r r e d food source. Adults are usually seen f r e q u e n t i n g flow­ e r i n g shrubs, often Acacia a n d Mimosa. T h e y are quick, n e r v o u s fliers, darting about in search of prey. T h e y are most easily recognized by their r a t h e r long waists (the swollen p o r t i o n of which is near the m i d d l e r a t h e r t h a n the apex, as in p o t t e r wasps). T h e a b d o m e n often has an additional constricted s e g m e n t next to the waist, m a k i n g the latter a p p e a r even longer. T h e terminal a b d o m e n segments are frequently telescoped into the basal seg­ m e n t s . T h e m a n d i b l e s a r e somewhat short for e u m e n i d s but are still significantly

Reference BOHART, R.

M.,

AND L. A. STANGE. 1965.

A

revision of the genus Zethus Fabricius in the Western Hemisphere (Hymenoptera ¡Eumeni­ dae). Univ. Calif. Publ. Entomol. 40: 1-208.

SOCIAL WASPS Vespidae. True social wasps, those that exhibit co­ operative b r o o d c a r e by specialized castes and overlap of g e n e r a t i o n s ( J e a n n e 1980), belong only to the family Vespidae, also called p a p e r wasps ( B e q u a e r t 1944; Dias Filho 1975; R i c h a r d s 1978; Richards a n d Richards 1 9 5 1 ; W e s t - E b e r h a r d 1975). (The g e n u s Microstigmus of the family Sphecidae is also social; see social sphecids, above.) In most social wasps, o n e egg-laying female is d o m i n a n t , a n d she is d u b b e d t h e "queen"; all o t h e r females usually have reduced o v a r i a n function. T h e n o n r e p r o ductive females forage, care for the b r o o d , and build the nest. T h e s e a r e the "work­ ers," a n d they may exhibit some a n a t o m i ­ cal differences from the q u e e n , usually smaller overall size. T h e r e may be multiple functional r e p r o d u c t i v e s (polygyny), how­ ever, sometimes very many, as in the honey wasps, with n o t h i n g to distinguish any dominant individual. All make nests, which a r e often large a n d elaborate ( J e a n n e 1975). T h e y are fabri­ cated from plant fibers, usually chewed wood, but a few form m u d houses or use hairs, floss, or vegetable wool of o n e kind or another for c o n s t r u c t i o n material. Colonies a r e f o u n d e d in t h r e e ways: in "le Polistinae, i n d e p e n d e n t l y by o n e or several q u e e n s w i t h o u t t h e aid of w o r k e r s from the p a r e n t colony or by o n e (usually)

or several q u e e n s a c c o m p a n i e d by swarms of workers, a n d in the Vespinae, by single q u e e n s alone ( J e a n n e 1980). A l t h o u g h ants a n d wasps are usually mortal enemies, certain wasps benefit by nesting in trees within ant territory, even on ant plants ( H e r r é et al. 1986). T h e y m a n a g e to evade p r e d a t i o n directly by the ants a n d find themselves freer of raids by their o t h e r enemies t h r o u g h the ant's p r e s ­ ence. Associations of nests of o n e ( J e a n n e 1978) or m o r e (Windsor 1972) wasp spe­ cies are also k n o w n , a n a r r a n g e m e n t that probably e n h a n c e s the defense effective­ ness of any o n e colony against v e r t e b r a t e p r e d a t o r s (ibid.). Birds nest also, but not always highly successfully, in the n e a r vicin­ ity of social wasps a n d likewise s h a r e in their n a t u r a l protection (Windsor 1976). Social wasps protect their nests from ant attack in a variety of ways: by direct attack, application of d e t e r r i n g sticky substances, t e m p o r a r y nest a b a n d o n m e n t , a n d even by effacing their own trail p h e r o m o n e s (WestE b e r h a r d 1989). Social wasps take a variety of insect prey. T h e y d o not sting the prey b u t p o u n c e o n it, cut it u p with their m a n d i b l e s , a n d chew it into a paste on which they a n d their larvae feed. T h e sting is used solely as a protective defense against v e r t e b r a t e p r e d a t o r s , a n d for that reason, its v e n o m contains sub­ stances to m a k e it especially painful (Edery e t a l . 1978). Because of the social s t r u c t u r e a n d nestbuilding abilities of these wasps, they hold religious significance for some I n d i a n g r o u p s . T h e Kayapó of Brazil give t h e m status as totem animals a n d respect t h e m as fellow inhabitants of their lands. T h e y also play a role in the tribe's pharmacology. Stings are considered to be beneficial, even if painful, a n d a r e r e c o m m e n d e d in the t r e a t m e n t of b o n e diseases (Overal 1984).

References BEQUAERT, J. C. 1944. T h e social Vespidae of the Guianas, particularly of British Guiana.

SOCIAL WASPS

423

Mus. Comp. Zool. (Harvard Univ.) Bull. 94: 249-304. DÍAS

FILHO,

M.

M.

1975.

Contribuicao

á

morfología de larvas de vespídeos sociais do Brasil (Hymenoptera, Vespidae). Rev. Brasil. Entomol. 19: 1-36. EDERY, H., J. ISHAY, S. GITTER, AND H. JOSHUA.

1978. Venoms of Vespidae. In S. Bettini, ed., Arthropod venoms. Springer, Berlin. Pp. 691-771. HERRÉ, E. A., D. M. WINDSOR, AND R. B. POSTER.

1986. Nesting associations of wasps and ants on lowland Peruvian ant-plants. Psyche 93: 321-330. JEANNE, R. L. 1975. T h e adaptiveness of social wasp nest architecture. Quart. Rev. Biol. 50: 267-287. JEANNE, R. L. 1978. Intraspecific nesting associa­ tions in the Neotropical social wasp Polybia rejecla (Hymenoptera: Vespidae). Biotropica 19: 234-235. JEANNE, R. L. 1980. Evolution of social behav­ ior in the Vespidae. Ann. Rev. Entomol. 25: 371-396. OVERAL, W. L. 1984. Wasp studies among the Kayapó Indians of Brazil. Sphecos 8: 19—22. RICHARDS, O. W. 1978. The social wasps of the Americas excluding Vespinae. Brit. Mus. Nat. Hist., London. RICHARDS, O. W., AND M. J. RICHARDS. 1951.

Observations of the social wasps of South America (Hymenoptera: Vespidae). Royal Entomol. Soc. London Trans. 102: 1-170. WEST-EBERHARD, M. J. 1975. Estudios de las

avispas sociales (Himenoptera, Vespidae) del Valle del Cauca. Cespedesia 4: 245-267. WEST-EBERHARD, M. J. 1989. Scent-trail diver­ sion, a novel defense against ants by tropical social wasps. Biotropica 21: 280-281. WINDSOR, D. M. 1972. Nesting association be­ tween two Neotropical polybiine wasps (Hy­ menoptera: Vespidae). Biotropica 4: 1-3. WINDSOR, D. M. 1976. Birds as predators on the brood of Polybia wasps (Hymenoptera: Vespi­ dae: Polistinae) in a Costa Rican deciduous forest. Biotropica 8: 111-116.

Polistes Paper Wasps Vespidae, Polistinae, Polistini, Polistes. T h i s g e n u s (and tribe) is immediately sepa­ rated from t h e similar Polybiini by t h e s h a p e of t h e slit ( p r o p o d e a l orifice, muscle slit) at t h e r e a r of t h e t h o r a x (actually t h e first a b d o m i n a l s e g m e n t fused t o t h e a b d o ­ m e n ) t h r o u g h which t h e major muscle

424

SAWFLIES, WASPS, ANTS, AND BEES

passes to connect with t h e first waist seg­ m e n t . In t h e Polybiini, t h e orifice U broadly r o u n d e d , never m o r e than twice as long as b r o a d ; in this g r o u p , it is narrow m u c h longer t h a n wide. T h e s e wasps other­ wise resemble most social vespids in their yellow a n d black color p a t t e r n s , moder­ ately long waist, a n d plaited wings. T h e cheeks by t h e eyes tend to be fairly wider a n d m o r e discretely m a r g i n e d than in o t h e r vespids, however. T h e nest is a single e x p o s e d c o m b with­ out a n envelope a n d s u p p o r t e d by a single (sometimes multiple o r b r a n c h e d ) stalk. Colonies a r e small, rarely consisting of m o r e t h a n two h u n d r e d adults. T h e y are usually f o u n d e d by a single female, but often several potentially fecund females may act as co-founders. In t h e latter case, after t h e new nest is established, a domi­ n a n t female assumes t h e q u e e n ' s function by a behavioral competition with the other females. Several may lay eggs, b u t these are eaten by c o m p e t i t o r s until o n e con­ sumes all t h e o t h e r s at a faster rate; this female then becomes t h e q u e e n . This m e t h o d of q u e e n d e t e r m i n a t i o n is the most c o m m o n in t h e g e n u s , b u t in some species (such as the varied p a p e r wasp, see below), o p e n direct struggle decides the o u t c o m e ( E b e r h a r d 1969). T h e g e n u s is fairly large a n d cosmopoli­ tan in distribution. I n t h e New World tropics, t h e r e a r e s o m e 80 species arranged in 5 s u b g e n e r a . T h e y r a n g e widely in all habitats t h r o u g h o u t t h e region, including the West Indies a n d some of t h e more isolated islands (P. dorsalis clarionensis in the Revillagigedo Islands; P. fuscatus o n Ascen­ sion, B e r m u d a , a n d C a p e Verde islands). P. carnifex is t h e largest m e m b e r of the g e n u s (BL to 3 cm) (Corn 1972), and P. canadensis (see below) is probably t h e most c o m m o n . Both a r e found t h r o u g h o u t the Neotropical Region. A general review of the g r o u p is given by W e s t - E b e r h a r d (1983). P. erythrocephalus biology is discussed by Nelson (1971).

References CORN, M. L. 1972. Notes on the biology oí Polistes carnifex (Hymenoptera, Vespidae) in Costa Rica and Colombia. Psyche 79: 150—157. EBERHARD, M. J. 1969. T h e social biology of polistine wasps. Mus. Zool., Univ. Michigan, Misc. Publ. 140: 1-101. NELSON, J. M. 1971. Nesting habits and nest symbionts of Polistes eiythrocephalus Latreille (Hymenoptera: Vespidae) in Costa Rica. Rev. Biol.Trop. 1 8 : 8 9 - 9 8 . WEST-EBERHARD, M. J. 1983. Polistes (quita calzón, lengua de vaca [name of nest], paper wasp). In D. H. Janzen, ed., Costa Rican natural history. Univ. Chicago Press, Chicago. Pp. 758-760.

Varied Paper Wasp Vespidae, Polistinae, Polistini, Polistes canadensis. Spanish: Chia (Costa Rica), panelera, p ú l a t e (Peru). Portuguese: Cavapita, caboclo (Brazil). Jack Spaniard (Trinidad). Populations of this wasp vary greatly in color p a t t e r n from place to place, a n d accordingly, m a n y varieties a n d subspecies have been n a m e d (West-Eberhard 1969). Most are of solid color, from nearly black to reddish-brown (described by B e q u a e r t [1943] as t h e "dirty color of d r i e d blood"). Different a m o u n t s of yellow a n d light brown i n t r u d e o n these solid b a c k g r o u n d s in other forms p r o d u c i n g blotched pat­ terns on t h e t h o r a x o r a b d o m i n a l b a n d s . The status of these entities is still in ques­ tion, although some a r e n o w t h o u g h t to

r e p r e s e n t separate species. All these pat­ tern variations occur o n structurally consis­ tent wasps of m e d i u m size (BL 1 7 - 2 5 mm), with a long, slender a b d o m e n with a pointed tip a n d constriction a b o u t the sixth s e g m e n t (fig. 12.6a). T h i s is t h e most widely d i s t r i b u t e d Polistes species a m o n g t h e a p p r o x i m a t e l y fifty in the New World, f o u n d from Mexico (not C a n a d a , despite its n a m e ) to Patago­ nia (although t h e g e n u s is p r e s e n t o n only a few of the Lesser Antilles). It lives in all habitats, except the driest. Its nest is a single, e x p o s e d c o m b , usu­ ally situated o n a vertical o r strongly in­ clined surface so that it e x t e n d s d o w n w a r d from the pedicel asymmetrically a n d paral­ lel to t h e s u b s t r a t u m (Rau 1943). Sites include m a n - m a d e structures, if available, but in t h e wild a r e normally large limbs, hollows in trees, a n d t h e u n d e r s i d e s of palm fronds. I n outline, t h e nest is a n elongate, oval s t r u c t u r e with n u m e r o u s cells, r e a c h i n g a length of 20 to 30 centime­ ters a n d a width of 10 to 11 c e n t i m e t e r s . T h e larvae line their cells with silk, a n d t h e adults cover t h e pedicel a n d roof of t h e nest with a black tarry substance that waterproofs a n d s t r e n g t h e n s it. T h e r e is a single q u e e n w h o initiates t h e nest b u t w h o is j o i n e d by o t h e r females soon after t h e building begins. Q u e e n d o m i n a n c e is d e t e r m i n e d by fighting, a r a r e p h e n o m e n o n in the g e n u s . Males t e n d

R9ure 12.6 SOCIAL WASPS (VESPIDAE). (a) Varied paper wasp (Polistes canadensis). i (b) Drumming wasp (Synoeca surinama). (c) Parasol wasp (Apoica pallens). (d) Polybia wasp [Polybia scutellaris).

SOCIAL WASPS

425

to stay o n t h e c o m b after e m e r g e n c e m u c h l o n g e r t h a n in most p a p e r wasps. T h e w o r k e r s a r e very protective of the colony a n d quick to attack a n y o n e o r a n y t h i n g c o m i n g too close a n d sting forcefully (Meneses 1969). T h e y s p e n d most of their time, however, seeking nectar, fruit juices, a n d caterpillars to feed to the larvae. In A m a z o n i a , a n u n n a m e d tineid m o t h infests t h e combs. Its larvae scavenge the liquid b o d y wastes that a c c u m u l a t e in the intestines of d e v e l o p i n g adults a n d are ejected w h e n they e m e r g e (meconia) a n d occasionally b u r r o w into cells with wasp p u p a e a n d destroy t h e m ( J e a n n e 1979).

References BEQUAERT.J. 1943. Color variation in the Ameri­ can social wasp Polistes canadensis (Linnaeus), with descriptions of two new forms (Hymenoptera, Vespidae). Bol. Entomol. Venezo­ lana 2: 107-124. JEANNE, R. L. 1979. Construction and utilization of multiple combs in Puli.st.es canadensis in relation to the biology of a predaceous moth. Behav. Ecol. Sociobiol. 4: 293-310. MENESES, B. 1969. Aspectos farmacológicos de la ponzoña de Polist.es canadensis. II. Simp. Foro Biol. Trop. Amazónica. Pp. 399—413. RAU, P. 1943. Nesting habits of Mexican social and solitary Vespidae. Entomol. Soc. Amer. Ann. 36: 515-536. WEST-EBERHARD, M. J. 1969. The social biology of polistine wasps. Mus. Zool. Univ. Mich. Misc. Publ. 140: 1-101.

Drumming Wasps Vespidae, Polistinae, Polybiini, Synoeca. Spanish: C a r a c h u p a avispas (Peru), g u i t a r r e r a (S. septentrionalis, Costa Rica). Portuguese: M a r i m b o n d o tatu, caba tatu (S. cyanea). T h e nests of these species a r e familiar objects a n d have given t h e wasps their c o m m o n n a m e in Brazil, marimbondo tatu, m e a n i n g " a r m a d i l l o " wasp, from the fan­ cied r e s e m b l a n c e of t h e e n v e l o p e to the a r m a t u r e of that m a m m a l . T h e c o m b s are a t t a c h e d directly to the s u b s t r a t u m , usually

426

SAWFLIES, WASPS, ANTS, AND BEES

an inclined tree t r u n k , a n d a r e covered with an elongate, half oval o u t e r envelope T h i s is ribbed transversely a n d has a small access hole at the u p p e r e n d . New combs a n d envelopes are a d d e d at the e n d from time to time, the b o u n d a r i e s of which r e m a i n plainly visible. T h e wasps themselves a r e formidable protectors of their domiciles. T h e y have a p o t e n t v e n o m dispatched with a barbed stinger that r e m a i n s i m b e d d e d as the wasp pulls away. T h e y are not p r o n e to immedi­ ate attack, however. Instead, they displaya w a r n i n g ritual, which may be related to the fact that their sting is b a r b e d a n d only used u n d e r e x t r e m e d u r e s s since it must be sacrificed along with the worker's life in a c o n s u m m a t e attack. W h e n disturbed, the wasps p r o d u c e a r h y t h m i c d r u m m i n g s o u n d by vibrating the envelope from within. T h e s o u n d may be h e a r d several m e t e r s away. If f u r t h e r a r o u s e d , hundreds of individuals r u s h o n t o the nest's outer surface, w h e r e they c o n t i n u e to produce t h u m p i n g s o u n d s a n d raise a n d lower their wings in synchrony. T h e threat is sufficient to scare off all but the most insistent or naive assailants. T h e s e are large (BL 20 m m ) , shiny blue wasps (fig. 12.6b), with a short waist and a constricted segment at the base of the a b d o m e n . T h e latter is h e a r t - s h a p e d , with a s h a r p a p e x ; the apical segments are also somewhat compressed. D r u m m i n g wasps belong to that group of social wasps that found new colonies by swarming. A mass of workers accompanies o n e q u e e n (sometimes several) to the new site. O n e egg-laying female always domi­ nates, repressing the o t h e r s with inhibitory p h e r o m o n e s . Q u e e n s are replaced in tan­ d e m as they expire in their perennial, long-lived colonies (one k n o w n to be 16 years of age). T h e g e n u s Synoeca. is restricted to the New World. T h e five similar species are f o u n d only in tropical, forested areas.

parasol Wasps Vespidae, Polistinae, Polybiini, Apoica. Portuguese: Beiju-cabas, m a r i m b o n d o s de c h a p é u , cabas d e l a d r á o (Brazil). The shape of its nest gives this g e n u s its common n a m e . T h e s t r u c t u r e is an u m ­ brella-shaped, single, o p e n p a p e r c o m b with a r u d i m e n t a r y e n v e l o p e partially cov­ ering the u p p e r surface. It is circular in outline a n d m e a s u r e s u p to 29 centimeters ¡n diameter. It is c o n s t r u c t e d of tightly packed b r a n c h e d plant hairs, giving it a feltlike t e x t u r e . During the day, the colony may b e seen sleeping o n t h e nest, each wasp side by side, f o r m i n g a tightly packed mass a n d covering the e n t i r e lower surface (pi. 4f). Peripheral females align themselves, fac­ ing o u t w a r d , eyes a n d a n t e n n a e f o r m i n g a defense p e r i m e t e r . At night, the workers are active in foraging a n d nest build­ ing and a r e frequently attracted to arti­ ficial light. A l t h o u g h n o c t u r n a l , Apoica will vehemently d e f e n d their nest in the daytime. Nocturnal habits are correlated with the pale colors of these wasps. In the most common species, A. pallens (fig. 12.6c), the abdomen is usually a light c r e a m or yellow hue, the t h o r a x buff with pale spots o n the shoulders a n d h i n d p o r t i o n . T h e body color in o t h e r species is light red to brown. The genus stands a p a r t from o t h e r vespids in having e n l a r g e d ocelli a n d c o m p o u n d eyes that nearly m e e t o n top of the h e a d . The entire b o d y is long a n d slender a n d the a b d o m e n r o u g h l y sausage-shaped with an abruptly n a r r o w e d , short waist. This g e n u s of just six species belongs to that g r o u p of social wasps in which colo­ nies are f o u n d e d by a s w a r m of workers accompanying multiple q u e e n s to the new nest site. Males seem to fly in the swarm as Well, indicating an u n u s u a l a n d not yet understood r e p r o d u c t i v e p a t t e r n . T h e general biology of the g e n u s is

discussed by van d e r Vecht (1973) a n d S c h r e m m e r (1972).

References SCHREMMER, F. 1972. Beobachtungen zur Biologie von Apoica pallida (Oliver, 1791), einer neotropischen sozialen Faltenwespe (Hymenoptera, Vespidae). Ins. Soc. 19: 343-357. VAN DER VECHT, J. 1973. The social wasps (Vespidae) collected in French Guyana by the Mission du Museum National d'Histoire Naturelle with notes on the genus Apoica. Soc. Entomol. France Ann. (n.s.) 8: 735—743.

Polybia Paper Wasps Vespidae, Polistinae, Polybiini, Polybia. Tupi-Guaraní: L a m b o r i n a s (Brazil). Boraseries (Guyana). T h e s e are the most c o m m o n a n d d o m i n a n t social wasps of the Neotropical Region (Richards 1978, W i n d s o r 1983). T h e g e n u s is large (54 species) a n d c o m p o s e d of wasps that are variable in color p a t t e r n a n d o t h e r features (fig. 12.6d). For this reason, they are identified with difficulty, a n d c o m m o n n a m e s must be applied with considerable caution. T h e vulgates of m a n y are d e r i v e d from the shapes of the nest, which a r e fairly diagnostic. T h e tendency for color variation is m a r k e d in most species. G r o u n d color tends to be d a r k (bluish to black) or shades of b r o w n a n d yellow. Often the a b d o m e n is b a n d e d with thin, yellow b o r d e r s to the segments a n d the r e a r p o r t i o n of the thoracic d o r s u m spotted with the same color. T h i s is especially c o n s p i c u o u s in d a r k species, in which the yellow contrasts sharply with the b a c k g r o u n d . T h e wings may be clear to smoky b r o w n . Structural characteristics that distin­ guish this g e n u s are the m o r e or less petiolate a b d o m e n a n d the waist s e g m e n t that is a b o u t as long as the h i n d femur. T h e t h o r a x a n d attached basal s e g m e n t form a smooth, r o u n d e d curve in profile, a n d the entire body is often long a n d s l e n d e r in general form.

SOCIAL WASPS

427

Nest shapes, t h o u g h varied t h r o u g h the g e n u s , a r e consistent a n d diagnostic within species. S h a p e , size, type of construction material, p l a c e m e n t o n the s u b s t r a t u m , c o m b s t r u c t u r e , a n d o t h e r criteria are all particular to each species of wasp (pi. 4c). Most nests are m a d e of p a p e r or carton, as is usual with social wasps, b u t a few species (P. singularis, P. emaciata) f o r m t h e o u t e r enve­ lope of m u d . T h i s may be highly polished a n d is r e f e r r e d to as t h e "ceramic" type. In these, t h e c o m b s a r e still m a d e of p a p e r , however. Nest shapes a n d e n g i n e e r i n g are a source of a d m i r a t i o n . Oval or bell-shaped forms a r e c o m m o n , b u t o n e also finds pears h a p e d , b u n - s h a p e d , s a u s a g e - s h a p e d , stel­ late, cylindrical, a n d o t h e r configurations. T h e o u t e r surface is usually s m o o t h but may also exhibit spiny processes, projec­ tions, flanges, a n d t e x t u r e s of m a n y types. Colors r a n g e widely, too, from buff to gray a n d b r o w n to rusts a n d yellows. T h e nest is n o r m a l l y s u s p e n d e d from tree b r a n c h e s by an apical fastening, b u t that of t h e largest species (P. dimidiala) is situated close to t h e g r o u n d a n d may be pierced by saplings t h a t s u p p o r t it inter­ nally. Many s u s p e n d e d nests a r e small, only a few c e n t i m e t e r s long, b u t o t h e r s may attain lengths of u p to 100 c e n t i m e t e r s or m o r e (P. scutellaris, pi. 4d). Nests of different species a r e s o m e t i m e s f o u n d in close proximity ( J e a n n e 1978). Prey consists of a wide r a n g e of softb o d i e d a r t h r o p o d s , mostly katydids, cater­ pillars, a n d beetle g r u b s . A few species store nectar in select cells, a n d this takes o n a similarity to h o n e y w h e n a b u n d a n t a n d c o n c e n t r a t e d . T h e n e c t a r of P. jurinei is said to be " p e r f u m e d . " Polybia a r e notoriously aggressive a n d p u g n a c i o u s . Most wasp attacks in tropical America a r e from these feisty d e f e n d e r s of their nest a n d territory. A n y o n e molesting a colony will suffer t h e stings of a m a d ­ d e n e d t h r o n g of relentless polybiine p u r s u ­ ers, w h o dive into the hair a n d clothing a n d attack any spot of skin they can reach.

428

SAWFLIES, WASPS, ANTS, AND BEES

H u m a n sweat, however, has been found to s u p p r e s s the stinging u r g e at least in some individuals (Young 1978). Colonies are f o u n d e d by swarming T h e y attain e n o r m o u s populations ¡ n older, larger p e r e n n i a l nests, u p to several t h o u s a n d individuals. Colonies o f f . scutel­ laris may persist as long as twenty-five to thirty years a n d , at any o n e time, may have 4,000 to 5,000 inhabitants. More com­ monly, nests are relatively small with only a few d o z e n or o n e h u n d r e d occupants Females are similar to q u e e n s , although generally larger. In o n e species, P. dimidiata, the reverse is t r u e . References JEANNE, R. 1978. Intraspecific nesting associa­ tions in the Neotropical social wasp Polybia rejecta (Hymenoptera: Vespidae). Biotropica 10: 234-235. RICHARDS, O. W. 1978. The social wasps of the Americas excluding Vespinae. Brit. Mus. Nat. Hist., London. WINDSOR, D. M. 1983. Polybia occidental (Co jones de toro [name of nest], paper wasp). In D. H. Janzen, ed., Costa Rican natural his­ tory. Univ. Chicago Press, Chicago. Pp. 760762. YOUNG, A. M. 1978. A human sweat-mediated defense against multiple attacks by the wasp Polybia diguelana in northeastern Costa Rica. Biotropica 10: 73-74. Bell Wasp Vespidae, Polistinae, Polybiini, Chartergus chartarius. Spanish: Campana-avispa. It is the e n o r m o u s , bell-shaped carton nests of this species that are most often seen, h a n g i n g from trees in a variety of forest habitats. T h e y are especially com­ m o n in the A m a z o n i a n hyalea on trees n e a r river courses. T h e i r beauty and pecu­ liarity have m a d e t h e m well-known tounst trophies that a r e sold in curio shops in South A m e r i c a n cities. Elaborate a n d beautifully engineered s t r u c t u r e s (pi. 4e), they may reach a length of nearly a meter, weigh 10 to 15 kilograms, a n d h a r b o r a colony of over 5,000 individu-

ngure 12.7 SOCIAL WASPS (VESPIDAE). (a) Bell wasp (Chartergus chartarius). (b) Honey wasp (Brachygastra lecheguana). (c) Long-waisted paper wasp (Mischocyttarus drewseni).

als. T h e nest e n v e l o p e is m a d e of a d u r a b l e , cardboardlike substance, t o u g h , thick ( 6 - 7 mm), smooth, a n d dirty white. T h e interior is intricately f o r m e d into several horizontal tiers of vertical cells. T h e fastening enve­ lope at the t o p is wide a n d completely encompasses the b r a n c h s u p p o r t i n g the nest. T h e r e is a single e n t r a n c e in the center of the slightly conical b o t t o m , leading to a single passageway r u n n i n g opposite the entrance t h r o u g h the m i d d l e of each of the stories of combs. T h e wasp (fig. 12.7a) is a fairly typical polybiine, recognizable by its m o d e r a t e size (BL 8 - 1 0 m m ) a n d color p a t t e r n , which is mostly black but with narrow, transverse, yellow b a n d s r u n n i n g across the front a n d h i n d edges of the t h o r a x and b o r d e r i n g the posterior m a r g i n of each a b d o m i n a l s e g m e n t ; the face is yel­ low spotted b e t w e e n the a n t e n n a l base and eye a n d o n the lower p a r t of the frons; and the wing has a clear m e m b r a n e with dark veins. Definitive structural fea­ tures are not c o n s p i c u o u s : r o u n d e d scutellum, short r i d g e s e p a r a t i n g the side of the head from its back, a n d b r o a d , first abjdominal s e g m e n t . This a n d two o t h e r similar species in the nus r a n g e t h r o u g h o u t most of midpical America ( B e q u a e r t 1938). »rence KQUAERT, J. 1938. A new Charter ginus from Costa Rica, with notes on Charterginus, Pseudochartergus, Chartergus. Pseudopolybia, Epi-

pona and Tatua (Hymenoptera: Vespidae) Rev. Entomol. 9: 99-117. Honey Wasps Vespidae, Polistinae, Polybiini, Brachygastra (= Nectarina). Tupi-Guaraní: Eixy, c h i g u a n a , capij (Brazil). Náhuatl: Yzaxalasmitl (Mexico). Mexican b e e . T h e nectar-storing habits of wasps have been perfected by this g r o u p of twelve species ( N a u m a n n 1968), the best k n o w n of which is B. lecheguana, to which most of the following discussion applies (Schwarz 1929). T h e cells of the nest of this species often contain large stores of a very palat­ able h o n e y that is widely exploited by h u m a n s (Bequaert 1933). N u m e r o u s early accounts speak of the use of t h e substance by the I n d i a n s of Mexico a n d Brazil a n d leave n o d o u b t a b o u t the identity of the insect because of its characteristic nest, which has a p a p e r e n v e l o p e a n d cells that are distinct from the quite different en­ claves of stingless bees, the only o t h e r c o m m o n source of native honey in early days. In some areas (e.g., Jalisco), t h e honey is g a t h e r e d regularly a n d even sold on the m a r k e t . It is very limpid a n d strongly scented a n d said to crystallize m o r e rapidly t h a n bees' honey. S o m e I n d i ­ ans k e e p the nests, cutting t h e m w h e n they are small a n d carrying t h e m to their gar­ d e n s . T h e source of the h o n e y is flower nectar g a t h e r e d by the wasps in bee fash­ ion. W h e n Datura or o t h e r toxic blooms a r e

SOCIAL WASPS

429

available, the h o n e y m a y be tainted, a n d cases of p o i s o n i n g are not r a r e . M a t u r e nests are large ( 4 0 - 5 0 cm long) a n d ovoid. W h e n filled with honey, they may weigh several kilograms. T h e r e are usually only a few l a r g e c o m b s (less than 10 b u t as m a n y as 20) a r r a n g e d concentrically within the thin e n v e l o p e . T h e e n t r a n c e is circular or slitlike a n d located t o w a r d the lower e n d . New colonies a r e f o u n d e d by s w a r m i n g , a n d fertile females (polygynous "queens") a r e n u m e r o u s , f o r m i n g u p to 17 p e r c e n t of the total p o p u l a t i o n . T h e n u m ­ ber oí nest i n h a b i t a n t s can b e c o m e very large, as high as 15,000 wasps. T h e wasps survive off-seasons by feeding on their stores of honey, t h e nest b e i n g p e r e n n i a l a n d persistent for several years. It a p p e a r s that the larvae a r e fed exclusively on the honey a n d pollen, a n u n u s u a l diet for wasps. Brachygastra is a typically Neotropical g e n u s , o c c u r r i n g in all b u t the driest habi­ tats, I r o m e x t r e m e s o u t h e r n Texas to n o r t h ­ e r n A r g e n t i n a . Its m e m b e r s are small (BL 7—9 m m ) a n d a p p e a r stubby because of the shape of the a b d o m e n (fig. 12.7b). T h e latter is n o n p e t i o l a t e , wider t h a n long, a n d its second s e g m e n t is greatly e n l a r g e d , often concealing the s u c c e e d i n g s e g m e n t s . T h e elevated scutellum, which often p r o ­ jects over t h e rest of the posterior of the t h o r a x , is also highly characteristic; the r e a r p o r t i o n of this f o r m s a flat, vertical surface. Color p a t t e r n s r a n g e from almost all black to solid yellow with m a n y i n t e r m e ­ diate p a t t e r n s of both. W o r k e r s are mild m a n n e r e d by polybiine s t a n d a r d s b u t sting h a r d w h e n suffi­ ciently p r o v o k e d . T h e sting is b a r b e d a n d stays in the w o u n d if t h e victim is a h u m a n or o t h e r large animal.

References BEQUAERT, J. 1933. T h e nearctic social wasps of the subfamily Polybiinae (Hymenoptera; Vespidae). Entomol. Americana 13: 87-150. NAUMANN, M. 1968. A revision of the genus

430

SAWFL1ES, WASPS, ANTS, AND BEES

Brachygastra (Hymenoptera: Vespidae) \jn4 Kans. Sci. Bull. 47: 929-1003. ' SCHWARZ, H. F. 1929. Honey wasps. Nat Hi L 39:421-426. '

Long-Waisted Paper Wasps Vespidae, Polistinae, Polybiini, Mischocyttarus. Tupi-Guaraní: C a p u x u (Brazil, M. ater). Mischocyttarus is the largest g e n u s of social wasps (186 species) a n d has achieved its diversity entirely in the New World, primar­ ily the tropical p o r t i o n s (Richards 1945 Zikán 1949). It is also a morphologically distinct g e n u s , characterized by a rather long waist a n d asymmetrical terminal seg­ m e n t s of the mid a n d h i n d tarsi, the inner lobes m u c h m o r e strongly projecting and p o i n t e d t h a n the outer. T h e latter is possi­ bly an a d a p t a t i o n to assist the wasp in searching for prey c a u g h t in spider webs. T h e d o r s u m of the t h o r a x is evenly curved a n d , in profile, slopes directly onto the attached basal a b d o m i n a l segment. There is also a characteristic conspicuous yellow spot p r e c e d i n g a d a r k patch at the apex of the hind tibia on the inside, which may serve as a recognition signal when the wasps fly n e a r the nest with hind legs dangling. T h e s e wasps otherwise generally resemble Polistes in b o d y coloring and shape but are consistently smaller (BL 1015 m m ) . All m e m b e r s of the g e n u s make small (rarely m o r e t h a n 100 cells), stalked, un­ covered combs s o m e w h a t like those of Polistes, a l t h o u g h in many, the combs may be t u r n e d to face the substrate and may be irregularly layered in series rather than horizontally. T h e r e also may be more than o n e s u p p o r t i n g stalk. T h e latter is var­ nished with a secretion from a gland on the a b d o m e n that has been shown in some species to be a n ant repellent. Nest sites of some species are carefully situated amid spiny plants or n e a r o t h e r wasps for maxi­ m u m protection (Gorton 1978).

Colonies are usually f o u n d e d by a single female (haplometrosis) or in c o m p a n y with few female n e s t m a t e s (pleometrosis) iPoltronieri a n d R o d r i g u e s 1976). Working 0 g e n y soon d e v e l o p a n d a d o m i n a n c e hierarchy e n s u e s . S u b o r d i n a t e females may eventually oust the q u e e n a n d take over the colony (Little 1981). A s t r o n g trophallactic b o n d exists between workers and q u e e n s in o n e species ( M a c h a d o a n d Wiendl 1976). T h e peculiar biology of M. drewseni (fig. j2_7c) has b e e n given s o m e direct attention (leanne 1972, J e a n n e a n d Castellón Berm ú d e z 1980).

References GORTON, R- E. 1978. Observations on the nest­ ing behavior of Mischocyttarus immarginatus (Rich.) (Vespidae: Hymenoptera) in a dry forest in Costa Rica. Ins. Soc. 25: 197-204. JEANNE, R. L. 1972. Social biology of the Neo­ tropical wasp Mischocyttarus drewseni. Mus. Comp. Zool. (Harvard'Univ.) Bull. 144: 6 3 150. IJEANNE, R. L., AND E. G. CASTELLÓN BERMÚDEZ.

" 1980. Reproductive behavior of a male Neotropical social wasp, Mischocyttarus drew­ seni (Hymenoptera: Vespidae). Kans. Ento­ mol. Soc. J. 53: 271-276. LITTE, M. 1981. Social biology of the Polistine wasp Mischocyttarus labiatus: Survival in a Co­ lombian rain forest. Smithsonian Contrib. Zool. 327: 1-27. CHADO, V L . L . , AND F. M . WlENDL. 1 9 7 6 .

Aspectos do comportamento de colonias de Mischocyttarus cassununga von lhering, trata­ das com alimento marcado por radioíósforo. Soc. Entomol. Brasil Ann. 5: 79-85. .TRONIERI, H .

S.,

AND V. M .

RoDRÍGUES.

). Vespídeos sociais: Estudo de algumas especies de Mischocyttarus Saussure, 1853 (Hymenoptera-Vespidae-Polistinae). Dusenia 9:99-105. XHARDS, O. W. 1945. A revision of the genus Mischocyttarus de Saussure (Hymen., Vespi­ dae). Royal Entomol. Soc. London Trans. 95: 295-462. ^N, J. F. 1949. O género Mischocyttarus aussure (Hym.-Vespidae), com a descricáo «e 82 novas especies. Par. Nac. Itatiaia Bol. 1: 1-251.

ANTS Formicidae. Spanish: H o r m i g a s . Portuguese: Formigas. Tupi-Guaraní: Q u e n q u e m . Náhuatl: A z c a m e h (sing. azcatl). Quechua: Sisi, cuki. If a n y o n e d o u b t s who is in possession of the Earth, let t h e m consider the ant (Hólldobler a n d Wilson 1990, W h e e l e r 1910, Wilson 1971: 2 7 - 7 4 ) . It is h a r d l y possible to find a place, particularly in the A m e r i c a n tropics, w h e r e ants a r e n o t visi­ ble, often a b u n d a n t l y a n d obnoxiously so (although they are conspicuously absent at high elevations). Because of their small size, social s t r u c t u r e , a n d adaptiveness, they have evolved into a fairly large n u m ­ ber of species in Latin America, w h e r e well over a third of the world's 7,600 k n o w n species are f o u n d ( K e m p f 1972). T h e s e are currently divided into 116 g e n e r a (Kusnezov 1978), r e p r e s e n t i n g nearly 50 p e r c e n t of the world's total. Many of these are k n o w n only to ant specialists, but m a n y a r e familiar pests. Yet, destructive t h o u g h some m a y be, most ants d e s e r v e o u r a p p r e ­ ciation for the roles they play in controlling o t h e r injurious insects, r e g u l a t i n g vegeta­ tive g r o w t h , a n d increasing the fertility of the soil ( B r a n n e r 1900). Resemblance of the social o r g a n i z a t i o n of ants to h u m a n society has e a r n e d t h e m special significance to the I n d i a n . I n d e e d , ants hold central, even r e v e r e d , positions in the mythologies of m a n y tribes. I n earlier days, such was the d o m i n a n c e of these insects in their forest e n v i r o n m e n t t h a t the K a i n g a n g of s o u t h e a s t e r n Brazil believed the glory of their p e o p l e to be derived from being t r a n s f o r m e d into ants after d e a t h (Lenko a n d P a p a v e r o , 1979: 225). A m o n g the Kayapó, even today, ants are s p o k e n of in t e r m s of their "power," or ability to inflict pain, which is u s e d by s h a m a n s to m a n i p u l a t e spirits to cause h a r m . Stinging ants are collected by the m e n , p o u n d e d into a paste with r e d u r u c u dye a n d painted o n h u n t i n g dogs to m a k e

ANTS

431

t h e m k e e p their noses to t h e g r o u n d a n d h u n t with d e t e r m i n a t i o n as t h e ants d o (Posey 1979: 143). Lest we be a m u s e d by such beliefs a m o n g simpler c u l t u r e s , c o n s i d e r t h e friars of t h e C o n v e n t of San A n t o n i o at M a r a n háo, Brazil, w h o p r o s e c u t e d s o m e local r e d ants u n d e r ecclesiastical law. T h e insects h a d d e v o u r e d t h e altar cloths a n d b r o u g h t u p into t h e chapel pieces of s h r o u d s from the c h u r c h y a r d graves (Cowan 1865: 168). Ants a r e easily distinguished from o t h e r H y m e n o p t e r a by t h e u n i q u e n a r r o w e d "waist" (petiole) c o n n e c t i n g t h e a b d o m e n a n d t h o r a x . T h e waist m a y consist of o n e or two s e g m e n t s , o n e o r both of which often have a s t r o n g , erect projection, o r " n o d e , " arising from it. T h i s a r r a n g e m e n t prevails w h e t h e r t h e a n t is a wingless, sterile female w o r k e r o r a winged, r e p r o ductively active a d u l t m a l e o r female. Ants a r e also recognized by t h e i r elbowed a n t e n ­ nae (the basal s e g m e n t of which is very long), usually well-developed m a n d i b l e s , a n d large mobile p r o t h o r a x . Equally characteristic of t h e family is their habit of living in colonies in a highly o r g a n i z e d social state. I n most species, this has led t o a division of labor correlated with differing m o r p h o t y p e s called "castes." T h e r e p r o d u c t i v e castes a r e t h e males, w h o die after m a t i n g , a n d t h e q u e e n s , w h o establish new colonies after m a t i n g a n d s h e d d i n g their wings. T h e bulk of t h e p o p u l a t i o n consists of sterile females w h o form a series of w o r k e r types from t h e smallest " m i n o r s " to t h e largest "majors." T h e latter, as a result of d i s p r o p o r t i o n a t e (allometric) b o d y g r o w t h , may d e v e l o p o u t sized h e a d s a n d j a w s used in defense of t h e colony a n d a r e functional "soldiers." Colonies live in nests c o n s t r u c t e d of a variety of materials a n d located in diverse sites, both terrestrial a n d a r b o r e a l (Wilson 1987). Most a r e excavations in t h e soil, sometimes simple c h a m b e r s at t h e e n d of a direct e n t r a n c e ; in o t h e r instances, they

432

SAWFIJES, WASPS, ANTS, AND BEES

are e n o r m o u s , complex, u n d e r g r o u n d labyrinths. Nests a r e also m a d e of s o ;i particles o r chewed wood fragments glu»H to vegetation (carton nests) o r a r e intri­ cately woven of silk (Camponotus senex). \ special category of t h e f o r m e r a r e those attached to leaves (Black 1987). Many spe­ cies live in hollowed-out plant structures a n d some even have d e v e l o p e d symbiotic relationships with their hosts (see ants and plants, following). Ants m a k e use of elaborate communica­ tion systems, based o n chemical messenger chemicals p r o d u c e d by various glands, to organize their foraging a n d o t h e r commu­ nal activities. Such a r e t h e trail-making substances laid d o w n from anal glands alarm o d o r s p r o d u c e d by t h e mandibular gland in t h e head a n d a b d o m i n a l Dufour's gland, noxious o d o r s o r antifungal com­ p o u n d s secreted by t h e m e t a p l e u r a l glands in t h e t h o r a x , a n d v e n o m s for subduing prey a n d defense against enemies which are g e n e r a t e d by v e n o m glands associated with t h e sting. Food often consists of special types of prey o r plant p r o d u c t s , t h e exploitation of which r e q u i r e s cooperation a n d initiative a m o n g colony m e m b e r s (Sudd 1967). Al­ most any edible material m a y be taken: living o r d e a d a r t h r o p o d s a n d small verte­ b r a t e animals, fungi grown o n leaf frag­ ments, seeds, flower pollen a n d nectar (including that from extrafloral nectaries), a n d honeydew, which they take from scale insects, aphids, a n d t r e e h o p p e r s . Food is s h a r e d with t h e whole colony by foraging workers. T h i s includes t h e help­ less larvae a n d t h e q u e e n , which delivers p h e r o m o n a l secretions in exchange. T h e m o v e m e n t of such substances t h r o u g h the colony (trophallaxis) acts as a b o n d , keep­ ing t h e colony m e m b e r s t o g e t h e r a n d regu­ lating t h e physiology a n d behavior of its m e m b e r s . A variety of diverse insects often s h a r e in t h e food e x c h a n g e a n d form symbiotic b o n d s with t h e colony. T h e r e are

the so-called a n t guests, o r " m y r m e c o ohiles," s u c n a s silverfish, crickets, cockióaches, beetles, flies, a n d even mites, spiders, a n d millipedes (see symbiosis, rhaP- 2)- C o o p e r a t i v e o r mutualistic associa­ tions a r e also c o m m o n between ants a n d h o n e y d e w - p r o d u c i n g a p h i d s , scale insects, gnd other H o m o p t e r a (Way 1963). Secondary only to their ubiquity a n d sociality, ants attract attention with their ability to inflict w o u n d s . Many have stings, gome, especially species a m o n g t h e primi­ tive tribes, very p o t e n t o n e s . C o n t r a r y to popular belief, formic acid is only o n e , minor, constituent of a n t v e n o m , except in the subfamily Formicinae w h e r e it p r e ­ dominates. I n most ants, various alkaloids and protein derivatives a r e m u c h m o r e important in p r o d u c i n g lasting a n d debili­ tating effects. T h e s e a r e only n o w being identified a n d u n d e r s t o o d by insect p h a r ­ macologists (Schultz a n d A r n o l d 1977). Because they sting o r a r e m a l o d o r o u s , ants serve as m o d e l s for Batesian mimics. They a r e closely r e s e m b l e d in form a n d behavior by stilt flies (Micropezidae), i m m a ­ ture assassin b u g s , a d u l t grass b u g s , wee­ vils, a n d o t h e r insects, as well as spiders (Jackson a n d D r u m m o n d 1974). T h e p h e n o m e n o m of " a n t mosaics," that is, colonies of a b u n d a n t , static species that compete to establish foraging territo­ ries in vegetation d o m i n a t e d by trees (first discovered in West Africa), is t h o u g h t t o occur in some tropical A m e r i c a n habitats (Jackson 1984; Leston 1978; W i n d e r 1978; Young 1983). Mosaics m a y b e i m p o r t a n t in the cultivation of tropical tree crops, for many pests a n d plant diseases a r e associ­ ated with o n e o r m o r e of t h e d o m i n a n t a n t •pecies. Most of t h e subfamilies of t h e Fornricidae a r e r e p r e s e n t e d in t h e A m e r i c a n tropics (Alayo 1974; Kusnezov 1963; Smith 1936; Snelling a n d H u n t 1976). T h e s e a r e e Ponerinae (primitive h u n t i n g ants), toninae (New World a r m y ants), Pseudo-

m y r m e c i n a e (fever ants), Myrmicinae (myrmicine ants), Dolichoderinae (odiferous ants), a n d Formicinae (formicine ants).

References ALAYO, D. P. 1974. Introducción al estudio de los himenópteros de Cuba, Superfamilia Formicoidea. Acad. Cien., Cuba, Inst. Zool. Ser. Biol. 53: 1-58. BLACK 111, R. W. 1987. T h e biology of leaf nesting ants in a tropical wet forest. Biotropica 19: 319-325. BRANNER, J. C. 1900. Ants as geologic agents in the tropics. J. Geology 8: 151-153. COWAN, F. 1865. Curious facts in the history of insects. LippincoU, Philadelphia. HOLLDOBLER, B., AND E. O. WILSON. 1990. T h e

ants. Harvard Univ. Press, Cambridge. JACKSON, D. 1984. Competition in the tropics: Ants on trees. Antenna 8(1): 19-25. JACKSON, J. F , AND B. A. DRUMMOND 111. 1974.

A Batesian ant mimicry complex from the Mountain Pine Ridge of British Honduras, with an example of transformational mim­ icry. Amer. Midi. Nat. 91: 248-251. KEMPF, W. W. 1972. Catálogo abreviado das formigas da Regiáo Neotropical (Hym. Formicidae). Studia Entomol. 15: 3-344. KUSNEZOV, N. 1963. Zoogeografía de las hormi­ gas en Sudamerica. Acta Zool. Lilloana 19: 25-186. KUSNEZOV, N. 1978. Hormigas argentinas-clave para su identificación. Min. Cult. E d u c , Fundación Miguel Lillo, Mise. 61: 1-147. LENKO, K., AND N. PAPAVERO. 1979. lnsetos no

folclore. Cons. Estad. Artes Cien. Hum., Sao Paulo. LESTON, D. 1978. A Neotropical ant mosaic. Entomol. Soc. Amer. Ann. 71: 649-653. POSEY, D. A. 1979. Ethnoentomology of the

Gorotire Kayapó of central Brazil. Ph.D. diss., Univ. Georgia. SCHULTZ, D. R., AND P. I. ARNOLD. 1977. Venom

of the ant Pseudomyrmex sp.: Further character­ ization of two factors that affect human complement proteins. J. Immunol. 119: 1690-1699. SMITH, M. R. 1936. T h e ants of Puerto Rico. Univ. Puerto Rico J. Agrie. 20: 819-875. SNELLING, R., AND J. H U N T . 1976. T h e ants of

Chile. Rev. Chilena Entomol. 9: 63-129. SUDD, J. 1967. An introduction to the behavior of ants. Arnold, London. WAY, M. J. 1963. Mutualism between ants and

ANTS

433

honeydew producing Homoptera. Ann. Rev. Entomol. 8: 307-344. WHF.F.LER, W. M. 1910. Ants, their structure, development and behavior. Columbia Univ., New York. WILSON, E. O. 1971. Insect societies. Belknap Press, Harvard Univ., Cambridge. See esp. pp. 27-74. WILSON, E. O. 1987. The arboreal ant fauna of Peruvian Amazon forests: A first assessment. Biotropica 19: 245-251. WINDER, J. A. 1978. The role of non-dipterous insects in the pollination of cocoa in Brazil. Bull. Entomol. Res. 68: 559-574. YOUNG, A. M. 1983. Patterns of distribution and abundance of ants (Hymenoptera; Formicidae) in three Costa Rican cocoa farm locali­ ties. Sociobiology 8: 52—76.

ANTS AND PLANTS Q u i t e a few u n r e l a t e d kinds of woody plants in the A m e r i c a n tropics have evolved s t r u c t u r e s a d a p t i n g t h e m for habitation by ants (Beattie 1985, W h e e l e r 1942). S o m e have existed so long in close association with their formicid guests that an obligatory m u t u a l i s m between p l a n t a n d insect exists ("myrmecophytes"). O t h e r associations are merely casual o r facultative, a l t h o u g h they may r e p r e s e n t incipient symbioses d e m o n ­ strating how the most highly evolved cases may have o r i g i n a t e d (Jolivet 1986). M y r m e c o p h y t e s a r e f o u n d a m o n g sev­ eral plant families a n d g e n e r a . All are similar in having at least s o m e p a r t of the stem, t r u n k , or leaf e n l a r g e d a n d hollowed to p r o v i d e living q u a r t e r s for ant colonies (formicarium, m y r m e c o d o m a t i u m ) . Some also have special tissues or glands that provide food (pearl bodies, Beltian bodies, etc.) o r nectar sources (extrafloral nec­ taries) for the ants a n d n o d o u b t assure t h e ants' d e p e n d e n c e o n , a n d jealous p r o t e c ­ tion of, the host plant. O n this level, the best e x a m p l e is the swollen t h o r n (bull's h o r n ) acacias (Mimosaceae, Acacia corní­ gera, a n d m a n y o t h e r species k n o w n by m a n y n a m e s , e.g., cornizuelas, palines, a n d

434

SAWFL1ES, WASPS, ANTS, AND BEES

guisadles corteñas in Mexico a n d cachitos in P a n a m a ) ( J a n z e n 1969). T h e s e are small trees or shrubs with delicately p i n n a t e c o m p o u n d leaves and large bifurcate t h o r n s that the ants enter by g n a w i n g an o p e n i n g j u s t below the tin of o n e of the pair a n d hollowing them out Extrafloral nectaries situated o n the upper side of the petioles a n d at the base of the p i n n a e w h e r e they j o i n the rachis provide sweet attractions for the ants, a n d there are elongate whitish food bodies (Beltian bod­ ies) p r o d u c e d at the tips of the youne leaflets, which the ants collect, store, and eat. Swollen t h o r n acacias a r e most abun­ d a n t in Mexico a n d C e n t r a l America; they d o not e x t e n d south b e y o n d Venezuela a n d Colombia. T h r o u g h o u t their range, they live in mutualistic associations with several species of ants of the genus Pseudomyrmex ( J a n z e n 1966, 1967). Work­ ers of these ants begin to patrol the limbs a n d leaves about n i n e m o n t h s following colonization by new q u e e n s a n d effectively protect it from the ravages of browsing m a m m a l s , vegetarian insects, and even vines whose tendrils a t t e m p t to entwine it. T h e y r e w a r d h u m a n i n t r u d e r s with a sting whose potency rivals that of some ponerine h u n t i n g ants. A similar relationship exists between several Azteca ants a n d the cecropia tree (Moraceae, Cecropia adenopus a n d other spe­ cies), also called the t r u m p e t tree, imbaúba (Brazil), cetico (Peru), g u a r u m o (Central America), a n d y a g r a m (Antilles). H e r e the jointed t r u n k s of small trees are the ant dwellings. T h e i n t e r n o d e s are hollow and s e p a r a t e d from each o t h e r by partitions. Externally, these j o i n t s are m a r k e d by a n o d e from which arises a leaf, and below each leaf base, t h e r e is a groove enclosing a d e e p pit. T h e f o u n d i n g q u e e n perforates the t r u n k by g n a w i n g t h r o u g h this pit and gaining e n t r a n c e to the c h a m b e r inside, t h e r e establishing a colony that grows to occupy adjoining c h a m b e r s when the sepa-

rating partitions a r e drilled t h r o u g h by orogeny w o r k e r s . I n the interior, the ants build a kind of c a r t o n nest for their larvae out of a b r o w n , waxlike substance. Like the acacias, the cecropias p r o v i d e nutritive bodies for the ants. At the base of each leaf petiole, t h e r e is a velvety brown cushion in which e g g - s h a p e d , food-rich structures (Müllerian bodies, "pearl bod¡ e s "; O ' D o w d 1982) a r e p r o d u c e d . T h e ants also coax h o n e y d e w from the white scale insects o r mealybugs that invariably also cohabit in the t r u n k s a n d stems of this tree. When a cecropia is d i s t u r b e d , its ant guardians boil o u t of their c h a m b e r s in attack. Bereft of a sting, they r u b noxious anal secretions in w o u n d s m a d e by their mandibles a n d j u s t as effectively discour­ age the plant's e n e m i e s as their a r m e d cousins. T h i s p a r t n e r s h i p is not ubiqui­ tous; it b r e a k s d o w n u n d e r some circum­ stances (de A n d r a d e a n d C a r a u t a 1982) at high elevations a n d o n some islands w h e r e large plant b r o w s e r s are absent ( J a n z e n 1973). Other ant plants have simpler relation­ ships with ants. Lacking nutritive bodies and extrafloral nectaries is the South American t a n g a r a n a , a n a m e applied in Peru by natives to both the tree a n d its protector ants. T h e r e p u t a t i o n of the ants has e a r n e d t h e tree (Polygonaceae, Triplaris americana a n d o t h e r species in the genus) a host of telling n a m e s : pau-sanlo, pau de novato, pau formigueiro (Brazil); tachizeiro (Para); itassi, L o n g J o h n (Guyana); jacuna (Arawak); hormigo, tabaco (Costa Rica); árvore de tachi (Amazonia); palo María, tarabas (Venezuela). During its flowering period in the dry season, Triplaris is a large a n d decorative tree that grows along w a t e r c o u r s e s in the wet forests of t h e n o r t h e r n half of the South A m e r i c a n continent. Its flowers are •mall and insignificant in themselves, but We conspicuous calyxes form d e n s e white •"asses that t u r n bright r e d , t r a n s f o r m i n g

the s l e n d e r - t r u n k e d trees into masses of flame. Flame would also describe the c h a r a c t e r of the ants associated with the t a n g a r a n a tree. T h e s e mostly belong to the g e n u s Pseudomyrmex (Latinodus G r o u p ) , but some Azteca are also k n o w n from this plant. Local n a m e s for the f o r m e r are fever ants (general), tachi (Amazonia), novato (Para), a n d jacuna sae (Arawak). T h e small t e r m i n a l b r a n c h e s a n d twigs are hollow a n d occupied by the ants, which obtain e n t r y by cutting t h r o u g h the thin tissue in the base of slitlike scars located toward the far e n d of each i n t e r n o d e . Tales of the attacks of these ants are legend a m o n g the I n d i a n s a n d were k n o w n even to the earliest jungle e x p l o r e r s . O n e of the earliest accounts is that of P. B e r n a b é C o b o in 1653, who wrote in his History of the New World ( 1 8 9 0 - 1 8 9 5 ) , "Since these ants a r e concealed within the tree, they a r e not seen a n d this is the reason why those w h o d o not know the secret, are not o n their g u a r d ; but if a single leaf be t o u c h e d , so m a n y of the ants swarm forth from all parts of the tree as to excite wonder, a n d they assail t h e person w h o touches the tree a n d , if h e does not withdraw in time, m a r t y r him with their stings." T h e s e same ant g e n e r a a n d o t h e r s (Solenopsis, Crematogaster) a n d still o t h e r plants have f o r m e d intricate p a r t n e r s h i p s , with different parts of the plant's a n a t o m y modified as formicaries. In Tachigaiia (Caesalpinaceae), the bases of the petioles of the large p i n n a t e leaves are swollen a n d hollow; access to the interior is t h r o u g h an elongate slit on o n e side (Wheeler 1921a). Bulbous swellings immediately below a n d s u r r o u n d i n g the n o d e s on the stems of certain Cordia species (Boraginaceae) are also the abodes of noxious ants (the g e n e r a already n a m e d plus Allomerus a n d Azteca; pi. 4g), which move in a n d out t h r o u g h a single o p e n i n g between the opposite leaves at o n e e n d of t h e n o d e . O n e side of the bases of the large velvety leaves of Tococa

ANTS AND PLANTS

435

(and o t h e r g e n e r a of Melostomaceae) are inflated, d o u b l e , globular c h a m b e r s har­ b o r i n g these formicids (pi. 3c). A pair of tiny holes at the bases of the forks of the t h r e e s t r o n g m a i n veins o n the u n d e r s i d e of the leaf p r o v i d e access to the c h a m b e r . A n o t h e r possibly coevolved mutualistic relationship exists b e t w e e n Pheidole ants a n d plants of the g e n u s Piper (Piperaceae) ( L e t o u r n e a u 1983, Risch et al. 1977). T h e ants live in petiolar cavities a n d in the stems, which they hollow out. T h e plant provides lipid-rich food bodies inside the petiolar cavities, a n d the ants a p p e a r to increase the survival of the plants by p r o ­ tecting t h e m from e n c r o a c h i n g vines. T h e plant may also receive nutritional benefit by a b s o r b i n g decay p r o d u c t s from the nest. Still o t h e r plants a r e k n o w n with similar but ill-defined symbiotic relationships to the Formicidae. For e x a m p l e , extrafloral nectaries on m a n y species (e.g., Bixa, Costus, Inga, Byttneria) attract ants whose pres­ ence may d i s c o u r a g e t h e attacks of herbi­ vores (Bentley 1977; H e s p e n h e i d e 1985; K o p t u r 1984; S c h e m s k e 1982). However, H e s p e n h e i d e (1984) thinks that the protec­ tion of the plant by parasitoid wasps at­ tracted to the nectaries may be m o r e i m p o r ­ tant. T h e s e wasps m i g h t t h e n significantly r e d u c e the incidence of p h y t o p h a g o u s cat­ erpillar attacks o n the plant hosts, but this has not yet b e e n amply d e m o n s t r a t e d . Ants may inhabit the stems of almost any hollow-stemmed plant a n d nest in all kinds of cavities in roots, b a r k , d e a d twigs, or parts of practically any woody type. But only those plants that h a v e a d a p t e d to the presence of ants by evolving formicaries, a n d some also by p r o v i d i n g nutritive bod­ ies, can be c o n s i d e r e d t r u e m y r m e c o p h y t e s . L a b y r i n t h i n e formicaries in the p s e u d o bulbs of orchids a n d g r o w i n g at the base of o t h e r e p i p h y t e s have b e e n described from the forests of Asia b u t h a v e not b e e n f o u n d with certainty in tropical America. T h e i r i m p o r t a n c e in the n u t r i t i o n of air plants on n u t r i e n t - p o o r substrates has b e e n d e m o n ­

436

SAWFLIES, WASPS, ANTS, AND BEES

strated a n d o u g h t to have evolved in the New World w h e r e it may yet be found. An additional curious intimacy between ants a n d plants are the so-called ant gar­ d e n s (pi. 4h). Species of Azteca, Crematogaster, Dolichoderus, a n d Camponotus in the A m a z o n i a n a n d o t h e r lowland forests form spongelike nests on tree branches from particles of soil. Sometimes pairs of species live t o g e t h e r in such structures ("parabiosis"; W h e e l e r 19216). Epiphytic plants of several types take root in these nests, which forms a kind of nutritional "potting mix," a n c h o r i n g a n d fostering their growth. Species of Codonanthe (Gesneriaceae) are the best-known benefactors of this a r r a n g e m e n t , even t e m p t i n g the ants to disperse their seeds by their shape a n d / o r o d o r that mimics those of ant lar­ vae. O t h e r plants in such associations are Peperomia, Anthurium, Philodendron, Epiphyllum, Markea, a n d various bromeliads. T h e ant benefits from a constant fruit and nectar supply as well as structural support of their nest by the plant's roots. T h e ants d o not a p p e a r to pollinate the plants but may protect t h e m from herbivores and possibly even actively plant their seeds in the nest matrix (Kleinfeldt 1978).

References BEATTIE, A. J. 1985. The evolutionary ecology of ant-plant mutualisms. Cambridge Univ. Press, New York. BENTLEY, B. L. 1977. Extrafloral nectaries and protection by pugnacious bodyguards. Ann. Rev. Ecol. Syst. 8:^407-427. COBO, B. 1890-1895. Historia del Nuevo Mundo. 4 vols. Marcos Jiménez de la Espada, Sevilla. DE ANDRADE, J. C , AND f. P. P. CARAUTA.

1982.

The Cecropia-Azteca association: A case oí mutualism. Biotropica 14: 15. HESPENHEIDE, H. A. 1984. Agrilu.s xanthonotus (yellow-spotted Byttneria borer). In D. H. Janzen, ed., Costa Rican natural history. Univ. Chicago Press, Chicago. Pp. (581—682. HESPENHEIDE, H. A. 1985. Insect visitors to extrafloral nectaries of Byttneria aculeata (Sterculiaceae): Relative importance and roles. Ecol. Entomol. 10: 191-204.

JANZEN, D. H. 1966. Coevolution of mutualism between ants and acacias in Central America. Evolution 20: 249-275. JANZEN, D. H. 1967. Interactions of the bull's horn acacia (Acacia cornígera L.) with an ant inhabitant (Pseudomyrmex femiginea F. Smith) in eastern Mexico. Univ. Kans. Sci. Bull. 47: 315-558. JANZEN, D. H. 1969. Allelopathy by myrmeco­ phytes: The ant Azteca as an allelopathic agent of Cecropia. Ecology 50: 147—153. JANZEN, D. H. 1973. Dissolution of mutualism between Cecropia and its Azteca plants. Bio­ tropica 5: 15—28. IOLIVET, P. 1986. Les fourmis et les plants—un example de coevolution. Soc. Nouv. Ed., Boubée, Paris. KLEINFELDT, S. E. 1978. Ant-gardens: The inter­ action of Codonanthe cmssifolia (Gesneriaceae) and Crematogaster longispma (Formicidae). Ecology 59: 449-456. KOPTUR, S. 1984. Experimental evidence for defense of Inga (Mimosoideae) saplings bv ants. Ecology 65: 1787-1793. LETOURNEAU, D. K. 1983. Passive aggression: An alternative hypothesis for the PiperPheidole association. Oecologia 60: 122—126. O'DOWD, D. J. 1982. Pearl bodies as ant food: An ecological role for some leaf emergences of tropical plants. Biotropica 14: 40—49. RISCH, S.,

M.

M C C L I R E , J. VANDERMEER, AND

S. WALTZ. 1977. Mutualism between three species of tropical Piper (Piperaceae) and their ant inhabitants. Amer. Midi. Nat. 90: 4 3 3 444. SCHEMSKE, D. W. 1982. Ecological correlates of a Neotropical mutualism: Ant assemblages at Costus extrafloral nectaries. Ecologv 63: 932— 941. WHEELER, W. M. 192 la. The tachygalia ants. Zoológica 3: 137-168. WHEELER, W. M. 19216. A new case of parabiosis and the "ant gardens" of British Guiana. Ecology 2: 89-103. WHEELER, W. M. 1942. Studies of Neotropical ant-plants and their ants. Mus. Comp. Zool. (Harvard Univ.) Bull. 90: 1-262.

q u a t r o s , formigas d e febre, formigas d e q u a t r o picadas (Brazil). T h e following two best-known a n d largest of the gigantic forest ants are often con­ fused as o n e a n d with o t h e r large, black species u n d e r the n a m e tocandira. Accord­ ing to Sampaio (Lenko a n d P a p a v e r o 1979), this appellation is a contraction of tucaha-ndy, or tucana-ngia, in Tupi for "the o n e that h u r t s m u c h with its u n d e r p a r t s , " an obvious reference to the e x t r e m e l y severe sting they can inflict. T h e smaller Paraponera a r e the m o r e aggressive a n d potent a n d can s u b d u e a healthy adult with a v e n o m o u s stab that is often described as "like a blow from a h a m m e r or bullet." T h e serious s y m p t o m s that may (but not invari­ ably) follow a sting a n d last a day or l o n g e r are p r o l o n g e d aching pain that rapidly s p r e a d s from the site of t h e w o u n d , some­ times labored b r e a t h i n g , cardiac palpita­ tions, a n d fever (Weber 1939, orig. obs.). It is alleged that d e a t h is an o u t c o m e in r a r e instances, a l t h o u g h n o such cases a r e d o c u ­ m e n t e d in the literature. T h e larger Dinoponera are gentler ants, less apt to sting a n d with less effect t h a n the fiercer Paraponera ( H e r m a n n et al. 1984).

GIANT HUNTING ANTS

Because of their ferocity a n d ability to inflict pain, these ants are e m p l o y e d in male initiation a n d virility rites a m o n g some A m a z o n i a n I n d i a n g r o u p s (Liebrecht 1886). Large n u m b e r s of ants a r e caught a n d tied to specially contrived wicke r w o r k panels a n d applied to the initiate's bare skin (see fig. 1.12). T h e m a d d e n e d mass of ants n e e d no i n d u c e m e n t to sting a n d d o so with impatience a n d repeatedly. Only youths w h o e n d u r e the excruciating e x p e r i e n c e without complaint (and sur­ vive) a r e d e e m e d worthy of passage to manhood.

Formicidae, P o n e r i n a e , Dinoponera a n d Paraponera. Spanish: C a l e n t u r a s , p e r r o s negros, j a u l a s (Peru). Portuguese: Tocandiras (vars. t u c a n d e r a s , tacanduiras, t o u c a n d e i r a s , etc.), vinte-e-

T h e tocandira are also the object of a curious belief t h r o u g h m u c h of A m a z o n i a . T h e usual story has the ant allegedly eating the tiny seed of a specific liana or aerial root, called tamshi (Carludovica divergens),

GIANT HUN TING ANTS

437

used by the I n d i a n s for b i n d i n g t o g e t h e r the f r a m e w o r k of h u t s . T h e seed germi­ nates in the ant's body, even while it lives, a n d grows from it into a new plant. In a n o t h e r version, the ant simply m e t a m o r ­ phoses directly into the plant. T h e basis for these myths u n d o u b t e d l y is f o u n d in the attacks of a c o m m o n parasitic fungus of the g e n u s Cordyceps o n these ants. Cordyceps australis, in fact, is a highly specialized p a t h o g e n infecting the a n t tribe Ponerini in tropical forest habitats. Before dying from an infection, the a n t attaches itself to a leaf or tree a n d b e c o m e s transfixed by the fungus while the fruiting bodies of the fungus g r o w o u t from it, in long filaments r e s e m b l i n g the early stages of a d e v e l o p i n g vine (pi. 3a). T h e s e t o c a n d i r a ants a r e r e p r e s e n t a t i v e of several large ants in t h e tribe Ponerini, all of which live in small colonies of a h u n d r e d or so w o r k e r s a n d a q u e e n (Zahl 1972). T h e i r nest is normally a b u r r o w situated in the g r o u n d at the base of trees; the nest e n t r a n c e is a g a p in the soil, usually c o n t i g u o u s with the tree t r u n k . W o r k e r s forage alone for a r t h r o p o d p r e y a n d plant materials a n d d o not exhibit c o o r d i n a t e d g r o u p behavior, a trait consid­ e r e d primitive in this insect family. T h e y are also virtually casteless, the workers not being well differentiated from the sexual females. T h e giant h u n t i n g ants may be e n c o u n ­ tered a n y w h e r e in the moister p o r t i o n s of tropical A m e r i c a b u t a r e absent from the Antilles a n d isolated islands.

References HERMANN, H. R., M. S. BLUM, J. W. WHEELER, W. L. OVERAL, J. O. SCHMIDT, ANDJ. T. CHAO.

1984. Comparative anatomy and chemistry of the venom apparatus and mandibular glands in Dinoponera granéis (Guérin) and Paraponera dai'afa(F.) (Hymenoptera: Formicidae: Ponerinae). Entorno!. Soc. Amer. Ann. 77: 272-279. LENKO, K., AND N. PAPAVERO. 1979.

Insetos no

folclore. Cons. Estad. Artes Cien. Hum., Sao Paulo.

438

SAWFLIES, WASPS, ANTS, AND BEES

LIEBRECHT, F. 1886. Tocandyraf estes. Zeit Ethnol. 18: 350-352. WEBER, N. A.

1939.

The

sting of

the

ant

Paraponera clávala. Science 89: 127-128. ZAHL, P. A. 1972. Giant ants oí the Amazon Nat. Geogr. Soc. Res. Repts. 1955—1960193-201.

Greater Giant Hunting Ant Formicidae, Ponerinae, Ponerini, Dinoponera gigantea. Spanish: Isula (Peru). T h i s ant a n d its few c o n g e n e r s (Kempf 1971) are primarily A m a z o n i a n but extend westward into the A n d e a n foothills, south­ ward to Paraguay, a n d across the Guiana a n d Brazilian h i g h l a n d s to the Atlantic coast. T h e worker is decidedly larger than all o t h e r large black p o n e r i n e s of the A m e r i c a n tropics; specimens with body lengths of over 3 centimeters are known, m a k i n g t h e m the largest ants in the world ( H e r m a n n et al. 1984). T h e females are shiny black ants with a s m o o t h cuticle, a l t h o u g h it is often clothed in a sparse, close-set, g o l d e n pile (fig. 12.8a) (Haskins a n d Zahl 1971). T h e n o d e of the petiole is n a r r o w a n d rectangular and h i g h e r t h a n long in outline, as viewed from the side. T h e r e are no overt projections or spines on the t h o r a x . Males are consider­ ably smaller t h a n females a n d chestnut b r o w n , with long silky pubescence. Colonies n u m b e r only a h u n d r e d or so individuals a n d live in shallow burrows at the bases of trees; Fowler (1986) records only thirty workers p e r nest of D. australis. Habitat is typically forest, but colonies survive in cut-over or second growth scrubby areas as well. T h e workers emerge d u r i n g the daytime to forage for food, normally on the g r o u n d , ascending vegeta­ tion only to take liquids from extrafloraJ nectaries. In A m a z o n i a n Peru, this ant is mim­ icked by an unidentified species of cerambycid beetle in the g e n u s Tillomorpha

Figure 12.8 ANTS (FORMICIDAE). (a) Greater giant hunting ant {Dinoponera gigantea). (b) Lesser giant hunting ant (Paraponera clavata). (c) Kelep (Ectatomma tuberculatum). (d) Trap jaw ant (Odontomachus sp.). (e) Cobra ant (Pachycondyla villosa). (Cerambycinae, tribe T i l l o m o r p h i n i ) (orig. obs.).

References FOWLER, H. G. 1986. Populations, foraging and territoriality in Dinoponera australis (Hymenoptera, Formicidae). Rev. Brasil. Entomol. 29: 443-447. HASKINS, C.

P., AND P. A.

ZAHL.

1971.

The

reproductive pattern of Dinoponera granáis Roger (Hymenoptera, Ponerinae) with notes on the ethology of the species. Psyche 78: 1 — 11. HERMANN, H. R., M. S. BLUM, J. W. WHEELER, W. L. OVERAL, J. O. SCHMIDT, ANDJ. T. CHAO.

1984. Comparative anatomy and chemistry of the venom apparatus and mandibular glands in Dinoponera granáis (Guérin) and Paraponera clavata (F.) (Hymenoptera: Formicidae: Pone­ rinae). Entomol. Soc. Amer. Ann. 77: 2 7 2 279. KEMPF, W. W. 1971. A preliminary review of the ponerine ant genus Dinoponera Roger (Hy­ menoptera: Formicidae). Studia F.ntomol. 14: 369-391.

Lesser Giant Hunting Ant Formicidae, P o n e r i n a e , E c t a t o m m i n i , Paraponera clavata. Spanish: C h a c h a , folofa ( P a n a m a ) ; isulilla (Peru); bala (Costa Rica). Portuguese: Tapiai (Brazil). This species ( J a n z e n a n d Carroll 1983) is much m o r e w i d e s p r e a d t h a n Dinoponera gigantea, living in forest habitats from Nica­ ragua t h r o u g h C e n t r a l A m e r i c a to A m a z o Bia. It is the sole m e m b e r of its g e n u s a n d ^distinguished from the o t h e r h u n t i n g ants jprimarily by its large size (BL 20 m m ) , b u t

it is smaller (fig. 12.8b) t h a n Dinoponera. It t e n d s to be hairier than the o t h e r s (stiff, bristly hairs generally) a n d r e d d i s h - b r o w n , with a coarsely c o r r u g a t e cuticle. T h e a n t e ­ rior c o r n e r s of the t h o r a x also b e a r large, conical projections. T h e n o d e of t h e peti­ ole is large, c o m p r e s s e d a n d r e c t a n g u l a r in outline. T h e s t r u c t u r e of the species m o u t h p a r t s (Whiting et al. 1989) a n d sting­ ing a p p a r a t u s has b e e n described in detail ( H e r m a n n a n d Douglas 1976, H e r m a n n a n d Blum 1966), as has the effects of the sting on h u m a n s (Weber 1939). T h e r e is some evidence of w o r k e r types with different tasks associated with allometric growth (Breed a n d H a r r i s o n 1988). Foraging workers actively p u r s u e prey, plant e x u d a t e s (Young 1981), a n d sap, usually at night, searching for insects from dusk to dawn on low vegetation ( a l t h o u g h daytime foraging occurs on overcast days) (McCluskey a n d B r o w n 1972, Young a n d H e r m a n n 1980). T h e y are c o m m o n l y seen o n tree t r u n k s a n d lianas n e a r the forest floor. Nest b u r r o w s a r e usually placed between tree buttresses a n d have multiple o p e n i n g s , the latter sometimes with a thin cone or weak chimney. Some a r b o r e a l nests have been observed (Breed a n d H a r r i s o n 1989). T h e r e a p p e a r s to be some d e g r e e of association between the ant a n d the gavilán tree (Pentaclethra macroloba) ( B e n n e t t a n d B r e e d 1985), but o t h e r trees are also se­ lected, possibly because they possess extrafloral nectaries (Belk et al. 1989). Work­ ers g u a r d the nest e n t r a n c e a n d d e f e n d the

GIANT HUNTING ANTS

439

nest against intrusion by vertebrates, usu­ ally stinging t h e m , a n d against o t h e r in­ sects, including m e m b e r s of o t h e r colonies of their o w n kind. T h e r e m a y also be aggression between ants from different colonies in foraging areas, indicating t h e possible existence of territoriality (Her­ m a n n a n d Y o u n g 1980). I n j u r e d workers may b e attacked by t h e parasitoid p h o r i d Hy Apocephalus paraponerae, which a p p a r ­ ently locates its hosts by o d o r s given off from their w o u n d s (Brown a n d Feener 1991). T h e species serves as a m o d e l for Batesian mimicry by t h e cerambycid bee­ tles Acyphoderes sexualis (Silberglied a n d Aiello 1976) a n d possibly also Stenygra contracta (orig. obs.).

References BEI.K, M. C , H. L. BLACK, AND C. D. JORGF.NSF.N.

1989. Nest tree selectivity by the tropical ant. Paraponera chivata. Biotropica 21: 173-177. BENNETT, B., AND M. D. BREED. 1985. On the

association between Pentaclethra macroloba (Mimosaceae) and Paraponera chivata (Hymenoptera: Formicidae) colonies. Biotropica 17: 253-255. BREED, M. D., AND J.

M. HARRISON.

M. HARRISON.

1989.

Arboreal nesting in the giant tropical anl, Paraponera clavata (Hymenoptera: Formici­ dae). Kans. Entomol. Soc. J. 62: 133-135. BROWN, B. V'., AND D. H. FEENER 1991. Behav­

ior and host location cues of Apocephalus paraponerae (Diptera: Phoridae), a parasitoid of the giant tropical ant, Paraponera clavata (Hymenoptera: Formicidae). Biotropica 23: 182-187. HERMANN, J R . H. R., AND M. S. BLUM. 1966.

The morphology and histology of the hymenopterous poison apparatus. 1. Paraponera clavata (Formicidae). Entomol. Soc. Amer. Ann. 59: 397-409. HERMANN, JR., H. R., AND M. DOUGLAS. 1976.

Sensory structures on the venom apparatus of a primitive anl species. Entomol. Soc. Amer. Ann. 69: 681-686. HERMANN, J R . H. R., AND A. M. YOUNG. 1980.

Artificially elicited defensive behavior and

440

JANZEN,

D.

H., AND C.

R. CARROLL.

1933

Paraponera clavata (bala, giant tropical ant). /„ D. H. Janzen, ed., Costa Rican natural history Univ. Chicago Press, Chicago. Pp. 752-753, MCCLUSKEY, E. S., AND W. L. BROWN, JR 1972

Rhythms and other biology of the giant tropical ant Paraponera. Psyche 79: 335-347 SILBERGLIED, R. E., AND A. AIELLO. 1976. Defen­

sive adaptations of some Neotropical longhorned beetles (Coleóptera, Cerambycidae): Antennal spines, tergiversation, and double mimicry. Psyche 83: 256—262. WEBER, N. A. 1939. T h e sting of the ant

Paraponera clavata. Science 89: 127-128. WHITING, JR., J. H., H. L. BLACK, AND C. D.

JORGENSEN. 1989. A scanning electron micros­ copy study of the mouthparts of Paraponera clavata (Hymenoptera: Formicidae). PanPacific Entomol. 65: 302-309. YOUNG, A. M. 1981. Giant Neotropical ant Paraponera clavata visits Heliconia pogonantha flower bracts in premontane tropical rain forest. Biotropica 13: 223. YOUNG, A. M., AND H. R. HERMANN, J R 1980.

Notes on the foraging of the giant tropical anl Paraponera clavata (Hymenoptera: For­ micidae: Ponerinae). Kans. Entomol. Soc. J. 53: 35-55.

1988.

Worker size, ovary development and division of labor in the giant tropical ant, Paraponera clavata (Hymenoptera: Formicidae). Kans. Entomol. Soc. J. 6 1 : 285-291. BREED, M. D., AND J.

reciprocal aggression in Paraponera cla­ vata (Hymenoptera: Formicidae: Ponerinae) Georgia Entomol. Soc. J. 15: 8-10.

SAWFL1ES, WASPS, ANTS, AND BEES

TRAP JAW ANTS Formicidae, P o n e r i n a e , Ponerini, Odontomachus. Spanish: T i n g o t e r o s (Peru). Portuguese: Bate-bicos, taracutingas (var. saracutingas), formigas p o r t a pincas (Brazil). Workers of this a n d related (Moffett 1989) g e n e r a a r e a m o n g t h e most easily recog­ nized of ants (fig. 12.8d). T h e i r head shape is uniquely widened in t h e anterior part, with t h e eyes situated at t h e broadest part o n p r o m i n e n c e s . T h e straight mandibles are elongate with a h o o k e d tip a n d lie nearly parallel at full closure b u t can be held o p e n to a full 180 d e g r e e s . T h e node of t h e petiole (waist) is also distinctive: a tall cone with a s h a r p , single point at its apex. T h e s e ants a r e slender bodied, long legged, a n d variously colored in dull

reddish-black h u e s to bright reds, blacks, and yellows. Like o t h e r p o n e r i n e s , Odontomachus a r e active h u n t e r s . T h e peculiar mandibles play an essential p a r t in this activity. While searching for prey, the j a w s a r e carried at an o p e n ready position, u n d e r tension by a locking m e c h a n i s m at their bases. Special sensory trigger hairs p o i n t i n g forward from t h e bases of t h e m a n d i b l e s initiate a sudden convulsive s n a p of the j a w s when something t o u c h e s t h e m . Insects a n d o t h e r small a r t h r o p o d s a r e thus c a u g h t a n d t h e n subdued with a sting. Empty m a n d i b l e s s n a p p i n g t o g e t h e r emit an a u d i b l e clicking s o u n d . If by c h a n c e when a n ant closes its jaws, they i m p i n g e o n some i m m o v a b l e large object like a steel instrument, t h e a n t m a y be flipped back­ ward several c e n t i m e t e r s . Such acrobatics have been described by a n u m b e r of observ­ ers and a r e testimony to t h e viciousness of the ant's ability to attack. Jaw s n a p p i n g is also used as a d e f e n s e (Carlin a n d Gladstein 1989). Like o t h e r fairly large p o n e r i n e s (BL 6— 20 mm), m a n y Odontomachus sting quickly and painfully. Trap j a w ants live in all parts of t h e American tropics, i n c l u d i n g t h e West I n ­ dies a n d Galápagos, a n d o t h e r Pacific is­ lands. H e r e they may be e n c o u n t e r e d usu­ ally d u r i n g t h e day walking to a n d fro o n vegetation, e n g a g e d in foraging activities. Their nests a r e situated in t h e soil, most commonly u n d e r r o t t i n g wood, an old stump, o r a log; nest t u n n e l s m a y even extend into t h e wood. A n o t h e r f r e q u e n t nesting site is in h u m u s a n d leaf litter at the base of large trees. T h r o u g h o u t t h e Neotropics, t h e r e a r e some twenty-four species, a n u m b e r that may be increased w h e n s o m e t a x o n o m i c complexes a r e b e t t e r k n o w n , for e x a m p l e , that going u n d e r t h e n a m e of "Odonto­ machus haematodus," a ubiquitous entity that some a u t h o r s c o n s i d e r a superspecies (Brown 1976).

References BROWN, J R , W. L. 1976. Contributions toward a reclassification of the Formicidae. Pt. VI. Ponerinae, tribe Ponerini, subtribe Odontomachiti. Sect. A. Introduction, subtribal char­ acters. Genus Odontomachus. Studia Entomol. 19:67-171. CARLIN, N. F , AND D. S. GLADSTEIN. 1989. T h e

"bouncer" defense of Odontomachus ruginodis and other odontomachine ants (Hymenop­ tera: Formicidae). Psyche 96: 1 — 19. MOFFETT, M. W. 1989. Trap-jaw ants: Set for prey. Nail. Geogr. 175(3): 395-400.

COBRA ANT Formicidae, P o n e r i n a e , Ponerini, Pachycondyla vülosa. Spanish: P u n g a r a (Peru). Portuguese: Saracutinga, beijo d e moca, c a c h o r r o m a g r o (Brazil). T h i s large, black, fiercely stinging species (fig. 12.8e) resembles t h e giant h u n t i n g ants. Workers a r e always smaller (BL 12— 13 m m ) , however, a n d have a distinctively s h a p e d petiole. Viewed from above, it is very wide a n d a p p e a r s like a n o t h e r a b d o m i ­ nal segment; in side view, it has straight a n t e r i o r a n d r o u n d e d posterior m a r g i n s . T h e cuticle is densely p u n c t a t e a n d set with golden yellow a n d white p u b e s c e n c e . T h e a b d o m e n is also m o r e o r less straight sided, without a s t r o n g constriction as in t h e giants. T h e cobra a n t h a s a very wide r a n g e , o c c u r r i n g from s o u t h e r n Texas to n o r t h ­ e r n A r g e n t i n a , t h r o u g h o u t continental, tropical America. Small colonies nest u n ­ d e r g r o u n d at t h e bases of trees o r in rotting logs a n d s t u m p s . Workers r u n r a p ­ idly o n t h e soil, often in t h e bright sun, in search of insect prey.

KELEP Formicidae, P o n e r i n a e , E c t a t o m m i n i , Ectatomma tuberculatum. Weevil a n t . T h e kelep (fig. 12.8c) is a n o t h e r fairly large solitary h u n t i n g a n t (BL w o r k e r 1 1 -

KELEP

441

12 m m ) b u t with a r a t h e r m i l d e r sting than its larger relatives. It is recognized by a coarsely s c u l p t u r e d , r e d d i s h - b r o w n cuticle a n d the s t r u c t u r e of the t h o r a x , the d o r s u m of which has a central convexity b o u n d e d o n all sides by distinct grooves a n d t h r e e p r o t u b e r a n c e s dorsally on its a n t e r i o r portion. A n o t h e r anatomical char­ acteristic is the large, c o m p r e s s e d petiole n o d e that, in side view, shows concave a n t e r i o r a n d convex p o s t e r i o r m a r g i n s . T h e r e a r e also s t r o n g bulbs at the bases of the a n t e n n a e . Viewed from above, the a b d o m i n a l s e g m e n t s a p p e a r swollen, the i n t e r s e g m e n t a l j o i n t s constricted. W i d e s p r e a d geographically ( s o u t h e r n Mexico to P a r a g u a y a n d s o u t h e r n Brazil), this ant is also c o m m o n in a wide variety of habitats, from arid lands to rain forest. Its nests a r e g r o u n d b u r r o w s with multiple o p e n i n g s at the base of trees. From these emerge prominent tubular runways made of thatch or m u d a p p r e s s e d against the side of t h e tree a few c e n t i m e t e r s to a m e t e r o r m o r e in height. W o r k e r s forage arboreally for small insects (especially so­ cial insects), nectar, a n d o t h e r food items. T h e y a r e often seen t e n d i n g extrafloral nectaries (pi. 3b). In P a n a m a , acrobat ants (Crematogaster limata) h a v e b e e n observed to file into kelep nests, p e r h a p s to steal food (Wheeler 1986). T h e species was the subject of an early, abortive biological control s c h e m e (Weber 1946). O n a trip to G u a t e m a l a in 1904, O. F. Cook, an e n t o m o l o g i s t from the U.S. De­ p a r t m e n t of A g r i c u l t u r e , f o u n d the ant associated with wild cotton. T h e natives of the area h e visited, w h o called t h e ant "kelep," knew of the association a n d con­ vinced Cook that the a n t p r e y e d on adult cotton boll weevils, k e e p i n g the plant free of that pest. T h e ants w e r e r e p u t e d to carry off the weevils a n d feed t h e m to their larvae, which have long flexible necks spe­ cially a d a p t i n g t h e m to r e a c h inside a n d c o n s u m e the soft tissues in the body of the prey (Cook 1905). In the same year, Cook

442

SAWFL1ES, WASPS, ANTS, AND BEES

g a t h e r e d keleps a n d i n t r o d u c e d t h e m into cotton fields in Texas, but they showed little inclination to colonize their new h o m e and died out after a short time, failing to control the weevils that h a d recently invaded the fields in that state. Unfortunately, Cook's observations on the kelep were superficial a n d m a n y of his conclusions r e g a r d i n g its biology fanciful, as p o i n t e d out by William M o r t o n W h e e l e r (1905, 1906), his m o r e sagacious c o n t e m p o r a r y .

References COOK, O. F. 1905. The social organization and breeding habits of the cotton-protecting kelep of Guatemala. U.S. Dept. Agrie. Tech Ser. 10: 1-55. WEBER, N. A. 1946. Two common ants of possible economic significance, Ectatomma tuberculatum (Olivier) and E. ruidum Roger. Entomol. Soc. Wash. Proc. 48: 1-16. WHEELER, D. E. 1986. Ectatomma tuberailatum: Foraging biology and association with Cremato­ gaster (Hymenoptera: Formicidae). Entomol. Soc. Amer. Ann. 79: 300-303. WHEELER. W. M. 1905. Dr. O. F. Cook's "Social organization and breeding habits of the cotton-protecting Kelep of Guatemala." Sci­ ence (n.s.) 22: 706-710. WHEELER, W. M. 1906. The kelep excused. Science (n.s.) 23: 348-350.

ARMY ANTS Formicidae, Ecitoninae, Eciton et al. Spanish: H o r m i g a s militares, guerreras, soldados (General); arrieras (Panama); r o n c h a d o r e s (Costa Rica); tepeguas, cazadoras, carniceras (Peru). Portuguese: Formigas correcóes, f. taiocas, f. m o r u p e t e c a s , etc. (Brazil). All the m e m b e r s of the subfamily Ecitoni­ nae a r e loosely r e f e r r e d to as "army ants," a category also including the tropical driver or legionary ants (Dorylinae, Dorylus) of Africa (Schneirla 1971). T h e name is m o r e a p p r o p r i a t e l y applied to the 130 or so New World species, especially those in the g e n u s Eciton ( B o r g m e i e r 1955, Franks

Figure 12.9 ARMY ANTS (ECITON: FORMICIDAE). (a) Major worker, (b) Physogastric queen, aided by minor worker, (c) Male.

1989). T h e s e a r e f o u n d t h r o u g h o u t the lowland moist a n d wet forests of t h e A m e r i ­ can tropics from coastal n o r t h e r n Mexico to n o r t h e r n A r g e n t i n a a n d s o u t h e a s t e r n Brazil. T h e r a n g e e x t e n d s to the larger islands of the West Indies as well. O t h e r , common, lesser a r m y ants b e l o n g to the genera Labidus a n d Neivamyrmex. Several species are primarily s u b t e r r a n e a n , e m e r g ­ ing to the soil's surface only at intervals to swarm a n d seek food. T h e larvae of a few species of t h e s e h a v e b e e n described (Wheeler a n d W h e e l e r 1984). T h e b e s t - k n o w n a n d most w i d e s p r e a d species of a r m y ants a r e Eciton hamatum and E. burchelli. Major w o r k e r s of the former species a r e light b r o w n to yellow, with shiny yellowish h e a d s ; the latter are dark b r o w n to blackish, with r e d d i s h a b d o ­ mens a n d dull-surfaced whitish h e a d s . A colony of e i t h e r consists of the q u e e n (fig. 12.9b), males (fig. 12.9c), w o r k e r s , and i m m a t u r e s . T h e males a r e winged a n d live long e n o u g h only to exit the colony to mate with wingless q u e e n s . Workers are graded in size a n d p r o p o r t i o n s from the very large majors, o r "soldiers," with outsized heads a n d gigantic h o o k e d mandibles (fig. 12.9a), to tiny m i n o r s , o n e q u a r t e r the size of the majors. T h e nest is n o t a fixed structure b u t a n enclave, called the biv­ ouac, f o r m e d a r o u n d the q u e e n a n d b r o o d by the interlocking bodies a n d limbs of the workers (pi. 2h). T h e life of Eciton colonies is o r g a n i z e d •nto a biphasic b e h a v i o r cycle by the r e p r o ­

ductive r h y t h m of the q u e e n . A stationary ("statary") p h a s e , lasting a r o u n d 20 days, alternates with a slightly s h o r t e r m i g r a t o r y ("nomadic") phase, typically of 17 days. In the stationary p h a s e , the colony lo­ cates the bivouac in a well-protected place, usually within a hollow log or s t u m p or u n d e r a partially b u r i e d fallen log. Within the bivouac, eggs are b e i n g laid by the q u e e n , a n d p u p a e from the previous b r o o d are developing. T h e p r e s e n c e of workers freshly e m e r g e d from their co­ coons (so-called callow workers) releases the n o m a d i c instinct, a n d the entire colony emigrates to a new bivouac site each day for the period of this p h a s e . D u r i n g these translocations, a r e t i n u e of w o r k e r s closely g u a r d s the q u e e n ( R e t t e n m e y e r et al. 1978). T h e bivouacs are placed in rela­ tively exposed places, such as the space b e n e a t h low o v e r h a n g i n g b r a n c h e s or be­ tween the buttresses of a tree. D u r i n g this phase, the eggs hatch, a n d the larvae are b r o u g h t to maturity. R e t u r n to the station­ ary phase ensues w h e n t h e larvae have s p u n cocoons a n d p u p a t e d . Virtually without fail, a r a i d i n g p a r t y issues from the bivouac each day to obtain animal prey, which is the only food of these ants. T h i s habit r e p r e s e n t s t h e most com­ plex instance of organized mass b e h a v i o r o c c u r r i n g regularly outside the nest in any social insect o r s u b h u m a n animal. Raids of n o m a d i c colonies of E. hamatum almost always start shortly after d a w n ; sta­ tionary colonies may raid in the m o r n i n g o r

ARMY ANTS

443

a f t e r n o o n . Raids consist of t h o u s a n d s of w o r k e r s r e a c h i n g o u t over t h e forest floor a n d lower p o r t i o n s o f t h e vegetation in search of any insects o r small animals that may be found a n d c a p t u r e d . With some exceptions (Young 1979), E. hamatum shows a p r e f e r e n c e for p a p e r wasp larvae a n d p u p a e ; E. burchelli is less specialized, taking cockroaches, spiders, a n d katydids. T h e a d v a n c i n g front of t h e raid also takes two forms d e p e n d i n g on t h e species. E. hama­ tum is a " c o l u m n raider," in which t h e files of workers r e m a i n m o r e o r less distinct. T h i s contrasts with t h e " s w a r m raid" of E. burchelli, in which t h e c o l u m n s a n a s t a m o s e at t h e front into a solid, fan-shaped mass. O t h e r species of Eciton d e m o n s t r a t e habits differing only in detail ( B u r t o n a n d Franks 1985).

T h e s e r e g u l a r " c a m p followers," and occasionally o t h e r insects, may be attracted to t h e swarm. Butterflies have been ob­ served mingling with a r m y a n t swarms possibly attracted by t h e latter's odor which is said to b e u n p l e a s a n t a n d detect­ able by t h e sensitive h u m a n nose. T h e o d o r may simulate t h e butterflies' sexual p h e r o m o n e s ( D r u m m o n d 1976) o r resem­ ble decay smells associated with proteinaceous food sources (Young 1977), in­ cluding t h e d r o p p i n g s of a n t birds (Lamas 1983, Ray a n d A n d r e w s 1980) o n which they commonly have b e e n seen to feed. Many m y r m e c o p h i l o u s m e m b e r s of the beetle families Staphylinidae, Histeridae a n d L i m u l o d i d a e , as well as silverfish and mites, have also b e e n f o u n d intimately associated with a r m y ants.

N u m e r o u s imaginative a n d sensational accounts of t h e ferocity of ants in these swarms a r e to be f o u n d in p o p u l a r litera­ t u r e . T h e m o d e l of these is t h e classic short story, " L e i n i n g e n versus t h e A n t s , " by Carl Stephenson. T h e a u t h o r created an image of an a n t capable of total devastation a n d destruction of a n y t h i n g in its p a t h (includ­ ing vegetation, an a p p a r e n t confusion with the leaf c u t t e r ants). T h i s myth is now widespread, except a m o n g natives of r e ­ gions w h e r e these ants a b o u n d . T h e s e people may actually welcome invasions by swarms for t h e i r value in e x t e r m i n a t i n g h o u s e h o l d cockroaches a n d o t h e r v e r m i n .

A m e r i n d s have capitalized on t h e tenac­ ity of t h e powerful "ice-tong" jaws of Eciton to aid in t h e healing of flesh w o u n d s . They pinch t h e edges of t h e injury together and hold t h e ant close so that it bites across the cut. T h e n they w r e n c h off t h e body, leav­ ing t h e h e a d intact a n d jaws fixed like sutures (Majno 1975). L a r g e leaf cutter ant workers may also be used this way (Gudger 1925).

Raids a r e a c c o m p a n i e d by m u c h c o m m o ­ tion a m o n g t h e o t h e r c r e a t u r e s of t h e forest. T h e w h i n i n g wings of several spe­ cies of tachinid (Calodexia a n d Androeuryops) a n d c o n o p i d flies (Stylogaster) min­ gle with t h e calls of a n t b i r d s a n d t h e rustle of fleeing insects, frogs, lizards, a n d o t h e r small animals ( R e t t e n m e y e r 1961). T h e flies take a d v a n t a g e of t h e panic to locate a n d parasitize certain of t h e insects r o u s e d by t h e ants. T h e insectivorous a n t birds a n d o t h e r birds (Willis a n d Oniki 1978, Gochfeld a n d T u d o r 1978) also find that h u n t i n g is m a d e easier within t h e melee.

BURTON, J. L, AND N. R. FRANKS. 1985.

444

SAWFL1ES, WASPS, ANTS, AND BEES

perú. Rev. Soc. Mexicana Lepidop. 8(2): 4 9 51. MAJNO, G. 1975. Ant saga. In G. Majno, The healing hand: Man and wound in the ancient world. Harvard Univ. Press, Cambridge. Pp. 304-309. J^Y. T. S., AND C. C. ANDREWS.

1980.

Ant

butterflies: Butterflies thai follow army ants to feed on ant bird droppings. Science 210: 1147-1148. RETTENMEYER, C. W. 1961. Observations on the

biology and taxonomy of flies found over swarm raids of army ants (Dipiera: Tachinidae, Conopidae). Univ. Kans. Sci. Bull. 42: 993-1066. RETTENMEYER, C. W.,

H. TOPOFF, AND J. M I ­

RANDA. 1978. Queen retinues of army ants. Entomol. Soc. Amer. Ann. 71: 519-528. SCHNEIRLA, T. 1971. Army ants, a study in social organization. Edited by H. Topoff. Freeman, San Francisco. WHEELER, G. C ,

AND J. WHEELER. 1984.

The

larvae of the army ants (Hymenoptera: Formicidae). Kans. Entomol. Soc. J. 57: 263-275. WILLIS, E. O., AND Y. ONIKI. 1978. Birds and

army ants. Ann. Rev. Ecol. Syst. 9: 243-263. YOUNG, A. M. 1977. Butterflies associated with an army anl swarm raid in Honduras: The "feeding hypothesis" as an alternative expla­ nation. J. Lepidop. Soc. 31: 190. YOUNG, A. M. 1979. Attacks by the army ant Eciton burchelli on nests of the social paper wasps Polisles erylhrocephalus in northeastern Costa Rica. Kans. Entomol. Soc. J. 52: 7 5 9 768.

References BORGMEIER, T. 1955. Die Wanderameisen der

neotropischen region. Studia Entorno!. 3: 1716, pis. 1-87. The

foraging ecology of the army ant Eciton rapax: An ergonomic enigma? Ecol. Entomol. 10: 131-141. DRUMMOND, B. A. 1976. Butterflies associated with an army ant swarm raid in Honduras. J. Lepidop. Soc. 30: 237-238. FRANKS, N. R. 1989. Army ants: A collective intelligence. Amer. Sci. 77: 139-145. GOCHFELD,

M.,

AND G.

TUDOR.

1978.

Ant-

following birds in South American subtropi­ cal forests. Wilson Bull. 90: 139-141. GUDGER, E. 1925. Stitching wounds with the mandibles of ants and beetles. J. Amer. Med. Assoc. 84(24): 1861-1864. LAMAS, G. 1983. Mariposas atraídas por hormi­ gas legionarias en la Reserva de Tambopata,

FIRE ANTS Formicidae, Myrmicinae, Solenopsidini, Solenopsis. Spanish: H o r m i g a s d e fuego, hormigas bravas (Cuba). Quechua: Pucacuro-kuna. Portuguese: Formigas d e fogo, f. brasas, lavapés, cagafogos, doceiras, q u e i m a - q u e i m a , etc. (Brazil). Indian: A r a c a r a s (Amazonia). This is a large, ubiquitous g e n u s with over one h u n d r e d species in t h e A m e r i c a n t r o p ­ ics, including t h e C a r i b b e a n a n d Pacific oceanic islands ( C r e i g h t o n 1930, Snelling pers. c o m m . ) . A c o u p l e of d o z e n species (especially S. saevissima a n d S. geminata) a r e particularly n o x i o u s pests, which, because

of their p o t e n t sting, a r e r e f e r r e d to as "fire ants." In spite of t h e w i d e s p r e a d o c c u r r e n c e of these nuisance species, little is known of their biology, save with t h e two South A m e r i c a n species that were i n t r o ­ d u c e d into t h e s o u t h e r n United States a n d created a furor that eventually r e a c h e d political significance in that nation's capitol (Lofgren et al. 1975, Williams a n d Whitc o m b 1973). T h e s e a r e t h e r e d i m p o r t e d fire ant (S. invicta) a n d t h e black i m p o r t e d fire ant (S. richteri). T h e f o r m e r (fig. 12.10a) is the m o r e i m p o r t a n t of the two, having s p r e a d over most of seven south­ eastern states since its accidental i n t r o d u c ­ tion in Mobile, Alabama, probably in t h e late 1930s. It is not a p p r o p r i a t e to review the fire ant saga h e r e . From intensive studies on these invaders, however, some generaliza­ tions can be m a d e about t h e genus's vital characteristics. All species nest in t h e g r o u n d , many f o r m i n g large m o u n d s a r o u n d the e n t r a n c e from soil castings. Funnels e x t e n d b e l o w g r o u n d to a m e t e r o r m o r e . Colonies a r e large ( u p to 230,000 workers) a n d often have two to m a n y q u e e n s (polygyny). T h e y a r e generally a d a p t e d to w a r m wet climates, a l t h o u g h some species a r e desert loving. T h e i r food habits a r e catholic, but they a r e primarily p r e d a t o r s on o t h e r insects. T h e y a r e highly competitive a n d often drive colonies of o t h e r ant species from their area. T h e y suffer from a variety of parasites, which k e e p populations u n d e r control in South America (Williams a n d W h i t c o m b 1973). T h e y a r e also fierce d e f e n d e r s of their own nests, using highly toxic stings as effective weapons, for which they a r e best known a n d to which most of t h e i r p o p u l a r n a m e s refer. T h e p h a r m a c o l o g y of t h e v e n o m has been analyzed a n d its composi­ tion possibly better known t h a n that of any o t h e r ant. Major active c o m p o n e n t s a r e p i p e r i d i n e alkaloids; p r o t e i n s a r e notice­ ably lacking (MacConnell et al. 1971). T h e effects of v e n o m injected into h u m a n s by

FIRE ANTS

445

America, Colombia); m o c h o m a s (Mexico); bachacos (Venezuela); c u r u h u i n s i (Peruvian A m a z o n ) ; coqui (Peru); bibijaguas (Cuba). Tupi-Guaraní: Saúva, icá (Brazil). Wee wee (Belize), parasol a n t s , town ants.

Figure 12.10 ANTS (FORMICIDAE). (a) Fire ant (Solenopsis invicta), (b) Leaf cutter ant (Atta cephalotes), dealate queen, (c) Leaf cutter ant, minor worker with leaf fragment, (d) Leaf cutter ant, major worker.

the ant's sting can be severe. Pustules form, a n d necrotic effects often follow. Allergic sensitivity frequently develops in r e s p o n s e to r a n d o m stings. D e a t h has b e e n k n o w n to occur in r a r e cases in highly sensitive individuals. S o m e a u t h o r s tell of fire ants in some areas so n u m e r o u s a n d o m n i p r e s e n t as to be a scourge. Such was how the residents f o u n d Aveyros on the Tapajós River in Bates's time. H e states (1892) that the village, "was d e s e r t e d a few years b e f o r e [his] visit o n account of this little t o r m e n ­ tor, a n d the inhabitants h a d only recently r e t u r n e d , t h i n k i n g its n u m b e r s h a d d e ­ creased." C a r v a l h o explains that the oldest h u n t i n g d o g s o n the Rio N e g r o were blind in f o r m e r days from c o r n e a s m a d e o p a q u e by c o n t i n u a l fire a n t stings. A local saying went, "the d o g with white eyes is always a good h u n t e r " ( L e n k o a n d P a p a v e r o 1979). Workers oí Solenopsis a r e recognizable by the s t r u c t u r e of the a n t e n n a e , which is u n i q u e a m o n g ants. T h e y have ten seg­ m e n t s with a t w o - s e g m e n t e d club. T h e body, w h e t h e r d a r k or light, has a r e d d i s h tint a n d is s m o o t h a n d shiny without spines but with sparse erect hairs. T h e petiole is two s e g m e n t e d , with distinct r o u n d e d n o d e s . T h e clypeus has a pair of elevated ridges t h a t d i v e r g e t o w a r d the a n t e r i o r e n d a n d (usually) distinct teeth that project b e y o n d the a n t e r i o r m a r g i n of this struc­ ture. T h e little fire ant, b e l o n g i n g to a related

446

SAWFLIES, WASPS, ANTS, AND BEES

g e n u s (Wasmannia auropunctata), is a wellknown, tropical m a i n l a n d t r a m p species. It has become established in P u e r t o Rico (where it is called abayalde) a n d in the Galápagos Archipelago w h e r e it is now a serious pest, displacing native species and preying on the i n d i g e n o u s fauna a n d sting­ ing h u m a n s (Silberglied 1972).

References BATES, H. W. 1892. The naturalist on the River Amazons. John Murray, London. CREIGHTON, W. S. 1930. The New World species of the genus Solenopsis (Hymenop. Formicidae). Amer. Acad. Arts Sci. Proc. 66: 39-151. LENKO, K., AND N. PAPAVERO. 1979.

Insetos no

folclore. Cons. Estad. Artes Cien. Hum., Sao Paulo. LOFGREN,

C.

S.,

W.

A.

BANKS, AND B.

M.

CLANGEY. 1975. Biology and control of im­ ported fire ants. Ann. Rev. Entomol. 20: 1-30. MACCONNF.LL, J. G.,

M. S. BLUM, AND H.

M.

FALES. 1971. The chemistry of fire anl venom. Tetrahedron 27: 1129-1139. SILBERGLIED, R. 1972. The "little fire ant," Wasmannia auropunctata, a serious pest in the Galápagos Islands. Noticias Galápagos 1920: 13-15. WILLIAMS, R. N., AND W. H. WHITCOMB.

1973.

Parasites of fire ants in South America. Tall Timbers Conf. Ecol. Anim. Hab. Mgmt. Proc. 5: 49-59.

LEAF CUTTER ANTS Formicidae, Myrmicinae, Attini, Atta, Acromyrmex, a n d o t h e r g e n e r a . Spanish: Z a m p o p o s , h o r m i g a s arrieras (Central

These a r e such conspicuous ants that they have inspired a large n u m b e r of fanciful indigenous n a m e s (Weber 1982). T h o s e given above a r e only a s a m p l e of the m o r e than fifty listed (Weber 1971: 6 - 7 ) . Only two g e n e r a of a b o u t two h u n d r e d species of " g a r d e n i n g ants" (ants that cultivate fungi for food), or Attini, actually cut leaves a n d c o m p r i s e the t r u e leaf cutters, Atta a n d Acromyrmex (Cherrett a n d C h e r rett 1989). T h e s e g e n e r a are distributed widely in the Americas, from the s o u t h e r n ­ most p o r t i o n s of the United States to northern A r g e n t i n a a n d Uruguay, al­ though they a r e absent from the Carib­ bean Islands e x c e p t C u b a . In this r a n g e , these ants p r e f e r relatively h u m i d habitats and are t h e r e f o r e also completely absent from the s o u t h e r n half of the A n d e s a n d South America's coastal deserts. Leaf cutters a r e fairly large, reddishbrown ants with strongly p o l y m o r p h i c work­ ers. Major w o r k e r s (fig. 12.10d) are often several times larger t h a n o t h e r w o r k e r castes (fig. 12.10c) a n d have d i s p r o p o r t i o n ­ ately e n l a r g e d , h e a r t - s h a p e d h e a d s . T h e body is spiny a n d t h e a p p e n d a g e s long a n d gangly. T h e base of the a n t e n n a is h i d d e n by an o v e r l a p p i n g frontal lobe. In the major genus, Atta, t h e males a n d females are much larger t h a n even the major workers. In this g e n u s , the female, especially, is an enormous a n t ( 3 0 - 4 0 m m long, h e a d to wing tips) with a h u g e , spherical a b d o m e n (fig. 12.10b). In o t h e r g e n e r a , such as Acromyrmex, t h e r e p r o d u c t i v e s are not the largest castes by m u c h . Because of the a b u n d a n c e of q u e e n s at emergence times a n d their size, they are exploited as food by aboriginals almost everywhere. T h e y have even f o u n d their

way into t r a d e as traditional a n d novelty foods. T h e d r i e d , packaged hormigas co­ lonas (or hormigas santandereanas) of Bucaramanga, Colombia, are famous. Nests of leaf cutters are highly c o m p l e x a n d varied in a r c h i t e c t u r e (Weber 1969, 1971). T h e surface m o u n d s of c o m m o n , well-known species of Atta (cephalotes, sexdens) may cover an area of 40 to 50 square meters. Nest density seems to be g r e a t e r in d i s t u r b e d habitats t h a n in pri­ m a r y forest (Jaffe a n d Vilela 1989). Ma­ t u r e nests have m a n y craters c o m p o s e d of soil fragments, b r o k e n down g a r d e n s , a n d o t h e r ejected debris (mostly from spent leaf substrate) a n d are conspicuous fea­ tures of the landscape in m a n y parts of tropical America, especially in o p e n c o u n ­ try like the p a m p a s . Colonies may b e fairly readily m a i n t a i n e d in artificial nests o n which detailed study is possible (Weber 1976). T h e s e ants a r e best k n o w n for their ability to cultivate specific types of fungi o n which they feed (Quinlan a n d C h e r r e t t 1978). T h e s e fungi a r e cultured on snips of leaves a n d o t h e r parts of plants carried to special nest c h a m b e r s by the workers. T h e ants tend these " g a r d e n s " continually, r e m o v i n g all alien bacterial a n d u n w a n t e d fungal growths, a n d they maintain a highly p u r e strain of p r e f e r r e d fungi (Boyd a n d Martin 1975). T h e tips of the h y p h a e p r o d u c e peculiar r o u n d swellings (gongylidia) that are plucked a n d eaten o r fed to larvae. Few of the latter have b e e n identi­ fied to species, but Leucocoprinus gongylophora a n d Lepiota sp. may be the most i m p o r t a n t of those known. T h e identities of the fungal species involved a r e still controversial a n d certainly differ a m o n g their ant "host" species. T h e source of the cuttings for t h e gar­ d e n s may be any t r e e or s h r u b g r o w i n g u p to several h u n d r e d meters from t h e nest (Rockwood 1977). T h e long trails of work­ ers carrying oval fragments over their h e a d s (fig. 12.10c), winding their way over

LEAF GUTTER ANTS

447

well-worked p a t h s o n t h e forest floor, a r e a familiar sight to tropical travelers (Fowler 1978). W o r k e r s maintain a trail by laying d o w n a m a r k i n g p h e r o m o n e from rectal glands, a l t h o u g h ants may be repelled o r attracted by o t h e r o d o r s a l o n g their forag­ ing p a t h s (Littledyke a n d C h e r r e t t 1978). Occasionally, a large w o r k e r is seen in these processions with a very tiny w o r k e r riding on its leaf cargo. Observations in Trinidad (Eibl-Eibesfeldt a n d Eibesfeldt 1967) have revealed t h e h i t c h h i k e r as p r o ­ tector of the l a r g e r a n t that is otherwise p r e o c c u p i e d with its load a n d u n a b l e to avoid endoparasitic p h o r i d flies seeking to oviposit on its neck. Milichiid flies of the g e n u s Pholeomyia ride leaves similarly but for u n k n o w n reasons (Waller 1980).

p u l p , which the ants relish ( M u d d et al 1978), o r o t h e r baits (Robinson et al. 1982), show promise. T h e i m p r o v e m e n t of tropi­ cal soils by their b u r r o w i n g a n d mixing activities, as well as the m u l t i t u d e s of micro­ organisms they i n t r o d u c e , p e r h a p s should e a r n t h e m m o r e appreciation t h a n scorn ( J o n k m a n 1978).

B e n e a t h t h e g r o u n d , the nest consists of n u m e r o u s oval c h a m b e r s c o n n e c t e d by a n a s t a m o s i n g t u n n e l s a n d passageways. T h e latter may p e n e t r a t e to considerable d e p t h s , u p to 4 o r 5 m e t e r s , a n d connect d o z e n s of c h a m b e r s . T h e latter a r e mostly used for fungus g a r d e n s , lesser n u m b e r s for b r o o d r e a r i n g , a n d a single royal c h a m ­ ber is reserved for the q u e e n . D e p e n d i n g on t h e species, t h e r e m a y be as m a n y as 200,000 to 300,000 in such colonies.

bibliography of the leaf-cutting ants, Atta spp. up to 1975. Overseas Devel. Adm., Nat. Res. lnst. Bull. 14: 1-58. Includes Acromyrmex.

Nests of Acromyrmex generally differ from those of Atta in that they a r e simpler a n d less extensive. I n p a r t of t h e seasonally flooded p a r t s of t h e A m a z o n Basin, this g e n u s is often f o u n d nesting in trees two o r m o r e m e t e r s above the g r o u n d . Because leaf-cutting ants often strip fo­ liage from cultivated vegetation a n d their m o u n d s spoil otherwise level f a r m l a n d , they have long b e e n c o n s i d e r e d pests (Blanton a n d Ewel 1985). Even as early as the sixteenth century, they e a r n e d t h e title "King of Brazil." F r e n c h naturalist St. Hilaire, i m p r e s s e d by t h e p r o b l e m d u r i n g his visit to that c o u n t r y in 1816—1822, said, "Either Brazil kills t h e saúva o r the saúva kills Brazil!" All sorts of control m e a s u r e s have b e e n applied against t h e m , most with little success, a l t h o u g h poison-laced citrus

448

SAWFL1ES, WASPS, ANTS, AND BEES

References BLANTON, C. M., AND J. J. EWEL. 1985.

Leaf-

cutting ant herbivory in successional and agricultural tropical ecosystems. Ecology 66' 861-869. BOYD, N. D., AND M. M. MARTIN. 1975. Faecal

proteinases of the fungus-growing ant, Atta texana: Properties, significance and possible origin. Ins. Biochem. 5: 619—635.

for leaf-cutting ant (Hymenoptera: For­ micidae) control. Bull. Entomol. Res. 72: 345-356. ROCKWOOD, L. L. 1977. Foraging patterns and plant selection in Costa Rican leaf cutting ants. New York Entomol. Soc. J. 85: 222-233. WALLER, D. A. 1980. Leaf-cutting ants and leafriding flies. Ecol. Entomol. 5: 305-306. WEBER, N. A. 1969. A comparative study of the nests, gardens and fungi of the fungus grow­ ing ants, Attini. 6th Congr. Int. Union Study Soc. Ins. (Bern 1969) Proc. Pp. 299-307. WEBER, N. A. 1971. Gardening ants, the Attines. American Philos. Soc, Philadelphia. WEBER, N. A. 1976. A ten-year laboratory col­ ony of Atta cephalotes. Entomol. Soc. Amer. Ann. 69: 825-829. WEBER, N. A. 1982. Fungus ants. In H. R. Hermann, Jr., ed., Social insects. 4: 255—363. Academic, New York.

CHERRETT, J. M., AND F. J. CHERRETT. 1989. A

ElBL-ElBESFELDT,

V.

1., AND E .

ElBESFELDT.

1967. Parasitenabwehren Minima-Arbeiterinen Blattschneide-Ameisen. Zeit. Tierpsychol. 24: 278-281. FOWLER, H. G. 1978. Foraging trails of leafcutting ants. New York Entorno!. Soc. J. 86: 132-136. JAFFE, K., AND E. VILELA. 1989. On nest densities

of the leaf-cutting ant Atta cephalotes in tropical primary forest. Biotropica 21: 234—236. JONKMAN, J. C. M. 1978. Nest of the leaf-cutting anl Atta volleniveiden as accelerators of succes­ sion in pastures. Zeit. Angewan. Entomol. 86: 25-34. LITTLEDYKE,

M.,

AND J.

M. CHERRETT.

1978.

Olfactory responses of the leaf-cutting ants Atta cephalotes (L.) and Acromyrmex octospinosus (Reich) (Hymenoplera: Formicidae) in the laboratory. Bull. Entomol. Res. 68: 273-282. MUDD, A., D. J. PEREGRINE, AND J. M. CHERRETT.

1978. The chemical basis for the use of citrus pulp as a fungus garden substrate by the leafcutting ants Alia cephaloles (L.) and Acromyrmex octospinosus (Reich) (Hymenoptera: Formici­ dae). Bull. Entomol. Res. 68: 673-685. QUINLAN,

R. J.,

AND

J. M.

CHERRETT.

1978.

Aspects of the symbiosis of the leaf-cutting ant Acromyrmex octospinosus (Reich) and its food fungus. Ecol. Entomol. 3: 221-230. ROBINSON, S. W., A. R. JUTSUM, J. M. CHERRETT,

AND R. J. QUINLAN. Í982. Field evaluation of methyl 4-melhylpyrrole-2-carboxylate, and anl trail pheromone, as a component of baits

CORK-HEAD ANTS Formicidae, M y r m e c i n a e , Cephalotini, Zacryptocerus. Portuguese: Cascudas, chiazinhas (Brazil). Turtle ants. T h e r e m a r k a b l e t h i n g a b o u t Zacryptocerus (Wilson 1976) is t h e large, disk-shaped head a n d widely e x p a n d e d p r o t h o r a x of the major w o r k e r s . Colonies nest in aban­ doned b u r r o w s of w o o d - b o r i n g beetles a n d other t u b u l a r hollows in d e a d twigs a n d branches of trees a n d large grasses. In some species, t h e h e a d is convex a n d lowered, so its long axis is a b o u t 120 degrees to t h e midline of the body a n d t h e prothoracic projections t h r u s t above it. With the a n t e r i o r p a r t of t h e body in this position at t h e e n t r a n c e , t h e nest is effec­ tively p l u g g e d , a n d u n w a n t e d intrusions are thwarted. In o t h e r species, the head is cup s h a p e d a n d very r o u n d , held at a 90degree angle, a n d neatly fitted into t h e circular e n t r a n c e hole of the nest without involvement of the t h o r a x . Cork-head ants may even m o v e against o p p o n e n t s , "bulldozing" t h e m o u t of the nest passageways. S o m e species emit a volatile chemical secretion from t h e tip of the a b d o m e n , which can be t i p p e d forward

over the t h o r a x ; o t h e r s rely o n their h a r d ­ e n e d , rigid body, heavy spines, a n d quick m o v e m e n t s for defense. T h i s h e a d p l u g g i n g ability (called "phragmosis") was described early in this century by the famous myrmecologist, William M o r t o n Wheeler, for the Carib­ b e a n species Z. varians a n d since con­ firmed by observations o n Z. texanus (Creighton a n d G r e g g 1954, C r e i g h t o n 1963) a n d o t h e r species (Wheeler a n d Hólldobler 1985). T h e s e were all once placed in Cryptocerus (and in Paracryptocerus), as were o t h e r s , but a r e now p r o p ­ erly assigned to Zacryptocerus. T h e most extremely modified h e a d s a r e f o u n d in the s u b g e n u s Cyathomyrmex. T h e surface of the h e a d is also often covered with a fine g r a n u l a r e n c r u s t m e n t that probably functions as camouflage. Zacryptocerus (fig. 12.11a) are small ants (BL 4—6 m m ) a n d hide in their nests most of the time, a l t h o u g h they occasionally a r e seen crawling o n the outsides of twigs a n d grasses. U n d e r a s t r o n g lens, the cephalic a n d prothoracic disks of the majors of some Zacryptocerus exhibit n u m e r o u s flat­ tened scalelike setae whose function is u n k n o w n b u t probably is associated with phragmosis. T h e feeding habits of Zacryptocerus a r e u n k n o w n , a l t h o u g h they have b e e n fre­ quently seen taking honeydew. O n e species (Z. maculatus) is a social parasite of Azteca ants (Adams 1990). T h e i r foraging activi­ ties a p p a r e n t l y a r e d i u r n a l , w h e n they a r e most often seen. A n u m b e r of species in Central America a r e believed to serve as Batesian mimicry m o d e l s for a wide variety of o t h e r insects, primarily b u p r e s t i d bee­ tles of the g e n u s Agrilus ( H e s p e n h e i d e 1986). T h e p r i m a r y defense of these ants seems to be distastefulness, their sting a n d bite being of little c o n s e q u e n c e . C o r k - h e a d ants of t h e related g e n u s Cephalotes (fig. 4.2d) a r e e n c o u n t e r e d in small g r o u p s on tree t r u n k s w h e r e t h e i r large size (BL 10—15 m m ) , d e e p black

CORK-HEAD ANTS

449

Figure 12.11 ANTS (FORMICIDAE). (a) Cork-head Ant (Zacryptocerus varians). (b) Aztec ant (Azteca instabilis). (c) Carpenter ant (Camponotus sericeiventris). (d) Acrobat ant (Crematogaster stolli). color (often with a silvery sheen), o d d shape, a n d lethargic m o v e m e n t s attract notice.

References ADAMS, E. S. 1990. Interaction between the ants Zacryptocerus maculatus and Azteca trígona: In­ terspecific parasitization of information. Biotropica 22: 200-206. CREIGHTON, W. S. 1963. Further studies on the habits of Cryptocerus texanus Santschi (Hymenoptera: Formicidae). Psyche 70: 133—143. CREIGHTON, W.

S.,

AND R.

E.

GREGG.

1954.

Studies on the habits and distribution of Cryptocerus texanus Santschi (Hymenoptera: Formicidae). Psyche 61: 41—57. HESPENHEIDE, H. A. 1986. Mimicry of ants of the genus Zacryptocerus (Hymenoptera: For­ micidae). New York Entomol. Soc. J. 94: 394-408. WHEELER, D.

E.,

AND B.

HÓLLDOBLER

1985.

Cryptic phragmosis: The structural modifica­ tions. Psyche 92: 337-353. WILSON, E. O. 1976. A social ethogram of the Neotropical arboreal ant Zacryptocerus varians (Fr. Smith). J. Anim. Behav. 24: 354-363.

AZTEC ANTS Formicidae, D o l i c h o d e r i n a e , T a p i n o m i n i , Azteca. All m e m b e r s of this g e n u s a r e arboreal. T h e y place t h e i r nests in s h r u b s a n d trees, sometimes in e x p o s e d situations but m o r e usually in n a t u r a l cavities, hollow stems, rot holes, insect b u r r o w s , a n d the like ( E i d m a n n 1948). T h e r e a r e f o u r g r o u p s , d e p e n d i n g o n t h e type of nest constructed.

450

SAWFLIES, WASPS, ANTS, AND BEES

Several species ( a m o n g t h e m the wellknown A. muelleri, A. alfari; L o n g i n o 19916) form spongy nests of a waxy material within the hollowed, j o i n t e d stems of trumpet trees (Cecropid) (Barnwell 1967, Harada a n d Benson 1988, L o n g i n o 1991a). Their association with the plant is mutualistic: the ant protects the tree from p r e d a t i o n by h e r b i v o r o u s animals, a n d the plant pro­ vides the ant with an a b o d e a n d nutritive substances (see ants a n d plants, above). Each tree s u p p o r t s a single colony of ants. A second g r o u p makes carton nests, usually o n the u n d e r s i d e of a large inclined or horizontal tree b r a n c h . T h e s e nests, c o m p o s e d of masticated wood mixed with saliva, have a p a p e r l i k e exterior; the inte­ rior is a labyrinth of cells a n d tunnels whose walls are m a d e of a substance resem­ bling c a r d b o a r d . Old nests of A. charlifex (called cacarema in Brazil) have external stringy excrescences, giving t h e m a shaggy appearance. Some nests a r e very large a n d may attain a length of 2 to 3 meters a n d stand away from the s u p p o r t i n g b r a n c h 30 to 40 centimeters. T h e nests of o n e Amazonian species (A. trígona; barba or casicero in the Peruvian A m a z o n ) h a n g d o w n in conical shapes that r e m i n d the natives of a man's beard (pi. 4g). T h e s e nests resemble those of arboreal termites a n d a r e often con­ fused with t h e m . However, the latter usu­ ally have runways leading from t h e m and a r e of a m o r e g r a n u l a r material than the structures m a d e by the ants.

T h e t h i r d assemblage (A. olitris, A. ulei, fr. traili, A. delpini) m a k e "ant g a r d e n s , " masses of soil a n d s p r o u t i n g y o u n g plants placed in t h e crotches of limbs in the forest (pi. 4h). A f o u r t h category consists of generalized cavity nesters. Azteca w o r k e r s are all small to very small ants (BL 1 — 1.5 m m ) a n d mostly dull b r o w n (fig- 12.11b). T h e y a r e highly aggressive and, a l t h o u g h lacking a sting, a r e capable of m a k i n g themselves exceedingly offen­ sive by d e s c e n d i n g o n their enemies in droves, biting a n d e m i t t i n g a r e p u g n a n t odor (resembling butyric acid). T h i s o d o r is quite noticeable even from a c r u s h e d individual. T h e a b d o m e n is held erect d u r i n g attacks, "cocktail" fashion like that oí Cremalogaster, but its a p e x is blunt r a t h e r than acute as in that g e n u s a n d less capable of total flexion. T h e r e a r e n o obvious structural characteristics to distinguish the genus. T h e w o r k e r s are p o l y m o r p h i c , the majors have a large, h e a r t - s h a p e d h e a d . T h e i n t e g u m e n t is sometimes soft, giving the body u n u s u a l flexibility. A b o u t 150 species a n d subspecies can be listed in t h e g e n u s from all parts of the American tropics, to which region the genus is restricted; certainly m a n y m o r e live t h e r e a n d will be found w h e n this poorly k n o w n g e n u s is p r o p e r l y studied. Because the application of m a n y n a m e s is uncertain a n d different species a r e often confused, l i t e r a t u r e r e c o r d s must be used with caution (especially with A. alfari).

References BARNWELL, F. H. 1967. Daily patterns in the activity of the arboreal ant Azteca alfari. Ecol­ ogy 48: 991-993. EIDMANN, H. 1948. Zur Kenntnis der Ókologie von Azteca muelleri Em. (Hym. Formicidae), ein Beitrág zum Problem der Myrmecophyten. Abt. Syst. Okol. Geog. Tiere 77: 1-48. HARADA, A.

Y.,

AND W.

W.

BENSON.

1988.

Especies de Azteca (Hymenoptera, Formici­ dae) especializadas em Cecropia spp. (Moraceae): Distribuicáo geográfica e consideracóes ecológicas. Rev. Brasil. Entomol. 32: 423-435. LONGINO, J. T. 199la. Azteca ants in Cecropia

trees: Taxonomy, colony structure, and behav­ iour. In C. R. Huxley and D. F. Cutler, eds., Ant-plant interactions. Oxford, Oxford. LONGINO, ¡. T. 1991ft. Taxonomy of the Ocro/Ha-inhabiting Azteca ants. J. Nat. Hist. 25: 1571-1602.

CARPENTER ANTS Formicidae, Formicinae, C a m p o n o t i n i , Camponotus. Spanish: H o r m i g a s agrias (Costa Rica). Portuguese: Sara sará (esp. C. rufipes), boca azedas, formigas d e c u p i m , jeja (C. abdominalis), tracuá (C. femora tus) (Brazil). Camponotus is the largest ant g e n u s in the world, with over 500 species a n d subspecies listed for the Neotropics. It is also the most widespread a n d ecologically tolerant g r o u p of species, r e a c h i n g all parts of the area including the most r e m o t e islands. U n f o r t u ­ nately, a n d u n d o u b t e d l y because of their g r e a t n u m b e r a n d confusing p o l y m o r ­ phism, the species a r e in a very sad state taxonomically, a n d little is k n o w n of t h e i r biology. Various subgeneric categories h a v e b e e n devised to b r e a k this e n o r m o u s ge­ neric taxon into workable divisions, but these have largely p r o v e d ineffectual in dealing with its complexity (Wheeler 1921). T h e s e ants nest in almost every situa­ tion, c o m m o n l y in s o u n d or r o t t i n g wood, even that used in construction (hence t h e n a m e c a r p e n t e r ants). Many live u n d e r rocks, logs, a n d o t h e r objects o n t h e g r o u n d a n d in ant g a r d e n s , b r o m e l i a d s , a n d o t h e r epiphytes. M e m b e r s of t h e subg e n u s Colobopsis nest in hollow twigs o r b r a n c h e s in trees, in insect galls a n d nuts, a n d have soldiers with peculiar flatfronted h e a d s (like Zacryptocerus) for block­ ing the single e n t r a n c e hole (phragmosis). A species in the s u b g e n u s Myrmobrachys, Camponotus senex, constructs large b a g nests in trees, b i n d i n g t o g e t h e r leaves with silk p r o d u c e d by the larvae in a fashion similar to that of the Old World weaver ants, Oecophylla ( S c h r e m m e r 1979).

CARPENTER ANTS

451

Camponotus a r e not e q u i p p e d with stings but are p u g n a c i o u s a n d capable of inflict­ ing c o n s i d e r a b l e pain with their mandibles a n d secretions of formic acid a n d o t h e r chemicals that they s p r a y into the bite. T h o s e living in silk nests r u s h out o n t o the surface of t h e nest at the slightest provoca­ tion, setting u p a rattle that resonates loudly in t h e d r y b a g a n d scares away any potential attacker. Species whose food habits a r e k n o w n are generally o m n i v o r o u s but prey to a large e x t e n t on o t h e r insects a n d a r a c h n i d s (Matthiesen 1980). Few actually b u r r o w into s o u n d wood a n d d o structural d a m ­ age. C. abdominalis is o n e such pest, which also attacks c o m m e r c i a l beehives, whose p o p u l a t i o n s it may d e c i m a t e . C. sericeiventris (fig. 12.11c) is a com­ m o n , well-known, p o l y m o r p h i c species that attracts attention because of its h a n d ­ some silver or gold p u b e s c e n c e ( B u s h e r et al. 1985). W o r k e r s a r e u n i q u e in t h e g e n u s in the possession of a s h a r p m e d i a n crest r u n n i n g a l o n g t h e d o r s u m of the mesoa n d m e t a t h o r a x a n d s h a r p spines project­ ing obliquely from the a n t e r o l a t e r a l an­ gles of the p r o t h o r a x . T h e species nests in decayed p o r t i o n s of s t a n d i n g tree t r u n k s some distance above t h e g r o u n d a n d is often seen d u r i n g t h e day s c r a m b l i n g u p a n d d o w n tree t r u n k s in search of prey. Major w o r k e r s a r e very aggressive a n d able to bite severely. C. sericeiventris is mimicked by a c e r a m b y c i d beetle, Eplophorus velutinus (W. M. W h e e l e r 1931), a n d a velvet ant, Pappognatha myrmiciformis (G. C. W h e e l e r 1983). T h e various s u b g e n e r a previously recognized (W. M. W h e e l e r 1921) have now mostly been found invalid. Camponotus a r e so diverse structurally that they defy diagnosis. O n e constant feature is t h e isolation of the a n t e n n a l sockets from the clypeus, b u t this is some­ times difficult to observe. The workers are p o l y m o r p h i c , a n d those of m a n y species are r a t h e r large (BL 1 0 - 1 8 m m ) .

452

SAWFLIES, WASPS, ANTS, AND BEES

References BUSHER,

C.

E.,

P.

CALABI,

AND

].

F.

A

TRANIELLO. 1985. Polymorphism and division of labor in the Neotropical ant Camponotus sericeiventris Guerin (Hymenoptera: FormiHdae). Entomol. Soc. Amer. Ann. 78: 221-228 MATTHIESEN, E. A. 1980. Sará-sará, forrniga predadora de escorpióes e opilioes. R ev Agrie. (Piracicaba, Sao Paulo) 55: 239-241. SCHREMMF.R, E 1979. Das nest der neotropischen Weberameise Camponotus (Myrmobrachys) senex Smith (Hymenoptera, Formicidae). Zool. Anz 203: 273-282. WHEELER, G. G. 1983. A mutillid mimic of an ant (Hymenoptera: Mutillidae and For­ micidae). Entomol. News 94: 143-144. WHEELER, W. M. 1921. Professor Emery's subgenera of the genus Camponotus Mayr Psyche 28: 16-19. WHEELER, W. M. 1931. T h e

ant Camponotus

(Myrmeponns) sericeiventris Guérin mimic. Psyche 38: 86-98.

and its

OTHER ANTS T h e acrobat ants form a large, nearly cosmopolitan g e n u s (Myrmicinae, Crematogastrini, Crematogaster) a n d a r e well repre­ sented in the Neotropics ( B u r e n 1958). T h e y occupy m a n y habitats a n d are com­ m o n mutualistic associates of myrmecophytes. Small, m o n o m o r p h i c , a n d some­ what r e s e m b l i n g aztec ants in the habit of elevating their a b d o m e n w h e n aroused, they have a sharply pointed, heart-shaped a b d o m e n that is flat on top, convex below (fig. 12.1 Id). T h e y lack any o d o r when d i s t u r b e d o r c r u s h e d . Nests often are of the carton type, placed in trees or shrubs, but are also situated in the g r o u n d , in termite nests, in n a t u r a l plant cavities, in rotting logs, in d e a d twigs a n d branches a n d tree t r u n k s , a n d in epiphytes. T h e s t r u c t u r e a n d use of the sting by acrobat ants is u n i q u e . T h e o r g a n is flat­ tened a n d b l u n t (spatulate) at the tip and quite unable to pierce the i n t e g u m e n t of prey or an enemy. W h e n a droplet of v e n o m is secreted, it clings to the tip of the

sting from which it is a p p l i e d topically to its target. To c o m p l e t e this process, the a b d o ­ men is elevated a n d a r c h e d all the way forward over t h e t h o r a x a n d h e a d in o r d e r to bring its a p e x in contact with the substra­ tum a h e a d of t h e ant's body. M e m b e r s of the g e n u s seem to have a particular proclivity for h o n e y d e w a n d are notorious p r o t e c t o r s of the h o m o p t e r o u s producers of this substance. S o m e construct succursal nests o r "tents" a r o u n d colonies of a p h i d s a n d coccids which they keep in good r e p a i r a n d which h i d e a n d protect their c h a r g e s (W. M. W h e e l e r 1906). O n e species raids the nests of the kelep (Eclatomma tuberculatum) (D. E. W h e e l e r 1986). Big-headed ants (Myrmicinae, Pheidolini, Pheidole; hormigas cabezones) are also small but by contrast form a large assem­ blage of several h u n d r e d species in Latin America (Kusnezov 1951). T h e y a r e ubiqui­ tous a n d varied in their nesting a n d feed­ ing habits, which a r e generally practiced along two lines, g r a m i n i v o r y a n d p r e d a tion. W o r k e r s of both are decidedly dimorphic, consisting of small m i n o r s with normal form a n d majors with e n o r m o u s heads a n d powerful, r i d g e d mandibles (fig. 12.12a). T h e s e formidable jaws a r e used for fighting in t h e p r e d a c e o u s g r o u p ; for cracking seeds a m o n g the graminivores. I n all w o r k e r s , t h e h e a d t e n d s to be longer than wide a n d the a n t e n n a e r a t h e r long. Nests of both types a r e usually situ­

ated in the g r o u n d but a r e s o m e t i m e s m a d e in decaying logs, termite nests, natu­ ral cavities in twigs, a n d even in the for­ micaries of m y r m e c o p h y t e s . T h e stinging habits of fever ants (Pseudom y r m e c i n a e , Pseudomyrrnex; tachi, formigasde-novato in Brazil, hormiga del cornizuelo in Costa Rica) have already been described in connection with ant plants. Not all, how­ ever, have close a n d specific associations with m y r m e c o p h y t e s , a l t h o u g h they always a r e arboreal to some d e g r e e , usually nest­ ing in p r e f o r m e d cavities, hollow stems, a n d the like ( K e m p f 1960). Colonies of some species have been f o u n d ensconced in termite nests. Colonies in acacias ("bull'sh o r n acacia ants") may have o n e q u e e n o r multiple q u e e n s . In the latter case, colonies may spread to m a n y plants in an area a n d e x p a n d to e n o r m o u s size, the largest of any ant in the world (as m a n y as 1.8—3.6 million workers) ( J a n z e n 1973). These are m e d i u m to large (5 to 11 m m BL), r a t h e r elongate, slender ants, with slightly down c u r v e d a b d o m e n s a n d a twos e g m e n t e d waist, usually r e d d i s h or yellow­ ish brown in color (fig. 12.12b). Each of the two segments of the waist bears a low r o u n d e d n o d e . Workers have a good-sized h e a d , with large eyes. They are active a n d agile a n d quick to sting. T h e sting a n d venom a p p a r a t u s is exceptionally well de­ veloped, a n d the w o r k e r s can inflict pain to a d e g r e e second only to the giant solitary h u n t i n g ants (Ponerini). Sensitive p e r s o n s

Rgure 12.12 ANTS (FORMICIDAE). (a) Big-headed ant {Pheidole fallax). (b) Bull's-horn acacia •nt [Pseudomyrrnex ferrugineus). (c) Argentine ant (Iridomyrmex humilis). (d) Stink ant (Tapinoma ftolanocephalum).

OTHER ANTS

453

react severely after t h e sting, the skin swelling a n d often blistering, occasionally with fever following. A polysaccharide recently isolated from this ant's v e n o m is k n o w n to be r e m e d i a l in the t r e a t m e n t of r h e u m a t o i d arthritis (Schultz a n d A r n o l d 1977). T h e g e n u s is found only in t h e New World, w h e r e it has evolved over o n e h u n d r e d fairly u n i f o r m species. A very well-known Neotropical a n t is t h e A r g e n t i n e ant, Iridomyrmex humilis (Dolichoderinae, T a p i n o m i n i ) (fig. 12.12c) (Mallis 1964). T h e species' fame derives from its particularly n o x i o u s n a t u r e in the s o u t h e r n half of t h e U n i t e d States, w h e r e it was i n t r o d u c e d via New O r l e a n s very late in t h e n i n e t e e n t h century. It is native a n d c o m m o n to most of tropical S o u t h America (described originally from n e a r B u e n o s Aires) a n d probably was t r a n s p o r t e d n o r t h in s h i p m e n t s of coffee. Colonies of this a n t m a y n u m b e r in the m a n y t h o u s a n d s of individuals a n d often m e r g e a n d have n u m e r o u s q u e e n s (poly­ gyny). Called t h e " G e n g h i s K h a n of t h e e m m e t world," it is aggressive a n d often displaces o t h e r g r o u n d - n e s t i n g species in its territory of invasion. W o r k e r s a r e small (BL 2 . 2 5 - 2 . 7 5 m m ) a n d dull b r o w n a n d h a v e an a b d o m e n capable of c o n s i d e r a b l e distension w h e n they a r e e n g o r g i n g with food, especially honeydew, of which they a r e fond. T h e single waist s e g m e n t has a well-developed node. O n e of t h e most objectionable traits of the species is its habit of fostering a p h i d s , mealybugs, a n d o t h e r h o m o p t e r o u s pests in o r c h a r d s a n d h o r t i c u l t u r a l plots. It is t h e most c o m m o n of t h e several h o u s e infecting types in S o u t h a n d N o r t h A m e r ­ ica, h e n c e its c o m m o n n a m e , "sugar a n t s " (ciganas acucareiras in Brazil). While its presence in t h e n o r t h h a s evoked a severe antipathy, it seems to be of little c o n c e r n in its native c o u n t r i e s . Stink

454

ants

(Dolichoderinae,

Tapino­

SAWFLIES, WASPS, ANTS, AND BEES

mini, Tapinoma) {hormigas hediondas) rather resemble Iridomyrmex in their thin flexible i n t e g u m e n t , small size, a n d single segment in the waist. T h e n o d e of the latter however, is u n d e v e l o p e d . Only a dozen or so species live in t h e Neotropics. Some of these are easily recognized by t h e type of nest they build, a loose e n c r u s t m e n t of e a r t h laid as a linear t u n n e l along twigs a n d as a flat p a n c a k e on t h e undersides of leaves. Excited workers b e c o m e aggressive a n d emit a characteristic o d o r like that of butyric acid. A widely known pest species is Tapinoma melanocephalum, t h e "crazy ant" (fig. 12.12d), whose wild gyrations on the table at mealtime always attract notice. O t h e r a n t g e n e r a characteristic of the A m e r i c a n tropics a r e Hypoclinea (formerly Monads; fig. 10.16b), a dweller of the rain forest canopy; Daceton, a large myrmicine with a g r o t e s q u e h e a d a n d spined thorax; a n d Pogonomyrmex, the familiar harvester ants that a r e restricted to d r i e r areas (Kugler 1984, González-Espinoza 1984).

References BURÉN, W. F. 1958. A review of the species of Crematogaster, sensu stricto, in North America (Hymenoptera: Formicidae). Pt. 1. New York Entorno! Soc. J. 66: 119-134. GONZÁLEZ-ESPINOZA,

M.

1984.

Patrones

de

comportamiento de forrajeo de hormigas recolectoras Pogonomyrmex spp. en ambientes fluctuantes (Hymenoptera Formicidae). Fol. Entomol. Mexicana 61: 147-158. JANZEN, D. H. 1973. Evolution of polygynous obligate acacia-ants in western Mexico. J. Anim. Ecol. 42: 727-750. KEMPF, W. W. 1960. Estudos sobre Pseudomyrmex I—III (Hymenoptera: Formicidae). (I) Rev. Brasil. Entomol. 9: 5-32. (II) Studia Ento­ mol. 1: 433-462. (Ill) Studia Entomol. 4: 369-408. KUGLER, C. 1984. Ecology of the ant Pogono­ myrmex mayri: Foraging and competition. Biotropica 16: 227-234.' KUSNEZOV, N. 1951. El género Pheidole en la Argentina (Hymenoptera, Formicidae). Acta Zool. Argentina 12: 5-88. MALLIS, A. 1964. Handbook of pest control. 4th ed. MacNair-Dorland, New York. Pp536-554.

SCHULTZ, D. R., AND P. I. ARNOLD. 1977. Venom

of the ant Pseudomyrmex sp.: Further character­ ization of two factors that affect human complement proteins. J. Immunol. 119: 1690-1699. WHEELER, D. E. 1986. Ectatomma tuberculatum: Foraging biology and association with Cremato­ gaster (Hymenoptera: Formicidae). Entomol. Soc. Amer. Ann. 79: 300-303. WHEELER, W. M. 1906. T h e habits of the tentbuilding ant (Crematogaster lineolata Say). Amer. Mus. Nat. Hist. Bull. 22(1): 1-18.

BEES Apoidea. Spanish: Abejas. Portuguese: Abelhas. Quechua: A m o , u r u n k u . TupiGuaraní: Eira. Náhuatl: Pipiyolmeh, sing, pipiyolin. Mayan: C a b . Bees ( S t e p h e n et al. 1969) c o m p r i s e a major g r o u p of e v o l u t i o n a r y a d v a n c e d H y m e n o p t e r a , derived from sphecid wasps through specialization for close association with flowers, from which they acquire pollen a n d nectar, their p r i m a r y food sources; in contrast, wasps mostly take food of a n i m a l origin. I n t h e process of collecting these substances, bees pollinate plants a n d t h u s a r e of major i m p o r t a n c e ecologically a n d economically to both natu­ ral plant c o m m u n i t i e s a n d c r o p s (Frankie and Coville 1979). T h i s process is aided by the most distinctive feature of bees, a thick body vestiture of finely b r a n c h e d (plumose) hairs, to which t h e pollen grains cling tenaciously a n d o n which they are dispersed a m o n g flowers. Bees usually collect pollen a n d nectar directly from t h e a n t h e r s a n d petal bases. Some however, have d e v e l o p e d special techniques for forcing these substances from difficult flowers. E d u c a t e d bees hover over blossoms with t u b u l a r a n t h e r s and buzz their wings violently, blowing t h e pollen o n t o their bodies ("buzz pollina­ tion"; B u c h m a n n 1974), o r bite holes n e a r the base of t h e corolla of d e e p - t h r o a t e d flowers, taking a s h o r t c u t to steal t h e

nectar from t h e side ("nectar r o b b i n g " ; Wille 1963). T h e latter t e c h n i q u e circum­ vents the a n t h e r s a n d fails to p r o v i d e t h e service of pollination (Barrows 1976). Dif­ ferent species may c o m p e t e directly for nectar sources ( H e d s t r ó m 1984). Most bees a r e solitary a n d m a k e t u b u l a r b u r r o w s for their nests in t h e soil o r in o t h e r locations (hollow twigs, t e r m i t e nests, rotting o r s o u n d wood, etc.) (Janvier 1955). H e r e they form cells in which to rear their larvae. A n unidentified species of Anthophora ( A n t h o p h o r i d a e ) is d o i n g considerable d a m a g e to Incan a n d r e c e n t a d o b e structures in t h e U r u b a m b a Valley of Peru by b o r i n g into t h e m in large n u m b e r s to m a k e their nests (orig. obs.). Many types a r e parasitic a n d d e v e l o p in the nests of o t h e r bees. T h e s e t e n d to be less hairy t h a n nonparasitic forms a n d to m o r e closely resemble wasps. Still o t h e r s are social (Michener 1974). Like wasps, female bees usually can sting b u t only d o so in defense. T h e venoms of some species, particularly those in t h e subfamily A p i n a e (honeybees a n d others), may cause serious c o n s e q u e n c e s when injected into h u m a n s ( O ' C o n n o r a n d Peck 1978). A curious habit of t h e males of m a n y bees is t h e formation of sleeping aggregations. Masses of d o z e n s or even h u n d r e d s of individuals lock m a n d i b l e s to leaves or twigs to pass t h e later a f t e r n o o n a n d night. H u n d r e d s of species of t h e several fami­ lies of bees live in Latin America (Michener 1954). Bees generally a r e most diverse in the arid a n d semiarid portions, a l t h o u g h m o r e highly evolved g r o u p s have a p p a r ­ ently arisen in lowland rain forests (Mich­ e n e r 1979, M o l d e n k e 1976).

References BARROWS, E. M. 1976. Nectar robbing and pollination of Lantana cámara (Verbenaceae). Biotropica 8: 132-135. BUCHMANN, S. L. 1974. Buzz pollination of Cassia quiedondilla (Leguminosae) of bees of

BEES

455

the genera Centris and Melipona. So. Calif. Acad. Sci. Bull. 73: 171-173. FRANKIF,, G. W., AND R. COVILLE.

1979. An

experimental study on the foraging behavior of selected solitary bee species in the Costa Rican dry forest (Hymenoptera: Apoidea). Kans. Entomol. Soc. j . 52: 591-602. v HEDSTRÓM, I. 1984. Interference competition between two species of Pliloglossa bees (Hymenoptera, Colletidae) in the central valley of Costa Rica. Brenesia 22: 219-231. JANVIER, H. 1955. Le nid et la nidification chez quelques abeilles des Andes tropicales. Ann. Sci. Natur. Zool. Biol. Anim. 17: 31 1-349. MICHENER, C. D. 1954. Bees of Panamá. Amer. Mus. Nat. Hist. Bull. 104: 1-175. MICHF.NER, C. D. 1974. T h e social behavior of the bees: A comparative study. Belknap Press, Harvard Univ., Cambridge. MICHF.NER, C. D. 1979. Biogeography of the

bees. Missouri Bol. Garden Ann. 66: 2 7 7 347. MOLDENKE, A. R. 1976. Evolutionary history and diversity of the bee faunas of Chile and Pacific North America. Wasmann J. Biol. 34: 147-178. O'CONNOR, R., AND M. L. PECK. 1978. Venoms

of Apidae. hi S. Bettini, ed., Venoms of arthropods. Springer, Berlin. Pp. 613—659. STEPHEN,

W.

P., G.

E. BOHART,

AND P. E

TORCHIO. 1969. T h e biology and external morphology of bees. Agrie. Exper. Sta., Ore. State Univ., Corvallis. WILLE, A. 1963. Behavioral adaptations of bees for pollen collecting from Cassia flowers. Rev. Biol. Trop. 11: 205-210.

SOLITARY BEES T h e majority of bees a r e solitary, conduct­ ing their lives i n d e p e n d e n t of o t h e r s of

their own kind a n d quite a p a r t from hives or c o m m u n a l nests. A m o n g these, a few forms have developed subsocial habits however, a n d live in aggregations wherein females may actually c o o p e r a t e with nestbuilding chores o r form primitive colonies with a few offspring w h o r e m a i n with her a n d m a k e u p a kind of primitive worker caste.

Leaf Cutter Bees Megachilidae, Megachilinae, Megachilini. Spanish: Ronsapitas (Peru). T h e females of m a n y species of this tribe, the most n u m e r o u s of which belong to the genus Megachile (fig. 12.13a), cut circular pieces out of leaves which they use to line their nests. The nests a r e almost always placed in preexisting cavities, in pockets or n a r r o w cracks a n d crevices in rocks and h a r d soil, a n d in hollow stems. T h e nests are elongate a n d t h e cells laid in tandem. O t h e r species form b r o o d cells from saw­ dust, plant down, leaf p u l p , a n d other materials c e m e n t e d t o g e t h e r with resin collected from plants. T h e s e d o not secrete wax o r cementlike substances as d o other bees. Leaf cutter bees a r e w i d e s p r e a d in the Neotropics in all habitats, from rain forest to coastal desert to m o u n t a i n s . T h e y are always recognized by t h e presence of only two s u b m a r g i n a l cells in t h e fore wing, extra-long tongues, a n d a large labrum, covered by t h e mandibles w h e n they are closed. T h e y a r e often d a r k , solid, black or

Figure 12.13 BEES, (a) Leaf cutter bee (Megachile leucographa, Megachilidae). (b) Sweat bee (Lasioglossum sp., Halictidae). (c) Centris bee (Centris inermis, Anthophoridae).

456

SAWFL1ES, WASPS, ANTS, AND BEES

brown colors. Many o t h e r s a r e buff with banded a b d o m e n s , a n d males of m a n y species sport "flowerlike" structural devel­ opments o n t h e forelegs which a r e used in mating. L e g s e g m e n t s a r e dilated a n d exca­ vated o n t h e i n n e r surface a n d fringed with e l o n g a t e hairs, a u n i q u e a r r a n g e m e n t a m o n g bees. Leaf cutters also differ from other bees by c a r r y i n g their pollen loads on t h e u n d e r s i d e of t h e a b d o m e n instead of on t h e legs; resting individuals often elevate t h e a b d o m e n characteristically. Megachile d o n o t e x t e n d their r a n g e south farther t h a n P a n a m a , b u t o t h e r gen­ era, like Cressomella, Pseudocentron, a n d Chrysosarus, occupy most all of t e m p e r a t e to tropical Latin America. T h e t a x o n o m y of leaf cutter bees h a s b e e n recently u p ­ dated by Mitchell (1980).

Reference MITCHELL, T. B. 1980. A generic revision of the megachiline bees of the Western Hemisphere (Hymenoptera: Megachilidae). Dept. Ento­ mol. N.C. State Univ. Contrib.

T h e latter also attract attention because of their brilliant metallic g r e e n o r blue colors ("green sweat bees") (Eickwort 19696). (Small stingless bees in t h e g e n u s Trígona also display t h e sweat-drinking habit a n d likewise a r e called "sweat bees.") Sweat bees a r e m e d i u m - s i z e d (BL 5—10 m m ) , c o m m o n , flower-visiting bees that nest in t h e soil. T h e nests a r e d u g in compact soil (a few hollow o u t wood; Eickwort a n d Eickwort 1973a, 19736) a n d consist of an oblique tube with b r o o d cells issuing directly from its walls (most A u g o ­ chlorini) o r at t h e e n d s of long, lateral tubes (Agopostemon a n d allies). Halictine bees a r e also r e m a r k a b l e for the primitively eusocial behavior exhibited by m a n y species: colonies of the most highly evolved forms have legitimate q u e e n s a n d w o r k e r castes, a l t h o u g h t h e latter a r e scarcely distinguishable from t h e former. Colonies a r e very small, never n u m b e r i n g m o r e t h a n several individuals. O t h e r spe­ cies of Augochlorella a n d Augochlora show habits r a n g i n g from completely solitary to subsocial o r semisocial.

Sweat Bees Halictidae, Halictinae, A u g o c h l o r i n i , Augochlora et al., Halictini, Lasioglossum and o t h e r g e n e r a . Spanish: Lameojos, morrocujes (Peru); c h u p a d o r e s (Costa Rica). Small bees of t h e subfamily Halictinae a r e called "sweat bees" because so m a n y have a fondness for h u m a n p e r s p i r a t i o n . A n over­ heated p e r s o n acts as a n irresistible attrac­ tion for these bees, which b e c o m e a n extreme nuisance because of their insis­ tence o n d r i n k i n g sweat, in so d o i n g enter­ ing the eyes, nose, a n d ears a n d generally pestering one's b o d y o n h o t , h u m i d days. There a r e two basic kinds of sweat bees (Eickwort 1969a): those b e l o n g i n g to t h e tribe Halictini, principally t h e g e n u s Lasio­ glossum (fig. 12.13b), a n d m e m b e r s of t h e tribe Augochlorini, primarily Agopostemon.

References EICKWORT, G. C. 1969CJ. Tribal positions of

Western Hemisphere green sweat bees, with comments on their nest architecture (Hy­ menoptera: Halictidae). Entomol. Soc. Amer. Ann. 62: 652-660. EICKWORT, G. C. 19696. A comparative morpho­ logical study and generic revision of the Augochlorine bees (Hymenoptera: Halicti­ dae"). Univ. Kans. Sci. Bull. 48: 325-524. EICKWORT, G. C , AND K. R. EICKWORT. 1973rt.

Aspects of the biology of Costa Rican halictine bees, V. Augochlorella edenlala (Hy­ menoptera: Halictidae). Kans. Entomol. Soc. J. 46: 3-16. EICKWORT, G. C , AND K. R. EICKWORT. 1973ft.

Notes on the nests of three wood-dwelling species of Augochlora from Costa Rica (Hy­ menoptera: Halictidae). Kans. Entomol. Soc. J. 46: 17-22. SAKAGAMI, S. E, AND C. D. MICHENER.

1962.

The nest architecture of the sweat bees. Univ. Kansas, Lawrence.

SOLITARY BEES

457

Centris Bees Anthophoridae, Anthophorinae, C e n t r i d i n i , Centris. Bees of the g e n u s Centris form a ubiquitous g r o u p in the Neotropics w h e r e some 150 species occur widely. T h e y a r e fairly large, robust bees (BL 1—2.5 cm) a n d a r e con­ spicuously m a r k e d with c o n t r a s t i n g black, yellow, a n d buff colors (fig. 12.13c). A c o m m o n p a t t e r n is a n all-red or all-black a b d o m e n a n d light b r o w n t h o r a x , al­ t h o u g h several species g r o u p s display b a n d e d , b u m b l e b e e l i k e pelages. O t h e r s ( s u b g e n u s Melanocentris) a r e very large, are mostly black (buff t h o r a x ) , a n d m u c h re­ semble c a r p e n t e r bees. Structurally, centris bees a r e characterized by m a n d i b l e s with p o i n t e d teeth, the basal tarsal s e g m e n t of the h i n d leg which is s h o r t e r t h a n the tibia, a n d a small m a r g i n a l cell in the fore wing. All a r e solitary, a l t h o u g h nesting colo­ nies a n d a g g r e g a t i o n s of sleeping males a r e often observed. Nesting habits vary greatly: nests a r e c o n s t r u c t e d in diverse substrates, such as clay b a n k s , r o t t e n wood, tree holes (Frankie et al. 1988), a n d even in a r b o r e a l t e r m i t e nests, b u t in all cases, they are t u b u l a r b u r r o w s with o n e to a few b r a n c h e s , each c o n t a i n i n g several b r o o d cells in series (Vinson a n d Frankie 1977). T h e y o u n g a r e r e a r e d entirely on pollen. S o m e species use e m p t y cells in a b a n d o n e d bees' nests. Preexisting b u r r o w s a r e used, or a new o n e is excavated (Coville et al. 1983). Males of m a n y Centris species a r e k n o w n to be highly territorial, patrolling specific areas, fighting off rival bees, a n d attacking o t h e r i n t r u d e r s . T h e y even m a r k their areas with a citrallike substance p r o d u c e d by the p h a r y n g e a l g l a n d s (Raw 1975). Males also display two diverse m a t i n g strategies c o r r e l a t e d with d i m o r p h i s m in body size. M u c h l a r g e r males, called " m e t a n d e r s , " fly over n e s t i n g areas a n d locate e m e r g i n g females by odor. O n locat­ ing the odor, they dig d o w n a n d m a t e with

458

SAWFL1ES, WASPS, ANTS, AND BEES

females as they e m e r g e . Smaller, "satellite" males hover within territories at the periph­ ery of nesting areas, waiting for females that escape the larger forms u n m a t e d ( A l c o c k e t a l . 1977, Frankie et al. 1980).

References ALCOCK, J., C. E. JONES, AND S. L. BUCHMANN

1977. Male mating strategies in the bee Centris patlida Fox (Anthophoridae: Hvmenoptera). Amer. Nat. I l l : 145-155. COVILLE,

R.

E.,

G.

W.

FRANKIE, AND S.

Figure 12.14 BEES, (a) Carpenter bee (Xylocopa fimbriata, Anthophoridae). (b) Orchid bee [Euglossa purpurea, Apidae). (c) Orchid bee (Eulaema meriana, Apidae).

B

VINSON. 1983. Nest of Centris segregala (Hymenoptera: Anthophoridae) with a review of the nesting habits of the genus. Kans Entomol. Soc.J. 56: 109-122. FRANKIE, G. W., S. B. VINSON, AND R. E. COVILLE

1980. Territorial behavior of Centris adani and its reproductive function in the Costa Rican dry forest (Hymenoptera; Anthophoridae). Kans. Entomol. Soc.J. 53: 837-857. FRANKIE, G. W., S. B. VINSON, L. E. NEWSTROM,

AND J. F. BARTHELL. f988. Nest site and habitat preferences of Centris bees in the Costa Rican dry forest. Biotropica 20: 30f-310. RAW, A. 1975. Territoriality and scent marking by Centris males (Hymenoptera, Anthopho­ ridae) in Jamaica. Behavior 54: 31 1-321. VINSON, S. B., AND G. VV. FRANKIE. 1977. Nest of

Centris aethyctera (Hymenoptera: Apoidea: Anthophoridae) in the dry forest of Costa Rica. Kans. Entomol. Soc.J. 50: 301-311.

Carpenter Bees A n t h o p h o r i d a e , Xylocopinae, Xylocopini, Xylocopa. Spanish: Ronsapas (Peru). Náhuatl: Xicotes (Mexico). T h e s e are the largest bees in the Neo­ tropics, the body length of some species (Xylocopa fimbriata or X. frontalis) measuring u p to 26 millimeters or m o r e . Thickly pubescent, with d a r k wings a n d ponderous flight, they somewhat resemble bumblebees but lack their well-defined yellow color b a n d s on the a b d o m e n or thorax (fig. 12.14a). T h e back of the t h o r a x is flattened, a n d they possess powerful, blunt mandi­ bles, characteristics not p r e s e n t in that o t h e r g r o u p of large bees. A n o t h e r special feature identifying the g e n u s is a triangular cell f o r m e d by the veins in the center of the

fore wings t o w a r d the leading e d g e . Usually all black, s o m e species possess g r e e n , orange, r e d , white, yellow, or o t h e r colored pelage; s o m e species a r e sexually d i m o r ­ phic, the males b e i n g s h a d e d entirely with tan, the females d a r k . T h e wings are often dark tinted a n d display violaceous, even resplendent metallic reflections (X. ornato). T h e biology of only a few species is known (Gerling et al. 1989, J a n z e n 1966, Sage 1968). T h e i r habit of b u r r o w i n g into wood gives these nonsocial bees their com­ mon n a m e . T h e b u r r o w s ramify a n d anas­ tomose t h r o u g h the wood to form c o m p l e x tubular galleries in which the larvae are reared. D u r i n g their d e v e l o p m e n t a l pe­ riod, the larvae a r e provisioned with pollen and sealed into linear rows of cells with disk-shaped partitions. O n e m e r g i n g , the neotenic bees m u s t wait their t u r n to es­ cape, since less-developed siblings from eggs laid late block their exit p a t h s . All sorts of d e a d wood serve as substrata for the nests, including tree t r u n k s , stumps, logs, large b r a n c h e s , a n d stems. Rarely d o the bees select living plant tis­ sues. A few species nest in t h e hollow, jointed stems of b a m b o o ( s u b g e n u s Stenoxylocopa; H u r d Í978) a n d some even in the ground, rock crevices, or e a r t h e n tunnels. Occasionally, s t r u c t u r a l timbers a n d con­ struction wood a r e attacked a n d weak­ ened, so that c a r p e n t e r bees are sometimes destructive. T h e Galápagos c a r p e n t e r bee (X. darwini) is the principal pollen vector

a m o n g plants in the Galápagos Islands (Linsley et al. 1966). T h e r e are about 150 New World tropical species of Xylocopa, g r o u p e d into 17 s u b g e n e r a ( H u r d a n d M o u r e 1963). T h e g r o u p is considered poorly d e v e l o p e d in c o m p a r i s o n to the Old World, w h e r e most species also occur in w a r m e r climes. C a r p e n ­ ter bees have been successful in r e a c h i n g distant oceanic islands, probably t h r o u g h their wood-nesting habits. Infested logs, rafting on ocean c u r r e n t s , likely have car­ ried colonizing females to the Galápagos, Revillagigedo, a n d West I n d i a n islands ( H u r d 1958).

References GERLING,

D.,

H.

H.

W.

VELTHUIS, AND

A.

HEFETZ. 1989. Bionomics of the large carpen­ ter bees of the genus Xylocopa. Ann. Rev. Entomol. 34: 163-190. HURD, JR., P. D. 1958. The carpenter bees of the eastern Pacific oceanic islands (Hymenop­ tera: Apoidea). Kans. Entomol. Soc. J. 31: 249-255. HURD, JR., P. D. 1978. Bamboo-nesting carpen­ ter bees (Genus Xylocopa Latreille) of the subgenus Stenoxylocopa Hurd and Moure (Hy­ menoptera: Anthophoridae). Kans. Entomol. Soc. J. 51: 746-764. HURD, JR., P. D.,

AND J. S. MOURE.

1963.

A

classification of the large carpenter bees (Xylocopini) (Hymenoptera: Apoidea). Univ. Calif. Publ. Entomol. 29: 1-365. JANZEN, D. H. 1966. Notes on the behavior of the carpenter bee Xylocopafimbriatain Mexico (Hymenoptera: Apoidea). Kans. Entomol. Soc. I. 39: 633-641.

SOLITARY BEES

459

LINSLEY, E. G., C. M. RICK, AND S. G. STEPHENS.

1966. Observations on the floral relationships of the Galápagos carpenter bee. Pan-Pacific Entomol. 42: 1-18. SAGE, R. D. 1968. Observations on feeding, nesting and territorial behavior of carpenter bees genus Xylocopa in Costa Rica. Entomol. Soc. Amer. Ann. 6 1 : 884-889.

Orchid Bees A p i d a e , B o m b i n a e , Euglossini, Euglossa, Eulaema et al. Spanish: Chiquizás (Costa Rica, Eulaema meriana). Gold Bees, e m e r a l d bees. O r c h i d bees (Dressier 1982) c o m p r i s e five g e n e r a (Kimsey 1987) of m o r e t h a n two h u n d r e d species restricted to the forested, tropical p o r t i o n s of the New World ( M o u r e 1967). T h e males of all a r e attracted to orchid flowers only by their o d o r s , n o food in the form of nectar or pollen being taken. O n arriving at t h e flower, the bee scrapes the petals with its front tarsi a n d sponges u p the e x ú d a t e from the w o u n d with masses of hairs o n t h e tarsal s e g m e n t s . It t h e n transfers the substance to storage tissues inside t h e e n l a r g e d h i n d tibia (Dodson 1966). In the process, the flower is pollinated, usually in a very specific m a n ­ ner. Males also collect fragrances from the blooms (and o t h e r parts) of plants in o t h e r families, timing their visits to coincide with the time of flowering ( A r m b r u s t e r a n d M c C o r m i c k 1990). T h e value of the substance to the b e e is not k n o w n . T h e o r i e s suggest a possible internal metabolic role or use as a n ac­ q u i r e d p h e r o m o n e . If t h e latter, the sub­ stance volatilizes a n d may call o t h e r males. A swarm of such males, with their brilliant colors a n d loud buzzing, could l u r e fe­ males a n d effect the m e e t i n g of the sexes m u c h like t h e lek a g g r e g a t i o n s of brightly colored birds (Kimsey 1980). It has also been suggested that males may also utilize the chemical as a metabolite (like a vitamin) o r as a scent to m a r k a m a t i n g territory. Many of the floral fragrances have b e e n

460

SAWFLIES, WASPS, ANTS, AND BEES

identified chemically a n d may be used in their p u r e state to attract male bees in the wild. Several are c o m m o n d r u g s t o r e com­ p o u n d s or p e r f u m e ingredients such as cineole (oil of eucalyptus), methyl salicylate (oil of w i n t e r g r e e n ) , benzyl acetate ("bubb l e g u m flavor"), a n d even vanillin a n d are used by entomologists to survey a n d study these bees (Dodson et al. 1969, J a n z e n et al 1982). Female euglossines d o not visit orchids but d o visit o t h e r kinds of flowers for nectar a n d pollen, like most bees. Their nests vary considerably in size, construc­ tion, a n d location (Young 1985). Nests contain only a relatively few ovoid cells and are of t h r e e types: (1) s u b t e r r a n e a n or in cavities a n d constructed of chewed or s h r e d d e d wood fragments, c e m e n t e d to­ g e t h e r with wax a n d resin; (2) attached to exposed leaves or twigs a n d m a d e entirely of resin (Young 1985); a n d (3) in hollow stems. A l t h o u g h not truly social, several females may form cells in a single globular mass. T h o s e of Eulaema may even cooper­ ate to some d e g r e e in nest building and may be considered subsocial (Zucchi et al. 1969). Euglossine bees a r e recognized morpho­ logically by their extremely long tongues, greatly swollen hind tibiae (males only), a n d small secondary lobe at the base of the main h i n d wing lobe, all features absent in their n e a r b u m b l e b e e relatives. T h e y also possess h i n d tibial s p u r s a n d are commonly bright metallic blue, g r e e n , or copper in color. T h e latter is t r u e of Euglossa (fig. 12.14b), but Eulaema (fig. 12T4c) and Eufriesea (Kimsey 1982) have metallic g r e e n only on p o r t i o n s of the integument a n d are large (BL 2.5 m m ) , hairier, and with a b d o m e n s colored with black, yellow, a n d o r a n g e (often in bands), m u c h resem­ bling bumblebees. Females of Eufriesea sting fiercely a n d have evolved Mullenan mimetic complexes in South America (Dressier 1979) a n d have Batesian mimics a m o n g the Díptera. T h e m u c h smaller (BL

12 m m ) , metallic Euglossa may be mim­ icked by the c o m m o n g r e e n " d r o n e flies," Ornidia s p p . Exaerete a n d Aglae generally resemble Euglossa with t h e i r shiny, bright colors but are generally twice as large (BL 25 m m ) and h a v e a m o r e s t r e a m l i n e d b o d y s h a p e with a n a r r o w e r h e a d a n d a t a p e r e d abdo­ men. T h e y also lack the contrasting white facial m a r k i n g s of that g e n u s . Exaerete has two tubercles o n the scutellum; in Aglae, this is a large, Hat, platelike s t r u c t u r e . Biologically, these two g e n e r a are still m o r e distinct, b o t h being kleptoparasites in the nests of their Eulaema cousins. A bibliography of orchid bees has been published (Williams 1978).

euglossine bees (Hymenoptera, Apidae). Atas Simposio Biota Amazónica 5 (Zool.): 395-415. WILLIAMS, N. H. 1978. A preliminary bibliogra­ phy on euglossine bees and their relation­ ships with orchids and other plants. Selbyana 2: 345-355. YOUNG, A. M. 1985. Notes on the nest structure and emergence of Euglossa turbinifex Dressier (Hymenoptera: Apidae: Bombinae: Euglos­ sini) in Costa Rica. Kans. Entomol. Soc. J. 58: 538-543. ZUCCHI, R-,

S. F. SAKAGAMI, AND J. M.

F. DE

CAMARGO. 1969. Biological observations on a Neotropical parasocial bee, Eulaema nigrita, with a review of the biology of Euglossinae: A comparative study. Hokkaido Univ. J. Fac. Sci., Ser. 6, Zool. Í7: 271-380.

SOCIAL BEES References ARMBRUSTER, W.

S.,

AND K.

D.

MCCORMICK.

1990. Diel foraging patterns of male euglos­ sine bees: Ecological causes and evolutionary response by plants. Biotropica 22: 160-171. DODSON, C. H. 1966. Ethology of some bees of the tribe Euglossini. Kans. Entomol. Soc. J. 39: 607-629! DODSON, C. H., R. L. DRESSLER, H. G. HILLS, R. M. ADAMS, AND N. H. WILLIAMS. 1969.

Biologically active compounds in orchid fra­ grances. Science 164: 1243-1249. DRESSLER, R. L. 1979. Eulaema bombiformis, E. meriana and Müllerian mimicry in related species (Hymenoptera: Apidae). Biotropica 11: 144-151. DRESSLER, R. L. 1982. Biology of the orchid bees (Euglossini). Ann. Rev. Ecol. Syst. 13: 373-394. JANZEN, D. H.,

P. J. DFVRIES, M. L. HIGGINS,

AND L. S. KIMSEY. 1982. Seasonal and site variation in Costa Rican euglossine bees al chemical baits in lowland deciduous and ever­ green forests. Ecology 63: 66-74. KIMSEY, L. W. 1980. The behavior of male orchid bees (Apidae, Hymenoptera, lnsecta) and the question of leks. Anim. Behav. 28: 996-1004. KIMSEY, L. S. 1982. Systematics of bees of the genus Eufriesea (Hymenoptera, Apidae). Univ. Calif. Publ. Entomol. 95: 1-125. KIMSEY, L. S. 1987. Generic relationships within the Euglossini (Hymenoptera: Apidae). Syst. Entomol. 12: 63-72. MOURE, J. S. 1967. A check list oí the known

Bees with large n u m b e r s of w o r k e r off­ s p r i n g that c o o p e r a t e in building a n d main­ t e n a n c e of complex colonies in nests or hives are considered t r u e social insects a n d are r e p r e s e n t e d in the A m e r i c a n tropics by the b u m b l e b e e s a n d stingless bees. T h e social h o n e y b e e (Apis mellifera), a l t h o u g h well established in this p a r t of the world, is an i n t r o d u c e d species living in domestic hives or as a feral insect in a foreign environment.

Reference MICHENER, C. D. 1974. T h e social behavior of the bees: A comparative study. Belknap Press, Harvard Univ., Cambridge.

Bumblebees A p i d a e , B o m b i n a e , Bombini, Bombus. Spanish: Abejorros, abejones (General). Portuguese: M a m a n g a b a s (var. m a n g a n g á s ) . Quechua: H u a y r o n c o k u n a . Náhuatl: Xicohtin, sing, xicohtli. B u m b l e b e e s (Alford 1975, Morse 1982) are large bees (BL 2 0 - 3 0 m m ) , distin­ guished from o t h e r large bees by their woolly hair that is colored in black a n d yellow (to o r a n g e ) b a n d s . T h e wings a r e

SOCIAL BEES

461

tinged with d a r k p i g m e n t , a n d t h e h i n d pair lack a small second lobe at t h e e x t r e m e base of t h e usual p o s t e r i o r lobe. T h e y also have very long t o n g u e s , a characteristic not f o u n d in o t h e r bees of great size. T h e y a r e widely d i s t r i b u t e d over t h e Americas, even to T i e r r a del Fuego, b u t a r e decidedly less a b u n d a n t in w a r m lowland wet a n d moist forests (Dias 1958) t h a n in m o r e t e m p e r a t e , drier, d e c i d u o u s forests a n d m o u n t a i n habitats ( M o u r e a n d Sakagami 1962). T h e y a r e a m o n g t h e few insects active o n cold days high in t h e m o u n t a i n s , o c c u r r i n g u p to 4,000 m e t e r s in t h e A n d e s a n d C e n t r a l A m e r i c a n cordillera, w h e r e they a r e i m p o r t a n t pollinators. It is postu­ lated by J a n z e n (1971) that nest sites a r e fewer a n d p r e d a t i o n h i g h e r in t h e f o r m e r habitats, c o n t r i b u t i n g to their relative scarceness. T h i s d e t e r m i n e s a t e n d e n c y to nest o n , r a t h e r t h a n in, t h e g r o u n d a n d even in trees, also n o t e d in t h e m o r e tropi­ cal areas (Laroca 1976). In m o u n t a i n o u s t e r r a i n , it is typical for q u e e n s e m e r g i n g from h i b e r n a t i o n after the r e t r e a t of t h e cold season to establish new nests in n a t u r a l hollows in t h e g r o u n d , including those m a d e by r o d e n t s a n d birds. T h e larvae a r e r e a r e d in saclike, waxen cells ("honey pots") a n d fed o n honey a n d pollen. Eventually, t h e colonies, which a r e t r u e insect societies, m a y c o m e to h a r b o r several h u n d r e d q u e e n s , sterile w o r k e r females, a n d males in a n n u a l rest­ ing species. Colonies of 2,000 to 3,000 a r e

k n o w n in some p e r e n n i a l tropical species T h e females sting readily a n d forcefully in defense of t h e nests. Males of m a n y species of b u m b l e b e e s fly along established routes hovering m o m e n t a r i l y at certain places (Stiles 1976) that they m a r k with scent to attract females (Blum 1981). An o u t d a t e d general t a x o n o m i c study indicates forty-five Neotropical species (Franklin 1913). T h e list has been modi­ fied some in m o r e recent works (Milliron 1971), b u t t h e overall n u m b e r is still about the same. Some a r e very w i d e s p r e a d , as the ubiquitous, typically m a r k e d Bombus tucumanus (fig. 12.15a) a n d t h e all-yellow Bombus dahlbomi, which ranges over most of s o u t h e r n South America. A useful bibliog­ r a p h y is available (Milliron 1970).

LAROCA, S. 1976. Sobre a bionomia de Bombus morio (Hymenoptera, Apoidea). Acta Biol. Paranaense 5(1-2): 107-127.

References

Stingless Bees

ALFORD, D. V. 1975. Bumblebees. DavisPaynter, London. BLUM, M. S. 1981. Sex pheromones in social insects: Chemotaxonomic potential. In P. E. Howse and L. L. Clement, eds., Biosystematics of social insects. 19: 163-174. Aca­ demic, New York. DÍAS, D. 1958. Contribuicáo para o conhecimento da biononiia de Bombus incarum Frank­ lin da Amazonia (Hymenoptera: Bombidae). Rev. Brasil. Entomol. 8: 1-20.

Apidae, M e l i p o n i n a e , Meliponini, Melipona, Trígona, a n d Lestrimelitta. Spanish: Abejas sin aguijón (General); zeganas ( P a n a m a ) ; abejas bobos, angelitos (Colombia); a r a m b a s a s (Amazonian Peru); p e g o n e s (Venezuela); culos d e vaca, abejas jicotes, abejas a t a r r á (Costa Rica). Portuguese: A b e l h a s sem ferráo, torcecabelos, i r a p u a , abelhas d e c u p i m , cupira, j a t a í , x u p é , abelhas bravas, cagafogos, etc. (Brazil). J u a n a t s (Melipona), p e g o n e s (Trigona) (Trinidad). Náhuatl: For specific types only, e.g., necutli, pipiolin.

FRANKLIN, H. J. 1913. The Bombidae of the

New World. Amer. Entomol. Soc. Trans. 39: 73-200. JANZEN, D. H. 1971. The ecological significance of an arboreal nest of Bombus putUatus in Costa Rica. Kans. Entomol. Soc. J. 44: 210-216.

Figure 12.15 SOCIAL BEES (APIDAE). (a) Bumblebee {Bombus tucumanus). (b) Stingless bee (Trígona fuscipennis). (c) Stingless bee (Trígona fulviventris). (d) Honeybee (Apis mellifera).

462

SAWFLIES, WASPS, ANTS, AND BEES

MILLIRON, H. E. 1970. A monograph of the

Western Hemisphere bumblebees (Hymenop­ tera: Apidae; Bombinae). Entomol. Soc. Can­ ada Mem. 65: 1-52. Bibliography only. MILLIRON, H. E. 1971. A monograph of the

Western Hemisphere bumblebees (Hymenop­ tera: Apidae; Bombinae). I. The genera Bombus and Megabombus subgenus Bombias. Entomol. Soc. Canada Mem. 82: 1-80. MORSE, D. H. 1982. Behavior and ecology of bumble bees. In H. R. Hermann, ed., Social insects. 3: 245-322. Academic, New York. MOURE, J. S., AND S. F. SAKAGAMI. 1962.

As

mamangabas sociais do Brasil (Bombus Latr.) (Hym. Apoidea). Studia Entomol. 5: 65—194. STILES, E. W. 1976. Comparison of male bumble­ bee flight paths: Temperate and tropical (Hymenoptera: Apoidea). Kans. Entomol. Soc. J. 49: 266-274.

Certainly m o r e can be said of t h e n a t u r a l history of this tribe t h a n any o t h e r Neotropical b e e g r o u p (Sakagami 1982, Schwarz 1948). T h a t they have b e e n p r o ­ foundly i m p o r t a n t in t h e region's h u m a n culture also is evidenced by t h e m a n y common n a m e s a p p l i e d to t h e m , not only as a g r o u p b u t to individual species (see above a n d L e n k o a n d P a p a v e r o 1979: 171). Before t h e i n t r o d u c t i o n of s u g a r c a n e

a n d t h e E u r o p e a n h o n e y b e e (Apis mellifera) to t h e New World, t h e chief s o u r c e of sweets was stingless bee h o n e y ( C r a n e 1983). To this day, t h e sweet p r o d u c t of these bees is p r e f e r r e d widely a m o n g Indi­ ans a n d c o u n t r y p e o p l e , w h o m a k e from it m a n y c o n d i m e n t s , beverages, a n d medici­ náis. T h e honey of specific bees was even s u p p o s e d to have value for particular ail­ ments, for e x a m p l e , h o n e y from Trigona jaty, a widely cultivated species, in s o u t h e r n Brazil, is a folk r e m e d y for a sore t h r o a t . Balche, m a d e from stingless b e e honey, was a p r i m e ceremonial d r i n k of t h e Maya a n d d r u n k e n n e s s from imbibing it was compulsory in religious rituals. H o n e y from some stingless bees, notably, t h e l e m o n b e e (Lestrimelitta limao), is p o i s o n o u s , a quality curiously c o n t r i b u t i n g to, r a t h e r t h a n detracting from, its t h e r a p e u t i c u s e . T h e G u a r a y o I n d i a n s of Bolivia w e r e r e ­ p u t e d to u s e h o n e y from this species for the c u r e of paralysis. Because of their long association with stingless bees, A m e r i n d s succeeded in d e ­ veloping a form of a p i c u l t u r e crudely par­ allel to that of Africa with t h e h o n e y b e e which is still practiced today ("meliponic u l t u r e " ; N o g u e i r a - N e t o 1953, Weaver a n d Weaver 1981). Native hives c o m m o n l y con­ sist of hollow logs that a r e s e e d e d with p o r t i o n s of c o m b from a wild nest. After the new colony develops to a healthy size, its h o n e y pots a r e harvested by r e m o v i n g the e n d plugs. A l t h o u g h a n u m b e r of species a r e occasionally kept in this m a n ­ ner, Melipona beecheii is most frequently domesticated a n d even r e f e r r e d to in early accounts as Melipona "domestica" (Weaver a n d Weaver 1981). No less i m p o r t a n t t h a n its honey, t h e wax of t h e stingless bees f o u n d n u m e r o u s applications a m o n g P r e - C o l u m b i a n p e o ­ ple, many of which c o n t i n u e a m o n g r u r a l s even today. Miscellaneous applications in­ clude m a k i n g candles, w a t e r p r o o f i n g arti­ cles, a n d f o r m i n g religious icons. It is also a c o m m o n adhesive, calking material,

SOCIAL BEES

463

filler, lubricant, a n d occasional medicinal in m i n o r t h e r a p e u t i c s such as t h e removal of c o r n s a n d warts. Its greatest historical significance, however, surely derives from its use in metallurgy, which all t h e clas­ sic P r e - C o l u m b i a n civilizations discovered without influence from t h e East. Ancient goldsmiths m o l d e d gold jewelry a n d o t h e r items of t h e finest quality with a "lost wax t e c h n i q u e " identical to that practiced by Old World a r t e s a n s b u t e m p l o y i n g wax from native m e l i p o n i n e s in place of that of t h e h o n e y b e e (Bird 1979). Such was the p r o m i n e n c e of stingless bees to primitives that they inevitably be­ c a m e e n t w i n e d in t h e c u l t u r e (Posey 1983). C e r t a i n P a r a g u a y a n tribes recognized p r o p ­ erty rights in wild honey. Tributes were often paid in h o n e y a n d wax; the C o d e x M e n d o z a text specifies quantities that were to be delivered to M o c t e z u m a by lowland Aztec c o m m u n i t i e s . Even a m o n g t h e Yuca­ tán Maya today, a major c e r e m o n y k n o w n as the u hanli cab is celebrated in which the ancient Maya bee gods a r e beseeched to bless the cultivated bees (Weaver a n d Weaver 1981). T h e s e bees a r e c o m m o n a n d conspicu­ ous t h r o u g h o u t the c e n t r a l Neotropics (Wille 1961), especially in moist lowland forest e n v i r o n m e n t s . T h e y a r e absent from the high A n d e a n valleys, coastal deserts, a n d Antilles except for the large islands a n d those close to the m a i n l a n d . T h e y are most directly recognized by their b o t h e r ­ some habits when a r o u s e d a n d their asso­ ciations with their nests, which are always densely p o p u l a t e d , aggressively d e f e n d e d , a n d uniquely c o n s t r u c t e d . Most are located in n a t u r a l cavities, usually in the g r o u n d or tree t r u n k s but s o m e t i m e s in o d d sites such as dry m a m m a l carcasses a n d bird, termite, or ant nests. T h e p a r t i c u l a r a r r a n g e m e n t of structural e l e m e n t s vary (Roubik 1979, Wille a n d M i c h e n e r 1973), bul the nest always contains b r o o d cells in a cluster (Trígona) or layered in horizontal combs (Melipona), these s u r r o u n d e d by a layered

464

SAWFL1ES, WASPS, ANTS, AND BEES

envelope, storage pots for h o n e y a n d po|. len located outside the e n v e l o p e , and waxen e n t r a n c e canal that often extends outside the nest as a freely projecting tube T h e whole c o m p l e x is walled off from the exterior by h a r d e n d plates o r a n outer shell called the b a t u m e n . Several kinds of building materials go into nest construc­ tion, primarily wax, but this is usually mixed with o t h e r matter, such as propolis plant resin, a n d g u m collected by the bees (Ramirez a n d G ó m e z 1978). T h i s habit may have led to the e n t o m b m e n t of indi­ viduals, several of which have been found as fossils in copal ( M o u r e a n d Camargo 1978). Mixed with wax ( c e r u m e n ) , this is t h e substance of t h e b r o o d cells. Mud, feces, plant fibers, a n d leaf fragments also are used in nest f o r m a t i o n . T h e larval provisions of some species are known to s u p p o r t a rich bacterial flora that may play a f u n d a m e n t a l role in the preservation and metabolic conversion (Gilliam et al. 1985) of these substances. T h i s type of nest is the most elaborate of all native social bees in the New World and identifies the stingless bees at once, as does their m e t h o d of" defense in the absence of a sting. T h e sting o r g a n is vestigial a n d of no use in inflicting w o u n d s o n large enemies, but these bees are by n o m e a n s impotent. In n u m b e r s , they h u r l themselves on those who t h r e a t e n the nest, crawling into nos­ trils, ears, hair, a n d eyes. A l t h o u g h most employ only mandibles to pinch, a few also deposit a caustic fluid from glands at the bases of the jaws. Many of the native names of these refer to their belligerence and potency (e.g., cagafogo.s, "spit fires"; torcecabellos, "hair twisters"). Structurally, stingless bees resemble o t h e r apids in h a v i n g broadly expanded h i n d tibia fringed with hairs, which form the pollen basket (but these are absent in Lestrimelitta). T h e y a r e recognized readily from o t h e r social bees by their usual smaller size (BL at most 15 m m ) , relative hairlessness (no pelage o n legs), a n d blunt

tip to the a b d o m e n . T h e wing venation is a lso u n i q u e , t h e m a r g i n a l cell of the fore wing b e i n g o p e n to t h e wing m a r g i n at the latter's a p e x . T h e i r biology (de C a m a r g o 1972, John­ son a n d H u b b e l l 1974, K e m p f 1962) also resembles that of the h o n e y b e e in m a n y respects, both b e i n g social a n d p r o d u c i n g large b r o o d s . T h e h o n e y b e e is mass provi­ sioned, however. Foragers also c o m m u n i ­ cate distance a n d direction to nestmates like h o n e y b e e s b u t with only s o u n d sig­ nals, a c o m p l e t e symbolic d a n c e not hav­ ing evolved, a l t h o u g h r e t u r n i n g foragers do move t h r o u g h the hive p e r f o r m i n g "buzzing r u n s " similar to the acoustic portion of t h e waggle r u n of h o n e y b e e s (Esch et al. 1965). O t h e r differences a r e found in the c o m p o s i t i o n of r e p r o d u c t i v e swarms, which i n c l u d e only y o u n g , never original, q u e e n s , a n d p r o d u c t i o n of wax from dorsal r a t h e r t h a n ventral glands o n the a b d o m e n . S o m e stingless bees forage for various f o r m s o r o r g a n i c matter, in­ cluding d e a d a n i m a l s ( B a u m g a r t n e r a n d Roubik 1989). T h e tribe includes t h r e e g e n e r a , each containing m a n y c o m m o n species, except Lestrimelitta, which has b u t two species. One, L. limao, is k n o w n for its l e m o n o d o r when c r u s h e d (from citral; B l u m et al. 1970). T h e s e lack a pollen basket o n the hind tibia, a r e shiny black with r o u n d heads, a n d a r e m e d i u m sized (BL 8 m m ) . Obligate r o b b e r s of h o n e y from the stor­ age c h a m b e r s of o t h e r stingless bees, the workers a r e n o t a d a p a t e d morphologically or behaviorally for n o r m a l foraging (Sakagami a n d Laroca 1963). W h e n scouts lo­ cate a suitable nest, they are probably killed at the e n t r a n c e , a n d the process releases large a m o u n t s of citral, the trailmarking p h e r o m o n e for this a n d o t h e r stingless bees. T h e o d o r p e r v a d e s the nest and diffuses into t h e air, attracting m o r e Lestrimelitta a n d confusing the victim bees (Blum et al. 1970), b u t this may not always be highly effective ( J o h n s o n 1987). Trígona

also p r o d u c e a volatile alarm p h e r o m o n e , c o m p o s e d of aliphatic alcohols, ketones, a n d b e n z a l d e h y d e (Luby et al. 1973). A l t h o u g h t h e r e a r e exceptions, Melipona species a r e generally the largest of the t h r e e g e n e r a (BL 6—15 m m ) , a r e relatively hairy, a n d have wings that d o not e x t e n d b e y o n d the tip of t h e a b d o m e n when folded (fig. 12.15b) (Schwarz 1932). T h e i n t e g u m e n t surface is dull a n d n o n r e flective. S o m e of the larger species a r e all black with white-tipped wings. T h e very w i d e s p r e a d Melipona beechii is pale b r o w n a n d superficially looks m u c h like the h o n ­ eybee. Trígona a r e mostly smaller (BL 2—8 m m ; Trígona duckei is the smallest bee known), a r e sparsely hairy, h a v e wings t h a t e x t e n d well beyond the a b d o m e n w h e n at rest, a n d are often shiny. T fulviventris (fig. 12.15c) is p e r h a p s the most c o m m o n ; it is recognized by its largeness for the g e n u s (BL 5—6.5 m m ) a n d c o n t r a s t i n g black tho­ rax a n d o r a n g e a b d o m e n ( J o h n s o n 1983). Stingless bees are generally beneficial t h r o u g h their pollination activities. How­ ever, they occasionally h a r m fruit c r o p s (especially citrus) by c u t t i n g the flowers into pieces, which they use in nest construction.

References BAUMGARTNER, D. L., AND D. W. ROUBIK.

1989.

Ecology of necrophilous and filth-gathering stingless bees (Apidae: Meliponinae) of Peru. Kans. Entomol. Soc. J. 62: 11-22. BIRD, J. 1979. Legacy of the stingless bee. Nat. Hist. 88(5): 4 8 - 5 1 . BLUM, M. S., R. M. CREWE, W. E. KERR, L. H. KEITH, A. W. GARRISON, AND M. M. WALKER

1970. Citral in stingless bees: Isolation and functions in trail-laying and robbing. J. Ins. Physiol. 16: 1637-1648. CRANE, E. 1983. The archaeology of beekeep­ ing. Cornell Univ. Press, Ithaca. DE CAMARGO, C. A. 1972. Mating of the social bee Melipona quadrifasciata under controlled conditions (Hymenoptera, Apidae). Kans. Entomol. Soc. J. 45: 520-523. ESCH, H.,

I.

Escn, AND W.

E.

KERR

1965.

Sound: An element common to communica­ tion of stingless bees and to dances of honey bees. Science 149: 320-321.

SOCIAL BEES

465

GlLLIAM,

M . , S.

L . BUCHMANN,

AND B . J .

LORENZ. 1985. Microbiology of the larval provisions of the stingless bee, Trígona hypogea, an obligate necrophage. Biotropica 17:28-31. JOHNSON, L. K. 1983. Trigona fulvivcntris (Abeja atarrá, abeja jicote, culo de vaca, trigona, stingless bee). In D. H. Janzen, ed., Costa Rican natural history. Univ. Chicago Press, Chicago. Pp. 770-772. JOHNSON, L. K. 1987. T h e pyrrhic victory oí nest-robbing bees: Did they use the wrong pheromone? Biotropica 19: 188—189. JOHNSON,

L.

K., AND S. P. HUBBELL.

1974.

Aggression and competition among stingless bees: Field studies. Ecology 55: 120-127. KEMPF, N. 1962. Mutualism between Trigona compressa Latr. and Crematogaster slolli Forel (Hymenoptera: Apidae). New York Entorno]. Soc.J. 70: 215-217. LENKO, K., AND N. PAPAVERO. 1979. lnsetos no

folclore. Cons. Estad. Artes Cien. Hum., Sao Paulo. LUBY, J. M., R. E. REGNIER. E. T. CLARKE, E. C. WEAVER, AND N. WEAVER. 1973. Volatile ce­

phalic substances of the stingless bees, Trigona mexicana and Trigona pectoralis. J. Ins. Physiol. 19: 1111-1127/ MOURE, J. S., AND J. M. E CAMARGO. 1978. A

fossil stingless bee from copal (Hymen­ optera: Apidae). Kans. Entorno!. Soc. J. 51: 560-566. NOGUEIRA-NETO, P. 1953. A criacao de abelhas indígenas sem ferráo. Ed. Chácaras e Quin­ táis, Sao Paulo. POSEY, D. A. 1983. Keeping of stingless bees by the Kayapó Indians of Brazil. J. Ethnobiol. 3: 63-73. RAMÍREZ, W., AND L. D. GÓMEZ. 1978. Produc­

tion of nectar and gums by flowers of Monstera deliciosa (Araceae) and of some species of Clusia (Guttiferae) collected by New World Trigona bees. Brenesia 14 — 15: 407—412. ROUBIK, D. W. 1979. Nesl and colony character­ istics of stingless bees from French Guiana (Hymenoptera: Apidae). Kans. Entomol. Soc. J. 52: 443-470. SAKAGAMI, S. E 1982. Stingless bees. In H. R. Hermann, ed., Social insects. 3: 361—423. Academic Press, New York. SAKAGAMI, S. E, AND S. LAROCA.

1963. Addi­

tional observation on the habits of the cleptobiotic stingless bees, the genus Lestrimelitta Friese (Hymenoptera, Apoidea). Hok­ kaido Univ. Fac. Sci., Ser. 6, Zool. J. 15: 3 1 9 339. SCHWARZ, H. F. 1932. T h e genus Melipona: The

466

SAWFL1ES, WASPS, ANTS, AND BEES

type genus of the Meliponidae or stine| e s s bees. Amer. Mus. Nat. Hist. Bull. 63: 231-460 SCHWARZ, H. F. 1948. Stingless bees of t n e Western Hemisphere. Amer. Mus. Nat Hist Bull. 90: 1-546. WEAVER, N., AND E. C. WEAVER 1981. Beekeep­

ing witli the stingless bee Melipona beeclieii, bv the Yucatecan Maya. Bee World 62: 7-19. WILLE, A., AND C. D. MICHENER. 1973. The nest

architecture of stingless bees with special refer­ ence to those of Costa Rica (Hymenoptera Apidae). Rev. Biol. Trop. 21(suppl. 1): 1-278* WILLE, A. 1961. Las abejas jicotes de Costa Rica Univ. Costa Rica Rev.22': 1-30.

Honeybee A p i d a e , A p i n a e , Apini, Apis mellifera. Spanish: Abeja d e miel. Portuguese: Abelha doméstica. A m o n g all of t h e d u b i o u s gifts bestowed by t h e O l d World o n t h e New World following its discovery, at least o n e was equal to t h e riches r e t u r n e d . T h i s was the h o n e y b e e (hg. 12.15d), from which a quantity a n d quality of honey (Crane 1979) is obtained m u c h s u p e r i o r to that of native m e l i p o n i n e (stingless) bees. Unfortu­ nately, early r e c o r d s of t h e establishment of Apis mellifera in tropical America are scanty. Apparently, its introduction came m u c h later t h a n into N o r t h America, w h e r e it a r r i v e d with t h e earliest colonists in t h e early 1600s. It was n o t until 1839 that honeybees w e r e b r o u g h t to Brazil ( N o g u e i r a - N e t o 1962) a n d 1857 that they first r e a c h e d Chile a n d Peru. O t h e r dates q u o t e d for t h e i n t r o d u c t i o n a n d spread of the species into various o t h e r countries of C e n t r a l a n d S o u t h America a r e at vari­ ance, a n d t h e history of apiculture in tropical America still n e e d s to be docu­ mented. T h e b e e industry today is well estab­ lished a n d a n i m p o r t a n t factor in rural economics in Latin A m e r i c a ( C r a n e 1978; O r d e t x 1952; O r d e t x a n d Pérez 1966; Smith 1960). Until a short time ago, the biggest e x p o r t e r a n d p r o d u c e r was cer­ tainly Mexico, followed closely by Argen-

tina. O t h e r c o u n t r i e s with major commit­ ments to b e e k e e p i n g a r e Brazil, Chile, Colombia, Costa Rica, a n d Venezuela. T h i s p i c t u r e c h a n g e d in t h e 1960s a n d 1970s as a result of the u n f o r t u n a t e , acci­ dental i n t r o d u c t i o n to t h e New World of the African subspecies, Apis mellifera scutellata (cited until most recently as A. m. adansonii), which is hardly distinguishable from o t h e r subspecies (Daly a n d Balling 1978, R u t t n e r 1976). T h i s u n e x p e c t e d c h a p t e r in the story of the h o n e y b e e in A m e r i c a began in 1956 with t h e intentional i n t r o d u c t i o n of fortysix q u e e n s from South Africa which were shipped to Rio Claro, Sao Paulo State, Brazil, w h e r e they w e r e t o b e carefully interbred with bees of E u r o p e a n origin already established in t h e c o u n t r y to im­ prove the latters' productivity. T h e follow­ ing year, a m i s f o r t u n e p e r m i t t e d t h e es­ cape of twenty-six of these q u e e n s to t h e wild, w h e r e they started a series of events that b e c a m e e x a g g e r a t e d into a h o r r o r story ( M i c h e n e r 1975, Taylor 1977, Taylor and Levin 1978). As a result of their intense d e f e n s e of their nests, t h e r e were mass attacks o n h u m a n s a n d even a few deaths. T h e s e h a p p e n i n g s received great notoriety, which led to an image of a h o r d e of so-called killer bees h e a d i n g across t h e land, t h r e a t e n i n g to cause great loss of life and wreak havoc o n established bee colo­ nies in its p a t h . T h e new entity was d u b b e d also variously as t h e " M a u M a u b e e , " "Brazilian bee," "African bee," a n d "kami­ kaze b e e " a n d even inspired incredible novels (The Swarm, by A r t h u r H e r z o g , 1974) a n d m o t i o n pictures (Savage Bees). Although n o t t h e m o n s t e r it was first purported t o be, t h e new m i x e d strain h a s persisted, nevertheless, a n d retains m a n y of its u n d e s i r a b l e traits as it slowly e x p a n d s its range o u t w a r d over S o u t h a n d C e n t r a l America. As of 1991, it h a d r e a c h e d n o r t h ­ ern Mexico a n d s p r e a d to most n o r t h e r n South A m e r i c a n countries (Venezuela, Trinidad) a n d t o u c h e d m u c h of Colombia,

Peru, Ecuador, a n d even points o n t h e n o r t h e r n coast of Chile. Some e n t o m o l o ­ gists predict that it will eventually occupy most of Latin America a n d e x t e n d to t h e s o u t h e r n fourth of the United States w h e r e it will displace t h e gentler E u r o p e a n bees now t e n d e d t h e r e by b e e k e e p e r s , seriously h i n d e r i n g h o n e y p r o d u c t i o n a n d pollina­ tion m a n a g e m e n t . T h e r e is evidence also that Africanized bees displace i m p o r t a n t native b e e pollinators (Roubik 1979). I n 1983, a p r o g r a m was initiated to q u a r a n ­ tine the bee to a control zone in Costa Rica, along t h e P a n a m a border, in a n effort to halt its n o r t h w a r d s p r e a d (Stibick 1984). Apiculturists will either have to a d a p t to t h e new bee (Roubik a n d B o r e h a m 1990), as they have in s o u t h e r n Brazil (Erickson et al. 1986, Goncalves 1982, Wiese 1977), o r be forced o u t of business, as h a s been t h e case in m a n y parts of South America. T h e biology of the h o n e y b e e is too well known to r e p e a t in detail h e r e (Butler 1955, D a d e 1962, Dietz 1982, Winston 1987). Some basic facts m a y be reviewed, however, especially as they differ in t h e Africanized strain a n d relate to t h e sur­ vival of a species primarily a d a p t e d to a t e m p e r a t e climate in its a d o p t e d , largely tropical, h o m e ( C r a n e 1980, O r d e t x 1952, Smith I960). T h e life of the colony follows a similar p a t t e r n w h e t h e r free in t h e wild o r p a m ­ p e r e d in a b e e k e e p e r ' s box. I t is largely r e g u l a t e d by t h e w o r k e r s whose p h e r o m o n e s help control t h e internal r e p r o d u c ­ tive instincts of the q u e e n , which, in t u r n , d e t e r m i n e s t h e n u m b e r of o t h e r w o r k e r s , timing of d r o n e p r o d u c t i o n , n e w q u e e n d e v e l o p m e n t , a n d o t h e r biological p h e ­ n o m e n a such as s w a r m i n g . T h e latter is the m e t h o d of f o u n d i n g new colonies a n d is of two basic types: " r e p r o d u c t i v e s w a r m i n g , " in which a p o r t i o n o f t h e p a r e n t a l hive leaves with a n e w q u e e n , a n d "abscond­ ing," which is t h e total removal of t h e entire colony to a new site. T h e t e n d e n c y to abscond a p p e a r s to b e m o r e c o m m o n in

SOCIAL BEES

467

Africanized t h a n in E u r o p e a n bees (Win­ ston et al. 1979). U n d e r h u m a n m a n a g e ­ m e n t , s w a r m i n g is u n w a n t e d a n d sup­ pressed b u t still occurs from commercial hives a n d results in feral p o p u l a t i o n s . T h e s e take u p r e s i d e n c e in hollow trees, in rock crevices, a n d in o t h e r p r o t e c t e d places. H e r e , w o r k e r s quickly construct a w a x e n c o m b with rows o f h e x a g o n a l cells placed back to back in h a n g i n g layers. T h e wax is secreted from g l a n d s o n t h e u n d e r ­ side of t h e a b d o m e n a n d m o l d e d into s h a p e by the m a n d i b l e s a n d legs. A p o r t i o n of the cells a r e used as r e a r i n g c h a m b e r s for t h e larvae o f new workers. T h e s e will p r o d u c e females similar in struc­ t u r e t o t h e q u e e n b u t smaller, sterile, a n d b u r d e n e d with a m u l t i t u d e of duties, in­ c l u d i n g the g a t h e r i n g of pollen a n d nectar from flowers. T h e pollen is fed to t h e y o u n g u n c h a n g e d ; t h e n e c t a r is c o n v e r t e d to h o n e y t h r o u g h enzymatic action in t h e worker's gut. Excesses a r e stored in o t h e r cells, a p a r t from t h e b r o o d cells, a n d a r e mainly for use d u r i n g h a r d times. W o r k e r larvae a r e fed almost exclusively o n pollen a n d honey. Africanized bees a r e consid­ ered more industrious than European, one of t h e desirable traits for which it was i m p o r t e d originally, t h o u g h in reality, it is not a better h o n e y p r o d u c e r . W o r k e r s of all h o n e y b e e s c o m m u n i c a t e t h e location of nectar sources with a " d a n c e l a n g u a g e , " scent m a r k i n g not b e i n g of p r i m a r y i m p o r ­ tance (Gould 1976). From time to time, t h e colony p r o d u c e s d r o n e s a n d / o r virgin q u e e n s . T h e f o r m e r d e v e l o p from unfertilized eggs a n d t h e r e ­ fore have half the c o m p l e m e n t of c h r o m o ­ somes in their tissues. T h e latter grow from n o r m a l eggs, b u t t h e i r larvae a r e fed a special diet o f "royal jelly," a protein-rich substance secreted by glands e m p t y i n g into the w o r k e r s ' p h a r y n x . Q u e e n s also d e v e l o p in oversized, irregularly s h a p e d cells. E m e r g e n c e of virgin q u e e n s , with fully functional r e p r o d u c t i v e o r g a n s , follows s w a r m i n g by a few days. T h e p a t t e r n varies

468

SAWFLIES, WASPS, ANTS, AND BEES

considerably b u t most c o m m o n l y goes as follows. After providing for a new genera­ tion of q u e e n s , t h e old q u e e n leaves the hive with a r e t i n u e of workers b o u n d for a new h o m e . From t h e q u e e n cells left be­ hind, o n e potential new q u e e n hatches soon, a h e a d of the o t h e r s , a n d destroys her rivals, often with the h e l p of workers. She t h e n issues from t h e hive, m a t e s with d r o n e s from o t h e r hives (who immediately die), a n d r e t u r n s to assume the role of her departed parent. I n s e m i n a t i o n lasts t h e q u e e n ' s lifetime; s p e r m stored in a special p o u c h off the oviduct a r e released as n e e d e d to fertilize eggs passing d o w n t h e tract. Q u e e n s may survive a year o r longer, laying u p t o 1,500 or m o r e eggs a d a y in favorable seasons, a n d p r o d u c e m a t u r e colonies of u p to 60,000 workers. Both q u e e n s a n d w o r k e r s have stings. T h e sting of the latter is b a r b e d so that it usually becomes a n c h o r e d in t h e wound when used o n large animals, a n d t h e at­ tached poison sac a n d o t h e r internal or­ gans a r e w r e n c h e d o u t w h e n the bee pulls free. T h e v e n o m is potent, a complex m i x t u r e of proteins that elicit varied, often severe, i m m u n e reactions in humans. T h e s e a r e sometimes f o u n d to remedy certain complaints, including arthritis and some n e u r a l d i s o r d e r s , a n d controlled v e n o m injections a r e occasionally pre­ scribed by physicians in t h e t r e a t m e n t of these ailments. T h e venom of Africanized bees is n o m o r e p o t e n t t h a n that of other honeybees, b u t a victim is likelier to be attacked in mass as a result of an irresist­ ible alarm p h e r o m o n e given off by en­ r a g e d workers. T h e species, in general, is adapted to semidry, t e m p e r a t e climates a n d lives best at h i g h e r elevations a n d latitudes in Latin America. For this reason, it has never been successfully kept in wet lowland tropical zones; it is largely absent from Amazonia in spite of a t t e m p t s to establish it there; the Africanized strain, however, is h a r d i e r and

seems t o b e m a k i n g i n r o a d s even into this warm a n d h u m i d e n v i r o n m e n t . T h e h o n e y b e e is susceptible to a n u m ­ ber of h a r m f u l parasites, including several parasitic mites (Camazine 1986, D e j o n g et al. 1982), t h e b e e louse, (Braula coeca) (Weems 1983), a n d p h o r i d flies of t h e genus Melaloncha (Ramirez 1984).

References BUTLER, C. G. 1955. The world of the honey­ bee. Macmillan, New York. CAMAZINE, S. 1986. Differential reproduction of the mite, Varroa jacobsoni (Mesostigmata: Varroidae), on Africanized and European honey bees (Hymenoptera: Apidae). Entomol. Soc. Amer. Ann. 79: 801-803. CRANE, E. 1978. Bibliography of tropical apiculture. Int. Bee Res. Assoc, London. CRANE, E. 1979 [1975]. Honey: A comprehen­ sive survey. Heinemann, London. CRANE, E. 1980. T h e scope of tropical api­ culture. Bee World 61: 19-28. DADE, H. A. 1962. Anatomy and dissection of the honeybee. Int. Bee Res. Assoc, London. DALY, H. V., AND S. S. BALLING. 1978. Identifica­

tion of Africanized honeybees in the Western Hemisphere by discriminant analysis. Kans. Entomol. Soc. J. 51: 857-869. DEJONG, D., R. A. MORSE, AND G. C. EICKWORT.

1982. Mite pests of honey bees. Ann. Rev. Entomol. 27: 229-252. DIETZ, A. 1982. Honey bees. In H. R. Hermann, Jr., ed., Social insects. I: 333-360. Academic, New York. ERICKSON, JR., E. H., B. J. ERK.KSON, AND A. M.

YOUNG. 1986. Management strategies for "Af­ ricanized" honey bees: Concepts strength­ ened by our experiences in Costa Rica. Pts. I, II. Glean. Bee Cult. 1986(Oct.): 456-459, 506-507. G0N5ALVES, L. S. 1982. The economic impact of the Africanized honey bee in South America. 9th Int. Cong. Int. Union Study Social Ins. (Boulder, Colo., 1982), Proc. Pp. 134-137. GOULD, J. L. 1976. The dance-language contro­ versy. Quart. Rev. Biol. 51:21 1-244.

MICHENER, C. D. 1975. T h e Brazilian bee prob­ lem. Ann. Rev. Entomol. 20: 399-416. NOGUEIRA-NETO, P. 1962. O inicio da apicultura no Brasil. Biol. Agrie. Sao Paulo 49: 5-14. ORDF.TX, G. S. 1952. Flora apícola de la América tropical. Ed. Lux, La Habana. [Not seen.] ORDETX, G. S., AND D. E. PÉREZ.

1966. La

apicultura en los trópicos. Priv. pubf, México. RAMÍREZ, W. 1984. Biología del género Mela­ loncha (Phoridae), moscas parasitoides de la abeja doméstica (Apis mellifera L.) en Costa Rica. Rev. Biol. Trop. 32: 25-28. ROUBIK, D. W. 1979. Africanized honeybees, stingless bees and the structure of tropical plant-pollinator communities. Proc. 4th Int. Symp. Pollination, Maryland Agrie. Exper. Sta., Spec. Misc. Publ. 2: 403-417. ROUBIK, D. W., AND M. M. BOREHAM.

1990.

Learning to live with Africanized honeybees. Interciencia 15: 146-153. RUTTNER, F. 1976. African races of honeybees. 25th Int. Apicul. Cong. (Grenoble, 1976). Pp. 325-347. SMITH, F. G. 1960. Beekeeping in the tropics. Longmans, Green, London. STIBICK, J. N. L. 1984. Animal and plant health inspection service strategy and the African honey bee. Entomol. Soc. Amer. Bull. 30(4): 22-26. TAYLOR, JR., O. R. 1977. T h e past and possible future spread of Africanized honeybees in the Americas. Bee World 58: 19-30. TAYLOR, J R . , O. R., AND M. D. LEVIN. 1978.

Observations on Africanized honeybees re­ ported to South and Central American gov­ ernment agencies. Entomol. Soc. Amer. Bull. 24:412-414. WEEMS, JR., H. V. 1983. Bee louse, Braula coeca Nitzsch (Díptera: Braulidae). Fia. Dept. Agrie. Consuni. Serv. Entomol. Giro 252: 1-2. WIESE, H. 1977. Apiculture with Africanized bees in Brazil. Amer. Bee J. 117: 166-168, 170. WINSTON, M. L. 1987. The biology of the honey bee. Harvard Univ. Press, Cambridge. WINSTON, M. L., G. W. O T I S , AND O. R. TAYLOR,

JR. 1979. Absconding behaviour of the Afri­ canized honeybee in South America. J. Apicul. Res. 18: 85-94.

SOCIAL BEES

469

13

INSECT STUDY

T h e study of insect life in Latin A m e r i c a has a long history a n d has a d v a n c e d today to a high level of excellence. T h e subject is t a u g h t in academic, agricultural, a n d medi­ cal curricula in all c o u n t r i e s , a n d research­ ers a n d n a t u r a l historians a r e p r o g r e s s i n g rapidly on all fronts. For this reason, in­ formation is readily available on most as­ pects, at least in the major cities. T h i s c h a p t e r is included to aid the s t u d e n t w h o may not have r e a d y access to entomological facilities.

References ANONYMOUS. 1983. The world of learning 198384. Europa, London. ANONYMOUS. 1987. Resources in entomology. Entomological Society of America, College Park, Md. (See pp. 199-232 for Latin Amer­ ica.) Teaching: Universities, and Other Schools

INFORMATION SOURCES I n f o r m a t i o n o n e n t o m o l o g y is available from a variety of sources (Gilbert a n d H a m i l t o n 1983). T h e s e a r e basically of two kinds: (1) p e r s o n a l consultation with au­ thorities, directly or t h r o u g h their institu­ tions, a n d (2) r e f e r e n c e to t h e r e c o r d s of authorities by r e a d i n g the literature, both p r i n t e d a n d in its o t h e r f o r m s .

Reference GILBERT, P., AND C. J. HAMILTON. 1983.

flect the variety of types a n d those that are well established or especially active cur­ rently. (See A n o n y m o u s [1983, 1987] for m o r e complete lists.)

Ento­

mology: A guide to information sources. Mansell, London.

Colleges,

Argentina División d e Entomología, Facultad de A g r o n o m í a ; Instituto d e Limnología, Facultad d e Ciencias Naturales y Museo Universidad Nacional d e La Plata, La Plata. D e p a r t a m e n t o d e Ciencias Biológicas, Facultad d e Ciencias Exactas, Físicas y Naturales; C á t e d r a d e Zoología Agrí­ cola, Facultad d e A g r o n o m í a : Universi­ d a d Nacional d e B u e n o s Aires, Buenos Aires.

Institutions

Bolivia Instituto d e Biología: Universidad Boli­ viana Mayor, Real y Pontificia d e San Francisco Xavier, Le Paz.

Listed below a r e selected institutions in Latin A m e r i c a that e m p l o y entomologists; they a r e s e g r e g a t e d a c c o r d i n g to adminis­ trative or organizational p u r p o s e ; some are n o t primarily entomological. T h e list is not c o m p l e t e ; entries a r e i n t e n d e d to re­

Brazil D e p a r t a m e n t o d e Zoología: Universid a d e d e Brasilia, Brasilia. Instituto d e Biología—Parasitología: Universidade Federal Rural d o Rio de Janeiro, Rio d e J a n e i r o .

470

D e p a r t a m e n t o d e Entomología, D e p a r ­ t a m e n t o d e Zoología: Escola S u p e r i o r d e A g r i c u l t u r a "Luiz d e Q u e i r o z , " Piracicaba. D e p a r t a m e n t o d e Biologia Geral, Facultade d e H i g i e n e e S a ú d e Pública, Insti­ tuto d e Biociéncias: Universidade d e Sao Paulo, Sao Paulo. D e p a r t a m e n t o d e Zoologia, Instituto d e Biología: Universidade Estadual d e C a m ­ pinas, C a m p i n a s . D e p a r t a m e n t o d e Zoologia: Universi­ d a d e Federal d o P a r a n á , Curitiba. Chile D e p a r t a m e n t o d e Zoología, Instituto d e Biología: Universidad d e Concepción, Concepción. D e p a r t a m e n t o d e Biología, Facultad d e Ciencias: Universidad d e Chile, Santiago. Colombia D e p a r t a m e n t o d e Biología: Universidad d e Antioquia, Medellín. Facultad d e A g r o n o m í a : Universidad d e Caldas, Manizales. D e p a r t a m e n t o d e Biología, D e p a r t a ­ m e n t o d e Microbiología: Universidad d e Los A n d e s , Bogotá. Facultad d e A g r o n o m í a , Instituto d e Ciencias N a t u r a l e s : Universidad Nacio­ nal d e Colombia, Bogotá. D e p a r t a m e n t o d e Microbiología, D e p a r ­ t a m e n t o d e Biología: Universidad del Valle, Cali.

El Salvador D e p a r t a m e n t o d e Ciencias Biológicas: Universidad d e El Salvador, San Salvador. Guatemala Escuela d e Biología: Universidad d e San Carlos, G u a t e m a l a City. Facultad d e Biología: Universidad del Valle, G u a t e m a l a City. Guyana Faculty of Natural Sciences: University of G u y a n a , G e o r g e t o w n . Haiti Faculté d e Science: Université d'Haiti, Port-au-Prince.

d'état

Honduras D e p a r t m e n t of Plant Protection: Escuela Agrícola P a n a m e r i c a n a , Tegucigalpa. D e p a r t a m e n t o d e Biología: Universidad Nacional A u t ó n o m a d e H o n d u r a s , Te­ gucigalpa. Jamaica Zoology D e p a r t m e n t : University of the West Indies, M o n a C a m p u s , Kingston.

Dominican Republic D e p a r t a m e n t o d e Biología: Universidad A u t ó n o m a d e Santo D o m i n g o , Santo Domingo.

Mexico L a b o r a t o r i o d e Acarología, Facultad d e Ciencias; D e p a r t a m e n t o d e Parasitolo­ gía, Facultad d e Medicina Veterinaria; Instituto d e Biología, D e p a r t a m e n t o de Zoología: Universidad Nacional A u t ó ­ n o m a d e México, Mexico City. Escuela Nacional d e Ciencias Biológicas: Instituto Politécnico Nacional, Mexico City. C e n t r o d e E n t o m o l o g í a y Acarología: Universidad A u t ó n o m a C h a p i n g o , Chapingo. Universidad A u t ó n o m a A g r a r i a "Anto­ nio N a r r o , " Saltillo, Coahuila. Escuela S u p e r i o r d e A g r i c u l t u r a " H e r ­ m a n o s Escobar," C i u d a d J u á r e z , C h i h u a ­ hua. Facultad d e Ciencias: Universidad A u t ó ­ n o m a d e Nuevo León, Monterrey.

Ecuador Instituto d e Ciencias: Pontificia Universi­ dad Católica del Ecuador, Q u i t o .

Nicaragua Facultad d e Ciencias: Universidad Nacio­ nal A u t ó n o m a d e Nicaragua, León.

Costa Rica D e p a r t a m e n t o d e Biología, D e p a r t a ­ m e n t o d e Microbiología, Facultad d e A g r o n o m í a : Universidad d e Costa Rica, San José. Organization for Tropical Studies, San José.

INFORMATION SOURCES

471

Panama Facultades d e Biología, A g r o n o m í a y Medicina; D e p a r t a m e n t o d e Ciencias Naturales y Farmacia: Universidad d e P a n a m á , P a n a m a City. Paraguay Facultad d e Ciencias y Tecnología: Uni­ versidad Católica " N u e s t r a S e ñ o r a d e La Asunción," Asunción. Peru D e p a r t a m e n t o d e Zoología, Facultad d e Ciencias Biológicas: Universidad Nacio­ nal Mayor d e San Marcos, Lima. D e p a r t a m e n t o d e Biología: Universidad Nacional d e Trujillo, Trujillo. D e p a r t a m e n t o d e E n t o m o l o g í a : Univer­ sidad Nacional A g r a r i a , Lima. D e p a r t a m e n t o d e Zoología: Universidad Nacional "San A n t o n i o A b a d , " Cuzco. Puerto Rico D e p a r t a m e n t o d e Biología: Universidad d e P u e r t o Rico, Río Piedras a n d Mayagüez. Trinidad and Tobago D e p a r t m e n t of Biological Sciences: Uni­ versity of the West Indies, St. A u g u s t i n e . Uruguay Facultad d e A g r o n o m í a ; Facultad d e H u m a n i d a d e s y Ciencias, D e p a r t a ­ m e n t o d e A r t r ó p o d o s : Universidad d e la República, M o n t e v i d e o . Venezuela Instituto d e Zoología Agrícola, Facultad d e A g r o n o m í a : Universidad Central d e Venezuela, Maracay. D e p a r t a m e n t o d e E n t o m o l o g í a , Facul­ tad d e A g r o n o m í a : Universidad del Zulia, Maracaibo. D e p a r t a m e n t o d e Entomología, Facul­ tad d e A g r o n o m í a : Universidad Cen­ tro-Occidental L i s a n d r o A l v a r a d o , Barquisimeto. Departamento de Entomología, Funda­ ción M u s e o d e Ciencias, Caracas. Instituto d e Zoología Tropical: Universi­ d a d C e n t r a l d e Venezuela, Caracas.

472

1NSF.CT STUDY

Government Agencies In every country, these exist primarily to serve the public a n d for the protection of n a t u r a l resources, crops, a n d public health. A sampling follows.

D e p a r t a m e n t o d e Entomología, Minis­ terio d e A g r i c u l t u r a y G a n a d e r í a , San José.

Panama Entomology Section, Servicio Nacional d e la Erradicación d e la Malaria. Instituto d e Investigaciones Agrícolas de Panamá.

Argentina Instituto Nacional d e Tecnología Agro­ pecuaria (INTA), various e x p e r i m e n t a l stations, B u e n o s Aires a n d o t h e r loca­ tions.

Ecuador Instituto Nacional d e Investigaciones A g r o p e c u a r i a s ( I N I A P ) , Q u i t o a n d sev­ eral b r a n c h locations. D e p a r t a m e n t o d e E n t o m o l o g í a , Insti­ tuto Nacional d e H i g i e n e " L e o p o l d o Izquieta Pérez," Guayaquil.

Barbados Entomology Division, Ministry of Agri­ culture, Food a n d C o n s u m e r Affairs, Bridgetown.

El Salvador C e n t r o Nacional d e Tecnología A g r o ­ pecuaria, Ministerio d e A g r i c u l t u r a y G a n a d e r í a , San A n d r é s .

Belize Ministry of A g r i c u l t u r e a n d Lands, Bel­ mopan.

Guadeloupe Instituí Nacional d e la R e s c h e r c h e Agr o n o m i q u e , Station d e Zoologie d e Lutte Biologique, D o m a i n e Duelos, Petit-Bourg.

Puerto Rico College of A g r i c u l t u r e a n d Mechanical Arts, University of Puerto Rico, Mayagüez.

Guyana Central E x p e r i m e n t a l Station, Ministry of A g r i c u l t u r e , East Coast D e m e r a r a .

Surinam C e n t r e for Agricultural Surinam, Paramaribo.

Honduras Ministerio d e Recursos Naturales, Tegu­ cigalpa.

Trinidad and Tobago D e p a r t m e n t of Biological Sciences, Uni­ versity of the West Indies, St. A u g u s t i n e .

Bolivia D e p a r t a m e n t o d e Sanidad Vegetal, Insti­ tuto Boliviano d e Tecnología Agropecua­ ria (IBTA), La Paz. Brazil E m p r e s a Brasileira d e Pesquisa Agro­ pecuaria (EMBRAPA), Brasilia and sev­ eral b r a n c h locations. Seccáo d e Entomología, Instituto Agro­ nómico d o Estado d e Sao Paulo, Campi­ nas. Chile Instituto Nacional d e Investigaciones A g r o p e c u a r i a s , Santiago. Colombia Instituto Nacional d e Salud, Bogotá. Servicio Nacional d e Erradicación de la Malaria (SEM), Bogotá. C e n t r o Internacional d e Agricultura Tropical (CIAT), Cali. Costa Rica D e p a r t a m e n t o d e Agricultura, Depar­ tamento de Ganadería, Departamento d e Ciencias Forestales, C e n t r o Agro­ nómico Tropical d e Investigación y Enseñanza (CATIE), formerly Institu­ ción Internacional d e Ciencias Agrí­ colas (IICA), Turrialba.

Mexico C e n t r o d e Investigaciones Biológicas d e Baja California Sur, La Paz, Baja Califor­ nia Sur, C O N O C Y T Comisión México A m e r i c a n a p a r a la erradicación del g u s a n o b a r r e n d a d o r del g a n a d o , S A R H - U S D A . Dirección G e n e r a l d e S a n i d a d Vegetal, Secretaria d e A g r i c u l t u r a y Recursos Hi­ dráulicos (SARH), P r o g r a m a Moscamed, C h i a p a s . Instituto d e Ecología, Asociación Civil, Mexico City. Instituto Nacional d e Investigaciones Agrícolas ( I N I A ) , Mexico City. Instituto d e S a l u b r i d a d y E n f e r m a d a d e s Tropicales, Mexico City. Nicaragua Instituto d e Recursos Naturales y del Ambiente ( I R E N A ) .

Paraguay Dirección d e Investigación y Extención A g r o p e c u a r i a y Forestal, Ministerio d e Agricultura y G a n a d e r í a , Asunción. Peru P r o g r a m a d e Erradicación d e la Malaria y Mal d e Chagas, Ministerio d e Salud, Lima. Instituto Nacional d e Investigaciones Agrarias, Ministerio d e A g r i c u l t u r a , Lima a n d o t h e r locations.

Research

in

Venezuela División d e Sanidad Vegetal, Ministerio d e Agricultura y Cría, Caracas. Fondo Nacional d e Investigaciones A g r o ­ pecuarias ( F O N A I A P ) , Maracay. C e n t r o Nacional d e Investigaciones A g r o p e c u a r i a s ( C E N I A P ) , Maracay. Several international g o v e r n m e n t a l or­ ganizations also o p e r a t e agencies in m a n y areas which are c o n c e r n e d with insects a n d employ entomologists. S o m e of these a r e the following: 1. U. S. Peace C o r p s . Volunteer e n t o m o l o ­ gists often participate in public health, agricultural, a n d academic p r o g r a m s . 2. Caribbean Research a n d D e v e l o p m e n t Institute (CARDI). Main office in Trini­ d a d , subsidiary offices on G r e n a d a , St. Vincent, St. Lucia, a n d o t h e r West In­ dian islands.

INFORMATION SOURCES

473

3. United Nations, World H e a l t h O r g a n i ­ zation, Pan A m e r i c a n Health Organiza­ tion. Societies Only the following few societies in Latin A m e r i c a a r e expressly d e d i c a t e d to insect study (Sabrosky 1956). Argentina Asociación A r g e n t i n a d e A r t r o p o d o logia, B u e n o s Aires, 1 9 4 4 - . Sociedad E n t o m o l ó g i c a A r g e n t i n a , Bue­ nos Aires, 1 9 2 5 - . Brazil Sociedade Brasileira d e Entomología, Sao Paulo, 1 9 3 7 - . Sociedade Entomológica d o Brasil, Rio de Janeiro, 1922-1945. Sociedade E n t o m o l ó g i c a d o Brasil, Rio G r a n d e d o Sul, 1 9 7 2 - . Chile Sociedad Chilena d e Entomología, Santi­ ago, 1 9 3 3 - . Sociedad E n t o m o l ó g i c a d e Chile, Santi­ ago, 1 9 2 2 - 1 9 2 9 . Colombia Sociedad C o l o m b i a n a d e Entomología, Bogotá, 1 9 7 3 - . Mexico Sociedad Mexicana d e Entomología, Mexico City, 1 9 5 2 - . Sociedad Mexicana d e L e p i d o p t e r o logía, Mexico City, 1974—. Peru Sociedad E n t o m o l ó g i c a del Perú (for­ merly Sociedad Entomológica Agrícola del Perú) Lima, 1 9 5 6 - (Aguilar 1987). Uruguay Sociedad U r u g u a y a M o n t e v i d e o , 1956—.

de

Entomología,

Venezuela Sociedad Venezolana d e Entomología, Maracay, 1964—.

References AGUILAR. P. G. 1987. Algunos apuntes sobre la Sociedad Entomológica del Perú, a los treinta

474

INSECT STUDY

años de su fundación. Rev. Peruana Entornol 29: 127-140. SABROSKY, C. W. 1956. Entomological societies Entornol. Soc. Arner. Bull. 2(4): 1-22.

Museo Territorial, Ushuaia, T i e r r a del Fuego. Instituto Patagónico d e Ciencias Natu­ rales, San M a r t í n d e Los A n d e s .

Directories Entomologists may be contacted personally to obtain information o n insects a n d other related a r t h r o p o d s . T h e i r addresses may be found in lists of m e m b e r s of societies a n d various directories (e.g., Yantko and Golley 1977, A r n e t t a n d A r n e t t 1985).

Bolivia Entomología, M u s e o Nacional d e Histo­ ria Natural, La Paz.

References ARNETT, JR., R. H., AND M. E. ARNETT. 1985.

The

naturalists' directory and almanac (interna­ tional). 44th ed. Flora and Fauna, Gainesville. YANTKO, J. A., AND F. B. GOLLEY. 1977. A world

census of tropical ecologists. Institute of Ecol­ ogy, Univ. of Georgia, Athens. Museums (Insect Collections) I n c l u d e d h e r e a r e all sizable or important collections of insects, a r a c h n i d s , a n d so on, regardless of status, that is, i n d e p e n d e n t m u s e u m s as well as those s u p p o r t e d by teaching institutions, g o v e r n m e n t agen­ cies, or privately. T h o s e in Brazil are dis­ cussed by P a p a v e r o (1985), in H o n d u r a s by O'Brien a n d W a r d (1987), a n d in Mex­ ico by Anaya et al. (1991); a world listing has b e e n compiled by A r n e t t a n d Samuelson (1986). S o m e collection managers have published lists of their holdings, espe­ cially of type material (see below). Abbre­ viations for most collections are available ( H e p p n e r a n d Lamas 1982). Argentina Sección d e Entomología, Museo Ar­ gentino d e Ciencias Naturales "Bernar­ d i n o Rivadavia," B u e n o s Aires. Museo d e Ciencias d e La Plata, La Plata. Insect collections, Fundación e Instituto "Miguel Lillo" e Instituto Superior de Entomología, Facultad d e Ciencias Natu­ rales, T u c u m á n . C e n t r o d e Entomología, Facultad de Ciencias Exactas, Físicas y Naturales, Córdoba.

Brazil D e p a r t a m e n t o d e Entomología, Museu Nacional, Rio d e J a n e i r o . Entomología, M u s e u d e Zoología, Universidade d e Sao Paulo, Sao Paulo. D e p a r t a m e n t o d e Entomología, M u s e u Paraense "Emilio Goeldi," Belém, Para. Colecáo sistemática d e Entomología, In­ stituto Nacional d e Pesquisas d a A m a z o ­ nia, M a n a u s . E n t o m o l o g y collection, D e p a r t a m e n t o d e Zoología, Universidade Federal d o P a r a n á , Curitiba, P a r a n á . Museu R i o - G r a n d e n s e d e Ciencias Naturais, Porto A l e g r e , Rio G r a n d e d o Sul. D e p a r t a m e n t o d e Entomologia, Insti­ tuto O s w a l d o C r u z , Rio d e J a n e i r o . Chile Insect collections, D e p a r t a m e n t o d e Zoo­ logía, Facultad d e Ciencias Biológicas y Reacursos Naturales, Universidad d e Concepción, C o n c e p c i ó n . Sección E n t o m o l o g í a , M u s e o Nacional d e Historia N a t u r a l , Santiago (Camousseight 1980). Museo E n t o m o l ó g i c o , Facultad d e A g r o ­ nomía, Universidad d e Chile, Santiago. Colombia Entomología, M u s e o d e Historia Natu­ ral, I n s t i t u t o d e Ciencias Naturales, Uni­ versidad Nacional d e Colombia, Bogotá. Costa Rica Museo Nacional d e Costa Rica, San José. Museo d e Insectos, Facultad d e A g r o ­ nomía, Universidad d e Costa Rica, Ciudad Universitaria, San José. Instituto Nacional d e Biodiversidad d e Costa Rica (INBio), C i u d a d Universi­ taria, San J o s é . ( J a n z e n 1991).

Cuba Museo Poey, Facultad d e Ciencias, Uni­ versidad d e La H a b a n a , La H a b a n a . Instituto d e Zoología, Instituto d e Eco­ logía y Sistemática, M u s e o Nacional d e Historia Natural: A c a d e m i a d e Ciencias d e Cuba, La H a b a n a . Dominican Republic Museo Nacional d e Historia Santo D o m i n g o .

Natural,

Ecuador Entomología, Museo E c u a t o r i a n o Ciencias Naturales, Q u i t o .

de

El Salvador Entomología, Museo d e Historia Natu­ ral, San Salvador. Guadeloupe Instituí d e Recherches E n t o m o l o g i q u e d e la Caribe, Pointe-a-Pitre. Guatemala Colección Nacional G u a t e m a l t e c a d e Art h r ó p o d a , Universidad del Valle, G u a t e ­ mala City. Guyana Guaraná Museum, Georgetown. Haiti Musée National, Port-au-Prince. Jamaica Institute of Jamaica, Kingston. Mexico Sección d e Entomología, Instituto d e Biología, Universidad Nacional A u t ó ­ n o m a d e Mexico, Mexico City (Vázquez 1981, Vázquez a n d Zaragoza 1979). D e p a r t a m e n t o d e Entomología, M u s e o d e Historia Natural d e la C u i d a d d e México, Mexico City ( B a r r e r a 1966, B a r r e r a a n d Martín 1968). L a b o r a t o r i o d e Acarología, M u s e o d e Zoología "Alfonso L. H e r r e r a " : Facul­ tad d e Ciencias, Universidad Nacional A u t ó n o m a d e México, Mexico City (Llórente 1984, Muñiz et al. 1981). Instituto Nacional d e Investigaciones Agrícolas, Servicio d e A g r i c u l t u r a y Ga­ nadería, C h a p i n g o (Carrillo et al. 1966).

INFORMATION SOURCES

475

Invertebrados Terrestres, Departa­ m e n t o d e Biología T e r r e s t r e , C e n t r o d e Investigaciones d e Baja California Sur, La Paz, Baja California Sur. Nicaragua Zoología, M u s e o Nacional d e Nicara­ gua, M a n a g u a . D e p a r t a m e n t o d e E n t o m o l o g í a , Minis­ terio d e Desarrollo A g r o p e c u a r i o y Re­ forma Agraria (MIDINERA), Managua. M u s e o Entomológico, Facultad d e Cien­ cias, Universidad Nacional A u t ó n o m a d e N i c a r a g u a , León. Panama Insect collections, L a b o r a t o r i o C o n m e ­ morativo G o r g a s , P a n a m a City. ( A r t h r o ­ p o d s of medical i m p o r t a n c e . ) S m i t h s o n i a n Tropical Research Insti­ tute, P a n a m a City. M u s e o d e I n v e r t e b r a d o s " G . B. Fairchild," Facultad d e Ciencias Naturales y Exactas, Universidad d e P a n a m á , P a n a m a City. Paraguay M u s e o d e Historia N a t u r a l , Sociedad Científica del Paraguay, Asunción. Peru Departamento d e Entomología, Museo d e Historia N a t u r a l , Universidad Naci­ onal Mayor d e San Marcos, Lima. M u s e o d e E n t o m o l o g í a , Universidad Nacional A g r a r i a la Molina, Lima (Ortiz a n d Raven 1972). Puerto Rico E n t o m o l o g í a , M u s e o d e Historia N a t u ­ ral, San J u a n . L a b o r a t o r i o d e E n t o m o l o g í a , Facultad d e Ciencias y Artes, Universidad d e P u e r t o Rico, Mayagüez. Uruguay D e p a r t a m e n t o d e Biología, M u s e o Naci­ onal d e Historia N a t u r a l , M o n t e v i d e o . Ministerio d e A g r i c u l t u r a y Pesca, Direc­ ción d e S a n i d a d Vegetal, M o n t e v i d e o (agricultural pests). Facultad d e A g r o n o m í a , Facultad d e H u ­ m a n i d a d e s y Ciencias, D e p a r t a m e n t o d e

476

INSECT STUDY

A r t r ó p o d o s , Universidad d e la R e p ú b ­ lica, Montevideo. Venezuela D e p a r t a m e n t o d e Entomología, Insti­ tuto d e Zoología Agrícola, Facultad d e A g r o n o m í a , Universidad C e n t r a l d e Venezuela, Maracay. C e n t r o Nacional d e Investigaciones A g r o p e c u a r i o s ( C E N I A P ) , Universidad Central d e Venezuela, Maracay. Entomología, Sociedad d e Ciencias Natu­ rales "La Salle," Caracas. D e p a r t a m e n t o d e Entomología, Facul­ tad d e A g r o n o m í a , Universidad del Zulia, Maracaibo. D e p a r t a m e n t o d e Entomología, Facul­ tad d e A g r o n o m í a , Universidad Cen­ tro-Occidental " L i s a n d r o Alvarado," Barquisimeto. Museo d e Biología, Universidad Central d e Venezuela Caracas. Collection of Sr. Carlos B o r d ó n , Mara­ cay, C o l e ó p t e r a . D e p a r t a m e n t o d e Entomología, Funda­ ción Museo d e Ciencias, Caracas.

References ANAYA, S., F. CERVANTES, R. PEÑA, N. BAUTISTA,

AND

R. CAMPOS,

eds. 1991. Colecciones

entomológicas de México: Objetivos y estado actual. Mem. Prim. Simp. Nac. Col. Entomol. Veracruz. ARNETT, JR., R. H., AND G. A. SAMUELSON. 1986.

The insect and spider collections of the world. E. J. Brill/Flora and Fauna, Gainesville. BARRERA, A. 1966. Primera lista de tipos de­ positados en el Museo de Historia Natural de la Ciudad de México. Acta Zool. Mexicano 8(4): 1-3. BARRERA, A. AND E. MARTÍN. 1968. Segunda

lista de tipos depositados en el Museo de Historia Natural de la Ciudad de México. Acta Zool. Mexicano 9(4): 1-5. CARRILLO S., J. L., A. ORTEGA C , AND W. C.

GIBSON. 1966. Lista de insectos en la colec­ ción entomología del Instituto Nacional de Investigaciones Agrícolas. Inst. Nac. lnves. Agrie. (Mexico) Fol. Mise. 14: 1-133. CAMOUSSF.IGHT M., A. 19*80. Catálogo de los tipos de insecta depositados en la colección del Museo Nacional de Historia Natural (San-

tiago, Chile). Mus. Nac. Hist. Nat. Publ. Occ. 32: 1-45. HEPPNER, |. B., AND G. LAMAS. 1982. Acronyms

for world museum collections of insects, with an emphasis on Neotropical Lepidoptera. Entomol. Soc. Amer. Bull. 28: 305-315. JANZEN, D. H. 1991. How to save tropical biodiversity. Amer. Entomol. 37: 158—171. LLÓRENTE, J. E. 1984. Las colecciones zoológicas de la Facultad de Ciencias, Acervo del Museo de Zoología "Alfonso L. Herrera." Univ. Nac. Autón. México, Mexico City. MUÑIZ, A. M., J. C. MORALES, R. A. BARAJAS,

AND J. L. BOUSQUETS. 1981. Primera lista de tipos depositados en el Museo de Zoología "Alfonso L. Herrera" de la Facultad de Cien­ cias de la Universidad Nacional Autónoma de México: Colección de insectos ectoparásitos "Alfredo Barrera." Fol. Entomol. Mexicana 49: 155-168. O'BRIEN, C. W., AND C. R. WARD. 1987. Current

state of insect collections in Honduras. Fol. Entomol. Mexicana 71: 87-101. ORTIZ,

M., AND K. RAVEN.

1972.

Catálogo

preliminar del Museo de Entomología de la Universidad Nacional Agraria. Univ. Nac. Agrar. La Molina, Mus. Entomol., Lima. PAPAVERO, N. 1985. Entomological collections and human resources in Brazil. Assoc. Syst. Colf.Newsl. 13(3): 21-24. VÁZQUEZ, L. 1981. Los tipos existentes en la colección entomológica del Instituto de Bio­ logía, de la Universidad Nacional Autónoma de México. Inst. Biol. Univ. Nac. Autón. México, Ser. Zool. 1, An. 52: 493-505. VÁZQUEZ, L., AND S. ZARAGOZA.

1979. Tipos

existentes en la colección entomológica del Instituto de Biología de la Universidad Nacional Autónoma de México. Inst. Biol. Univ. Nac. Autón. México, Ser. Zool. 1, An. 50: 575-632.

Bellair's Research Institute, McGill Uni­ versity, St. J a m e s (Peck a n d Peck 1980). Bolivia C e n t r o d e Investigaciones d e Mejora­ m i e n t o d e C a ñ a d e Azúcar, Santa C r u z . Brazil Instituto Nacional d e Pesquisas d e A m a ­ zonia (INPA), M a n a u s . F u n d a c á o Instituto Oswaldo C r u z , Rio de Janeiro. Instituto B u t a n t a n , Sao Paulo. Instituto A g r o n ó m i c o d e C a m p i n a s , Campinas. Costa Rica Tropical Science Center, San José. Dominican Republic J a r d í n Botánico y P a r q u e Nacional, Santo D o m i n g o .

Zoológico

Ecuador Asociación Nacional d e Cultivadores d e Palma Africana, Q u i t o . French Guiana Institut Pasteur, C a y e n n e . Office d e la R e c h e r c h e Scientifique e t Technique d'Outre-Mer (ORSTOM), Cayenne. Guyana G u y a n a Rice B o a r d , G e o r g e t o w n . Honduras Division of Tropical Research, United Fruit C o m p a n y , Cortés. Jamaica T h e Institute of Jamaica, Kingston.

Miscellaneous Institutions Other institutions with entomological d e ­ partments exist for private industry o r commercial p u r p o s e s (mainly control of crop pests), purely for e x p e r i m e n t a l a n d research aims, for special projects, o r to serve o t h e r e n d s . S o m e major institutions appear in t h e following list.

Panama Smithsonian Tropical Research Insti­ tute (STRI), P a n a m a City. Maintains field research station o n B a r r o Colo­ r a d o Island. Gorgas Memorial L a b o r a t o r y of T r o p i ­ cal a n d Preventive Medicine, P a n a m a City. Entomology Unit, P a n a m a Canal C o m ­ mission.

Barbados T h e B a r b a d o s S u g a r P r o d u c e r s Associa­ tion, St. Michael.

Peru Fundación p a r a el Desarrollo d o n e r o ( F U N D E A L ) , Lima.

Algo­

INFORMATION SOURCES

477

Surinam F o u n d a t i o n for Scientific Research in S u r i n a m a n d the N e t h e r l a n d s Antilles, Paramaribo. Trinidad and Tobago C o m m o n w e a l t h I n s t i t u t e of Biological Control, Curepe. Venezuela Instituto Pedagógico d e Caracas (Dr. González-Sponga), Caracas. A r a c h n i d s . Virgin Islands C a r i b b e a n Research Institute, College of the Virgin Islands, St. J o h n .

Reference PECK, S. B., AND J. PECK. 1980, Insect field work opportunities in Barbados, Lesser Antilles. Entomol. News 91: 63-64.

Literature T h e entomological l i t e r a t u r e is vast a n d c o m p l e x . N o g e n e r a l text o t h e r t h a n the p r e s e n t treats the subject of Latin A m e r i ­ can insects a n d e n t o m o l o g y in a c o m p r e ­ hensive way, a l t h o u g h p o r t i o n s of some g e n e r a l n a t u r a l history books refer to a significant n u m b e r of regional species ( C e n d r e r o 1 9 7 1 , von I h e r i n g 1968, Shel­ ford 1926). Access to p e r t i n e n t references r e q u i r e s a k n o w l e d g e of i n f o r m a t i o n sources ( B l a n c h a r d a n d Farrell 1981, T r a u g e r et al. 1974) a n d library sources (Davis 1989).

References BLANCHARD, J. R., AND L. FARRELL. 1981. Guide

to sources for agricultural and biological research. Univ. California, Berkeley. CENDRERO, L., ed. 1971. Zoología hispano­ americana. Vol. 2. Ed. Porrúa, Mexico City. DAVIS, T. J., compiler. 1989. Latin American research libraries in natural history: a survey. American Ornithol. Union, Washington, D.C. SHELFORD, V E. 1926. Naturalist's guide to the Americas. Williams and Wilkins, Baltimore. TRAUGER, S. C ,

R. D. SHENEFELT, AND R.

H.

FOOTE. 1974. Searching entomological litera­ ture. Entomol. Soc. Amer. Bull. 20: 303-315. VON IHERING, R. 1968. Dicionário dos animáis do Brasil. (Ed. Univ. Brasilia, Sao Paulo.

478

INSECT STUDY

Monographs and Serials According to their frequency a n d finality of issue, publications may be classified into two categories, m o n o g r a p h s a n d seri­ als. M o n o g r a p h s are individual a n d selfcontained t r e a t m e n t s of a particular sub­ ject. T h e y may be c o m p o s e d of a single v o l u m e or several volumes, all published at o n e time. An e x a m p l e of a single v o l u m e m o n o g r a p h is: Price, P. W., 1984 (2 ed.), Insect ecology (Wiley, New York). A multivolume m o n o g r a p h is: Kerkut, G. A., a n d L. I. Gilbert, eds., 1985, Comprehensive insect physiology, biochemistry and pharmacol­ ogy ( P e r g a m o n , O x f o r d ) , vols. 1 - 1 3 . Serial publications are characterized by issue over a n interval of time. Periodicity may be r e g u l a r or occasional, continuing ( m e a n t to be o p e n - e n d e d , a l t h o u g h some have ceased publication after a period) or t e m p o r a l (complete after a preset n u m b e r of issues). Many are o r g a n s of learned societies (journals), b u t o t h e r s a r e indepen­ d e n t outlets for original r e p o r t s of scien­ tific investigation. Major c o n t i n u i n g serials published in Latin America which are strictly entomological or m o r e general but with significant space d e d i c a t e d to regional entomology are the following (see Anony­ m o u s 1983, H a m m a c k 1970, a n d King 1986 for m o r e complete lists). Argentina Acta Scientífica: Institutos d e Investi­ gación d e San Miguel, Instituto de Ciencias Naturales, San Miguel, 1 9 5 5 - . Agro: Ministerio d e Asuntos Agrarios, La Plata. 1 9 5 9 - . Arthropoda: Asociación A r g e n t i n a de A r t r o p o d o l o g í a , B u e n o s Aires, 1 9 4 7 - . Bibliografía Entomológica Argentina, Supp l e m e n t o (Augusto A. Piran), 1 9 6 1 - . Boletín d e la Sociedad Entomológica Ar­ gentina, B u e n o s Aires, 1 9 2 5 , 1 9 3 1 . Boletín Técnico del Instituto Científico de Medicina Veterinaria, B u e n o s Aires, 1957-. Comunicaciones, Entomología del Museo

A r g e n t i n o d e Ciencias Naturales "Ber­ n a r d i n o Rivadavia," B u e n o s Aires, 1964-. Physis: Asociación A r g e n t i n a d e Ciencias Naturales, B u e n o s Aires, 1915—. Revista Argentina de Entomología: Museo A r g e n t i n o d e Ciencias Naturales, Bue­ nos, Aires, 1935—. Revista d e la Sociedad Entomológica Ar­ gentina, B u e n o s Aires, 1 9 2 6 - . Brazil Acta Amazónica: Instituto Nacional d e Pesquisas d a A m a z o n i a , M a n a u s , 1 9 7 1 - . Anais d a Sociedade d e Biología d e P e r n a m b u c o , Instituto d e Antibióticos, U n i v e r s i d a d e d e Recife, Recife, 1941—. Arquivos de Entomología, Series A a n d B, Escola d e A g r o n o m í a "Eliseu Maciel," Instituto A g r o n ó m i c o d o Sul, Pelotas, Rio G r a n d e d o Sul, 1 9 5 8 - . Arquivos d o Instituto Biológico, De­ p a r t a m e n t o d e Defensa Sanitaria d a A g r i c u l t u r a , Sao Paulo, 1 9 2 8 - . Boletim d o I n s t i t u t o Biológico da Bahia, Bahia, 1 9 5 4 - . Boletim d o M u s e u Nacional d e Rio d e J a n e i r o (Zoología), Brazil, 1 9 4 2 - . Boletim, Nova Serie, Zoología, M u s e u P a r a e n s e "Emilio Goeldi," Instituto Naci­ onal d e Pesquisas d a A m a z o n i a , 1956—. Boletim d o Servico d e Entomología, Sec­ retaria d a A g r i c u l t u r a , I n d u s t r i a e Comercio, Rio G r a n d e d o Sul, 1956—, Boletim d a Sociedade Brasileira d e Entomologia, 1 9 4 8 - . Dipan: Directoria d a P r o d u c á o Animal, Secretaria d e A g r i c u l t u r a , Rio G r a n d e d o Sul, Brasil, I 9 4 8 - . Entomologista Brasileiro, Sao Paulo, 1908-09. Iheringia, Zoología, M u s e o Rio-Grandense d e Ciencias Naturais, Porto Alegre, Rio G r a n d e d o Sul, 1 9 5 7 - . Memorias d o Instituto B u t a n t a n , Sao Paulo, 1 9 2 1 - . Revista Brasileira de Entomología: Socie­ d a d e Brasileira d e Entomología, Sao Paulo, 1 9 5 4 - .

Revista Brasileira de malariologia e doencas tropicais: D e p a r t a m e n t o Nacional d e E n d e m i a s Rurais, Divisáo d e C o o p e r acáo e Divulgacáo, Rio d e J a n e i r o , 1951-. Revista de Entomología, Rio d e J a n e i r o , 1931-1951. Studia Entomológica (Revista I n t e r n a c i ­ onal d e Entomología), Petropolis, Rio d e Janeiro, 1952-. Revista Brasileira de Biología: A c a d e m i a Brasileira d e Ciencias, Rio d e J a n e i r o , 1941-. Chile Boletín Chileno de Parasitología: D e p a r t a ­ m e n t o d e Parasitología, Universidad d e Chile, Santiago, 1 9 5 4 - . Acta Entomológica Chilena: Instituto d e Entomología, Universidad Metropoli­ tana d e Ciencias d e la Educación, Santi­ ago, 1 9 7 5 - . Investigaciones Zoológicas Chilenas: C e n t r o d e Investigaciones Zoológicas, Universi­ d a d d e Chile, Santiago, 1 9 5 0 - . Publicaciones del C e n t r o d e Estudios Entomológicos, Facultad d e Filosofía y Educación, Universidad d e Chile, Santi­ ago, 1 9 5 8 - . Revista Chilena de Entomología: Sociedad Chilena d e Entomología, Santiago, 1951-. Colombia Anales d e la Sociedad d e Biología d e Bogotá, Bogotá, 1 9 4 5 - . Caldasia: Instituto d e Ciencias N a t u ­ rales, Universidad Nacional d e Colom­ bia, Bogotá, 1 9 4 0 - . Revista d e la Facultad d e Medicina Veterinaria, Universidad Nacional d e Co­ lombia, Bogotá, 1 9 2 8 - . Agricultura Tropical: Asociación Colom­ biana d e I n g e n i e r o s A g r o n ó m i c o s , Bo­ gotá, 1 9 4 5 - . Costa Rica Revista de Biología Tropical, San José, 1953-. Brenesia: Museo Nacional d e Costa Rica, San José, 1 9 7 2 - .

INFORMATION SOURCES

479

Cuba Boletín a n d o t h e r series: Secretaría d e A g r i c u l t u r a , I n d u s t r i a y Trabajo, Sec­ ción d e S a n i d a d Vegetal, La H a b a n a , 1916-. Poeyana, Series A a n d B, Comisión Nacional d e la A c a d e m i a d e Ciencias d e la República d e C u b a , Instituto d e Biología, La H a b a n a , 1964—. Ecuador Revista Ecuatoriana de Entomología y Para­ sitología: C e n t r o E c u a t o r i a n o d e Investi­ gaciones Entomológicas, Q u i t o , 1953—. El

Salvador Boletín Técnico d e la Dirección G e n e r a l d e Investigaciones a g r o n ó m i c a s , Sección d e E n t o m o l o g í a , San Salvador, I960—.

Guyana Entomological Bulletin: D e p a r t m e n t of A g r i c u l t u r e , British G u i a n a (Guyana), Georgetown, 1930-. Jamaica Entomological Bulletin: D e p a r t m e n t of Agriculture of Jamaica, Kingston, 1921-1932. Entomology Circular: D e p a r t m e n t of Ag­ r i c u l t u r e of Jamaica, Kingston, 1921 — 1934. Mexico Boletín de Divulgación: Instituto p a r a el M e j o r a m i e n t o d e la P r o d u c c i ó n d e Azúcar, B a l d e r a s , 1956—. Folia Entomológica Mexicana: Sociedad Mexicana d e E n t o m o l o g í a , Mexico City, 1961-. Revista d e la Sociedad Mexicana d e E n t o m o l o g í a , Mexico City, 1955—. Revista d e la Sociedad Mexicana d e L e p i d o p t e r o l o g í a , Mexico City, 1975—. Nicaragua Circular Entomológica: D e p a r t a m e n t o d e E n t o m o l o g í a , Servicio T é c n i c o Agrícola, Managua, 1952-. Revista Nicaragüense de Entomología, pri­ vately p u b l i s h e d , L e ó n , 1987—.

480

INSECT STUDY

Peru Boletín d e la Sociedad Entomológica Agrícola del Peru, Lima, 1959—. Informe mensual d e la Estación Experi­ mental Agrícola "La Molina," Lima 1927-. Revista Peruana de Entomología (formerly Revista P e r u a n a d e Entomología Agrí­ cola), Sociedad Entomológica del Peru Lima, 1 9 5 8 - . Puerto Rico Journal of Agriculture: University P u e r t o Rico, Río Piedras, 1917—. Uruguay Revista d e la Sociedad U r u g u a y a Entomología, Montevideo, 1956—.

of

de

Venezuela ActaBiológica Venezuelica: Caracas, 1 9 5 1 - . Boletín de Entomología Venezolana: De­ p a r t a m e n t o d e Entomología, Instituto d e Higiene, Caracas, 1941—. Boletín Técnico del Instituto Nacional de Agricultura d e Venezuela, Maracay, 1951-1956. Revista d e la Facultad d e Agronomía, Universidad Central d e Venezuela, Mara­ cay, 1 9 6 8 - . Revista de Medicina Veterinaria y Para­ sitología: Facultad d e Medicina Veteri­ naria, Universidad Central d e Vene­ zuela, Maracay, 1 9 3 9 - . In addition, t h e r e are n u m e r o u s interna­ tional or foreign serials, purely entomologi­ cal a n d m o r e inclusive, that contain signifi­ cant n u m b e r s of articles o n Latin American insects a n d related a r t h r o p o d s : Acta Trópica, Basel, 1 9 4 4 - . Amazoniana, Instituto Nacional de Pes­ quisas d a Amazonia, M a n a u s , 1965—. Beitráge zur Neotropischen Fauna: Humboldt-Hauses in Miraflores u n d des Insti­ tuto C o l o m b o - A l e m á n in Santa Marta, Stuttgart, Germany, 1956—. Biotropica, Association for Tropical Biol­ ogy, Florida, 1963—. Boletín d e la Oficina Sanitaria Pan-

america, Pan A m e r i c a n Sanitary Bu­ r e a u , W a s h i n g t o n , D.C., 1923—. Boletín de Patología Vegetal y Entomología Agrícola, L a b o r a t o r y of Entomology, Estación Nacional A g r o n ó m i c a , Madrid, Spain, 1 9 2 7 - . Ceiba, Escuela Agrícola Panamerica, Tegucigalpa, 1950—. International Journal of Tropical Insect Sci­ ence, P e r g a m o n , O x f o r d , 1980—. Neotropica, Notas Zoológicas S u d a m e r i ­ canas, B u e n o s Aires, 1954—. Revista Sudamericana de Entomología Apli­ cada, Serie A, Entomología Agrícola, 1946-. Studies on Neotropical Fauna and Environ­ ment, 1956—. C o n t i n u e s Beitráge zur N e o t r o p i s c h e n F a u n a a n d Studies on the Neotropical Fauna. Tropical Ecology I n t e r n a t i o n a l Society for Tropical Ecology, Allahabad. I 9 6 0 - . C o n t i n u e s Bulletin of the I n t e r n a t i o n a l Society for Tropical Ecology. Tropical Titles and Pest Management, Cen­ tre for Overseas Pest Research, L o n d o n . 1955—. C o n t i n u e s Pest Articles News S u m m a r y (PANS). T e m p o r a l serials are too n u m e r o u s to list.

References ANONYMOUS. 1983. Serial sources for the Biosis data base. Biosciences Information Service, Philadelphia. HAMMACK, G. M. 1970. The serial literature of entomology: A descriptive study. Entomol. Soc. America, College Park, Md. KING, A. H. 1986. Latin American entomologi­ cal serials. Fla. Entomol. 69: 30-45.

T h e entomological literature can also be classified into a n u m b e r of categories ac­ cording to its p u r p o s e . Only a few of the more i m p o r t a n t areas can be m e n t i o n e d (see T r a u g e r et al. 1974 for an extensive review).

Reference TRAUGER, S. C ,

R. D. SHENF.FELT, AND R.

H.

FOOTE. 1974. Searching entomological litera­ ture. Entomol. Soc. Amer. Bull. 20: 303-315. Reference Works T h e s e are p r i m a r y guides to i n f o r m a t i o n sources, citing publications a n d availability of all o t h e r kinds of literature. T h e y also may provide access to o t h e r r e f e r e n c e works. An e x a m p l e is: A r n e t t , R. H., Jr., 1970, Entomological information a n d re­ trieval (Bio-Rand Fd., Baltimore). Catalogs T h e s e are listings of taxa, usually species, with complete or partial citations of appli­ cable publications a n d o t h e r i n f o r m a t i o n . An e x a m p l e is: Trichopterorum catalogus. A simple file of t a x o n o m i c n a m e s , or "check­ list," is often used by collectors a n d m u ­ seum c u r a t o r s to a r r a n g e their collections a n d is a vital entry point into the technical literature. Bibliographies and Literature Indexes C o m p e n d i a of citations of literature con­ c e r n i n g a particular subject are often p u b ­ lished for the convenience of investigators, reviewers, a n d writers. T h e y r a n g e from very limited, individual t r e a t m e n t s of a single species to very elaborate series cover­ ing a large taxonomic a n d / o r g e o g r a p h i c area. T h e y may be simple listings of pa­ pers, with or without subject analyses a n d a n n o t a t i o n s . N u m e r o u s limited bibliogra­ phies to particular taxa are cited in the text references. An e x a m p l e is: Atchley, W. R., et al., 1981, A bibliography and keyword index of the biting midges (Díptera: Ceratopogonidae), U.S. Dept. A g r i e , Bibl. Lit. Agrie. 13: 1 544. Geographically o r i e n t e d g e n e r a l e n t o ­ mological bibliographies a r e the following: Argentina Piran A. A. 1946. Bibliografía e n t o ­ mológica A r g e n t i n a . Min. Agr. Dir. Gen. Invest. Inst. San. Veg., Ser. B 2(5): 1-144.

INFORMATION SOURCES

481

Costa Rica J i r ó n , L. F., a n d M. E. S a n c h o d e B a r q u e r o . 1983. Índice d e publicaciones entomológicas d e Costa Rica. Consej. Nac. Inves. Cien. T é c . a n d O r g . T r o p . Stud., San José. Mexico Trujillo, P. 1967. Bibliografía e n t o m o ­ lógica d e Baja California. Ed. Californidad, T i j u a n a . Several c o m p r e h e n s i v e , world bibliogra­ phies are available which p e r t a i n to Latin A m e r i c a n entomology. T h e single most i m p o r t a n t c o n t i n u i n g work is the Zoological Record (BioSciences I n f o r m a t i o n Service, Philadelphia, a n d Zoological Society of L o n d o n , L o n d o n ) , which a t t e m p t s to i n d e x all the entomological l i t e r a t u r e from 1849 to d a t e (see T r a u g e r et al. 1974 for others).

Reference TRAUGER, S. C ,

R.

D. SHENEFELT, AND R.

H.

FOOTE. 1974. Searching entomological litera­ ture. Entomol. Soc. Amer. Bull. 20: 303-315. General Works and Textbooks T h e most significant i n f o r m a t i o n o n a subject is ultimately digested a n d orga­ nized into works primarily for teaching a n d p r i m a r y r e f e r e n c e . An e x a m p l e is: C o r o n a d o , R. A., a n d A. M á r q u e z , 1976, Introducción a la entomología (Ed. Limusa, Mexico City). Natural History and Travel Books Valuable i n f o r m a t i o n o n insects is often c o n t a i n e d in books of a g e n e r a l n a t u r e , especially those o n n a t u r a l history a n d travel. It may be necessary to r e a d the entire book to find p e r t i n e n t passages, especially o l d e r works, because they com­ monly have i n a d e q u a t e or missing indexes. A n e x a m p l e is: H o g u e , C. L., 1972, Armies of the ant (World, New York) 2 3 4 p . Dictionaries Semantics is t h e subject of a variety of glossaries, lexicons, a n d dictionaries com­

482

INSECT STUDY

piled especially for entomology. Useful a r e : (1) The Torre-Bueno Glossary of Entomol­ ogy, S. W. Nichols, compiler, a n d R. p Schuh, editor. Revised edition of a glossary of entomology by J. R. d e la Torre-Bueno, including S u p p l e m e n t A by G. S. Tulloch (New York Entomol. Soc. a n d American Mus. Nat Hist., New York, 1989); (2) Dictionnaire des termes techniques d'Entomologie élémentaire by E. Seguy (Lechevalier, Paris 1967); (3) Entomologisches Worterbuch, by S. von Kéler (Akademie, Berlin, 1963); (4) Glossário de Entomología by M. B. d e Carvalho, E. Carvalho d e A r r u d a , a n d G. Pereira d e A r r u d a (Univ. Fed. Rur. Pernambuco, Recife, 1977). (5) Spanish-EnglishSpanish Lexicon of Entomological and Related Terms with Indexes of Spanish Common Names oj Arthropods and Their Latin and English Equivalents, by M. Grieff ( C o m m o n w e a l t h lost. Entomol., Slouth, E n g l a n d , 1985). T h r e e parts, pages separately. Key words for entomological topics and n a m e s a r e listed a n d classified in Thesaurus of Entomology, by R. H. Foote (Entomol. Soc. America, College Park, Md., 1977). Compendia of Research Papers Research p a p e r s o n related topics are fre­ quently collected u n d e r o n e cover a n d the e d i t o r s h i p of o n e or m o r e specialists. T h e y d i l l e r from m o n o g r a p h s in lacking an integrated coverage of the topic. An exam­ ple is: Nault, L. R., a n d J. G. Rodriguez, eds., 1985, The leafhoppers and planthoppers (Wiley, Chichester).

Research Papers T h e majority of scientific literature con­ sists of the original r e p o r t s of research. T h e y are usually narrowly specialized and cover only a small p a r t of a subject and n u m b e r in the h u n d r e d s of t h o u s a n d s . An e x a m p l e is: Bullock, S. H., a n d A. Pes­ cador, 1983, W i n g a n d proboscis dimen­ sions in a sphingid fauna from western Mexico, Biotropica 15: 2 9 2 - 2 9 4 .

Review Papers T h e s e a r e s u m m a r i e s of a subject, usually to discuss c u r r e n t t h o u g h t a n d b r i n g u p to date an analysis of the literature. Some j o u r n a l s are d e d i c a t e d entirely to this type of paper, for e x a m p l e , Annual Review of Entomology. Popular Articles Many scientific subjects a r e of interest to the g e n e r a l public a n d lay r e a d e r s . T h e y are discursive, often a c c o m p a n i e d by n u ­ m e r o u s illustrations, a n d published in p o p u ­ lar magazines. An e x a m p l e is: H o g u e , C. L., 1982, La V i b o r u g a : El E x t r a ñ o insecto q u e se p a r e c e a u n a víbora, Geomundo 6(10): 308-309. Field and Identification Guides Very few of these kinds of publications, which a r e very useful for b o t h the ama­ teur a n d professional alike, a r e available for the Latin A m e r i c a n e n t o m o f a u n a , a n d these t e n d to cover only the m o r e p o p u l a r and b e t t e r - k n o w n g r o u p s , such as butter­ flies. T h e y a r e usually profusely illus­ trated. O n e such work is D ' A b r e r a , B., 1984, Butterflies of South America (Hill House, Victoria, Australia). Faunal Surveys and Species Lists This category includes publications o n the kinds of insects or o t h e r terrestrial a r t h r o ­ pods f o u n d in a particular g e o g r a p h i c area or r e p o r t s of faunistic studies (see faunistics, c h a p . 2). T h e y serve for identi­ fication p u r p o s e s a n d often include a n n o ­ tations a n d o t h e r d a t a useful for identi­ fying potential e c o n o m i c pests, for ecol­ ogy, or for b i o g e o g r a p h i c studies. T h e y are seldom even close to c o m p l e t e a n d generally e x t r e m e l y limited in coverage. Some e x a m p l e s follow for whole insect faunas a n d s o m e locally i m p o r t a n t , smaller areas. Argentina Brewer, M. M., a n d N. V d e Argiielo. 1980. Guía ilustrada d e insectos c o m u n e s

d e la A r g e n t i n a . Min. Cult. E d u c . F u n d . Miguel Lillo. Mise. 67: 1 - 1 3 1 . Havrylenko, D. 1949. Insectos del P a r q u e Nacional N a h u e l H u a p i . A d m . G e n . Parq. Nac. Tur., B u e n o s Aires. Brazil da Costa Lima, A. 1 9 3 9 - 1 9 6 2 . Insetos d o Brasil. Escuela Nac. A g r o n . , Rio d e J a n e i r o , Ser. Didac. Vols. 2 - 5 , 7 - 1 0 , 12-14. Zikáan, J. F., a n d W. Zikán. 1967. Insetofauna d o Itatiaia e d a Mantiqueira. Rev. Brasil. Entomol. 12: 1 1 7 - 1 5 4 . Zikán, J. F , a n d W. Zikán. 1968. Insetofauna d o Itatiaia e d a M a n t i q u e i r a . 3. L e p i d o p t e r a . Pesq. A g r o p e c . Brasil. 3 : 45-109. Central America G o d m a n , F. D., a n d O. Salvin, eds. 1 8 7 9 - 1 9 1 5 . Biología C e n t r a l i - A m e r i cana. 41 vols. Dulau, L o n d o n . Selander, R. B., a n d P. Vaurie. 1962. A gazetteer to a c c o m p a n y the "Insecta" volumes of the "Biología CentraliA m e r i c a n a . " A m e r i c a n Mus. Nov. 2 0 9 9 : 1-70. Chile Irwin, M. E., a n d E. 1. Schlinger. 1986. A gazetteer for the 1966—67 University of California-Universidad d e Chile ar­ t h r o p o d expedition to Chile a n d p a r t s of A r g e n t i n a . Calif. Acad. Sci. Occ. P a p . 144: 1 - 1 1 . Cuba Alayo, P. various dates. Catálogo d e la fauna d e C u b a . T r a b . Divulg. Mus. "Felipe Poey," Acad. Cien. C u b a , La H a b a n a . Several sections, mostly o n Hemiptera. d e Zayas, F. 1974. E n t o m o f a u n a Cu­ bana. Vol 3. Ed. Cien.-Tech., Insto. Cu­ b a n o Libro, La H a b a n a . Polyneoptera. El Salvador Berry, P. A., a n d M. S. Vaquera. 1957. Lista d e insectos clasificados d e El Salva­ dor. Min. Agrie. G a n a d . El Salvador Bol. T é c . 2 1 : 1-134.

INFORMATION SOURCES

483

Galápagos Islands Linsley, E. G. 1977. Insects of the Galá­ pagos ( S u p p l e m e n t ) . Calif. Acad. Sci. Occ. Pap. 125: 1-50. Linsley, E. G., a n d R. L. Usinger. 1966. Insects of the G a l á p a g o s Islands. Calif. Acad. Sci., Ser. 4., P r o c . 3 3 : 1 1 3 - 1 9 6 . Peck, S. B. 1990. Eyeless a r t h r o p o d s of the Galápagos Islands, E c u a d o r : C o m p o ­ sition a n d origin of t h e cryptozoic fauna of a y o u n g , tropical, oceanic archipel­ ago. Biotropica 22: 3 6 6 - 3 8 1 . Roth, V. D., a n d P. R. Craig. 1970. V I I . A r a c h n i d a of the Galápagos Islands. Miss. Zool. Belgique Galápagos, Ecua­ dor. N. a n d J. L e l e u p . 1 9 6 4 - 6 5 . Paris. Vol. 2. Haiti Wolcott, G. N. 1927. Entomologie d'Haiti. Service t e c h n i q u e d u D é p a r t e m e n t d e 1'Agriculture et d e Tenseignem e n t professionnel, Port-au-Prince. Antilles Beatty, H. A. 1944. T h e insects of St. Croix, V I . J. Agrie. Univ. P u e r t o Rico 28: 1 1 4 - 1 7 2 . Bonfils, J. 1969. C a t a l o g u e r a i s o n n é des insects des Antilles francaises. 2. Dictyo p t e r a : Blattaria et M a n t i d a . A n n . Zool. Ecol. A n i m . 1: 1 0 7 - 1 2 0 . G r u n e r , L., a n d J. Riom. 1977. Insectes et papillons des Antilles. Ed. Pacifique, Papeete, Tahiti. Miskimin, G. W., a n d R. M. B o n d . 1975. T h e insect f a u n a of St. Croix, United States Virgin Islands. Science Surv. P u e r t o Rico 13(1): 1 - 1 1 4 . Stiling, P. D. 1986. Butterflies a n d o t h e r insects of the e a s t e r n C a r i b b e a n . Macmillan, L o n d o n . Tucker, R. W. E. 1952. T h e insects of B a r b a d o s . J. Agrie. Univ. P u e r t o Rico 36: 3 3 0 - 3 6 3 . Jamaica Gowdey, C. C. 1928. Catalogus ins e c t o r u m Jamaicensis. Dept. Agrie. Ja­ maica E n t o m o l . Bull. 4: 1-47.

484

INSECT STUDY

Panama Weber, N. A. 1972. T h e entomology o f P a n a m á . Biol. Soc. Wash. Bull. 2: 187 197. Puerto Rico Drewry, G. E. 1970. A list of insects from El Verde, P u e r t o Rico. In H . T. Odurn ed., A tropical rain forest: A study of irra­ diation a n d ecology at El Verde, Puerto Rico. U.S. AEC, Washington, D.C. M a l d o n a d o , J., a n d C. A. Navarro. 1967 Additions a n d corrections to Wolcott's InsectsofPuertoRico.Carib.J.Sci.7:45-64.1 Various a u t h o r s . 1919—. T h e insects of Porto Rico a n d the Virgin Islands. New York Acad. Sci., Scientific survey of P u e r t o Rico a n d the Virgin Islands. Several volumes published in this incom­ plete series. Wolcott, G. N. 1936. "Insectae Borinquenses." A revised a n n o t a t e d check-list of the insects of P u e r t o Rico with a hostplant index by José I. O t e r o . J. Agrie. Univ. P u e r t o Rico, 20: 1-627. Wolcott, G. N. 1941. S u p p l e m e n t to "Insectae B o r i n q u e n s e s , " J. Agrie. Univ. P u e r t o Rico 2 5 : 3 3 - 1 5 8 . Wolcott, G. N. 1951 [1948]. T h e insects of P u e r t o Rico. J. Agrie. Univ. Puerto Rico 32: 1 - 9 7 5 . Surinam Geijskes, D. C. 1967. De insektenfauna van S u r i n a m e , ook vergeleken met die van d e Antillen. Speciaal wat betreft de O d o n a t a . E n t o m o l . Ber. A m s t e r d a m 27: 69-72. Various islands Alvarenga, M. 1962. A e n t o m o f a u n a do A r q u i p é l a g o d e F e r n a n d o d e Noronha, Brasil. Mus. Nac. Rio d e J a n e i r o Arq. 52:21-25. C a m p o s , L., a n d L. E. Peña. 1973. Los insectos d e la Isla d e Pascua. Rev. Chilena E n t o m o l . 7: 2 1 7 - 2 2 9 . Duffy, E. 1964. T h e terrestrial ecology of Ascension Island. J. A p p l . Ecol. 1: 219-251.

Clarke, J. F. G. 1965. M i c r o l e p i d o p t e r a of the J u a n F e r n a n d e z Islands. U.S. Nati. Mus. Proc. 117: 1-106. G r a h a m , J. B., ed. 1975. T h e biological investigation of Malpelo Island, Colom­ bia. S m i t h s o n i a n C o n t r i b . Zool. 176: 1 — 98. H o g u e , C. L., a n d S. E. Miller. 1981. E n t o m o f a u n a of Cocos Island, Costa Rica. Atoll Res. Bull. 250: 1-29. Palacios-Vargas, J. G., J. Llam pallas, a n d C. L. H o g u e . 1982. Preliminary list of the insects a n d related terrestrial a r t h r o ­ pods of S o c o r r o Island, Islas Revillagigedo, Mexico. So. Calif. Acad. Sci. Bull. 8 1 : 1 3 8 - 1 4 7 . Ramos, J. A. 1946. T h e insects of M o n a Island (West Indies). J. Agrie. Univ. P u e r t o Rico 30: 1-74. Robinson, G. S. 1984. Insects of the Falkland Islands: A check list a n d bibli­ ography. Brit. Mus. Nat. Hist., L o n d o n . Schiapelli, R. D., a n d B. S. G e r s c h m a n de Pikelin. 1974. A r a ñ a s d e las Islas Malvinas. Rev. Mus. A r g e n t i n o Cien. Nat. B e r n a r d i n o Rivadavia, Insto. Nac. Invest. Cien. Nat., E n t o m o l . 4: 7 9 - 9 3 . Skottsberg, C , ed. 1 9 2 0 - 1 9 5 6 . T h e natural history of J u a n F e r n a n d e z a n d Easter Island. Vols. 1 - 3 . Almquist a n d Wiksells, U p p s a l a . A few articles o n insects in vol. 3 (Zoology). Smith, D. S., S. J. Ramos, F. McKenzie, E. M u n r o e , a n d L. D. Miller. 1988. Biogeographical affinities of the butterflies of a " f o r g o t t e n " island: M o n a (Puerto Rico). Allyn M u s . Bull. 121: 1-35. Venezuela Martorell, L. F. 1939. Insects observed in the state of A r a g u a , Venezuela, South America. J. Agrie. Univ. P u e r t o Rico 2 3 : 177-264. Expedition Reports A special type of r e s e a r c h r e p o r t covers the results of e x p e d i t i o n s . These a r e often taxonomic in f o r m a t a n d may i n c l u d e the description of new species. Localities,

dates, a n d routes of travel a r e also usually covered a n d b e c o m e a useful r e s o u r c e for the p r e p a r a t i o n of faunal surveys. An e x a m p l e is: Vaurie, C , a n d P. Vaurie, 1949, Insect collecting in G u a t e m a l a 65 years after C h a m p i o n , New York E n t o m o l . Soc. J. 57: 1 — 18. S o m e a r e very elaborate, multivolume series such as the f a m o u s Biología Centrali-Americana (see faunal surveys, above). Much insect a n d a r a c h n i d material was collected by entomologist H. W. Foote o n the Yale Peruvian E x p e d i ­ tion of 1911 w h e n the Inca citadel of Machu Picchu was discovered by H i r a m B i n g h a m (several r e p o r t s by taxonomists on various g r o u p s w e r e published). Computerized Data Banks A new a n d rapidly e x p a n d i n g m e t h o d of storing information, offering quick access to m a n y specialized fields, is by c a p t u r e o n electromagnetic tapes a n d disks. These data banks are available commercially to subscribers (on-line via t e l e p h o n e m o ­ d e m ) , either directly or t h r o u g h libraries. In the field of e n t o m o l o g y only a few are currently available a n d are of limited use­ fulness for Latin America. S o m e of these are (1) Agricultural O n - L i n e Access (AGRÍCOLA), U.S. D e p a r t m e n t of Agri­ c u l t u r e (National Agricultural Library), Science a n d Education A d m i n i s t r a t i o n , Technical I n f o r m a t i o n Systems, Washing­ ton, D. C. (= electronic version of Bibliog­ r a p h y of Agriculture); (2) Biosciences In­ formation Service ( B I O S I S = Biological Abstracts); a n d (3) C A B abstracts, C o m ­ m o n w e a l t h Agricultural B u r e a u , Royal Slough, E n g l a n d . Recent volumes of the Zoological Record are also available on com­ puter. All index m a n y entomological j o u r ­ nals, internationally. A listing of these a n d o t h e r such data banks is available (Kruzas a n d Schmittroth 1981).

Reference KRUZAS. A. T., AND J. SCHMITTROTH, JR.,

eds.

1981. Encyclopedia of information systems and services. 4th ed. Gale Res. Co., Detroit.

INFORMATION SOURCES

485

Government Publications The nature of the publisher is yet another way the entomological literature can be classified. Of special importance in this category are government documents. These are issued for all purposes and normally for the benefit of a wide audi­ ence. Many are often very current and of practical utility. Their availability tends to be limited and short-lived, however, and it is often hard to find older issues. Those of more active departments, commonly those concerned with agriculture and public health, may be numerous and complex bibliographically. Agencies, for example, are often the authors and series can be broken by long periods of inactivity or changes in authority. The special assis­ tance of librarians in the different coun­ tries may have to help researchers in these areas. Access to Literature The primary source of entomological litera­ ture is libraries. Municipal public libraries seldom include extensive technical hold­ ings, and it is usually necessary to consult university, national, or even private librar­ ies to find all but the most popular works. Individual articles from serials, if recently published, often are available as reprints (also called separates or offprints) from the authors. Items may also be purchased from their publishers if recent or from used or antiquarian book dealers if out of print. Quite a few of the latter are in business around the world which include entomological items in their inventories. The best known of these are the following: Antiquariaat Junk, Van Eeghenstraat 129, 1071 GA Amsterdam, The Nether­ lands. A. Asher and Company, Kaisergracht 489, 1017 DM, Amsterdam, The Neth­ erlands. Australian Entomological Press, 14 Chisolm Street, Greenwich, 2065 New South Wales, Australia.

486

INSECT STUDY

Bioquip Products, 17803 La Salle Ave Cárdena, CA 90248, U.S.A. E. W. Classey Ltd., P.O. Box 93, p a r k Road, Faringdon, Oxon, SN7 7DR England. Entomological Reprint Specialists, P.O. Box 77224, Dockweiler Station, Los An­ geles, CA 90007, U.S.A. Dieter Schierenberg BV, Prinsengracht 485-487, 1016 HP Amsterdam, The Netherlands. Sciences Nat, 2 rue André Méllenne, Venette, F-60200 Compiégne, France. Wheldon and Wesley, Ltd., Lytton Lodge, Codicote, Hitchin, Herts, SG4 8TE, England.

RESEARCH Research is investigative activity leading to the discovery and recording of new knowl­ edge. A tremendous amount of such activ­ ity is taking place in the field of entomology. Hundreds of insect scientists in academic as well as applied areas of the subject are rapidly unveiling new facts and interpreta­ tions to be added to humanity's store of information. The process is well ordered and follows the logical steps of the scientific method, which works as well on those problems subject to "proof" by repeatable results (experimental method) and those demonstrable only by application of logical principles (deductive method). Both, how­ ever, require first the collection of facts (data, specimens, in laboratory and field), their subsequent analysis, and finally, the making of conclusions that are published.

Fields of Study Entomologists, arachnologists, myriapodologists, and acarologists pursue research in many different areas. Basically, they work in either the academic or applied (economic) realms, although some cross

over between both. Particular approaches are numerous and increasing in number as the tendency for greater and greater spe­ cialization progresses. The major special­ ties at present, and references to their practices, are: Taxonomy (syslemalics): classification and nomenclature of species and other cate­ gories (Papavero 1983). Evolution: reconstruction of phylogeny and analysis of the evolutionary pro­ cesses. Morphology: structure, cellular to organismal levels. Physiology: function. Genetics: cytology and heredity. Toxicology: properties and use of pesti­ cides. Behavior: insect comportment. Ecology: insects in relation to their envi­ ronment. Agriculture: pests of crops. Medicine: vectors and agents of human diseases. Veterinary medicine: agents and vectors of diseases of domestic animals. Sometimes, specialists combine these in various ways, for example, functional anat­ omy or physiological ecology.

Reference N. 1983. Fundamentos práticos de taxonomía zoológica: colecóes, bibliografía, nomenclatura. Mus. Paraense Emilio Goeldi, Belém.

PAPAVERO,

Fieldwork The collection of facts to apply to studies of insect ecology and many aspects of behavior, taxonomy, functional morphol­ ogy, and other areas, takes place in the field. The entomologist must go to the arena of natural occurrence of the species under investigation and observe uninhib­ ited activity or manipulate the environ­

ment and/or organisms experimentally. Of­ ten considerable planning and logistical preparation are necessary before the work can commence, especially if it involves travel and extended stay in remote locali­ ties. Considerable benefit is to be gained from the use of established field stations, where technical facilities are already avail­ able and where lodging necessities are handled by resident staff, freeing the scien­ tist to concentrate on the research. In Latin America, there are a number of such field stations, run by various agencies, private and governmental, for academic and ap­ plied work. Many are associated with forest reserves, wildlife preserves, nature centers, or are part of national parks. A list of the better known of these that have provided service to entomologists in the recent past follows (some may not be in operation at present). See Castner (1990) for an exhaus­ tive list. Argentina Museo Territorial, Ushuaia, Tierra del Fuego. Belize Desmond Slattery Research Station, Blue Creek River, near Dangriga, sea level, rain forest. Brazil Estacáo Biológica de Boracéia, Museu de Zoología, Universidade de Sao Paulo. Near Mogi das Cruces, Sao Paulo, 850— 900 meters, subtropical wet forest (Travassos and de Camargo 1958). Estacáo de Campo, Parque Nacional de Itatiaia, 20 kilometers north of Itatiaia, Paraná, 300-1,000 meters, mixed mon­ tane forest. Reserva Campinas, Instituto Nacional de Pesquisas Amazonia, 60 kilometers north of Manaus on Boa Vista Highway, 25 meters, mixed lowland wet forest (white sand area). Ducke Forest Reserve, Instituto Nacional de Pesquisas da Amazonia, 26 kilometers northeast of Manaus on Manaus-Ita-

RESEARCH

487

coatiara Highway, 25 m e t e r s , m i x e d low­ land wet forest. Reserva H u m b o l d t , University of Matto Grosso, M a t t o Grosso, at A r i p u a n á , 200 m e t e r s , lowland wet forest. Colombia Los Llanos Tropical Research Station, Centro Internacional de Agricultura Tropical, n e a r Villavicencio, 4 0 0 m e t e r s , dry forest. El Refugio, private c o n s o r t i u m , C a n y o n of Río Claro, 20 kilometers west of D o r o d a l , Antioquia D e p a r t m e n t , 1000 m e t e r s , wet forest. M e r e n b e r g Preserve, n e a r La Plata (road between P o p a y á n a n d Neiva), 230 m e t e r s , wet forest (Buch n.d.). Reserva La Planada, n e a r Ricaurte, Narifio, 2,000 m e t e r s , c l o u d forest. El Rufugio Biological Station, Chocó D e p a r t m e n t , 23 kilometers west of Cali on r o a d to B u e n a v e n t u r a , 1,600—1,900 meters, wet forest ( C a l d e r ó n 1989). Costa Rica ( A n o n y m o u s 1972) La Selva Field Station, O r g a n i z a t i o n for Tropical Studies, n e a r confluence of Ríos P u e r t o Viejo a n d Sarapiqui, H e r e d i a Province, 37—150 m e t e r s , p r e m o n t a n e tropical wet forest (Clark 1988). Palo V e r d e Field Station, O r g a n i z a t i o n for Tropical Studies, h e a d of Golfo d e Nicoya, G u a n a c a s t e Province, 3 - 1 8 3 me­ ters, lowland tropical d r y forest. Las C r u c e s Field Station, O r g a n i z a t i o n for Tropical Studies, 7 kilometers south of San Vito d e Java, C a r t a g o Province, 1,200 m e t e r s , p r e m o n t a n e rain forest. M o n t e v e r d e C l o u d Forest Reserve Field Station, Tropical Science Center, 1,500 m e t e r s , cloud forest. Ecuador La Chiquita, E c u a d o r i a n D e p a r t m e n t of Forest Resources, a p p r o x i m a t e l y 11 kilo­ m e t e r s southeast a n d inland from coastal city of San L o r e n z o , sea level, rain forest (Peck a n d Kulakova-Peck 1980). C e n t r o Científico d e Río P a l e n q u e , Uni­

488

INSECT STUDY

versity of Miami a n d Universidad Cató­ lica d e Q u i t o , 47 kilometers south of Santo D o m i n g o d e Los Colorados on the road to Q u e v e d o , 220 meters, rain forest. T i n a l a n d i a , private, 5 kilometers south of Santo D o m i n g o d e Los Colorados 700 meters, cloud forest. J a t u n Sacha Biological Station, private 8 kilometers east of P u e r t o Misahualli u p p e r Rio Napo, 400 meters, tropical wet forest (Neill a n d Neill 1988). Charles Darwin Research Station Charles Darwin F o u n d a t i o n , Santa Cruz Island, Galápagos Islands, sea level, tropi­ cal scrub. Jamaica Blue M o u n t a i n Field Station, Irish Town, Foothills of Blue Mountains, 700 meters, m o n t a n e d r v forest (Freeman 1986).

search Institute ( S T R I n.d., Ingles 1954, Leigh et al. 1982), G a t ú n Lake, P a n a m a Canal, 100 m e t e r s , tropical rain forest. Peru T a m b o p a t a Reserved Zone, private, at the j u n c t i o n of the Ríos La Torre a n d T a m b o p a t a , M a d r e d e Dios, 290 meters, mixed lowland wet forest (Erwin 1985). Puerto Rico El Verde Station, U.S. D e p a r t m e n t of Energy, Luquillo E x p e r i m e n t a l Forest, 510 m e t e r s , m o n t a n e forest ( O d u m 1970). Biological Field Station, University of P u e r t o Rico (Río Piedras C a m p u s ) , Biol­ ogy D e p a r t m e n t , Sierra Luquillo, R o u t e 191, 600 m e t e r s , m o n t a n e forest. Toro N e g r o Station, University of P u e r t o Rico (Mayaguez C a m p u s ) , Biology De­ p a r t m e n t , Cordillera Central, Villalba, 800 m e t e r s , m o n t a n e forest.

ANONYMOUS. 1972. Field stations of the Organi­ zation for Tropical Studies in Costa Rica. OTS, So. Miami. BUCH, G. n.d. "Merenberg," un santuario selvático en Los Andes de Colombia. López, Popayán. Pamphlet. CALDERÓN, E. 1989. Announcement: "El Refu­ gio" biological station (Chocó biogeographical region, Colombia): An invitation to tropi­ cal biologists. Biotropica 21: 177. CASTNER, J. L. 1990. Rainforests: A guide to research and tourist facilities al selected tropi­ cal forest sites in Central and South America. Feline, Gainesville. CLARK, D. A. 1988. Research on tropical plant biology at the La Selva Biological Station, Costa Rica. Evol. Trends Plants 2: 75-78. ERWIN, T. L. 1985. Tambopata Reserved Zone, Madre de Dios, Perú: History and descrip­ tion of the reserve. Rev. Peruana Entomol. 27: 1-8. FREEMAN, B. 1986. Blue Mt. Field Sta., Irishtown, Jamaica, West Indies. Entomol. News 97: 197. INGLES, L. G. 1954. Barro Colorado—tropical island laboratory. Smithsonian Ann. Rep. 1953: 361-366, pis. 1-6.

Mexico Estación d e Biología C h a m e l a , Instituto d e Biología, Universidad Nacional Autó­ n o m a d e México, 125 kilometers north­ west of Manzanillo (5 km north of P u e r t o Careyes), n e a r sea level, decidu­ ous dry forest. Estación d e Biología Tropical Los Tuxtlas, Instituto d e Biología, Universidad Naciorfal A u t ó n o m a d e México, Sierra d e San Martin, Veracruz state, 33 kilo­ m e t e r s n o r t h e a s t of Catemaco, near coast, h u m i d seasonal forest (Anony­ mous n.d.). Vermillion Sea Field Station, San Diego M u s e u m of Natural History, Bahía de Los Angeles, Baja California, sea level, desert. Reserva Ecológica "El M o r r o d e la Mancha," Instituto Nacional de In­ vestigaciones y Recursos Biológicos. Near Jalapa, Veracruz, sea level, tropical forest.

Trinidad Asa W r i g h t N a t u r e Center, private, Springhill Estate, 4.2 kilometers n o r t h of A r i m a , 365 m e t e r s , wet forest.

References

Collection and Preservation

Panama B a r r o C o l o r a d o Island Biological Re­ search Station, Smithsonian Tropical Re-

ANONYMOUS, n.d. Estación de biología tropical Los Tuxtlas. Cent. Univ. Común. Cien., Univ. Nac. Auton. México, Mexico City. Pamphlet.

T h e collection a n d preservation of speci­ m e n s a n d data a r e basic to all areas of

Venezuela Estación Biológica R a n c h o G r a n d e , P a r q u e Nacional H e n r i Pittier, 15 kilo­ meters n o r t h of Maracay, 1,100 meters, cloud forest. H a t o M a s a g u r a l , private, 40 kilometers south of Calabozo, A n z o á t e g u i , 200 m e ­ ters, llanos. Estación C u l e b r a , F u n d a c i ó n T e r r a m a r , Caracas, P a r q u e Nacional D u i d a - M a r a waka, T e r r i t o r i o Federal A m a z o n a s , 250 m e t e r s , lowland forest. Virgin Islands Virgin Islands Ecological Research Sta­ tion, C a r i b b e a n Research Institute, Col­ lege of the Virgin Islands, L a m e s h u r Bay, St. J o h n , sea level, coastal scrub.

LEIGH, J R ,

E.

G.,

A.

S.

RAND, AND D.

M.

WINDSOR, eds. 1982. T h e ecology of a tropical forest: Seasonal rhythms and long-term changes. Smithsonian Inst., Washington, D.C. Several articles on the environment of Barro Colorado Island. NEILL, D., AND D. NEILL. 1988. Jatun Sacha Bio­ logical Station, Amazonian Ecuador: An invita­ tion to tropical biologists. Biotropica 20: 59. ODUM, H. T. 1970. The El Verde study area and the rain forest systems of Puerto Rico. In H. T. Odum, ed., A tropical rain forest: A study of irradiation and ecology at El Verde, Puerto Rico. U.S. AEC, Washington, D.C. Pp. B3-B32. PECK, S. B., AND J. KULAKOVA-PECK.

1980.

A

guide to some natural history field localities in Ecuador. Stud. Neotrop. Fauna Environ. 15: 35-55. STRI. n.d. Smithsonian Tropical Research Insti­ tute, Information for visitors. Typewritten brochure. TRAVASSOS, L., AND H. F. A. DE CAMARGO.

1958.

A Eslacáo Biológica de Boracéia. Arq. Zool. 11: 1-21.

RESEARCH

489

entomological r e s e a r c h (Kim 1978). It is a truism that living or d e a d insects must be acquired before any investigations can be­ gin, either for e x p e r i m e n t a l o r deductive studies. Extensive collections of specimens are t h e very f o u n d a t i o n of taxonomy, a n d vouchers s h o u l d be kept for any ecological, physiological, o r o t h e r studies to p e r m i t future verification of identifications. T h e r e ­ fore, considerable a t t e n t i o n m u s t be paid to the processes of a c q u i r i n g a n d p r o p e r han­ dling, p r e s e r v i n g , a n d processing of mate­ rial a n d d a t a (Gibson 1960; O l d r o y d 1958; Pastrana 1985; Peterson 1955; South wood 1966; Steyskal et al. 1986; Valenzuela n.d.).

References GIBSON, W. W. 1960. Cómo manejar y usar la colección de insectos. Secr. Agrie. Ganad. Of. Estud. Espec, México. KIM, K. C., ed. 1978. T h e changing nature of entomological collections: Uses, functions, growth and management. Entomol. Scand. 9: H5-177. OLDROYD, H. 1958. Collecting, preserving and studying insects. Macmillan, New York. PASTRANA, J. A. 1985. Caza, preparación y conservación de insectos. 2d ed. Libr. "El Ateneo," Buenos Aires. PETERSON, A. 1955. A manual of entomological technique. 8th ed. Edwards Bros., Ann Arbor. SOUTHWOOD, T R. E. 1966. Ecological methods with particular reference to the study of insect populations. 2d ed. Methuen, London. STEYSKAL, G. C ,

W. L. MURPHY, AND E.

M.

HOOVER, eds. 1986. Insects and mites: Tech­ niques for collection and preservation. U.S. Dept. Agrie. Misc. Publ. 1443: 1-103. VALENZUELA, G. O. n.d. Recolección, montaje y classificación de insectos. Agrie. Trop., Bo­ gotá. Acquiring Material A l t h o u g h t h e r e a r e c o m m e r c i a l insectaries from which living s p e c i m e n s may be p u r ­ chased a n d insect d e a l e r s w h o sell p r e ­ served specimens (some listed below), it is m o r e often necessary for t h e entomologist to find his o r h e r o w n material in t h e field. Of course, if a specific t y p e is sought, it is necessary to look for it in its p r o p e r locale a n d habitat. After these have been deter­

m i n e d , various m e t h o d s for location of specimens a n d their c a p t u r e may then be employed. T h e success of t h e search will depend o n t h e abilities of t h e collector, who should go forth a r m e d with as much knowledge of t h e insect's microhabitat a n d habits as possible. It may take much time a n d detective work to locate rarities in this pursuit, locals familiar with their n a t u r a l s u r r o u n d i n g s a r e often a great help. T h e q u a r r y may be located in its h o m e a n d forced o r enticed from it in different ways. A useful p r o c e d u r e for finding those forms that live hidden a m o n g s h r u b b e r y is to knock them off o n t o a clean piece of cloth o r p a p e r by b l u d g e o n i n g t h e main stems with a stick. Grass- a n d herb-dwelling types may be swept into a n insect net. T h e latter is the s t a n d a r d i m p l e m e n t of t h e collector and comes in a variety of types for special p u r p o s e s (sweeping nets, aerial nets, aquatic nets, etc.). Many kinds of traps for catching insects have b e e n d e v e l o p e d . O n e of the most useful is t h e Malaise t r a p (Malaise 1937). T h e r e a r e different designs (e.g., Townes 1962), b u t all basically c o m b i n e a vertical baffle to stop flying insects a n d a tentlike u m b r e l l a to funnel t h e m into a killing chamber. T h e t r a p works passively or in combination with lures. It can be placed on the g r o u n d o r in vegetation, even sus­ p e n d e d from trees. A device for extracting soil- a n d litterdwelling m i c r o a r t h r o p o d s is t h e Berlese (or Tullgren) funnel (Allison 1983, Mer­ chant a n d Crossley 1970). It consists sim­ ply of a metal o r plastic funnel suspended over a collecting c h a m b e r (killing bottle, alcohol reservoir) over which is placed a s t r o n g light or heater. T h e sample is put into t h e funnel (prevented from falling t h r o u g h by a screen over t h e m o u t h of the stem), a n d t h e a r t h r o p o d s , seeking refuge from t h e light a n d heat, travel downward, eventually d r o p p i n g into t h e chamber.

Lures also w o r k a n d include chemicals a s well as light for nocturnally active forms. Volatile o r a r o m a t i c substances of m a n y kinds (eugenol, feces, r o t t i n g fruit a n d meat, etc.) attract insects; they may be specific in their effect, especially t h e p h e r o mones that h a v e b e e n identified chemically and a r e available in a bottle. Such is Medlure, used to catch a n d m o n i t o r Medi­ terranean fruit fly infestations. O t h e r s a r e more g e n e r a l , as eucalyptol o r oil of winterorreen, which will d r a w in t h e males of many species of o r c h i d bees, o r phenylacetaldehyde, which attracts various kinds of moths. Naturally o c c u r r i n g scents from blossoms o r d r i e d plants can be used also, a prime e x a m p l e b e i n g h e l i o t r o p e , which is irresistible to ithomiid butterflies a n d other L e p i d o p t e r a . L u r e s a r e used in traps (colored p a n s with liquid to d r o w n t h e insects, enclosures, o r sticky surfaces), o r to attract specimens directly to t h e collector. Light, especially that in t h e nearultraviolet p o r t i o n of t h e s p e c t r u m , draws many night insects to its source. Specimens may be collected as they c o n g r e g a t e around street l a m p s o r t h e outside of windows o r even by a gas l a n t e r n placed o n a reflecting b a c k g r o u n d . However, sophisti­ cated e m i t t e r s a n d t r a p s of m a n y kinds have b e e n i n v e n t e d to take direct advan­ tage of this p h e n o m e n o n a n d provide special c o n v e n i e n c e for t h e entomologist. Some a r e m a d e to o p e r a t e u n d e r w a t e r . Currently, most e m p l o y m e r c u r y v a p o r o r fluorescent bulbs that o p e r a t e o n electric­ ity. Portable b a t t e r y - p o w e r e d units a r e available commercially o r can be easily

built.

tractor for soil microarthropods. Georgia Entomol. Soc. J. 5: 83-87. TOWNES, H. 1962. Design for a Malaise trap. Entomol. Soc. Wash. Proc. 64: 253-262. Commercial Dealers T h e r e a r e n u m e r o u s dealers in business to sell living a n d p r e s e r v e d insect specimens a n d / o r e q u i p m e n t a n d supplies for collec­ tors, researchers, a n d teachers. T h o s e cur­ rently active in t h e different c o u n t r i e s can be ascertained by r e f e r e n c e to t h e o t h e r entomological sources listed in this chapter. Protocol It is imperative that t h e collector comply not only with t h e laws of t h e c o u n t r y o r district w h e r e t h e collecting is b e i n g d o n e but also with c o m m o n courtesies r e g a r d ­ ing e n t r y of private property. Permissions may have to be o b t a i n e d in a d v a n c e of fieldwork. Collecting a n d e x p o r t p e r m i t s are r e q u i r e d by most countries a n d special d o c u m e n t s especially n e e d e d to work in n a t u r e preserves, national p a r k s , a n d o t h ­ erwise protected o r militarily sensitive areas. I n t e r n a t i o n a l laws, such as those a d m i n i s t e r e d by t h e C o n v e n t i o n o n I n t e r ­ national T r a d e in E n d a n g e r e d Species of Flora a n d Fauna ( C I T E S ) , also apply to the taking of certain species of insects a n d spiders a n d their t r a n s p o r t a r o u n d t h e world. Many species a r e c o n s i d e r e d threat­ e n e d o r e n d a n g e r e d a n d may b e a c q u i r e d only u n d e r t h e most stringent restrictions. Often, different regulations apply to liv­ ing versus p r e s e r v e d specimens (Fuller a n d Swift 1985).

Reference FULLER, K. S., AND B. SWIFT. 1985. Latin Ameri­

References ALLISON, A. 1983. An inexpensive, portable Tullgren extractor suitable for the tropics. Int. J. Entomol. 25: 321-323. MALAISE, R. 1937. A new insect-trap. Entomol. Tidsskr. 58: 148-160. MERCHANT, V. A., AND D. A. CROSSLEY, JR. 1970.

An inexpensive, high-efficiency Tullgren ex­

can wildlife trade laws. World Wildlife Fund, Washington, D.C. Handling Material Specimens to be kept very differently from ately p r e s e r v e d . T h e y vided with sustenance

alive will be t r e a t e d those to be i m m e d i ­ will have to be p r o ­ and proper environ-

RESEARCH 490

INSECT STUDY

491

mental conditions to e n s u r e their survival both o n their way from t h e field a n d in the laboratory. Special climate-controlled insectaries or vivaria may have to be con­ structed for types with n a r r o w r e q u i r e ­ m e n t s . O t h e r s a r e m o r e easily m a i n t a i n e d with a m i n i m u m of care, a l t h o u g h all should be t r e a t e d with t h e u t m o s t c o n c e r n d u e any living thing. I n s t r u c t i o n s for rear­ ing a n d c u l t u r i n g m a n y kinds of insects, spiders, a n d like c r e a t u r e s a r e available in the materials a n d m e t h o d s sections of re­ search p a p e r s a n d in special t r e a t m e n t s (Siverly 1962; Smith 1966; Singh a n d M o o r e 1985; Singh 1977). T h o s e specimens to be d i s p a t c h e d i m m e ­ diately a r e best killed quickly in a tight gas c h a m b e r or, if large, by injection. T h e most c o n v e n i e n t c h a m b e r s a r e bottles that use dry cyanide crystals, h e l d in place in the b o t t o m by a plug of plaster, c a r d b o a r d , or o t h e r material. Volatile toxic liquids, such as ethyl acetate, b e n z e n e , a n d so on, may be e m p l o y e d likewise. A fraction of a cubic c e n t i m e t e r of ethyl acetate or isopropyl alcohol from a h y p o d e r m i c syringe, in­ jected into the t h o r a x of big beetles, o r t h o p t e r a n s , or m o t h s , will kill t h e m in­ stantly. Soft-bodied a n d m a n y small types, a n d those d e s i r e d for m o r p h o l o g i c a l study, a r e collected directly into fluids (com­ monly 75—90% ethyl or isopropyl alcohol or various fixatives) w h e r e they will die a n d r e m a i n for t r a n s p o r t . Dead specimens quickly b e c o m e brittle or will d e t e r i o r a t e , especially in h u m i d climes, unless c a r e d for properly. T h e y s h o u l d be placed in an airy, rigid c o n t a i n e r (e.g., c a r d b o a r d box) for t r a n s p o r t to the laboratory or field base. Plastic bags a r e a p p r o p r i a t e only for very t e m p o r a r y stor­ age of living or d e a d material. If time is available for i m m e d i a t e m o u n t i n g of speci­ m e n s , this s h o u l d be d o n e while they a r e still flaccid a n d body p a r t s can be m o v e d to desired positions. If m o u n t i n g is to be delayed, t h e specimens s h o u l d be d r i e d to p r e v e n t d e c o m p o s i t i o n . T h i s can be d o n e

by placing t h e m in delicate p a p e r (glassine tissue) envelopes or layering t h e m between sheets of similar material. T h e y should not be placed in contact with cotton because they will b e c o m e e n t a n g l e d in the fibers a n d will be difficult to extract later. O n c e back in the laboratory, the speci­ m e n s can be m o u n t e d or p e r m a n e n t l y preserved in o n e of t h r e e ways, d e p e n d i n g on their body s t r u c t u r e a n d later use. T h e soft-bodied a n d m a n y small types that w e r e taken originally in fluids will be t r a n s f e r r e d to t h e same or other, but clean, fluids. Most dry, h a r d - b o d i e d insects will be p i n n e d . L e p i d o p t e r a may also have their wings s p r e a d . Specific instructions for p i n n i n g a n d s p r e a d i n g a r e available in the references listed above. It is important to point out h e r e that only pins made especially for insect m o u n t i n g must be used. T h e s e are high-quality steel and coated with shellac or m a d e of stainless steel, so that they d o not easily bend or c o r r o d e . T h e y also a r e extra-long (approxi­ mately 30 m m ) to a c c o m m o d a t e the bodies of the specimens a n d allow labels to be fixed b e n e a t h t h e m .

References SINGH, P. 1977. Artificial diets for insects, mites, and spiders. Plenum, New York. SINGH, P., ANO R. F. MOORE, eds.

1985.

Hand­

book of insect rearing. 2 vols. Elsevier, Am­ sterdam. SIVERLY, R. E. 1962. Rearing insects in schools. W. C. Brown, Dubuque. SMITH, C. N. 1966. Insect colonization and mass production. Academic, New York. Data Collection and Labeling I n f o r m a t i o n associated with specimens is as i m p o r t a n t as the specimens themselves, a n d considerable care m u s t be given to collecting a n d r e c o r d i n g it. At the very least, for taxomonic p u r p o s e s , the precise location a n d d a t e of c a p t u r e , plus the collector's n a m e , should be r e c o r d e d and attached. Very useful also will b e notes on observations of ecological variables, behav-

tions, are mechanized (movable m o d u l e storage facilities, "compactors"). Placing specimens u n d e r glass in p i c t u r e frames (Riker m o u n t s ) is not r e c o m m e n d e d for scientific collections. T h e y a r e often b r o k e n by such t r e a t m e n t a n d a r e impossi­ ble to m a n i p u l a t e for close e x a m i n a t i o n . Fumigation is often necessary to p r e v e n t the ravages of m u s e u m pests such as d e r m e s t i d beetles (Anthrenus a n d Thylodrius), psocids ("book lice"), silverfish, a n d the like (Edwards et al. 1981, Story 1985). T h i s can be d o n e on a c o n t i n u o u s basis with n a p t h a l e n e , thymol, or p a r a d i c h l o r o benze, which act mainly as repellents, or intermittently by use of lethal gases such as dimethyl b r o m i d e in controlled c h a m b e r s or u n d e r tents. T h e a t m o s p h e r e must also be kept dry, to p r e v e n t the g r o w t h of molds that also destroy insect specimens. Light should be kept off specimens as well, References because it fades colors a n d contributes to ERWIN, L.J. M. 1976. Application of a computer­ specimen d e t e r i o r a t i o n . ized general purpose information manage­ Liquid ("wet") collections s h o u l d be ment system (SELGEM) to a natural history m a i n t a i n e d in glass, n o t plastic, vials. T h e research data bank (Coleóptera: Carabidae). practice of using individually s t o p p e r e d Coleop. Bull. 30: 1-32. HODGES, R. W., AND R. H. FOOTE. 1982. a n d separately stored vials s h o u l d be Computer-based information system for in­ avoided for large collections. No truly sect and arachnid systematists. Entomol. Soc. perfect seal that p e r m i t s easy removal has Amer. Bull. 28(1): 31-38. yet been devised, a n d such vials dry u p HOGUE, C. L. 1966. A field note form for without fail, r u i n i n g the specimens they special collecting. Entomol. Soc. Amer. Ann. contain. C u r a t o r s a r e w a r n e d against r u b ­ 59: 230-233. ber stoppers, especially, because chemicals they contain dissolve in the fluid a n d Permanent Preservation d a m a g e the specimens. T h e r e c o m m e n d e d Final h a n d l i n g of specimens involves their m e t h o d is to use simple shell vials, stop­ placement into s o m e sort of p e r m a n e n t p e r e d with cotton plugs a n d i m m e r s e d in a storage facility. P i n n e d , dry insects a r e large reservoir (200—300 m i j a r ) . Vigilance normally k e p t in tight-closing w o o d e n con­ is still necessary to k e e p such reservoirs tainers of various sorts. Most m o d e r n col­ filled, b u t considerable time m u s t pass lections e m p l o y small c a r d b o a r d "unit before the vials themselves completely trays" with soft material in t h e b o t t o m to dessicate. receive the pin which, in t u r n , fit into shallow d r a w e r s with glass lids. T h i s m e t h o d p e r m i t s t h e m a x i m u m conve­ References nience for a d d i n g or r e a r r a n g i n g speci­ EDWARDS, S. R., B. M. BELLAND, AND M. E. mens with a m i m i m u m of d a n g e r of break­ KING, eds. 1981. Pest control in museums: A status report (1980). Assoc. System. Coll., age. T h e d r a w e r s are stored in cabinets or Lawrence, Kans. Paginated by sections. racks, which, in some f o r t u n a t e institu­

ior, or o t h e r p e r t i n e n t details that were manifest at t h e time of the insect's cap­ ture. T h e s e a r e best r e c o r d e d in some s t a n d a r d i z e d w r i t t e n format or p r i n t e d form. T h e latter can b e specialized for particular insect types (e.g., aquatic) or general for any taxon ( H o g u e 1966). Ex­ perimental studies or well-defined re­ search projects may r e q u i r e extensive a n d highly o r g a n i z e d d a t a c a p t u r e m e t h o d s . T h o u g h t s h o u l d be given to m a k i n g forms and data c o m p u t e r compatible (Erwin 1976, H o d g e s a n d Foote 1982); devices are even available for directly r e a d i n g data in the field o n t o electronic storage tapes or disks. T h e i n f o r m a t i o n also must be e n c o d e d to e n s u r e correlation with t h e specimens, various a l p h a n u m e r i c systems being most useful.

RESEARCH 492

INSECT STUDY

493

STORY, K. O. 1985. Approaches to pest manage­ ment in museums. Conserv. Anal. Lab., Smithsonian Inst., Washington, D.C.

Illustration and Photography

p h o t o m i c r o g r a p h y . T h e first can be accom­ plished with almost any c a m e r a a n d elemen­ tary knowledge of p r o p e r focusing and lighting. T h e o t h e r s r e q u i r e s o m e special e q u i p m e n t a n d specialized training.

As a n integral p a r t of b o t h the r e c o r d i n g M a c r o p h o t o g r a p h y is practiced fairly of data a n d for p r e s e n t a t i o n of published close to the subject a n d with the intent of results, graphics a r e a n i m p o r t a n t p a r t of p r o d u c i n g an i m a g e usually in the range of research. Pictures o r illustrations can be 0.3 to 2.0 times its actual size. A n absolute m a d e in a variety of ways, i n c l u d i n g h a n d essential is a single-lens-reflex (SLR) cam­ r e n d e r i n g s (drawings, paintings, sketches), era, with t h r o u g h - t h e - l e n s viewing, to per­ p h o t o g r a p h y , a n d with c o m p u t e r imaging. mit accurate focusing a n d avoid the paral­ Illustrations p r e p a r e d by h a n d have lax effect of c a m e r a s with view finders. long b e e n a n d will c o n t i n u e to be essential Lenses with focal lengths from 30 to 200 to research. A variety of m e d i a are used. millimeters a r e used; 50- or 55-millimeters T h e simplest a n d most direct is pencil a n d zoom o r " m a c r o " (long throw) types serve paper, but this suffers from i m p e r m a for the most c o m m o n magnifications and n e n c e . I n k a n d p a i n t a r e m o r e d u r a b l e but a r e the most versatile. Extension tubes, also r e q u i r e m o r e time a n d care to p r o ­ "tele-extenders," or bellows give further d u c e . All types may be r e p r o d u c e d readily choices of magnification. by copy m a c h i n e s a n d p r i n t i n g processes. Pictures are usually taken at small aper­ Some basic principles apply to the p r e p a ­ tures (f 16 to f32) to maximize d e p t h of ration of biological illustrations ( H o d g e s focus. T h e s e r e q u i r e extra light for proper 1989, Wood 1979). Unless o n e is an accom­ e x p o s u r e , which only stroboscobic lights plished artist, for the sake of accuracy a n d provide. efficiency, mechanical aids a r e usually nec­ I n a n i m a t e objects or d e a d organisms are essary to obtain p r o p e r size, p r o p o r t i o n s , subjects easily m a n i p u l a t e d a n d lighted. a n d s h a p e . Such a r e t h e c a m e r a lucida a n d P h o t o g r a p h y of living insects, spiders and o t h e r m i r r o r a n d prism devices that attach the like, is m u c h m o r e difficult because of to microscopes. Grids m a y also be s u p e r i m ­ their usual u n c o o p e r a t i v e behavior. Many posed o n microscope fields or o n objects kinds a r e very timid a n d seldom remain directly a n d d r a w i n g m a d e by m a t c h i n g quiet or in a p r o p e r a t t i t u d e for the photog­ line by line o n a c o r r e s p o n d i n g grid o n the r a p h e r . T h e y may be s t u n n e d o r stilled with paper. To allow c h a n g e s , pencil s h o u l d be cold or chemicals, b u t this destroys their used for p r e l i m i n a r y figures a n d these n o r m a l a p p e a r a n c e a n d may even give either inked or painted over directly o r e r r o n e o u s information to viewers of the t r a n s f e r r e d to a second, m o r e d u r a b l e , p h o t o g r a p h . Skill, patience, a n d experi­ final surface. Size of the subject should ence with insect behavior a r e all necessary always be indicated o n technical d r a w i n g s , prerequisites to this kind of photography. the best way by a scale line to o n e side. T h e Very good pictures are possible using elec­ size of the illustration s h o u l d fit its p u r ­ tronic flash units c o n n e c t e d to SLR cameras pose: very large for exhibit, two to t h r e e with a bracket to hold a constant distance times the p a g e p r i n t e d length for publica­ from light to subject. T h e extremely short tion, smaller for r e c o r d . flash d u r a t i o n stills all motion a n d allows P h o t o g r a p h y for r e s e a r c h p u r p o s e s (Blasmall a p e r t u r e s to be used which give sharp ker 1989, Lefkowitz 1979) generally falls images over a wide r a n g e of magnifications. into o n e of t h r e e types: g e n e r a l p h o t o g r a ­ Photomicrography is photography phy, m a c r o p h o t o g r a p h y (close-up), a n d t h r o u g h a microscope, which in effect takes

494

INSECT STUDY

the place of the c a m e r a ' s lens. Microscope lenses a r e m u c h s h o r t e r in focal length than those of c a m e r a s a n d a r e used for magnifi­ cation m u c h h i g h e r t h a n two or t h r e e times actual size. Almost any microscope can be fitted with a c a m e r a body, b u t lighting must be carefully controlled to give good color o r contrast. Detail is often p o o r because of the very shallow focusing r a n g e of microscope lenses. Often, only a p o r t i o n of a subject can be shown clearly, a n d multiple p h o t o g r a p h s are necessary to tell the whole story. Special types of p h o t o m i c r o g r a p h s may be taken with modified microscopes such as the transmission electron (EM) a n d s c a n n i n g electron (SEM) microscopes, p h a s e con­ trast, X-ray, diffraction microscopes, a n d others. T h e y all have their own applications in entomological research, particularly with work in m o r p h o l o g y , histology, a n d physiol­ ogy. SEM is now used extensively to depict surface details of very small insect struc­ ture, because of its great clarity a n d d e p t h of focus. C i n e m a t o g r a p h y is particularly useful in many insect studies, especially the behavior of living, active specimens. T h e same basic principles that g o v e r n still p h o t o g r a p h y apply, with t h e a d d e d restriction of a fixed shutter s p e e d . Special effects a r e available, however, such as stop action a n d time lapse exposures, of m u c h utility to behavioral and o t h e r analyses. C a m e r a s using film a r e being rapidly replaced by v i d e o t a p e cam­ eras because of the latter's utility at lower ambient light levels a n d reusable r e c o r d i n g surfaces. I m a g e s are also instantly viewable (not n e e d i n g chemical developing) on a c o m m o n television monitor. C o m p u t e r s a r e available which also have graphics capabilities. C h a r t s , g r a p h s , ta­ bles, a n d even high resolution pictures can be d o n e rapidly with p r o g r a m s d e s i g n e d for this p u r p o s e .

References BLAKER, A. A. 1989. Handbook for scientific photography. 2d ed. Focal, Boston.

HODGES, E. R. S., ed. 1989. The Guild hand­ book of scientific illustration. Van Nostrand Reinhold, New York. LEFKOWITZ, L. 1979. Manual of close-up photog­ raphy. Amphoto, Garden City, N.Y. WOOD, P. 1979. Scientific illustration. Van Nos­ trand Reinhold, New York.

Identification I n h e r e n t to taxonomic research a n d essen­ tial to all o t h e r areas of e n t o m o l o g y is t h e correct identification of specimens u n d e r study. Many e r r o r s have been c o m m i t t e d in both academic studies a n d in the applica­ tion of information to control as a result of incorrect species recognition. T h e process of identifying specimens is difficult, owing to the vast n u m b e r s of insect species a n d the special knowlege n e e d e d to work o u t their identities, a n d falls p r o p e r l y in the purview of the taxonomist. Even they trust themselves only within their own specialty areas. Workers are assisted by well-written a n d illustrated taxonomic p a p e r s . T h e s e con­ tain various kinds of s u m m a r i e s of identifi­ cation characteristics, o r d e r e d in some way to p e r m i t step-by-step analysis of the di­ verse features often used. T h e most univer­ sal system for this is t h e t a x o n o m i c key, a series of mutually exclusive s t a t e m e n t s about the organisms, with o n e of which the specimen must agree. T h e s t a t e m e n t s a r e progressively m o r e a n d m o r e exclusive a n d refined until a t e r m i n u s is r e a c h e d , which is the n a m e of the taxon. T h e s a m e result may b e h a d with pictorial keys, diagnostic tables, matrices, a n d o t h e r forms of systematizing characters a n d their states. Lately, c o m p u t e r p r o g r a m s have b e e n developed for identification as well. N o n t a x o n o m i s t s a r e u r g e d to submit p r o p e r l y p r e p a r e d a n d p r e s e r v e d speci­ m e n s to taxonomic authorities. To d o this, they must d e t e r m i n e who a n d w h e r e the best person is, for which t h e r e a r e few directories, unfortunately. Otherwise, they m u s t inquire of their taxonomist col-

RESEARCH

495

leagues, w h o can m a k e r e c o m m e n d a t i o n s . Sometimes n a m e s a n d a d d r e s s e s can be found by r e f e r e n c e to p a p e r s on the gen­ eral g r o u p to which t h e specimens belong. To obtain h e l p with identification, o n e should also follow accepted protocol, first contacting a prospective collaborator a n d asking permission to s u b m i t specimens. T a x o n o m i s t s have the p r e r o g a t i v e of retain­ ing some material in e x c h a n g e for d e t e r m i ­ nations a n d may p r o p e r l y c h a r g e a fee for the service in s o m e cases. Many technical guides exist primarily for identification. Only a few a r e generally useful in Latin A m e r i c a ( B a c h m a n n 1966, Brues et al. 1954, Hollis 1980).

References BACHMANN, A. O. 1966 [1967]. Nueva clave para determinación de los órdenes de in­ sectos sudamericanos. Soc. Eiitomol. Argen­ tina Rev. 29: 11-16. BRUF.S, C.

T.,

A.

L.

MELANDER,

AND F.

M.

CARPENTER. 1954. Classification of insects. Mus. Compar. Zool. Harvard Univ. Bull. 108: 1-917. HOLLIS, 1)., ed. 1980. Animal identification. 3. Insects. Brit. Mus. Nat. Hist., London.

Publication T h e final step in t h e r e s e a r c h process is publication of results (Alley 1987, Trelease 1982). T h i s r e q u i r e s the u t m o s t care a n d ability a n d r e p r e s e n t s t h e goal of all the foregoing activities. Writing a n d g r a p h i c talents as well as k n o w l e d g e of the writings of o t h e r authorities are called for. Research p a p e r s have different formats, d e p e n d i n g on p u r p o s e a n d m e t h o d o l o g y . T h e results of e x p e r i m e n t a l studies are usually re­ p o r t e d u n d e r a series of h e a d i n g s : i n t r o d u c ­ tion (outlining history, significance, a n d aims or h y p o t h e s e s of t h e study), material a n d m e t h o d s (telling precisely how the study was c a r r i e d out, so that it may be r e p e a t e d by others), results (which data resulted from t h e e x p e r i m e n t ) , discussion ( p r e s e n t i n g varied or c o n t r a d i c t o r y aspects of the results), a n d conclusions (what the

496

INSECT STUDY

motivated p e o p l e , a r e displays that i n c o r p o ­ rate actual s p e c i m e n s , p h o t o g r a p h s , a n d data into an i n t e g r a t e d exposition on some topic or p h e n o m e n o n . Particularly well re­ ceived a r e so-called insect zoos w h e r e the public can see at close h a n d e x a m p l e s of living s p e c i m e n s of spectacular, colorful, or

results mean toward p r o v i n g or disproving the initial hypothesis or goal of the study). T h e literature cited section at the e n d gives the references used to g u i d e a n d substanti­ ate the research. T a x o n o m i c p a p e r s follow o t h e r outlines (Mayr a n d Ashlock 1990), generally with headings such as i n t r o d u c t i o n (as in experi­ mental papers), synonymies (listings of var­ ied n a m e s used for the taxa covered), material e x a m i n e d (inventory of specimens used for the study), systematics (presenta­ tion of new g r o u p i n g s a n d descriptions of existing species or newly discovered spe­ cies). Keys a n d tables for identification of new a n d old species are also usually in­ cluded. A u t h o r s of good t a x o n o m i c papers have the responsibility of m a k i n g clear the identity of the taxa included; this is accom­ plished with precisely written descriptions a n d keys as well as good illustrations and the placing of v o u c h e r specimens in multi­ ple public m u s e u m s .

i m p o r t a n t species. Many such zoos a r e o p e n in various parts of the world, a l t h o u g h n o n e to d a t e in Latin America. Education of t h e lay public on insect life is immensely i m p o r ­ tant to the future of agriculture, health, a n d conservation of n a t u r a l resources.

O t h e r deductive studies are written up u n d e r whatever h e a d i n g s best explain the hypothesis a n d application of logic to sup­ port or disclaim t h e m .

References ALLEY, M. 1987. The craft of scientific writing. Prentice-Hall, Englewood Cliffs, N.J. MAYR, E., AND P. D. ASHLOCK. 1990.

Principles

of systematic zoology. 2d ed. McGraw Hill, New York. TRELEASE, S. F. 1982. How to write scientific and technical papers. MIT, Cambridge.

Entomological education Research on insect material forms a fund of knowledge that ultimately will b e c o m e avail­ able to e v e r y o n e t h r o u g h the education process. T h i s is formally achieved by teach­ ing in schools, universities, a n d colleges but is also accomplished t h r o u g h m o r e popular a n d informal media, such as museums, television, n e w s p a p e r s , a n d even in amuse­ m e n t p a r k s . Effective in the process of conveying information, especially to un-

I

RESEARCH

497

Included Insect and Arthropod Taxa

T h e following table will aid the r e a d e r in locating taxa by their classification. All n a m e s or o r d e r s , families, g e n e r a , a n d species that a p p e a r in the book a r e in­ cluded plus a few i n t e r m e d i a t e or h i g h e r categories as n e e d e d . T h e a r r a n g e m e n t of o r d e r s is the s a m e as given in c h a p t e r 1; lower taxa within o r d e r s a r e in the a p p r o x i ­ m a t e s e q u e n c e of their a p p e a r a n c e in the text t h r o u g h o u t t h e book or g r o u p accord­ ing to g e n e r a l evolutionary relationships. C o m m o n n a m e s a r e p r o v i d e d only w h e n in g e n e r a l usage or if given in t h e book. Synonyms a r e e x c l u d e d . Asterisk (*) indi­ cates that taxon is figured. TERRESTRIAL A R T H R O P O D S O T H E R T H A N INSECTS

Phylum O n y c h o p h o r a — o n y c h o p h o r a n s Order Onychophora Peripatopsidae Metaperipatus Peripatidae Macroperipatus torquatus* Peripatus heloisae Speleoperipatus speloeus Phylum A r t h r o p o d a — a r t h r o p o d s S u b p h y l u m Biramia Class C r u s t a c e a — c r u s t a c e a n s Subclass P e r c a r i d a Order Isopoda—isopods Tylidae Ligiidae—sea r o a c h e s Ligia exotica* Ligidium Trichoniscidae

Porcellionidae—woodlice Porcellio laevis* Oniscidae Trichorhina Armadillidiidae—pillbugs Armadillidium vulgare* Armadillidae—pillbugs Order Amphipoda—amphipods Talitridae—beach h o p p e r s Hyale Orchestia platensis—sandflea* S u b p h y l u m Chelicerata Class A r a c h n i d a Order Araneae—spiders Suborder Orthognatha Theraphosidae—tarantulas Acanthoscurria Brachypelma smithi—Mexican redlegged t a r a n t u l a Hapalopus Theraphosa lablondi* Trechona Suborder Labidognatha Salticidae—jumping spiders Aphantochilus* Ctenidae Ctenus Phoneutria fera* P. nigriventer Lycosidae—wolf spiders Lycosa raptoria* A r a n e i d a e — o r b web spiders Araneus Argiope argéntala—silver orb weaver* A. aurantia—golden o r b weaver

499

A. trifasciata—banded o r b weaver Eustala anistera* Mastophora—bolas spiders M. dizzydeani* M. gasteracanthoides Nephila clavipes—golden silk spider* Gasteracanthinae—spiny orb weavers Gasteracantha cancriformis* G. tetracantha Micrathena H e t e r o p o d i d a e — g i a n t c r a b spiders Heteropoda venatoria—huntsman spider* S e l e n o p i d a e — g i a n t c r a b spiders T h e r i d i i d a e — c o m b - f o o t e d spiders Anelosimus eximius Argyrodes Conopistha Latrodectus—widow spiders L. curacaviensis L. geometricus L. mactans—black widow* Mallos gregalis Loxoscelidae—loxoscelid spiders Loxosceles—violin spiders L. laela* Clubionidae Castianeira rica Order Opiliones—harvestmen Suborder Cyphopalpitores Gagrellidae Prionostemma* Suborder Laniatores Cosmetidae Vonones sayi Gonyleptidae Gonyleptus janthinus* Zygopachylus albomarginis S u p e r o r d e r A c a r i — m i t e s a n d ticks O r d e r Astigmata Analgidae Chirodiscidae Chirorhynchobiidae Dermoglyphidae Pyroglyphidae

500

Dermatophagoides faunae— A m e r i c a n h o u s e d u s t mite* D. neotropicalis D. pteronyssinus—European house dust mite Acaridae Tyrophagus putrescentiae—mold mite* Carpoglyphidae Carpoglyphus laclis—dried fruit mite Sarcoptidae Sarcoptes scabiei—scabies mite* O r d e r Prostigmata Cheyletidae Halacaridae Trombiculidae Eutrombicula—chiggers E. alfreddugesi g r o u p E. batatas—sweet p o t a t o chigger* Iguanacarus Leptotrombidium Parascoschoengastia nunezi Pseudoschoengastia Myobiidae Archemyobia latipilis Eriophyidae—gall mites Eriophyes guerreronis—coconut mite E. sheldoni—citrus b u d mite* T e t r a n y c h i d a e — s p i d e r mites Eotetranychus sexmaculatus—sixspotted mite Metatetranychus citri Mononychellus Tetranychus bimaculatus—twospotted mite T. cinnabarinus T. telarius* Pyemotidae Pyemotes ventricosus Demodicidae—follicle mites Demodex bovis D. canis D. caprae D. cati D. equi

INCLUDED INSECT AND ARTHROPOD TAXA

D. folliculorum—human mite* D. ovis D. phylloides O r d e r Mesostigmata Laelaptidae Hypoaspis dasypus Varroa jacobsonii—varroa Macrochelidae Macrocheles Arrhenuridae Arrhenurus Antennophoridae Ophiomegistus Halarachnidae Pneumonyssus Dermanyssidae Dermanyssus Spelaeorhynchidae Spinturnicidae Ascidae Proctolaelaps Rhinoseius

follicle

mite

Order Cryptostigmata Oribatulidae Oribatula minuta* O r d e r Metastigmata—ticks I x o d i d a e — h a r d ticks Amblyomma cajennense—Cayenne tick* A. variegatum—tropical bont tick Aponomma Boophilus microplus—southern cattle tick Dermacentor nitens—tropical h o r s e tick* Haemaphysalis Ixodes pararicinus Rhipicephalus Argasidae—soft ticks Antricola Argas miniatus* A. moreli A. persicus—fowl tick A. transversus Nothoaspis Ornitiiodorus darwini

O. galapagensis O. talaje O. rudis Otobius O r d e r U r o p y g i — w h i p scorpions Elyphonidae Mastigoproctus giganteus— vinegarroon* Thelyphronellus Hypoctonidae A rnauromastigon O r d e r Amblypygi—whipless whip scorpions S u b o r d e r Apulvillata Phrynidae Acanthophrynus Heterophrynus longicornus* Paraphrynus Phrynus Damonidae Trichodamon S u b o r d e r Pulvillata Charontidae Charinides Chirinus Paracharon Tricharinus O r d e r Pseudoscorpionida— pseudoscorpions Chernetidae Cordylochernes scorpioides Lustrochernes Cheliferidae Chelifer cancroides* Cheiridiidae Cheiridium muesorum, Withiidae Withius piger O r d e r Scorpionida—scorpions Buthidae Centruroides limpidus C. suffusus—Durango scorpion* Tityus serrulatus* Order Solpugida—sunspiders Daesiidae Amacata penai

INCLUDED INSECT AND ARTHROPOD TAXA

501

Syndaesia mastix Ammotrechidae Eremobatidae Eremobates* Subphylum Uniramia Class M y r i a p o d a — m y r i a p o d s Subclass C h i l o p o d a — c e n t i p e d e s Order Scolopendromorpha Scolopendridae Scolopendra gigantea—giant centipede* Order Geophilomorpha Order Lithobiomorpha Order Scutigeromorpha Scutigeridae Scutigera coleoptrata—house centipede* Subclass D i p l o p o d a — m i l l i p e d e s Superorder Helminthomorpha O r d e r Spirostreptida Spirostreptidae Orthoporus* Vilcastreptus hoguei O r d e r Polydesmida Platyrhacidae Amplinus Barydesmus* Nyssodesmus python Polylepiscus Pycnotropis Psammodesrnus Chelodesmidae Chondrodesmus O r d e r Spirobolida Rhinocricidae Eurhinocricis Rhinocricus iethifer INSECTS

Class H e x a p o d a — i n s e c t s Subclass Parainsecta—subinsects O r d e r Collembola— springtails Sminthuridae Temeritas surinamensis* Entomobryidae

502

Ctenocyrtinus Coenaletidae

prodigus*

Subclass I n s e c t a — t r u e insects Infraclass A p t e r y g o t a — p r i m i t i v e wingless insects Order Thysanura—thysanurans S u b o r d e r Zygentoma—silverfish Lepismatidae Ctenolepisma longicaudata—longtailed h o u s e silverfish* Lepisma saccharina L. wasmanni Stylifera gigantea Maindroniidae Maindronia Nicoletiidae S u b o r d e r Microcoryphia—bristletails Meinertellidae Meinertellus Neomachillelus scandens* Machilidae Machilinus Machiloides Infraclass P t e r y g o t a — w i n g e d insects S u p e r o r d e r Paleoptera—ancient-winged insects Order Ephemeroptera—mayflies Baetidae Callibaetis Heptageniidae Epearus Tricorythidae Tricorythodes Leptophlebiidae Nousia Thraulodes* Polymitarcyidae Campsurus albicans* Tortopus Siphlonuridae Chaquihua Chiloporter Metamonius Siphlonella Order Odonata—dragonflies and damselflies

INCLUDED INSECT AND ARTHROPOD TAXA

Suborder Anisoptera—dragonflies Petaluridae Phenes raptor Corduliidae Gomphomacromia chilensis Petaliidae Libellulidae Diastatops—black wings D. dimidiata* Libellula hercúlea—ruby tail Orthemis ferruginea—ferruginous skimmer Pantala flavescens—globetrotter* Perithemis—amber wings P. indensa* Zenithoptera—butterfly dragonflies S u b o r d e r Zygoptera—damselflies Calopterygidae Hetaerina americana—ruby spot* Coenagrionidae Acanthagrion Argia vivida* Telchasis Heliocharitidae Perilestidae Polythoridae Pseudostigmatidae Mecistogaster Megaloprepus coerulatus* Superorder Neoptera—modern-winged insects Orthopteroids O r d e r Plecoptera—stoneflies Austroperlidae Eustheniidae Gripopterygidae Araucanioperla Pelurgoperla Notonemouridae Neonemura illiesi Perlidae Anacroneuria* Nemouridae Amphinemura Diamphipnoidae

O r d e r G r y l l o p t e r a — k a t y d i d s a n d crickets Tettigoniidae—katydids Decticinae—shield-backed katydids Eremopedes colonialis Pseudophyllinae—broad-winged katydids Ancistrocercus Celidophylla albimacula Cocconotus* Cyclopteru speculata* Mimetica Panoploscelus Pterochroza ocellata* Tanusia Thliboscelus hypericifolius—Tananá Typophyllum Phaneropterinae—narrow-winged katydids Aganacris* Championica Dysonia fuscifrons* Scaphura Steirodon* Vellea Conocephalinae—cone-headed and m e a d o w katydids Coniungoptera Conocephalus* Copiphora* Neoconocephalus* Panacanthus* Gryllidae—crickets Phalangopsinae Amphiacusta annulipes* A. maya Eneopterinae Eneoptera surinamensis* Oecanthinae Neoxabea Oecanthus* Gryllinae Acheta domesticus—house cricket Grylloides supplicans—Indian h o u s e cricket Gryllus assimiiis Gryllotalpidae—mole crickets Neocurtilla Scapteriscus*

INCLUDED INSECT AND ARTHROPOD TAXA

503

5. S. S. S.

abbreviatus didactylus imitatus oxydactylus

Order Orthoptera—grasshoppers and allies Pauliniidae Cornops aquaticum Marellia remipes Paulinia acuminata* Eumastacidae Eumastax* Acrididae—grasshoppers Achurum sumichrasti* Melanoplus Schistocerca americana S. cancellata S. piceifrons—American locust* Sphenarium Trimerotropis pallidipennis* Romaleidae—lubber grasshoppers Brachystola magna Chromacris speciosa— independence grasshopper* Taenopoda eques T. varipennis* Titanacris gloriosa* T. velazquezii Tropidacris cristata* P r o s c o p i i d a e — j u m p i n g sticks Apioscelis* Order Blattodea—cockroaches Euthyrrhaphidae Holocampsa Nyctiboridae Paratropes Plectoptera Blattidae Blatta orientalis—Oriental cockroach* Litopeltis Neostylopyga rhombifolia— h a r l e q u i n cockroach* Periplaneta americana—American cockroach* P. australasiae—Australian cockroach*

504

Blatellidae Blatella germánica—German cockroach* Megaloblatta Pseudomops* Supella longipalpa—brownb a n d e d cockroach* Oxyhaloidae Leucophaea maderae—Madeira cockroach* Nauphaeta cinérea—lobster cockroach* Atticolidae Achroblatta luteola* Attiphila Myrmecoblatta* Epilampridae Epilampra* Blaberidae Blaberus colosseus B. craniifer B. giganteus—death's-head cockroach* B. parabolicus Pycnoscelididae Pycnoscelus surinamensis— S u r i n a m cockroach* P a n c h l o r i d a e — g r e e n cockroaches Panchlora nivea—Cuban cockroach* Order Mantodea—mantids Mantidae Acanthops falcataria—dead leaf mantid* Angela Choeradodis rhombicollis—leaf mantid* Liturgusa—bark mantids* Mantoida maya Stagmomantis Stagmotoptera* Vates* O r d e r Phasmatodea—walkingsticks Bacteriidae Bactridium grande Bostra scabrinota Otocrania aurita

INCLUDED INSECT AND ARTHROPOD TAXA

Clothoda urichi* Anisembiidae Chelicera Oligotomidae Oligotoma saundersii

Bacunculidae Libethra minúscula Phibalosomatidae Phibalosoma phyllinum* Anisomorphidae Paradoxomorpha crassa— chinchemoyo* Pseudophasmatidae Pseudophasma* Pterinoxylus spinulosus

Hemipteroids

Order Dermaptera—earwigs Anisolabiidae Anisolabis marítima—maritime earwig Carcinophora americana* Euborellia annulipes—ring-legged earwig Metresura ruficeps* Forficulidae Doru lineare—lined earwig* Forfícula auricularia—European earwig Labiduridae Labidura riparia—shore earwig* Sparattidae Spar alia pelvimetra* Labiidae Marava Pygidicranidae Order Isoptera—termites Termitidae Amitermes Cornitermes cumulans Mimeutermes Nasutitermes corniger N. costalis* N. fulviceps Neocapritermes braziliensis* Syntermes dirus* Rhinotermitidae Coptotermes havilandi C. niger Kalotermitidae Cryptotermes brevis Incisitermes snyderi Order Embiidina—web spinners Clothodidae

O r d e r Psocoptera—psocids Liposcelidae Belaphapsocus Liposcelis bostrychophila— booklouse* Asiopsocidae Notiopsocus Psocidae Ceratipsocus Graphocaecilius Poecilopsocus iridescens* Thrysophorus Elipsocidae Drymopsocus Trogiidae O r d e r M a l l o p h a g a — c h e w i n g lice S u b o r d e r Amblycera Abrocomophagidae Trochiliphagidae G y r o p i d a e — g u i n e a pig lice Gliricola porcelli—slender guinea pig louse Gyropus ovalis—oval g u i n e a pig louse* Menoponidae Bovicola Columbicola columbae—pigeon louse Menacanthus stramineus—chicken louse* Menopon gallinae—shaft louse Piagetiella bursaepelecani—pelican louse Ricinidae Trimenoponidae Laemobothriidae Laemobothrion opisthocomi S u b o r d e r Ischnocera Philopteridae Paragoniocotes mirabilis*

INCLUDED INSECT AND ARTHROPOD TAXA

505

T r i c h o d e c t i d a e — m a m m a l chewing lice Cebidicola Felicola felis—cat louse* Geomydoecus Lymeon Neotrichodectes Trichodectes O r d e r A n o p l u r a — s u c k i n g lice Haematopinidae Haematopinus suis—hog louse* Pecaroecus javalii* Solenopotes Linognathidae Linognathus peááalis Microthoracicus mazzai* M. minor M. praelongiceps Hoplopleuridae Hoplopleura Polyplax P e d i c u l i d a e — p r i m a t e lice Pediculus humanus capitis—head louse P. h. corporis—body louse* Phthirus pubis—crab louse* Order Hemiptera—true bugs Suborder Heteroptera-heteropterans Coreidae—big-legged bugs Anasa Diactor bilineatus* Pachylis pharaonis* Paryphes blandus Thasus acutangulus Pentatomidae—stinkbugs Antiteuchus tripterus Chlorochroa ligata—conchuela* Edessa* Eus chis tus Mormidea Oebalus poecilus—rice stinkbug* Lygaeidae—seed bugs Blissus leucopterus—chinch bug* Geocoris punctipes Lygaeus Oncopeltus—milkweed bugs

506

0. fasciatus—large milkweed bug* M i r i d a e — p l a n t bugs Barberiella Engytatus Lygus lineolaris—tarnished plant bug* Monalonion Paracarnus Aradidae—flat bugs Dysodius lunatus—lunate flat bug* Tingidae—lace bugs Corythucha gossypii—cotton lace bug* Reduviidae—assassin bugs Apiomerus lanipes* A. pictipes Arilus carinatus—cogwheel bug* Empicoris rubromaculatus* Graptocleptes Hiranetix Notocyrtus vesiculosus Salyavata variegata* Spiniger* S. ater Triatominae—kissing b u g s Panstrongylus megis tus* Rhodnius pallescens R. prolixus* Triatoma dimidiata T. infestens Polyctenidae—bat bugs Hesperoctenes* Cimicidae—bedbugs Cimex hemipterus—tropical bedbug C. lectularius—bedbug* Haematosiphon inodorus—Mexican chicken b u g Ornithocoris toledoi—Brazilian chicken b u g Psitticimex Pyrrhocoridae—red bugs Thaumastaneis montandoni Alydidae Hyalymenus* Largidae Arhaphe

INCLUDED INSECT AND ARTHROPOD TAXA

C y d n i d a e — b u r r o w i n g bugs Amnestus—pepper flies Nabidae—damsel bugs Arachnocoris albomaculatus B e l o s t o m a t i d a e — g i a n t water b u g s Lethocerus granáis L. maximus* N a u c o r i d a e — c r e e p i n g water bugs Cryphocricos Notonectidae—backswimmers Buenoa pallens* Martarega Notonecta unifasciata Corixidae—water boatmen Corisella Trichocorixa reticulata—salt marsh water b o a t m a n * G e r r i d a e — w a t e r striders Gerris remigus—common water strider* Halobates micans* H. robustas Rheumatobates Hydrometridae—water measurers Bacillometra woytkowskii Saldidae Suborder Homoptera—homopterans, wax b u g s Cicadidae—cicadas Eiáicina chlorogena F. manijera Quesada gigas* Zammara smaragdina* Membracidae—treehoppers Bocyáium* Combophora* Cyphonia Eucyphonia Hemihyptha Heteronotus flavomaculatus* Membracis* Metcalfiella monogramma— p e r i q u i t o del aquacate Polyglypta Spongophorus* Umbonia spinosa* Aetalionidae Aetalion reticulaturn

Cicadellidae—leafhoppers Amblyscartidia albofasciata* B aleja flavoguttata* Dalbulus Empoasca kraemeri* Saccharosyáne saccharivora—cane leafhopper Cercopidae—spittlebugs, froghoppers Aeneolamia varia saccharina— sugarcane froghopper* Tomapsis inca* Aleyrodidae—whiteflies Dialeuroáes Trialeuroáes Aphididae—aphids Acyrihosiphon pisum—pea aphid* Aphis fabae—bean aphid A. gossypii—cotton aphid* A. sacchari Sip ha flava—yellow s u g a r c a n e aphid* Therioaphis maculata—spotted alfalfa a p h i d * Toxoptera aurantii—black citrus aphid Superfamily Coccoidea—scale insects a n d mealybugs Coccidae—tortoise scales Icerya purchasi—cottony cushion scale* Neolecanium sallei M a r g a r o d i d a e — g i a n t coccids Llaveia. axin—axin* Ma rga roáes formicaru m—ground pearls* M. vitium Termitococcus D i a s p i d i d a e — a r m o r e d scales Aoniáiella aurantii—California r e d scale* Aspiáiotus áestructor—coconut scale Lepidosaphes beckii—purple scale Qyadraspidiotus perniciosus—San Jose scale Sclenaspidus articulatus—West I n d i a n r e d scale

INCLUDED INSECT AND AR'LHROPOD TAXA

507

Pseudococcidae—mealybugs Dysmicoccus brevipes—pineapple mealybug Planococcus citri Pseudococcus adonidum P. comstocki P. longispinus—longtailed mealybug* Eriococcidae—mealy scales Dactylopidae—cochineal scales Dactylopius coccus* D. opuntiae D. tomentosus Superfamily F u l g o r o i d e a Fulgoridae—planthoppers Cathedra serrata* Cerogenes auricoma—flying mouse* Fulgora laternaria—dragonheaded bug* Lystra strigata* Phenax variegata* Phrictus diadema* Pterodictya reticularis—reticulate planthopper* Acanaloniidae Derbidae Cixiidae Myndus crudus Flatidae Order Thysanoptera—thrips Phlaeothripidae Leptothrips mali—black h u n t e r * Thripidae Arachisothrips Chaetanaphothrips—banana thrips Dasythrips regalis Frankliniella párvula—banana flower t h r i p s F. tritici—wheat thrips Heliothrips haemorrhoidalis— greenhouse thrips* Hercinothrips bicinctus—banana thrips Scirtothrips Selemnothrips rubrocinctus—cacao thrips Taeniothrips simplex—gladiolus thrips 508

Thrips Uzelothripidae Franklinothrips vespiformis— vespiform thrips Neuropteroids O r d e r Megaloptera—alderflies a n d dobsonflies Sialidae—alderflies Corydalidae-dobsonflies Archichaulides Chloronia Corydalus armatus C. cornutus* Platyneuromus Protochauliodes O r d e r N e u r o p t e r a — n e r v e - w i n g e d insects C o n i o p t e r y g i d a e — d u s t y wings Myrmeliontidae—antlions Brachynemurus Dimares Glenurus peculiaris* Maracandula Morocordula apicalis Myrmeleon* Navasoleon Mella Mantispidae—mantispids Anchieta Climaciella* Drepanicus gayi Chrysopidae—lacewings Ceraeochrysa Chrysopa slossonae Chrysoperla* Leucochrysa H e m e r o b i i d a e — b r o w n lacewings Ascalaphidae—owlflies Albardia fur cata Ameropterus Corduleceris maclachlani* Ululodes Panorpoids O r d e r Diptera—flies a n d midges T i p u l i d a e — c r a n e flies Tipula* Blephariceridae C h i r o n o m i d a e — w a t e r midges Chironomus*

INCLUDED INSECT AND ARTHROPOD TAXA

Siolimyia amazónica P s y c h o d i d a e — m o t h flies Clogmia albipunctata—bathroom fly* Maruina Psychoda altérnala P h l e b o t o m i n a e — s a n d flies Lutzomyia* L. colombiana L. longipalpis L. verrucarum C e r a t o p o g o n i d a e — p u n kies Atrichopogon Bezzia Culicoides* C. furens Dasyhelea Forcipomyia Lasiohelea Leptoconops Microhelea Palpomyia Pterobosca Simuliidae—blackflies Simulium* S. amazonicum S. callidum S. metallicum S. ochraceum Culicidae—mosquitoes Anophelinae Anopheles—malaria mosquitoes A. albimanus A. bellator A. cruzii A. darlingi* A. gambiae A. pseudopunctipennis Chagasia Toxorhynchitinae Toxorhynchites—giant mosquitoes* T. haemorrhoidalis T. theobaldi Culicinae Aedes—Aedes mosquitoes A. aegypti—yellow fever mosquito* A. taeniorhynchus—salt marsh mosquito

Coquillettidea Culex—Culex mosquitoes C. bahamensis C. opisthopus C. quinquefasciatus—southern h o u s e mosquito* Culiseta particeps Deinocerites—crabhole mosquitoes D. cancer* Galindomyia leei Haemagogus—blue devils* Limatus Mansonia Orthopodomyia Phoniomyia Psorophora—gallinippers Sabethes* Trichoprosopon digitatum Uranotaenia Wyeomyia Bibionidae Cecidiomyiidae—gall m i d g e s Latrophobia brasiliensis—manioc gall m i d g e Tabanidae—horseflies Chlorotabanus Chrysops Dichaelacera Fidena Lepiselaga crassipes—mosca c o n g o * Scaptia lata—colihuacho Scione Tabanus dorsiger* C o n o p i d a e — c o n o p i d flies Stylogaster M y d i d a e — m y d a s flies My das* M. rubidapex Asilidae Bombyliidae P a n t o p h t h a l m i d a e — t i m b e r flies Opetiops Pantophthalmus* Phoridae Apocephalus paraponerae Melaloncha Syrphidae—flower flies Copestylum Fristalis tenax—drone fly*

INCLUDED INSECT AND ARTHROPOD TAXA

509

Metasyrphus americanus* Ornidia obesa—green flower fly* S t r a t i o m y i d a e — s o l d i e r flies Hermetia illuscens—wasp fly* Merosargus E p h y d r i d a e — s h o r e flies Dimecoenia D r o s o p h i l i d a e — p o m a c e flies Drosophila carcinophila D. endobranchia D. melanogaster* L o n c h a e i d a e — l o n c h a e i d flies Neosilva perezi—cassava shoot fly T e p h r i t i d a e — f r u i t flies A nastrepha fraterculus—South A m e r i c a n fruit fly A. ludens—Mexican fruit fly A. suspensa—Caribbean fruit fly Ceratitis capitata—Mediterranean fruit fly* Euxesta Rhagoletis lycopersella Toxotriparta curvicauda—papaya fruit fly Milichiidae Pholeomyia Phyllomyza Braulidae Braula coeca C h l o r o p i d a e — f r u i t flies Hippelates Liohippelates pusio c o m p l e x — e y e gnats* Pseudogaurax Coelopidae Chamaemyiidae Paraleucopis mexicana Micropezidae—stilt-legged flies Plocoscelus arthriticus Taeniaptera* Agromyzidae Anthomyiidae Fucellia—kelp flies* F. marítima M u s c i d a e — m u s c i d flies Fannia canicularis—lesser house fly* Haematobia irritans—horn fly* Limnophora 510

Musca domestica—house fly* Muscina stabulans—green house

fly Neivamyia Ophyra aenescens—black g a r b a g e fly* Philornis Stomoxys calcitrans—stable fly* Sarcophagidae—flesh flies Bercaea haemorrhoidalis* Dexosarcophaga Doringia acridiorum Peckia Sarcophaga C a l l i p h o r i d a e — c a r r i o n flies Calliphora Chrysomya Cochliomyia hominovorax— screwworm* C. macellaria—secondary screwworm Luc ilia illustris—greenbottle fly Phoenicia cuprina P. eximia P. sericata—green blowfly* Phormia regina—black blowfly T a c h i n i d a e — t a c h i n i d flies Androeuryops Calodexia Lixophaga diatraeae—Cuban fly Metagonistylum mínense—Amazon fly \ C u t e r e b r i d a e — r o b u s t botflies Alouattamyia Cuterebra Dermatobia hominis—human botfly* O e s t r i d a e — b o t flies Oestrus ovis—sheep botfly* H y p o d e r m a t i d a e — c a t t l e grubs Hypoderma bovis—northern cattle grub H. lineatum—common cattle grub* G a s t e r o p h i l i d a e — h o r s e botflies Gasterophilus haemorrhoidalis— nose botfly G. intestinalis—horse botfly* G. nasalis—throat botfly

INCLUDED INSECT AND ARTHROPOD TAXA

H i p p o b o s c i d a e — l o u s e flies Hippobosca equina—horse louse fly Lipoptena mazamae Melophagus ovinus—sheep ked Olfersia fassulata* Pseudolynchia canariensis—pigeon louse fly S t r e b l i d a e — b a t flies Trichobius dugesii* N y c t e r i b i i d a e — b a t tick flies Basilia ferrisi* Order Siphonaptera—fleas Pulicidae Ctenocephalides canis—dog flea C. felis—cat flea* Leptopsylla segnis—mouse flea Nosopsyllus fasciatus—northern rat flea Pulex irritans—human flea Xenopsylla cheopis—Oriental rat flea* Tungidae Echidnophaga gallinácea— sticktight flea Tunga penetrans—burrowing flea* Ceratophyllidae Ceratophyllus Dolichopsyllidae Dasypsyllus lasius* Ischnopsyllidae Malacopsyllidae Rhopalopsyllidae Pygiopsyllidae S t e p h a n o c i r c i d a e — h e l m e t e d fleas O r d e r Trichoptera—caddisflies Anamolopsychidae Helicophidae Helicopsychidae Helicopsyche Hydrobiosidae Atopsyche callosa* Hydropsychidae Leptonema albovirens* Hydroptilidae Kokiriidae

Lepidostomatidae Leptoceridae Atanalolica Grumichella Hudsonema Nectopsyche punctata* Notalina Triplectides Limnephilidae Philorheithridae Sericostomatidae Grumicha Phylloicus Stenopsychidae Tasimiidae O r d e r Lepidoptera—butterflies and moths Moths Saturniidae—wild silk m o t h s Ceratocampinae Githeronia—regal moths C. laocoon* Lacles—imperial moths E. imperialis decoris* Arsenurinae Arsenura A. ponderosa Gopiopteryx semiramis* Dysdaemonia Loxolornia Paradaernonia Rhescyntis Saturniinae Copaxa—copaxas

C. cydippe C. decrescens C. lavendera* G. rnoinieri Lonornia achelous Rothschildia—window-winged saturnians R. aurota R. erycina* R. orizaha* Hemileucinae Automerella Automeris—eyed saturnians

INCLUDED INSECT AND ARTHROPOD TAXA

511

A. illustris* Dirphia—dirphias D. avia* Camelia Hylesia—hylesias H. canitia H. linéala* H. metabus Hyperchiria Leucanella Paradirphia Pseudautomeris Sphingidae—sphinx moths Macroglossinae Erynnis—ashy sphingids E. ello—ashy s p h i n x * Eumorpha—harlequin sphingids E. fasciata* E. labruscae Hemeroplanes—viper worms H. ornatus* Isognathus Pachylia ficus—fig s p h i n x * Pseudosphinx tetrio—frangipani sphinx* Sphinginae Agrius cingulata Amphimoeca walkeri Cocytius antaeus—giant sphinx* Manduca quinquemaculata— tomato hornworm M. sexta—tobacco hornworm* Sematuridae—eyetails Nothus luna* Uraniidae—rainbow moths Urania fulgens U. leilus* Arctiidae—arctiids Arctiinae—tiger moths Bertholdia Cratoplastis diluta Eucereon Hypercompe decora* Idalus herois* Leuconopsis Opharus bimaculatus Paranerita Viviennea moma*

512

Ctenuchinae—wasp moths Antichloris viridis Correbia* C. lycoides Correbida assimilis Macrocneme chrysitis* Pseudopompilia Pseudosphex Pericopinae—flag m o t h s Che tone angulosa* Daritis howardi* Dyssckema jansoni D. leucophaea* Lithosiidae Ptychoglene coccínea

P. phrada Zygaenidae—smoky moths Harrisina tergina* Seryda constans Dioptidae—dioptid moths Dioptis restricta* Josia Castniidae—giant day-flying moths Castnia cyparissias C. licoides* Microcastnia Agaristidae—forester m o t h s Noctuidae—owlet m o t h s Erastrinae Cydosia Agrotinae Agrotis Ípsilon—black c u t w o r m A. malefida—palesided cutworm A. subterránea—granulate cutworm* Euxoa Mamestra Mods Peridroma saucia—variegated cutworm* Polia Prodenia Spodoptera exigua—beet armyworm* 5. frugiperda—fall armyworm S. latifascia—lateral lined armyworm

INCLUDED INSECT AND ARTHROPOD TAXA

S. ornithogalli—yellow-striped armyworm Heliothidinae Helicoverpa zea—corn e a r w o r m * Catocalinae Alabama argillacea—cotton leaf worm* Plusiinae Pseudoplusia includens Rachiplusia ou—upsilon looper* Trichoplusia ni—cabbage looper* Ophiderinae Ascalapha odorata—black witch* Dipthera festiva — h i e r o g l y p h i c moth* Thysania aggrippina—birdwing moth* T. zenobia—Zenobia's birdwing moth* Hadeninae Pseudaletia adultera P. unipuncta—armyworm* Xanthopastis timáis—Spanish moth* Notodontidae—prominents Cliara croesus* Lasiocampidae—lappet moths Euglyphis cribraria* Gloveria psidii Geometridae—measuring worm moths Atyria dicroides Pantherodes pardalaria—polka dot moth* Megalopygidae—flannel moths Endobrachus revocans Megalopyge lanata* Poda lia Trosia Limacodidae—shag moths Acharia nesea* Phobetron hipparchia—monkey slug* Stenoma cecropia Dalceridae Dalcerina tijucana Lymantriidae—tussock moths Elnoria noyesi

Bombycidae—silk m o t h s Bombyx mori—domestic silk m o t h Tineidae Tinea pellionella—case-bearing clothes m o t h Tineola bisselliella—webbing clothes m o t h Tischeriidae Nepticulidae Psychidae—bagworm moths Oiketicus kirbyi* O. platensis Tortricidae—tortricids Cydia deshaisiana—Mexican jumping bean moth* Lyonetiidae—lyonetiids Leucoptera coffeella—coffee leaf miner Pyralidae—pyralid m o t h s Epipaschiinae Chilozela Macalla thrysisalis—mahogony webworm Chrysauginae Cryptoses choloepi—sloth m o t h * Crambinae Diatraea centrella D. considerata D. grandiosella D. magnifactella D. saccharalis—sugarcane borer* Myelobia smerintha* Phycitinae Anagasta kuehniella— M e d i t e r r a n e a n flour m o t h * Cactoblastis cactorum—cactus moth* Ephestie Plodia interpunctella—Indian meal m o t h * Galleriinae Gallería mellonella—greater wax moth* Gelechiidae Pectinophora gossypiella Phthorimaea operculella Sitotroga cerealella—Angoumois grain m o t h

INCLUDED INSECT AND ARTHROPOD TAXA

513

Cossidae—cossid m o t h s Comadla redtenbacheri—agave worm moth* Butterflies Papilionidae—swallowtail butterflies Battus Eurytides—kites E. bellerophon* E. philolaus* Papilio—true swallowtails P. anchisiades P. andraemon P. cresphontes—giant swallowtail P. homerus P. multicaudata P. thaos—giant swallowtail* P. zagreus Parides—aristolochias P. ascanius P. iphidamas* L y c a e n i d a e — b l u e s , hairstreaks, a n d metalmarks Riodininae—metalmarks Amarynthis menaria* Chlorinea faunus* Helicopis acis* Juditha molpe* Thisbe irenea Lycaeninae—blues and hairstreaks Arawacus aetolus* Chitaría Tmolus basilides—pineapple hairstreak P i e r i d a e — w h i t e s a n d sulfurs Ascia monuste—great southern white* Catasticta semiramis* Colias lesbia Dismorphia amphiona* Eucheira socialis—madrone butterfly Phoebis sennae—cloudless sulfur* Pieris brassicae—European cabbage butterfly Nymphalidae—brush-footed butterflies

514

Satyrinae—satyrs Argyrophorus argenteus—silverwinged butterfly* D a n a i n a e — m o n a r c h butterflies Danaus cleophile—Jamaican monarch D. eresimus—soldier D. erippus—southern monarch D. gilippus—queen D. plexippus—monarch* Lycorea cleobaea—large tiger L. halia* L. ilione Nymphalinae—nymphalines Anaea Anartia amathea—red anartia* A. chrysopelea A. fatima—fatima A. jatrophae—white peacock A. lytrea Callicore Callidula Catacore Colobura dirce—head-for-tail butterfly* Consul fabius* Diaethria asíala D. clymena* Dynamine Eresia phillyra* Hamadryas—crackers H. feronia* Historis odius—cecropia butterfly* Memphis M. arachne Paulogramma Perisama Siproeta stelenes—malachite green* Acraeinae Ithomiinae—ithomiines Creta Hypoleria andromica* Hypolhyris Mechanitis polymnia* Melinaea ethra* Oleria

INCLUDED INSECT AND ARTHROPOD TAXA

Heliconiiae—passion vine butterflies Agraulis vanillae—gulf fritillary* Dryas iulia—Julia* Heliconius charitonius—zebra butterfly H. erato* H. ismenius* H. melanops* Laparus doris Philaethria dido*—green heliconius Morphinae—morphos Morpho achillaena—Achilles morpho* M. hecuba*—hecuba M. peleides M. polyphemus M. rhetenor Brassolinae Caligo—owl butterflies Dynastor darius—Darius* Hesperiidae—skippers Calpodes ethlius—canna skipper* Hylephila phyleus—fiery skipper Urbanus proteus—bean leafroller Chioides C. eurilochus* Elbella polyzona* Phocides thermus Tarsoctenus papias Jamadia gnetus Megathymidae Aegiale hesperiaris—agave worm butterfly* O r d e r s of U n c e r t a i n Affinities Order Coleóptera—beetles C a r a b i d a e — g r o u n d beetles Agra* Calosoma alternans* Enceladus gigas Eurycoleus Lebia Notiobia peruviana* Selenophorus* C i n c i n d e l i n a e — t i g e r beetles Cincindela carthagena*

Ctenostoma Megacephala* Odontocheila* Pseudoxychila bipustulata* Cucujidae—flat bark beetles Oryzaephilus Nitiduidae Haptoncus Mystrops D e r m e s t i d a e — d e r m e s t i d beetles Anthrenus Dermestes Thylodrius Bostrichidae—branch borers Apate A n o b i i d a e — d e a t h - w a t c h beetles Lasioderma serricorne—cigarette beetle Stegobium paniceum—drugstore beetle L y c t i d a e — p o w d e r p o s t beetles Lyctus T e n e b r i o n i d a e — d a r k l i n g beetles Cuphotes* C. irnmaculipes Gyriosomus* Mylaris* Nyctelia* Poecilesthus Proacis bicarinatus* Scotobius gayi* Strongylium* Tauroceras* Tenebrio Tribolium Trogoderma granarium Zophobas—attic beetles* Zopherinae Zopherus chilensis—ma'kech* Erotylidae—giant fungus beetles Cypherotylus dromedarius* Erotylus* Priotelus Pselaphicus giganteus Dytiscidae—predaceous diving beetles Cybister* Megadytes giganteus*

INCLUDED INSECT AND ARTHROPOD TAXA

515

H y d r o p h i l i d a e — w a t e r scavenger beetles Berosus* Gillisius Hydrophilus insularis* Tropisternus laleralis* H e l o d i d a e — m a r s h beetles G y r i n i d a e — w h i r l i g i g beetles Dineutus Gyretes* Gyrinus* H i s t e r i d a e — h i s t e r beetles Ptiliidae—feather-winged beetles Nanosella fungi Silphidae S t a p h y l i n i d a e — r o v e beetles Amblyopinus* Bledius* Cryptomimus Dioploeciton Ecitophya* Odontolinus Paederus irritans—whiplash beetle* Quichuana Spirachtha Termitogaster* Termitonannus Limulodidae—horseshoe crab beetles P s e p h e n i d a e — w a t e r - p e n n y beetles Dryopidae Elmidae Elateridae—click beetles Aeolus Chalcolepidius bonplanni* Conoderus* Pyrophorus—headlight beetles P. nyctophanus* Semiotus* Lampyridae—fireflies Aspisoma Cratomorphus Lucidota* Photinus* Photuris* C a n t h a r i d a e — s o l d i e r beetles Phengodidae—glowworms

516

Phrixothrix* Passalidae—passalid beetles Passalus* Ptichopus L u c a n i d a e — s t a g beetles Chiasognathus granti—Chilean stag beetle* S c a r a b a e i d a e — s c a r a b beetles Aphodiinae Ataenius S c a r a b a e i n a e — d u n g scarabs Canthidium Canthon—dung rollers C. smaragdulum* Coprophanaeus lancifer* Dichotomius carolinus—black dung beetle* Eurysternus deplanatus* Glaphyrocanthon Liatongus Onlhophagus Phanaeus—dung diggers P. demon* Scarabaeus Trichillum, Uroxys U. gorgon D y n a s t i n a e — h o r n e d scarabs Cyclocephala*

INCLUDED INSECT AND ARTHROPOD TAXA

Dynastes hércules—Hercules beetle* D. hyllus D. neptunus D. satanás Enema pan—pan beetle* Golofa aegeon G. eacus G. porteri—caliper beetle* Megaceras jasoni—great horned scarab Megasoma aclaeon M. elephas—elephant beetle* M. mars Oryctes Strategus—ox beetles 5. aloeus* S. oblongus—coconut rhinoceros beetle

S. talpa—sugarcane rhinoceros beetle Cetoniiae—flower scarabs Cotinus mutabilis—green fruit beetle* Gymnetis holocericae circumdata* R u t e l i n a e — s h i n y scarabs Chrysina* Chrysophora chrysochlora—greengold beetle* Heterosternus Macropoidelimus Macropoides Paraheterosternus Pelidnota sumptuosa* P. virescens Plusiotus batesi—gold beetle* P. chtysargyrea—silver beetle M e l o l o n t h i n a e — J u n e beetles Ma crodactylus—cockchafers* Phyllophaga portiricensis* B u p r e s t i d a e — m e t a l l i c wood b o r e r s Agrilus Euchroma gigantea—giant metallic ceiba b o r e r * Coccinellidae—ladybird beetles Cycloneda sanguínea* Epilachna paenulata—melon beetle E. tredecimnotata—southern squash beetle* E. varivestis—Mexican bean beetle Rodolia cardinalis—vedalia beetle L y c i d a e — n e t - w i n g e d beetles Galopteron brasiliense* Lycus arizonensis L. fernandezi O e d e m e r i d a e — f a l s e blister beetles Meloidae P l a t y p o d i d a e — a m b r o s i a beetles S c o l y t i d a e — b a r k beetles Hypothenemus hampei—coffee borer* Platypus parallelus Xyleborus ferrugineus—cacao borer* C e r a m b y c i d a e — l o n g - h o r n e d beetles

Acrocinus longimanus—harlequin beetle* Acyphoderes sexualis* Ancistrotus cummingi Callichroma velutinum Gallipogon—imperious sawyers C. armillatum—giant imperious sawyer* G. barbatum—bearded imperious sawyer* C. senex Elytropeltus apicalis Eplophorus velutinus Hypocephalus armatus—mole beetle* Lycoplasma Macrodontia cervicornis—giant j a w e d sawyer* M. dejeani—big jawed sawyer* M. fiavipenriis Neoptychodes trilineatus—threelined fig t r e e b o r e r * Oncideres sara* Plinthocoelium* Psalidognathus friendi* P. modes tus Schwarzerion Stenygra Taeniotes scalaris* Thelgelra* Tillomorpha Titanus giganteus—titanic longhorn* C h r y s o m e l i d a e — l e a f beetles Alticinae Donaciinae Eumolpinae Colaspis hypochlora Cassidinae—tortoise beetles Acromis spinifex* Gharidotis circumducta* Goptocycla arcuata Cyclosoma mirabilis* Omaspides pallidipennis Omocerus eximius* Polychalca Stolas cyanea* Tauroma

INCLUDED INSECT AND ARTHROPOD TAXA

517

H i s p i n a e — m i n i n g leaf beetles Cephaloleia Chelobasis—rolled-leaf hispine beetles C. bicolor* Pseudocalaspidea cassidea* Xenarescus monocerus Galerucinae Diabrotica undecernpunctata— s p o t t e d c u c u m b e r beetle* B r u c h i d a e — s e e d beetles Acanthoscelides* Callosobruchus Pachymerius nucleorum—bicho de coco Curculionidae—weevils Anthonomus grandis—cotton boll weevil Cosmopolites sordidus—banana weevil Entimus—jeweled weevils E. imperialis—jeweled weevil* E. nobilis Rhinostotnus barbirostris—bearded weevil* Rhynchophorus cruentatus R. palmarum—palm weevil* Sitophilus Brentidae—brentids Brentus anchor ago* O r d e r H y m e n o p t e r a — w a s p s , ants, a n d bees Suborder Symphyta P e r g i d a e — p e r g i d sawflies Acordulecera A r g i d a e — a r g i d sawflies Orussidae T e n t h r e d i n i d a e — c o m m o n sawflies Syzgonia cyanocephala Waldheimia ochra* S i r i c i d a e — h o r n tails Sirex Urocerus californicus* U- gigas fpgas U. patagonicus S u b o r d e r Apocrita I c h n e u m o n i d a e — i c h n e u m o n wasps

518

Rhynchophion Tetragonochora Thyreodon* B r a c o n i d a e — b r a c o n i d wasps Apanteles congregatus* Iphiaulax Superfamily Chalcidoidea—chalcids Torymidae Idarnes Myrmaridae

Alaptus Eurytomidae Bruchophagus plaiyptera Desantisca Trichogrammatidae Trichogramma minutum—minute egg parasite* Eulophidae Aphelinus mali* Eupelmidae Encyrtidae Tetracnernus peregrinus Pteromalidae Chalcididae Agaonidae—fig wasps Blastophaga dugesi* Tetrapus O t h e r Superfamiles Cynipidae—gall wasps Atrusca spinuli* Scelionidae C h r y s i d i d a e — c u c k o o wasps Neochrysis carina* Trichrysis Mutillidae—velvet ants Hoplocrates Hoplomutilla xanthocerata Leucospilomutilla Pappognatha myrmiciformis Traumatomutilla indica* S c o l i i d a e — m a m m o t h wasps Campsomeris ephippium* P o m p i l i d a e — s p i d e r wasps Pepsis* S p h e c i d a e — d i g g e r wasps A mmophila Bembix—sand wasps

INCLUDED INSECT AND ARTHROPOD TAXA

B. americana B. citripes* Larra Microstigmus comes* Sceliphron—mud daubers S. asiaticum S. assimile* S. fistularium Trypoxylon albitarsi* Eumenidae—caterpillar hunters Eumenes—potter wasps E. consobrinus* Montezumia azurescens* Packodynerus galapagensis P. nasidens—wanderer* Zethus matzicatzin* Zeta Vespidae—social wasps Polistinae Apoica—parasol wasps A. pallens* Brachygastra—honey wasps B. leche guana* Chartergus chartarius—bell wasp Mischocyttarus—long-waisted p a p e r wasps M. ater M. drewseni* Parischnigaster Polistes—Polistes p a p e r wasps P. canadensis—varied paper wasp* P. carnifex P. dorsalis clarionensis P. erythrocephalus P. fuscatus Polybia—polybia p a p e r wasps P. dimidiata P. emaciata P. jurinei P. rejecta P. scutellaris* P. singularis Synoeca—drumming wasps 5. cyanea S. septentrionalis S. surinama* Vespinae

Formicidae—ants P o n e r i n a e — g i a n t h u n t i n g ants Dinoponera australis D. gigantea? Ectatomma tuberculatum—kelep: Odontomachus—trap jaw ants* Pachycondyla villosa—cobra ant : Paraponera clavata* E c i t o n i n a e — a r m y ants Eciton E. burchelli E. hamatum* La Indus Neivamyrmex Myrmicinae Acromyrmex Allornerns Atta—leaf cutter ants A. cephalotes A. sexdens Cephalotes* Crematogaster—acrobat ants C. limata C. stolli* Daceton Pheidole—big-headed ants P. fallax* Pogonomyrmex Solenopsis—fire ants S. geminata S. invicta* S. richteri S. saevissima Wasmannia auropunctata Zacryptocerus—cork-head ants Z. maculatus Z. texanus Z. varians* Dolichoderinae Azteca—aztec ants A. alfari A. chartifex A. delpini A. instabilis* A. muelleri A. olitris A. traili A. trigona

INCLUDED INSECT AND ARTHROPOD TAXA

519

A. ulei Dolichoderus Hypoclinea Iridomyrrnex humilis—Argentine ant* Tapinoma—stink ants T. melanocephalum Formicinae Camponotus—carpenter ants C. abdominalis C. femoratus C. planatus C. rufipes C. senex C. sericeiventris* Pseudomyrmecinae Pseudomyrmex—fever ants P. ferrugineus* Superfamily A p o i d e a — b e e s Megachilidae—leaf c u t t e r bees Chrysosarus Cressoniella Megachile leucographa* Pseudocentron Halictidae Agopostemon Augochlora

520

INCLUDED INSECT AND ARTHROPOD

Augochlorella Lasioglossum* Anthophoridae Anthophora Centris—centris bees C. inermis* Xylocopa—carpenter bees X. darwini X. fimbriata* X. frontalis X. ornata Apidae Aglae Apis mellifera mellifera—honeybee* A. m. scutellata—African bee Bombus—bumblebees B. dahlbomi B. tucumanus* Eufriesea Euglossa purpurea* Eulaema meriana* Exeaerete Lestrimelitta limao Melipona beecheii Trígona duckei T. fulvivetitris* T. fuscipennis* T. jaty

TAXA

Index

All taxa h a v e b e e n i n d e x e d at t h e g e n e r i c o r species level, a n d all a r t h r o p o d s a n d plants a r e i n d e x e d by family. C o m m o n n a m e s for a r t h r o p o d species a r e i n d e x e d , but v e r n a c u l a r n a m e s a n d c o m m o n n a m e s at t h e family level o r h i g h e r a r e g e n e r a l l y not i n d e x e d . R e f e r e n c e s to illustrations a r e p r i n t e d in boldface. A b r o c o m o p h a g i d a e , 207 Acacia, 2 8 7 , 4 2 2 , 4 3 5 ; cornígera, 4 3 4 ; decurrens, 2 8 1 , 3 2 3 A c a n a l o n i i d a e , 237 Acanthaceae, 347, 349 Acanlhagrion, 199 Acanlhophrynus, 139 Ac anthops fale ataría, 176—177 Acanlhoscelides, 2 8 7 , 2 8 8 Acanthoscurna, 115 Acari, 3 8 , 56, 5 7 , 5 8 , 5 9 , 70, 76, 8 2 , 9 5 , 9 6 , 9 7 , 9 9 , 1 1 1 , 126— 137, 1 8 7 . 2 6 3 , 2 6 9 , 2 8 3 , 2 9 4 , 409, 433, 444, 469 A c a r i d a e , 129, 1 3 0 - 1 3 1 Adiaría, 3 2 6 , 3 2 7 ; nesea, 3 2 6 Acheta domeslicus, 159 Achroblalta luleola, 170, 171, 2 9 3 , 312,326 Achurum sumichrasli, 162, 163 Acordulecera, 4 0 7 Acrdidae, 63, 64, 91, 161-165, 397 Acrocinus longimanus, 8 3 , 140, 282-284, pl.2a Acrocomm, 2 8 1 , 2 8 9 Acromis, 2 8 6 ; spimfex, 2 8 5 Acromyrmex, 9 1 , 4 4 6 - 4 4 9 Acyphoderes sexualis, 2 7 9 , 2 8 0 , 4 4 0 Acyrthosiphon pisum, 2 4 4 Aedes, 5 9 , 3 7 4 , 3 7 5 , 3 7 6 , 3 7 9 380, 38\;aegypti, 97, 375, 376, 3 8 0 ; taenwrhyndius, 5 8 , 3 7 4 , 379 Aegiale hesperiaris, 3 3 3 Aeneolamia i'aria saccharina, 92— 93, 242, 243 Aeolus, 2 5 9 Aetalion reliculatum, 241 A e t a l i o n i d a e , 241 Aganacris, 1 5 6 . 4 0 9 , 4 1 7 A g a o n i d a e , 84, 4 0 6 , 4 1 0 - 4 1 2 Agaristidac, 2 9 3 , 3 1 1 , 3 1 7 Agavaceae, 272, 333 Agave, 2 7 2 , 3 3 3 Aglae, 461

Agopostemon, 457 Agra, 2 4 7 , 2 4 8 Agrauhs, 354; vanillae, 351 A g r i c u l t u r a l e n t o m o l o g y , 11 — 12, 86,90,91-94,487 Agnlus, 4 4 9 Agnus cingulatus, 306 A g r o m y z i d a e , 72 Agrolis: ípsilon, 3 1 8 ; malefida, 3 1 8 ; subterránea, 3 1 8 Alabama argillacea, 12, 3 2 0 - 3 2 1 Alaptus, 30, 4 0 9 Albardia furcala, 191 Aldrovandi, U , 7 Aleurocanlhus woglumi, 103 A l e y r o d i d a e , 92^ 103 Algae, 392 Allamanda, 3 0 8 AUomerus, 4 3 5 Alnus, 2 8 4 Aloualtamyia, 401 Alydidae,'217 Amacala penai, 143 Amanoa caribaea, 2 6 9 A m a r y l l i d a c e a e , 324 Amaryllis, 324 Amarynthis menaria, 339, 341 Amblyanthera versicolor, 309 Amblyomma, 135; cajennense, 132, 135, 136; variegatum, 136 Amblyopinus, 2 5 6 A m b l v p y g i , 1 8 , 3 7 , 3 8 , 5 6 , 111, 138-139 Amblyscartidia albofasciala, 2 4 2 , 243 A m e r i c a n locust, 163, 164 Ameropterus, 192 Amitermes, 180 Arnmophila, 3 3 , 81 Amnestus, 2 1 7 A m m o t r e c h i d a e , 143 Ampelopsis, 309 Amphiacusta, 5 6 ; annulipes, 158, 159; maya, 158 A m p h i b i a n ( s ) : as host o r prey,

115, 135. 147, 3 7 0 , 3 7 5 , 4 4 4 ; as m i m e t i c m o d e l . 3 5 7 ; as p r e d a ­ tors, 76, 1 0 3 , 2 5 6 , 312 Amphimoeca walkerx, 307 Amphinemura, 201 A m p h i p o d a , 3 8 , 109, 1 1 0 - 1 1 1 Amplinus, 145 Anacardiaceae, 234, 297, 298, 2 9 9 , 301 Anacardium, 301 Anacroneuria, 201 Anaea, 349 Anagasta, 9 9 ; kuehmella, 3 3 2 , 3 3 3 A n a l g i d a e , 128 Anamolopsychidae, 204 Anarlia, 3 4 8 - 3 4 9 ; amalhea, 3 4 8 , 3 4 9 ; chrysopelea, 3 4 9 ; falima, 3 4 9 ; jatrophae, 3 4 9 ; lytrea, 349 Anasa. 218, 219 Anaslrepha, 92, 3 9 0 ; fralerculus, 3 9 0 ; ludens, 102, 3 9 0 ; suspensa, 390 Anchiela, 189 Ancislrocercus, 154 Ancislrolus cummingi, 281 Androeuryops, 4 4 4 Anelosimus eximius, 112 Angela, 177 A n i s e m b i i d a e , 185 Anisolabiidae, 60, 178, 179 Anisolabis marítima, 179 Anisomorphidae, 168-169 A n i s o p t e r a , 196, 1 9 7 - 1 9 8 Anobiidac, 99 Anolis, 2 3 9 Annona, 3 0 4 ; glabra, 3 0 7 Annonaceae, 304, 307, 338 Anopheles, 9 6 , 3 7 4 , 3 7 6 , 3 7 8 3 7 9 ; albimanus, 3 7 4 , 3 7 8 ; bellalor, 3 7 8 ; cruzii, 3 7 8 ; darling!, 59, 3 7 5 , 3 7 8 ; gambiae, 3 7 8 ; pseudopunctipennis, 378 A n o p l u r a , 38, 39, 206, 2 0 9 - 2 1 2 Ant(s): a c r o b a t , 4 3 5 , 4 3 6 , 4 4 2 , 450, 451, 452; Argentine, 453,

521

Ant(s) (continued) 454; army, 76, 82, 256, 257, 336, 351, 442-445, 482, pl.2h; aztec, 183, 434, 435, 436, 449, 4 5 0 - 4 5 1 , 452, pl.3c, pl.4g; bigheaded, 436, 453; carpenter, 177,217,241,432,436,451452; cobra, 439, 441; corkhead, 449-450, 451; fever, 434, 435, 453; fire, 435, 445-446; kelep, 439, 441-442, 453, pl.3b; leafcutter, 91, 171, 263, 265, 385, 446-449; stink, 117, 453, 454; trap jaw, 439, 4 4 0 441. See also Formicidae Ant gardens, 436, 451, pl.4h Antennophoridae, 128 Anthomyiidae, 58, 72, 392-393 Anlhonomus granáis, 92, 288, 442 Anthophora, 455 Anthophoridae, 455, 456. 4 5 8 460 Anthrenus, 493 Anthurium, 436 Antichloris viridis, 314 Antiteuchus Iripterus, 219 Antricola, 136 Aonidiella aurantii, 92, 235 Apúnteles congrégalas, 407 Apate, 91 Aphantochilus, 116, 117 Aphelinus mali, 410 Aphid: bean, 93; black citrus, 91; cotton, 92, 244; pea, 244; spotted alfalfa, 93, 244; yellow sugarcane, 93, 244. See also Aphididae Aphididae, 64, 74, 83, 91, 92, 93, 187,232,243-245,386, 409,410,432,433,454 Aphis, 244; fahae, 93; gossypü, 92, 244; sacchari, 93 Apidae, 77, 81, 86, 95, 105. 128, 333, 455, 459, 460-469 Apiomerus, 222; lanipes, 223; pictipes, 223 Apioscelis, 167, 168 Apis, 95; mellifera adansonii, 467; mellifera mellifera, 77, 86, 105, 128^ 333, 388, 461, 462, 463, 466-469; mellifera scutellata, 77, 467 Apocephalus paraponerae, 440 Apocynaceae, 293, 307, 308, 309,351 Apoica, 427; pallens, 425, 427, pl.4f Apoidea, 74, 77, 84, 455-469 Aponomma, 135 Apterygota, 37, 38 Aquatic insects, 54, 57-60, 66, 70, 76, 106, 128, 162, 170, 1 9 3 205,227-231,253-256,285,

522

INDEX

365-385, 393; fossil, 40; respi­ ration, 28-29, 58-59, 375, 382 Araceae, 85, 382, 384, 436 Arachisothnps, 188 Arachnida, 37, 38, 58, 111-144 Arachnocoris albomaculalns, 217 Aradidae, 220, 221 Araneae, 38, 54, 56, 57, 58, 59, 81, 111, 112-125,200,409, 416,417,418,433,444 Araneidae ( = Argiopidae), 119122, 200 Araneus, 119 Araucanioperla, 58 Araujia sericofera, 85, 293 Arawacus aelolus, 3413-341 Arbutus, 328 Archemyobia latipilis, 127 Archichauhdes, 202 Arctiidae. 1, 79, 276, 277, 292, 2 9 3 , 2 9 6 , 3 1 0 , 3 1 1 , 312-315, 326, 417,pl.2e, pl.3f Arctiinae, 292, 310, 31 1, 3 1 2 313, 326 Arecaccae ( = Palmae), 86, 115, 270, 272,281, 287.289,290. 316,327,358, 383 Argas, 136; miniatus, 132, 136; moreli, 136; persicus, 136; transversus, 136 Argasidae, 132, 134-137 Argia, 199; vivida, 199 Argidae, 407 Argiope, 119; argentata, 119, 120; aurantia, 119; trijasciala, 1 19 Argyrodes, 121 Argyrophorus argenteus, 342, 343 Arhaphe,2\l Arilus carinatus, 222, 223 Anstolochia, 85. 315, 339 Aristolochiaceae, 85, 315, 339 Armadillidae, 110 Armadillidiidae, 109, 110 Armadillidium vulgare, 109, 110 Armyworm: beet, 318, 319; fall, 320; lateral-lined, 319-320; yellow-striped, 319 Arnaurotnastigon, 138 Arrhenuridae, 128 Arrhenurus, 128 Arribálzaga, E., 10 Arribálzaga, F., 10 Arsenura, 304; ponderosa, 297 Arthropoda, 37, 38 Artocarpus, 283 Ascalapha odorala, 322-324 Ascalaphidae, 189, 191-192 Ascia monuste, 5, 342 Ascidae, 127 Asclepiadaceae, 85, 220, 293, 307, 343, 344 Asclepias, 343; curassavica, 220, 344

Asilidae, 277 Asiopsocidae, 63 Aspidiotus destructor, 93 Aspisoma, 170, 171 Asleraceae ( = Compositae), 293, 299,312,313, 314,344 Astigmata, 127 Ataemus, 56 Atanatolica, 203 Áteles, 211 Atopsyche callosa, 204 Atnchopogon, 370 Atrusca, 412; spinuli, 410 Atta, 91, 171, 263, 265,385. 446-449; cephalotes, 446, 447; sexdens, 447 Attalea, 281,289 Atticolidae, 170, 171, 293, 312, 326 Attinae, 73, 91, 171, 263, 265, 385, 446-449 Atliphila, 171 Atyria dicroides, 325 Augochlora, 457 Augochlorella, 457 Austroperlidae, 201 Autographa brassicae, 321 Automerella, 301 Automeris, 298, 301-302, pl.2d; illustris, 301 Avocado, 241,300 Axin. 233, 234 Azteca, 183, 434, 435, 436, 449, 450-451, 452, pl.3c; alfan, 450, 451; chartifex, 450; delpini, 451; inslabilis, 450; muelleri, 450; olilris, 451; tradi, 451; trigona, 450, pl.4g; ulei, 45\ Babesia bigemina, 136 Baccharis, 299 Bacillometra woytkowskii, 57 Bacillus thuringiensis. 75. 103 Bacopa monmeri, 349 Bacteria, 75, 96, 103, 116, 362, 363,394 Bacteriidae, 57, 168 Bactridium grande, 168 Bacunculidae, 57, 169 Bean leafroller, 358 Baetidae, 195 Bagassa guianensis, 283 Baleja ftavoguttata, 242, 243 Banana, 92, 187. 314, 327, 357. 392 Barbenella, 217 Barydesmus, 145 Bas'dia, 403; ferrisi, 402 Bates, Henry Walter, 8-9, 78, 155,352,446 Bats, 56, 123, 127, 131, 136, 153, 207.212,213, 226,227, 296, 312,403-404

Battus, 339 Bedbug, 37, 216, 224, 226, 227 Bee; African, 77, 467; honey. 77, 86, 105, 128, 333,388,461', 462, 463, 466-469; Beebe, William, 9, 268 Beetle: bearded imperious saw­ yer, 281, 282; big jawed sawyer, 281; black dung, 265, 267; ca­ cao borer, 276; caliper, 271; Chilean stag, 8, 261, 263-264; cigarette, 99; coconut rhinoc­ eros, 272; coffee borer, 91, 276, 278; drugstore, 99; elephant, 30, 269, 270; giant imperious sawyer, 281, 282; giant jawed sawyer, 280, 281; giant metallic ceiba borer, 274, 275, pl.lh; gold, 273; great horned scarab, 274; green fruit, 271; greengold, 273-274; harlequin, 83, 140, 282-284, pl.2a; headlight, 4, 6, 23, 46, 239, 256, 259-260, 293, 312; hércules, 30, 246, 268, 269; khapra, 99; melon, 276; Mexican bean, 276; mole, 279, 284; pan, 274; silver, 273; southern squash, 276; spotted cucumber, 285; sugarcane rhi­ noceros, 272; titanic longhorn, 279, 280; three-lined fig tree borer, 283, 284; vedalia, 276; whiplash, 256. See also Coleóptera Behavior, 31-34 Belaphapsocus, 63 Belbenoit. Rene, 334 Belostomatidae, 228-229 Belt, Thomas, 6 Bembix,&\, 405, 413, 419; ameri­ cana, 419; citripes, 418, 419 Bercaea haemorrhoidalis, 396, 397 Berosus, 254, 255 Bertholdia, 312; excelsa, 86 Betulaceae, 284 Be7.ua, 59, 370 Bibionidae, 79 Bicho de coco, 287 Bignoniaceae, 273, 299, 302 Biogeography, 6 2 - 6 8 Biolley, P., 10 Bioluminescence, 23, 259, 2 6 0 261 Biovularia, 76 Biramia, 38 Birds, 56, 183, 263, 304, 336, 405, 414, 423; ant bird, 76, 351, 444; as competitors, 77; as host or prey, 115, 127, 128, 133, 134, 135, 136,207,208,212. 2 1 3 , 2 2 6 , 3 6 9 , 3 7 0 , 3 7 5 , 391, 393, 403; oil bird, 56, 263; as

predators, 76,84, 103,312, 327,334,343,351,444 Bixa, 436 Bixaceae, 436 Blaberidae, 56, 171, 174 Blaberus, 56, 171, 174-175; co/avseus, 17'4; craniifer; \75; giganteus, 174, 175; parabólicas, 175 Black witch, 322-324 Blastophaga, 411; dugesi, 410 Blatella germánica, 174 Blatellidae, 171, 173, 174, 1/5 Blatta orientalis, 174 Blattidae, 56, 60, 172-173, 174 Blattodea, 39, 56, 60, 70, 73, 82, 100, 123, 169-176,286,433, 444 Blechum, 347, 349 Bledius, 256 Blephariceridae, 58, 79 Blissus leucopterus, 220 Bocydium, 241 Bodkin, G. E., 12 Bombacaceae, 275, 281, 283, 291,304 Bombacopsis, 275, 304 Bombax, 291, 304 Bombus, 461-463; dahlbomi, 462: tucumanus, 462 Bombycidae. 298, 327-328 Bombyliidae, 84 Bombyx man, 298, 327-328 Bonpland, Aimé, 8, 147, 362, 377 Booklouse, 185, 186 Boophilus, 135; microplus, 135, 136 Boraginaceae, 287, 293, 312, 313.343, 350, 435, pl.3c Borrelia, 97, 136 Bostra scabrinota, 57, 169 Bostrichidae, 91,99 Botfly: horse, 98, 398, 400: hu­ man, 96, 98, 401-403; sheep, 364, 398, 400 Bothrops, 309; bilineatus, 309 Bovicola, 208 Brachygastra, 429-430; lecheguana, 422, 429 Brachynemurus, 190 Brachypelma smithi, 88, 115 Brachystola magna, 166 Braconidae, 407, 408-409 Bradypus, 330 Braula coeca, 469 Braulidae, 469 Brazilian chicken bug. 226 Brentidae, 288, 291 Brentus anchorago, 288, 291 Bromeliaceae, 59, 76, 86, 115, 170,203, 247,365,371,375, 378,379, 383,384,436,451 Bruchidae, 287-288

Bruchophagus platyptera, 410 Buenoa, 229; pallens, 228, 229 Buprestidae, 72, 95, 274-275, 449, pl.lh Burmeister, H., 9 Bursera, 291 Burseraceae, 291 Buthidae, 95, 141-142 Butterfly: Achilles morpho, 356, pl.2f; agave worm, 333; cecropia, 348, 350; cloudless sul­ fur, 342-343; Darius, 358-359; European cabbage, 342; fatima. 349; giant swallowtail, 338; green heliconius, 347, 354; great southern white, 5, 342; gulf fritillary, 351; head-for-tail, 344. 346-347; hecuba, 30, 356; Jamaican monarch, 344; Julia, 351, 354; large tiger, 344; madrone, 328; malachite green, 344, 346, 354; monarch, 88, 344; owl, 358, pl-2g; pineapple hairstreak, 341; queen, 344; red anartia, 348, 349; silverwinged, 342, 343; soldier, 344; southern monarch, 344; white peacock, 349; zebra, 354 Byrstmima, 302 Byttneria, 436 Cacao, 86, 91, 188,221,272, 278, 370, 375, 383 Cactaceae, 105, 332, 436 Cactoblastis caclorum, 105, 331, 332 Caesalpiniaceae, 40, 239, 301, 303, 323, 342, 435 Calathea, 357,371, 383,389 Caligo, 357-358, pl.2g; eunlochus, 358 Calhandra, 84, 342 Callibaetis, 195 Callichroma, 282; velutinum, 282 Callicore, 348 Callidula, 348 Calliphora, 397 Calliphoridae, 72, 98, 364, 393, 396, 397-400 Callipogon, 282; armillatum, 281, 282; harbatum, 281, 282; senex, 282 Catlosobruchus, 287 Calodexia, 444 Calopteron, 170, 276; brasiliense, 276 Calopterygidae, 199 Calotropis procera, 344 Calpodes ethlius, 358, 359 Calosoma, 248; alternans, 248 Calven, A. S., 9 Camponotus, 177, 217, 241, 436, 451-452; abdominalis, 451, 452;

INDEX

523

Camponotus {continued) femoralus, 4 5 1 ; planalus, 2 1 7 ; rufipes, 4 5 1 ; senex, 4 3 2 , 4 5 1 ; sericeiventns, 4 5 0 , 4 5 2 Campsomens ephippium, 4 1 5 Campsums, 194, 195; albicans, 195 Cane leafhopper, 93 Canna, 3 1 4 . 357 C a n n a c e a e , 3 1 4 , 357 Cantharidae, 277, 280 Canthidium, 2 6 5 Canlhon, 2 6 5 , 2 6 6 ; smaragdulum, 265 C a r a b i d a e , 56, 5 9 . 70, 2 4 7 - 2 5 0 ,

253, pl.lg Carcinophora, 6 0 ; americana, Cardiacephala myrrnex, 392 Cardiospermum, 3 4 8 Cardisoma, 3 8 2 Carludomca divergens, 437 Carpoglyphidae, 130-131 Carpoglyphus lactis, 131 Cassia, 342; fistula, 3 2 3 Castianeira rica, 113 CastHia, 3 1 5 - 3 1 6 , pl.3f;

178

cyparissias, 3 1 6 ; licoides, 3 1 3 , 3 1 6 C a s t n i i d a e , 72, 2 9 3 , 3 1 1 , 3 1 3 , 3 1 5 - 3 1 6 , pl.3f Catacore. 3 4 8 Calagramma, 3 4 8 Calasticta, 3 4 2 ; semiramis, 3 4 2 Cathedra serrata, 2 3 8 , 2 4 0 Cattle g r u b , 9 8 , 3 9 8 , 4 0 0 - 4 0 1 C a v e n d i s h , Sir T h o m a s , 2 5 9 Ceanothus, 301 Cebidicola, 2 0 8 C e c i d i o m y i i d a e , 72 Cecropia, 3 4 7 , 3 7 5 , 4 3 5 , 4 5 0 ; adenopus, 4 3 4 Cedrela odorata, 303 O d r a s , 282 CWia, 2 9 1 ; pentandra, 2 7 5 , 281 Celidophylla albimacula, 155 C e n t i p e d e , 20, 1 4 6 - 1 4 8 ; giant, 30, 145, 147; h o u s e , 145, 147 Cenlris, 4 5 8 ; mermis, 4 5 6 Centruroides, 9 5 , 1 4 1 - 1 4 2 ; //)«p¡í/m, 142; suffusus, 1 4 1 - 1 4 2 Cephaloleia, 60 Cepbaloles, 116, 117, 4 4 9 Ceraeochrysa, 189 C e r a m b y c i d a e , 72, 78, 8 3 , 9 5 , 140, 2 4 6 , 2 4 7 , 2 7 6 , 2 7 7 , 2 7 8 284, 2 8 5 . 4 3 8 . 4 4 0 , 4 5 2 , p l . 2 a Ceratipsocus, 186 Ceralitis capitata, 92, 102, 3 8 9 , 390 Ceratophyllidae, 213 Ceratophyllus, 2 1 3 C e r a t o p o g o n i d a e , 19, 58, 5 9 , 84, 86, 9 7 , 100, 102, 169, 3 6 2 , 3 6 3 , 369, 3 7 0 - 3 7 1 , 4 8 1

524

INDEX

Cercopidae, 9 2 - 9 3 , 242, 243 Cerogenes auricoma, 237—238 Chaelanaphothrips, 188 Chagas, C , 11,224 C h a g a s ' disease, 96, 9 9 , 2 2 4 Chagasia, 3 7 8 Chalcididae, 298, 410 C h a l c i d o i d e a , 84, 4 0 8 , 4 0 9 - 4 1 2 Chale olepidius, 2 5 8 ; bonplanni, 256 C h a m a e m y i i d a e , 391 Championica, 156 Chaquihua, 194, 195 Charidotis circumducta, 285, 286 Charinides, 139 C h a r o n t i d a e , 139 C h a r r i é r e , H e n r i , 334 Chartergus charlarius, 4 1 3 , 428— 429, pl.4e C h e i r i d i i d a e , 140 Cheiridium museoruni, 140 Chelicera, 185 C h e l i c e r a t a , 3 7 , 3 8 , 111 Chetifer cancroides, 138, 140 C h e l i f e r i d a e , 138, 140 Che/obasis bicolor, 285, 286 C h e l o d e s m i d a e , 145 C h e r n e t i d a e , 140, 2 8 3 Chetone angulosa, pl.3f C h e y l e t i d a e , 128 Cliiasognatlius granti, 8, 2 6 1 . 263— 264 ' C h i g g e r ( s ) , vii, 4. 132, 1 3 3 - 1 3 4 ; sweet p o t a t o . 132, 134 C h i g o e , 4. See also Tunga peuetrans C h i l o p o d a . 20, 3 8 , 56, 145, 1 4 6 148 Chiloporter, 194, 195 Chilozela, 9 2 Chineh bug, 220 Chinchemoyo, 168-169 Chwides, 3 5 9 Cliinnus, 139 C h i r o d i s c i d a e , 127 C h i r o n o m i d a e , 23, 5 7 - 5 8 , 59, 63, 365, 3 6 6 - 3 6 7 Chironomus, 2 3 , 5 9 , 3 6 5 , 366 C h i r o r h y n c h o b i i d a e , 127 Chliaria, 341 Chlorinea, 3 4 0 ; faunus, 3 4 1 Chlorochroa ligata, 217. 219 Chloronia, 202 Chlorophora, 284 C h l o r o p i d a e , 124, 3 6 3 , 3 8 9 , 391 Chlorotabanus, 384 Choeradodis rhombicollis, 177 Choloepus, 3 3 0 Chondrodesmus, 145 Chonsia, 2 8 3 , 2 9 1 , 304 Chromacris, 166; speciosa, 165, 166 Chrysididae, 4 1 4 - 4 1 5

Chrysina, 2 7 3 , 2 7 4 Chrysomelidae, 60, 78, 92, 280, 285-287, pl.2b Chrysomya, 397 Chysopa slossonae, 189 Chrysoperla, 189 Chrysophora chrysocldora, 2 7 3 - 2 7 4 C h r y s o p i d a e , 189 Chrysops, 385 Clnysosarus, 457 Chuspi, 43 Citadellidae, 78, 93, 232, 2 4 2 243, 420, 482 C i c a d i d a e , ii, 9 3 , 2 3 2 - 2 3 4 Cicindela, 2 4 8 , 2 4 9 ; carthagena, 248 Cicindelinae, 58, 181, 248, 2 4 9 250, p l . l g Cimex: hemipterns, 2 2 6 , 2 2 7 ; lectulanus. 37, 2 1 6 , 2 2 4 , 2 2 6 227 Cimicidae, 37, 1 0 0 , 2 1 6 , 2 1 7 . 224, 2 2 6 - 2 2 7 Cirplus, 3 1 9 Citheronia, 2 9 8 - 2 9 9 ; laocoon, 299 C i t r u s , 92, 2 7 4 , 3 1 9 , 3 2 7 , 3 9 0 . 465 Cixiidae, 9 3 Classification, 3 8 - 3 9 Chara croesus, 3 2 4 , 3 2 5 Clifford, Sir G e o r g e , 3 8 0 Climaciella, 189 Climate, 4 8 - 5 0 Clogmia albipunclala, 3 6 5 , 367 Clolhoda urichi, 185 C l o t h o d i d a e , 185 C l u b i o n i d a e , 113 Cnidoscolus, 307 C o b o , P. B e r n a b é , 4, 4 3 5 C o c c i d a e , 2 9 , 2 3 5 , 341 Coccinellidae, 2 7 5 - 2 7 6 Coccoidea, 9 3 , 187, 2 3 2 , 2 3 4 237, 386, 409, 432, 4 3 3 , 435 Cocconotus, 154, 155 C o c h i n e a l b u g , 4, 5, 2 1 0 , 2 3 6 Cochhomyia: hominovorax, 9 8 , 364, 3 9 7 . 3 9 8 - 4 0 0 ; macellana, 399 Cochlospermaceae, 299 Cochlosperrnum, 299 C o c k r o a c h : A m e r i c a n , 172, 173; A u s t r a l i a n . 172, 173; b r o w n b a n d e d , 1 7 3 ; C u b a n , 174, 176; d e a t h ' s - h e a d , 174, 175; Ger­ m a n , 174; h a r l e q u i n , 1 7 3 ; lob­ ster, 174; M a d e i r a , 1 7 3 ; O r i e n ­ tal, 174; S u r i n a m , 173 Coco I s l a n d , 198, 3 5 0 , 4 1 9 , 4 2 2 , 485 Cocos, 2 7 0 , 2 7 2 , 2 8 1 , 2 8 9 , 2 9 0 , 358 Cocytms anlaeus, 3 0 6 , 307 Codonanthe, 4 3 6 C o e l o p i d a e , 58

C o e n a g r i o n i d a e , 199 Coenaletidae, 1 4 8 - 1 4 9 Coffee, 9 1 , 2 3 3 , 2 7 4 , 2 7 8 Coffee leaf m i n e r , 91 Cogwheel bug, 222, 223 Colaspis hypochlora, 9 2 C o l e ó p t e r a , 3 8 , 3 9 , 4 6 , 5 7 , 60, 72, 1 0 5 . 2 2 7 , 2 4 6 - 2 9 1 , 3 1 3 , 409,415,428,433,492 Colias lesbia, 342 C o l i h u a c h o , 384 Collection a n d p r e s e r v a t i o n methods, 4 8 9 - 4 9 3 Collembola, 3 8 , 54, 5 7 , 5 9 , 70, 82, 1 4 8 - 1 4 9 , 4 2 0 Colobura dirce, 3 4 4 , 346—347 Columbicola columbae, 2 0 8 Columbus, Cristopher, 4 Comadla redtenbacheri, 46, 3 3 3 Combophora, 241 C o m p e t i t i o n . 7 5 , 7 7 , 287 Conchuela, 2 1 7 . 2 1 9 C o n i o p t e r y g i d a e , 188, 189 Coniungoptera, 158 Conocephalus, 157 Conoderus, 2 5 6 , 2 5 9 Conopidae, 444 Conopistha, 121 C o n s e r v a t i o n , 87—89 Consul fabias, p l . 3 f Cook, O . F., 4 4 2 Copaxa, 3 0 0 - 3 0 1 ; cydippe, 3 0 0 ; decrescens, 3 0 0 ; lavendera, 299; moinieri, 3 0 0 Copestylum, 60 Copiopteiyx, 3 0 4 ; semiramis banghaasi, 3 0 1 Copiphora, 157 Coprophanaeus, 2 6 7 ; lancijer, 2 6 5 Coptocycla arcuata, 2 8 6 Coplotermes, 9 5 , 9 9 ; havilandi, 181; niger, 181 Cordia, 2 8 7 , 4 3 5 , p l . 3 c Coquillettidea, 3 7 5 , 382 Corduleceris, 1 9 1 ; maclachlani, 189 C o r d u l i i d a e , 198 Cordyceps, 7 5 , 4 3 8 ; auslralis, 4 3 8 Cordylochernes, 2 8 3 ; scorpioides, 140, 2 8 3 C o r e i d a e , 1, 2 1 7 , 2 1 8 Consella, 2 3 0 Conum: solará, 2 4 4 ; viatorum, 244 Corixidae, 58, 6 1 , 2 2 8 , 230 C o r n , 9 2 , 2 7 1 , 2 7 4 , 3 1 9 , 331 Corn e a r w o r m , 92, 320, 321 Cornilermes cumula ns, 180 Cornops aquaticum, 162 Correbia, 2 7 6 , 3 1 3 ; lycoides, 2 7 6 Correbidia assimilis, 3 1 3 Corvanthes, 8 4 - 8 5 C o r y d a l i d a e . 5, 2 0 1 - 2 0 2 Corydalus: armatus, 2 0 1 ; cornutus, 5,201

Corythucha gossypii, 220, 222 C o s m e t i d a e , 126 Cosmopolites sórdidas, 92 Cossidac, 46, 72, 3 3 3 - 3 3 4 Costas, 4 3 6 Cotinus, 2 7 2 ; mutabilis, 271 C o t t o n , 9 2 , 101, 222. 2 4 4 , 3 2 0 , 442 C o t t o n b o l l w o r m , 9 2 . See also Helicoverpa zea C o t t o n lace b u g , 2 2 0 , 222 C o t t o n leaf w o r m , 12, 3 2 0 - 3 2 1 Cowdria ruminantium, 136 Coya, 4 3 Cratomorphus, 170, 2 6 1 , 312 Cratoplastis dilata, 2 9 3 , 3 1 2 , 326 Crematogaster, 4 3 5 , 4 3 6 , 4 5 1 , 4 5 2 : límala, 4 4 2 ; slolli, 4 5 0 Cressoniella, 457 Cricket: h o u s e , 159; I n d i a n h o u s e , 159. See also Gryllidae Crotalana, 2 9 3 , 312 Crotón, 2 9 9 C r u s t a c e a , 37, 38, 1 0 9 - 1 1 1 . 3 8 2 Crypliocncos, 58 Ciyptocerus, 449 Cryptomimas, 256 Ciyptoses choloepi. 7 3 , 329, 3 3 0 331 C r y p t o s t i g m a t a , 54, 127 Ciyptotermes brevis. 181 C t e m d a e , 9 5 , 113, 116, 1 1 8 - 1 1 9 Clenocephalides: cams, 2 1 3 ; jells, 213 Clenocyrlinus prodigas, 149 Ctenolepisma longicaudala, 149, 150 Clenosoma, 249 C t e n u c h i n a e , 79, 2 7 6 , 2 9 2 , 2 9 3 , 296,311,313-314,417 Cieñas, 119 C u c u j i d a e , 99 Cucurbitaceae, 218, 353 C u c u y o , 4, 2 5 6 , 2 5 9 - 2 6 0 . See also Pyrophorus C u l e x , 60, 3 7 4 , 3 7 6 , 3 8 1 , 382; bahamensis, 3 7 4 ; opislhopus, 374; quinquejascialus, 376, 381 Culicidae, 5 8 , 5 9 - 6 0 , 76, 9 5 , 9 6 , 9 7 , 9 8 , 100, 102, 1 2 8 . 2 5 9 , 3 6 2 , 370,373-383,401 Cu/tcoides, 3 6 2 , 3 6 9 , 370; furens, 370 Culisela parliceps, 3 8 3 C u n o n i a c e a e , 264 Cuphotes, 2 5 0 , 2 5 2 ; immaca/ipes, 25.3 Curculionidae, 78, 86, 92, 95, 99,247,264,287,288-291, 433, 442 Cuterebra, 4 0 0 Cuterebridae, 96. 98. 364, 4 0 0 403

C u t w o r m : black, 3 1 8 ; g r a n u l a t e , 318; palesided, 318; variegated, 318 Cybister, 2 5 4 Cyclanthaceae, 85, 437 Cyclanthus, 85 Cyclocephala, 8 5 , pl.3h Cycloneda sanguínea, 2 7 6 Cycloptera, 155; speculala, 154 Cyclosoma mirabilis, p i . 2 b Cydia deshumana. 3 2 9 - 3 3 0 C y d n i d a e , 217 Cydosia , 3 1 7 Cyenwmys, 2 5 8 C y n i p i d a e , 72, 4 0 6 , 4 1 0 , 4 1 2 Cypherotylus, 2 5 2 ; dromedarius, 250, 2 5 3 Cyplioma, 218, 241 Daceton, 454 Dactylopidae, 235, 2 3 6 - 2 3 7 Daclylopius: coccus, 2 3 5 ; opuntiae, 2 3 6 , 2 3 7 ; tomentosas, 2 3 6 Daesiidae, 143 Dahlberg, C , 7 Dalbulus, 2 4 3 D a l c e r i d a e , 327 Dalcerina tijucana, 327 Dalechampia, 3 4 8 ; scandeus, 346 D a m o n i d a e , 139 Danainae, 292, 3 4 3 - 3 4 5 , pl.3f Danaus, 343—345; cleophi/e, 3 4 4 ; eresimus, 3 4 4 ; erippus, 3 4 4 ; gilippus, 3 4 4 ; plexippas. 8 8 , 3 4 4 Daritis howardi, 3 1 3 , 3 14 D a r w i n , C h a r l e s , 8, 164, 2 6 3 , 335 Dasyhelea, 3 7 0 Dasypsyl/us lasius, 2 1 3 Dasylhnps regalis, 187 Datura, 4 2 9 d e Acosta, | o s é , 4 d e S a h a g ú n , Fray B e r n a r d i n o , 5, d e Ulloa, A n t o n i o , 7 Deinocentes, 3 7 4 , 3 7 6 , 3 8 1 , 3 8 2 ; cancer, 381 Demodex: boiñs, \'5l; canis, 1 3 1 ; caprae, 1 3 1 ; cali, 1 3 1 ; equi, 1 3 1 ; folliculorum, 129, 1 3 1 - 1 3 2 ; mis, Ví\; phylloides, 131 D e m o d i c i d a e , 7 1 , 129, 1 3 1 - 1 3 2 D e n g u e , 9 7 , 362 D e r b i d a e , 237 Dermacenlor, 135; nitem, 132, 135 D e r m a n y s s i d a e , 127 Dermanyssus, 127 D e r m a p t e r a , 37, 39. 6 0 , 1 7 8 179 Dermatobia hominis, 96, 9 8 , 4 0 1 403 Dermatophagoides: jarinae, 1 2 9 130; neotropicalis, 129; pteronyssinus, 129

INDEX

525

Dermestes, 99 Dermestidae, 72, 99 Dermoglyphidae, 128 Desantisca, 124 Desmodium, 320 Dexosarcophaga, 397 Diabrotica, 285; undecempanetata, 285 Diactor bitineatus, 217, 218 Diaethna, 347-348; asíala, 348; clymena, 348 Dia/eurodes, 92 Diamphipnoidae, 201 Diapause, 22 Diaspididae, 92, 93, 235 Diastatops, 198; dimidiata, 196 Diatraea, 92, 93, 103; centrella, 332; considérala, 332; grandiosella, 332; magnifactella, 332; saccharalis, 93, 331-332 Diaz, Bernal, 4 Dichaelacera, 385 Dichotomius, 266; carolinus, 265, 267 Didelphis, 258 Dieffenbachia, 85 Dimares, 189 Dimecoenia, 61 Diueutus, 255 Dinoponera, 45, 437-438, 439; australis, 438; gigantea, 438— 439, pl.3a Dioploeciton, 256 Dioptidae, 293,311,313, 315 £>!0/;fe restricta, 313, 315 Diplopoda, 23, 38, 56, 59, 78, 82,95, 144-146,433 Diplura, 38 Diptera, 38, 39, 57, 60, 62, 72, 76, 113,360-403,409,412, 415,433,460 Dipthera festiva, 322, 324 Dirofilaria irnitis, 96, 98 DiV^m, 298, 303; «wa, 301. 303 Dismorphia amphiona, pl.3f Dispersal, 19, 6 3 - 6 5 , 70, 83, 335, 390. Sff a/io Migration Diversity, 68, 7 4 - 7 5 , 204, 246, 294, 334, 406. 408 Dolichoderinae, 433, 4 5 0 - 4 5 1 , 454 Dolichoderus, 436 Dolichopsyllidae, 213 Doringia acridiorum, 397 Doru lineare, 178 Dorylus, 442 Dragon-headed bug, ii, 1,6,7, 238, 239-240, pl.le Drake, Sir Francis, 378 Drepanicus gayi, 189 Drosera, 76 Droseraceae, 76 Drosophila, 30, 388-390;

526

INDEX

carcinophila, 389; endobranclua, 389; melanogasler, 106, 388, 389 Drosophilidae, 30, 60, 106, 3 8 8 390 Dryasiulia, 351, 354 Drymopsocus, 187 Dryopidae, 58 Durango scorpion, 141-142 Dynamine, 348 Dynastor darius, 358-359 Dynastes, 21, 78, 246, 268-269; hércules, 30, 246, 268, 269; hyllus, 269; neptunus, 269; mtona.s, 269 Dysdaemonia, 304Dysdercus, 12 Dysmicoccus, 91; brevipes, 92, 235 Dysodius lunatus, 220, 221 Dysonia Juscifrons, 156 Dysschema: jansoni, 314; leucophaea, 313 Dytiscidae, 253, 254-255 Eac/es, 298-299; imperialis decoris, 299 Earwig: European, 179; lined, 178; maritime, 179; ringlegged, 179; shore, 178, 179. S«" tf/vo Dermaptcra Echidnophaga gallinácea, 213 £rtócs, 309 Eaton, 257, 336, 442-445; bvrchelli, 443, 444; hamatum, 443, 444, pl.2h Ecitoninae, 433, 442-445 Eatophya, 256, 257 Ecpanlhena, 312 Ectatomma luberculatum, 439, 441-442, 453, pl.3b Edessa, pi. Id Education, 470-474, 496-497 Eichornia, 382 Elaeis guineensis, 86, 289, 316, 327 Elateridae, 4, 6, 78, 239, 256, 258-260,277,293,312 £7¿W/o polyzona, 358, 359 Elipsocidae, 187 Elizabeth, Queen (of England), 356 Elmidae, 58 Elnoria noyesi, 105-106 F.lyphonidae, 137-138 Elytropeltus apicalis, 277 Embiidina, 39, 184-186 Empicoris rubromaculalus, 224 Empoasca, 243; fabae, 243; kraemeri, 243 Enceladus gigas, 248 Encephalitises, 97, 362. 376 Encyrtidae, 103 Endobrachys revocans, 293, 326 Enema pan, 274

Eneoplera surinamensis, 159 Engytalus, 221 Enterolobium, 283, 287 Enlimus, 290-291; imperialis, 288; nobilis,29i Entomobryidae, 149 Entomology: literature, 4 7 8 486; museums and collections, 474-477; research, 486-497; schools, 470-474 Fnyaliodes, 239 Eotetranychus sexmaculatus, 92 Epeorus, 58 Ephemcroptcra, 37, 38, 58, 193 194-196 Ephestia, 99, 332 Ephydra, 61 Ephydridae, 58, 61 Epüachna, 276; paenulata, 276; Iredeamnotala, 276; varivestis, 276 Epilampra, 54, 170, 171 Epilampridae, 54, 170, 171 Fpiphyllum, 436 Eplophorus velutinus, 452 Erebus odoru, 323 Eremobates, 141, 143 Eremobatidae, 141, 143 Eremopedes colonialis, 154 Eresia plullyra, pl.3f Ericaceae, 328 Eriococcidae, 235 Eriophyes: guerreronis, 93, 129; sheldom, 129 Eriophyidae, 72, 93, 127, 129, 130 Enstalis tenax, 386-387 Erotylidae, 170,250,253 Erotylus, 170, 250, 253 Ermnis, 306-307; f //o, 92, 306, 307 Erythnna, 167,235,301 Etiborellia annuhpes, 179 Eucalyptus, 94 Eucereon, 312, 313 F.ucheira socialis, 328 Euchroma gigantea, 274, 275, pl.lh Eucyphonia, 241 Eufriesea, 460 Eugeropteron, 39 Euglossa, 460-461; purpurea, 459 Euglyphis cribraria, 325 Eulaema, 460-461; meriana, 459, 460 Eulophidae, 410 Eumastacidae, 163, pl.lc Eumaslax, 163, pl.lc Eumenidae, 420-423 Eumenes, 420—421; consobnnus, 420, 421 Eumorpha, 305, 309; fasciata, 308, 309; labruscae, 309

Eupatorinm. 293, 343 Eupelmidae, 410 Euphorbia, 312 Euphorbiaceae. 269, 299, 307, 310,312,329, 346, 348, 356 Eurhinocricis, 56 Eurycoleus, 247, 253 Eurysternus, 265; deplanatus, 265 Eurytides, 338; bellerophon, 338; philolaus, 338 Eurytomidae. 124, 410, 411 E use his tus, 219 Eustheniidae, 201 Eustala anistera, 119, 120 Euthyrrhaphidae, 170 Futrombicula, 133—134; «/freddugesi group, 134; batatas, 132, 134 Euxesla, 93 Euxoa, 317 Evolution, 3 6 - 3 8 Exaerete, 461 Extinction, 63 Extrafloral nectaries, 340, 353, 434, 436. 439, 442, pl.3b Fabaceae. Sec Caesalpiniaceae; Mimosaceae; Papilionaceae Fagaceae, 158, 238, 264, 273, 301,413 Fabricius, J. C , 7 Fannia, 393; camcularis, 393, 395 Fawcett, Col. P. H., 362-363 Felicolafelis, 208 Fftoo, 318 Ferruginous skimmer. 198 Rfw, 84, 220, 283,284, 301, 306,410-411 fVrfraa, 383 Fidicina: chlorogena, 233; mamfera, 233 Filariasis, 96, 362, 376, 382 Finlay, Carlos. 11, 380 Fish, 103, 254, 256, 353, 366 Flacourtiaceae, 350 Flatidae, 237 Flea: burrowing, 213, 214-215; cat, 213; dog, 213; human, 213; mouse, 213; northern rat, 213; Oriental rat, 213; sticktight, 213. See also Siphonaptera Fly: Amazon, 12, 103, 331-332, 361; bathroom, 365, 367; black blowfly, 397; black garbage, 396; cassava shoot, 92; Cuban, 361; drone, 386-387; green blowfly, 396, 397; green flower, 387; green house, 395; greenbottle, 397; horn, 98, 362, 393, 395-396; house, 393, 394; lesser house, 393, 394; pigeon louse, 404; stable, 98, 362, 393, 395,401; wasp, 387, 388

Flying mouse, 237—238 Food: insects as food for hu­ mans, 46, 106, 147, 160-161, 203,219.229,230, 247,264. 272, 275, 287, 289, 333, 447; re­ lations of insects, 71-74 Forcipomyia, 19, 86, 169, 370 Forensic entomology, 72. 397 Forest entomology. 90, 94-95, 106 Forfícula aiiricu/aria, 179 Forhculidae, 178, 179 Formicidae, 44. 45, 54, 57, 59, 70, 71, 73, 7 6 , 8 1 , 9 1 , 9 5 , 9 9 , 100, 183,208, 236, 336,352. 406. 431-455,482, pl.2h, pl.3c; as biological control agents, 136: fossil, 40; as host or prev, 397, 409; as mimetic model! 79, 113, 116, 117, 177, 188, 217, 279, 280, 392; in sym­ biosis. 8 2 - 8 3 , 171,232,234, 241, 244-245, 335, 340, 341, 353, 423, pl.3b, pl.3c. pl.4g Formicinae. 208, 433, 451-452 Fossil insects, 3 9 - 4 1 , 407 Frankliniella, 188; párvula, 187; tritici, 93 Franklinothnps vespiformis, 188 Fruit flv: Caribbean, 390; Medi­ terranean, 92, 102, 389, 390; Mexican, 102, 390; papaya, 390; South American, 390 Fucellia, 58, 392-393; marítima, 393 Fucus, 392 Fulgora, ii, 1. 6, 7, 237, 238, 239-240; ¡alemana, 238, 2 3 9 240, pl.le Fulgoridae, ii, 1, 6, 7, 232. 2 3 7 240, 482, pl.le Fulgoroidea, 237-240 Fungi, 75, 83, 92, 93, 263, 278. 283. 438, 447 Gagrellidae, 123, 125 Galápagos Islands, 64, 128, 165, 231,372,380,422,441,446, 459, 484, 488 Galindomyia leei, 382 Gallería mellonella, 71, 333 Galls, 72.92,244, 410,412 Gamella, 301 Gardner, G., 6 Gasteracanlha, 122, 200; canenformis. 120, 122; letraeanlha, 122 Gasterophilidae, 98, 364, 398, 400-401 Gasterophilus, 98, 400; haemorrhoidalis, 400; intestinalis, 398, 400; nasalis, 400 Gastropoda, 397

Gay, C , 8 Geíechiidae, 92, 93, 99, 102, 332 Genetics, 3 0 - 3 1 , 103, 106, 389, 487 Geochelone, 181 Geocoris punctipes, 220 Geography, 48-52 Geometridae, 311, 321, 325-326 Geomydoecus, 208 Geophilomorpha, 147 Geropteron, 39 Gerridae, 57, 228, 230-231 Gems remigus, 228, 231 Gesner, K., 6 Gesneriaceae, 436 Gillisius, 60 Glaphyrocanthtm, 265 Glenurus, 189, 190, \9\; pecu­ lio ris. 189 G/iricola porcelli, 208 Globetrotter, 196, 198 Glossina, 362 Gloveria psidii, 325, 328 Gmelina arbórea, 94 Godman, F.. 7 Goeldi, E., 10 Golofa, 246, 270-272; aegeon, 271: eacus. 271: porten. 271 Gomphomacromia chilensis, 198 Gonyleptidae, 123, 125-126 Gonyleptus janthinus, 123 Gorgas, William, 11 Graphocaecilius, 1 S() Graptocleptes, 223 Greta, 351 Grew. Nehemiah, 7, 239 Gripopterygidae, 58, 201 Ground pearls, 233, 234 Growth and development, 22. 34-36 Grumicha, 203 Grumichella, 203 Gryllidae, 56, 158-160, 433 Grylloides supplicans, 159 Grylloptera, 37. 39. 153-161 Gryllotalpidae, 159, 160-161 Gtyllus, 159; assimilis, 159 Guenther, Konrad, 6 Gymnetis, 272; holocericea circumdata, 271 Gymnocladus dioica, 323 Gynerium, 289 Gyretes, 254, 256 Gyrinidac, 253, 254, 255-256 Gynnus, 254, 256 Gyriosomus, 250, 251—252 Gyropidae, 208 Gyropus ova/is, 208 Haemagogus, 376, 380-381 Haemaphysa/is, 135 Haemalobia irntans, 98, 362, 393, 395-396

INDEX

527

Hacmatopinidae. 209, 210 Haematopinus suis, 2 0 9 Haematosiphon inodorus, 226 H a l a c a r i d a e , 58 H a l a r a c h n i d a e , 127 Halictidae, 4 5 6 , 4 5 7 Halobates. 2 3 1 ; micans. 228, 2 3 1 ; robustus, 231 Halysidola, 3 1 2 Hamadryas, 345—'A46;feronia, 3 4 4 Hapalopus, 115 Haptoncus, 86 Hamsina tergina, 3 1 3 Hawks, H „ l l H e a r t w o r m , 96, 98 Hedycluum, 357 Heliamphora, 5 9 , 76, 3 7 5 Helicoma, 6 0 , 115, 2 8 6 , 2 8 7 , 3 5 7 , 371,375,383,389 H e l i c o n i a c c a c , 60, 115, 2 8 6 , 2 8 7 , 357,371,375,383,389 Heliconiinae, 347, 351, 3 5 2 - 3 5 5 , pl.3f Helicomus, 352—355; charitonius, 354; erato, 3 5 4 ; ismenius, pl.3f; melanops,354 H e l i c o p h i d a e , 204 Helicopis, 3 3 9 ; acts, 341 Helicopsyche, 2 0 3 Helicopsychidae, 203 Helicoverpa, 9 2 ; zea, 92, 3 2 0 , 3 2 1 H c l i o c h a r i t i d a e , 199 Heliuthis, 3 2 0 Heliothrips haemorrhoidalis, 185, 188 Heliotropiam, 3 1 2 . 3 1 3 , 3 4 3 , 3 5 0 ; fedegoso, 2 9 3 Helodidae, 59 H e m e r o b i i d a e . 189 Hemeroplanes, 1, 3 0 8 - 3 0 9 ; ornatus, 3 0 8 , p l . 3 e Hernikyptha, 241 H e m i p t e r a , 38, 39, 58, 2 1 6 - 2 4 5 H c p t a g e n i i d a e , 58 Hercinothrips bicinclus, 92 Hermetic, illuscens, 3 8 7 , 3 8 8 H e r n á n d e z , F. G., 5, 11 Herrania, 85 Herse angulatus, 3 0 6 H e s p e r i i d a e , 7 7 , 3 5 8 , 359 Hesperoctenes, 2 2 4 , 2 2 6 Helaerina, 199; americana, 199 Heteronotusflavomaculalus, 241 Heterophrynus longicurnus, 138, 139 Heteropoda venatoria, 1 2 2 - 1 2 3 Heteropodidae, 122-123 H e t e r o p t e r a , 39, 2 1 6 - 2 3 1 Heterosternus, 274 H e x a p o d a , 3 7 , 3 8 , 148 Hibiscus, 301 H i n g s t o n , R., 6 Hippelates,36'A, 391

528

INDEX

Hippobosca equina, 4 0 3 Hippoboscidac, 98, 402, 4 0 3 404 Hiranetix, 2 2 3 Histeridae, 444 Historis odius, 3 4 8 , 350 History, of Latin A m e r i t a n Ento­ mology, 3 - 1 3 Holocampsa, 170 H o m o p t e r a . 57, 70, 78, 8 3 , 216, 231-245,433,453,454 Honey, 463, 464, 466 Hoplocrales, 4 1 5 Hop/omutilla, 4 1 5 ; xanlhucerala. 415 Hoplopleura, 210 Hoplopleuridae, 210 H o r m o n e s , 24, 102 Hudsonema, 2 0 4 Hyale, 111 Hyalymenus, 2 1 7 Hydrobiosidae, 58, 204 H y d r o m e t r i d a e , 57 H y d r o p h i l i d a c , 60, 254, 2 5 5 Hydroplulus, 254, 2 5 5 ; insulans, 254 Hydropsychidae. 204 H y d v o p t i l i d a e , 203 Hyleplula phyleus, 359 Hylesia, 2 9 8 , 3 0 1 , 3 0 2 - 3 0 3 ; canina, 3 0 2 ; linéala, 3 0 1 , 3 0 3 ; melabas, 302 Hymenaea, 40, 2 3 9 ; courbanl, 303 Hymenitis, 351 Hymenolepis, 99 H y m e n o p t e r a , 3 8 , 3 9 , 74, 78, 8 1 , 9 5 , 154, 2 8 0 , 2 9 6 , 3 1 3 , 3 9 7 , 405-469 Hyperchina, 301 Hypoaspis dasypus, 127 Hypocephalus armalus, 279, 284 Hypoclinea, 3 4 1 , 454 Hypercompe, 3 1 2 ; decora, 3 1 0 H y p o c t o n i d a e , 138 Hypoderma: bovis, 4 0 0 ; lineatum, 398, 4 0 0 H v p o d e r m a t i d a e , 98, 398, 4 0 0 401 Hypoleria, 3 5 0 ; andromica, 351 Hypotlienemus hampei, 9 1 , 276, 278 Hypothyris, 351 / m - w purchasi, 2 9 , 2 3 5 , 276 l c h n e u m o n i d a c , 223, 277, 407, 408-409 Ma/iM, 3 1 2 ; /¡croi'j. 3 1 0 Idarues, 411 Identification, 4 9 5 - 4 9 6 Iguanacarus, 128 Illustration, 4 9 4 - 4 9 5 ¡ncisitennes snyderi, 181

I n d e p e n d e n c e g r a s s h o p p e r , 165, 166 /»#«, 2 6 9 , 3 0 2 , 3 1 0 , 3 2 3 , 3 4 2 , 4 3 6 , pl.3b Iphiaulax, 4 0 8 Iridomyrmex, 4 5 4 ; humilis, 4 5 3 . 454 ' l s c h n o p s y l l i d a e . 213 Isognalhiis, 3 0 5 l s o p o d a , 3 7 , 38, 109, 110 l s o p t e r a , 3 9 , 56, 5 9 , 7 3 , 8 1 , 8 3 , 100, 109. 1 8 0 - 1 8 4 , 187, 397 I t h o m i i n a e , 292. 2 9 3 , 3 1 4 , 3 5 0 352, pl.3f Itzpapálotl, 4 5 , 300 Ixodes, 135; pararicinus, 136 l x o d i d a e , 132, 1 3 4 - 1 3 7 Jacaranda, 2 9 9 Jamadia gne/us. 359 Jalropha, 299 jessenia weberbauri, 281 '/own, 315 V«rf