Costa Rican Natural History 0226393321, 0226393348

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Citation preview

COSTA RICAN NATURAL HISTORY EDITED BY

Daniel H. Janzen WITH

174 CONTRIBUTORS

THE UNIVERSITY OF CHICAGO PRESS CHICAGO AND LONDON

ร

at the University of PennsylH JANZEN is professor of biology Nacional de Costa Rica. Museo the of member associate vania and an

CONTENTS

DANIEL

Preface ix TUB UNIVERSITY OF CHICAGO PRESS. CHICAGO 60637 THE UNIVERSITY OF CHICAGO PRESS LTD LONDON © 1983 by The University of Chicago All rights reserved . Published 1983 Printed in the United States of America

Acknowledgments xi

.

.

1 Searchers on That Rich Coast: Costa Rican Field Biology, 14๓- 1980 1

5432 1

90 89 88 87 86 85 84 83

2 The Central American Dispersal Route: Biotic History and Palaeogeography 12 3 Climate 35

10 13

12 16

. Identification of cox’er illustrations ( all photographs by Daniel H ) Janzen 1 Female Rothschildia lebeau ( saturniid moth ) in copulo Santa

. Rosa National Park 2. Last - instar caterpillar of Rothschildia Santa Rosa National Park .

8 Reptiles and Amphibians 351

.

.

erycina

( saturniid

moth ).

Pentaclethra macroloba ( legume tree ) 3. Dehisced frail and seeds ofSelva . La Finca . on forest floor Park . 4. Adult Felis wiedii ( margay ). Santa Rosa National ( speckled racer ) swallowing 5. Adult Drymobius margaritiferusNational Park . adult Rana ( frog ). Santa Rosa palm ) , with leaves 6 to 8 meters 6. Subadult Welfia georgii ( welflasecondary succession . Finca La long , growing through late Selva .

sloth ). La Lola . 7. Adult Bradypus variegatus ( three(-toed pygmy ). Limon . 8. Adult Glauadium minulissimum leastsapiemumowl( banana ). Mot 9. Infructescence/inflorescence of Musa te verde . 10. Last -instar caterpillar of Automeris rubrescens ( io moth ). San Rosa National Park . ). Finca La Selv 11. Adult Citherias menander (satyrid butterfly swa ) pterocarpus in tree legume ( officinalis Pterocarpus Adult . 12 . Park National Corcovado Rosa National Pal 13. Adult Ganoderma ( bracket fungus ). Santa 14 . Adult female Odocoileus virgimanus ( white - tailed deer ). Sal Rosa National Park . ) with crab ร 15. Inflorescence of Asclepias curassavicaant( milkweed der that has just seized queen Azieca as prey. Palo Verde.

4 Geology 47

Introduction 351 Checklist of Reptiles and Amphibians 367

Species Accounts 374

9 Mammals 426 Introduction 426 Checklist of Mammals 443

Species Accounts 448

10 Birds 502 Introduction 502 Checklist of Birds 530 Species Accounts 544

11 Insects 619 Introduction 619 Checklists of Insects 645 Species Accounts 678

5 Soils 63

6 Agriculture 66 Introduction 66 Species Accounts 73

Addresses of Contributors 781 Index 789

7 Plants 118 Introduction 118 Checklist of Trees 158

Species Accounts

184

Library of Congress Cataloging in Publication Data Main entry under title:

Costa Rican natural history.

Includes bibliographical references and index . 1 Natural history — Costa Rica . I . Janzen ,

.

Daniel H. 508.7286 QHI 08.C6C67 1983 ) ( cloth 393321 226 ISBN 0ISBN 0 -226 393348 ( paper )

82-17625

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vii

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but March. ๖ ฺ8๑ problem continues. BaiTett , o. 1935. Notes concerning manatees and ด 16:216-20. dugongs. 8ด Frantzius, A . von. 1869. Die saugethiere Costaricas. ด ( Berlin ) 35:247-325. ด ๑ ไ ์ Hartman , D. 1979. ิ ้ Publica ด Special ๖ ๑ tion no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee ้

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals ิ ด ด ด separated from their group , for example , for foraging , ด ด ด are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ไ such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds , feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufvampire. One reported that individuals of fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ิdo not fan their seems a distinct possibility. It is clear that these bats have ฺand ฺ pires ้ complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees , with the exception of ฺwhich ๖may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIESACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. 1% 1. A review for fear that some small museum will feel the need bats of Trinidad and Tobago. M. ด ด . ด complete its skin collection at the expense of the remnantto of the 122 : 191 301 . populations. Fieldworkers are encouraged to observe and Greenhall , A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. 8ด ด49:337-40. of the and collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1%9. Notes on ไ ฺthe false 8ด ด47:140 15. vampire bat , in Panama. ด , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , and of the Neotropical carnivorous ฺin Costa Rica. 8 ไ ด ด44:186-89. ด 8ด ด58:469 78. bat ,

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501

PREFACE

book . Special thanks are due Peter H . Raven and Donald E. Stone for their strong encouragement and their work to secure contributions to support publication . My deepest indebtedness in the production of this book is to those students in past OTS courses who have wanted to know about this or that organism , and to all those who have cared enough to learn something rather than merely be entertained by Costa Rican natural history. I could never have undertaken this project without the certain knowledge that people out there would contribute to it. I owe a special acknowledgment to the writers of chapters 1 - 5 and later chapter introductions, and to Luis Gomez, who not only volunteered many pages of writing but found more contributors. Most important of all , he served as counsel during the tortuous journey through the offices of three publishing houses that backed out on their promises, resulting in a full two years delay in the publication date. The authors of the chapter on agri cultural species planned and contributed that chapter in dependent of my efforts. พ. Hallwachs, G . Stevens, M. Johnston , and L . Gomez served as volunteer editorial assistants. D. Gill , G. Stevens, R . Hallwachs, M . Hall*

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wachs , and R. Foster carried edited manuscript and proofs to me in Costa Rica to help ensure safe delivery and avoid loss of time in the mails. The secretarial staff of the Department of Biology at the University of Penn sylvania helped enormously. Traditionally the University of Chicago Press does not allow its staff to be thanked by name in such acknowl edgments as this, but I thank the Press for its great interest in this book and for extraordinary efforts on its behalf . In a manner difficult to define , the presence of the Servicio de Parques Nacionales de Costa Rica , an organization that frequently goes far out of its way to support Costa Rican field biology studies, served as a major inspiration to me to organize and carry out this task. Likewise , the Museo Nacional de Costa Rica provided a logistical and psychological home for the project far beyond the call of duty. Finally, I must recognize that it was Jay Savage who first brought me to Costa Rica in 1963, and that the National Science Foundation paid for it .

-

D. H . Janzen

A list of the contributors and their mailing addresses at the time of publication begins on page 781.

ACKNOWLEDGMENTS Further research in tropical biology is essential if we are to understand and appreciate the biological complexity of the tropics and contribute to their preservation and wise use as well as to the preservation of the biological di versity of life on earth. Thanks are due the following individuals and institutions , whose commitment to the importance of this work is manifest in their generous support of the publication of this book: The Animal Research and Conservation Center of the New York Zoological Society The Association for Tropical Biology George R . Cooley Stanley R . and Lynn พ. Day Fundacion de Parques Nacionales de Costa Rica The Hershey Fund Philip and Margaret Hess Lewis F. Kibler Marion Lloyd Mr. and Mrs . J. Daniel Mitchell Museo Nacional de Costa Rica The Organization for Tropical Studies E. Lisa Smith The Smithsonian Tropical Research Institute Catherine H . Sweeney Typoserv ice Corporation Universidad de Costa Rica The World Wildlife Fund United States

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XI

MAMMALS

SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich roosts are now kept s ecret , G. G., andGreenhall Goodwin . 8 March . continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . . Theyappearto be highlysocial cluster . complete itsskincollection attheexpense oftheremnant :191301. 122 20. 16:216 . dugongs forlongperiods stable mayremain group c omposition . . populations Fieldworkers areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe intheroost disturbed enjoytheselargebats , butunnecessary false v ampire bat. . ) 35:247 325 (Berlin disturbance : 49 and 337 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . whenever , p. 1%9. Notes Hartman possible . on forthe presence marker a good andconstitutes cies of Publica the Special falsevampire bat, in Panama . t ree groups Captive . h ollow in given a have 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup separated on the foraging behavior phyllostomid and , avian bats p rey and of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. carnivorous bat, inCosta Rica . 44:186 -89. . A rapidhigh inwolves . ; ofmouths manatee muchlikethatfound 58:46978. , J. E. 1979 . Thesemisocial Reynolds , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group Vampyrum spectrum ( Vampiro Falso, times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear 8ด , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld isthelargest that , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects feedoninsects taken match thebats atnight in groups . Someofthespecies mixlarger and forms suchas thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects , feedeverynight , thata foraging cansome suchasbirds favors substantial vertebrates arestrik timesfillitsstomach within the anhourafterleaving , andother . Thejawsof bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking in theflooroftheroost insuf of : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning . Itisclearthatthesebatshave donotfantheir seems adistinct possibility and pires with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian roosts ingroups . carnivores suchascanids oftheAmerican occurs overa widerange ofhollow batspe other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably tropics of ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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SEARCHERS ON THAT RICH COAST

hardly allowed for the scant and elementary education of her subjects , while the iron hand of the church treated intellectual endeavors as works of the devil , fatal to the souls of her charges particularly if these endeavors were contaminated by the ideas of the French Revolution and other demoniacal inspirations , such as Freemasonry. The condition of public education in Costa Rica during the colonial period is well documented by Gonzalez F. ( 1978) , and the capital importance of French liberalism in the movement for independence is presented by Lascaris ( 1964 ). Debilitated from its very inception ( Gonzalez F. 1978; Gonzalez V. 1977; Soley 1940 ) , the economy of the area could barely sustain the first twenty -odd years of republican life. But then , despite the fashionable efforts of the Duchess of Bedford and her cup of tea , England settled for a cup of coffee. And quite

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unexpectedly, too. In 1843 William Le Lacheur, an English merchant , docked the Monarch in the port of Caldera on the Pacific coast of Costa Rica . The holds of the brig were almost empty for his homeward voyage , and Le Lacheur wished to obtain some cargo that would compensate him for the long and hazardous trip around Cape Horn . He rode a mule into San Jose, where he met Santiago Fernandez , dealer in coffee. Fernandez trusted Le Lacheur, and on his credit Le Lacheur departed carrying 5,505 hundredweight of Costa Rican coffee. He returned in 1845 to pay Fernandez , and this time he brought more vessels and a good supply of sterling . A new industry had been bom . Ships arriving to load the coffee brought holds full of goods , including Manchester cotton fabrics , which were to open entire con tinents to British imperialism . And , of course , they brought people. The cash flow turned a dilapidated region into a prosperous one , changing many things as the criollos became able to barter for cultural goods as well . Coflfee was responsible for the initiation of diplomatic relations between Costa Rica and England , France, and the Hansa in 1849 and for the expansion of agricultural areas and many of the subsequent movements of the population , as well as for the opening of access roads and railways. Coflfee was perhaps also responsible for the first officially recognized coup d 'etat in Costa Rica; J . R . Mora Fernandez and R . Aguilar, partners in a coflfee export firm , parted on bad terms owing to a financial disagreement . Later, Mora was given pecuniary satisfaction and eventually became president of Costa Rica, but he was deposed by a vengeful Aguilar in 1859. The importance of coflfee in the development of the history and geography of Costa Rica is documented by Hall ( 1978 ) and Estrada ( 1965). The TVaveler- Naturalists

Why did people come to Costa Rica aboard the growing coffee fleet ? A look at the history of continental Europe

in the last third of the nineteenth century would provide many of the answers , but even a brief overview lies outside the scope of this chapter. Immigration of Euro I peans to Latin America in general , and Central America in particular, was primarily a response to the socio political situation of Europe and the apparent Utopian conditions of the New World . The relative stability of Central American politics , coupled with niches for the crafts and professions in the flourishing regional econ omy, enticed foreign entrepreneurs and scholars. Cities that had lacked elementary schools suddenly had private “ academies" catering to a clientele eager to leam , be it bookkeeping or fine arts. The streets resounded with for eign names as the general stores gave way to specialized shops and cottage manufactures yielded to incipient in dustries. This birth of cosmopolitanism is both the cause and the effect of immigration. The Gold Rush ot 18481 also fueled the demographic explosion in Central Amer ica; the way to California was either around Cape Horn , a lengthy and hazardous enterprise , or by the much faster route across southern Nicaragua ( Houwaldt 1975; Folk mann 1972) , along the present border between Nicaragual and Costa Rica . come? Certainly But why did the traveler- naturalists ' ' and pol nor economy s findings area the neither Sutter s this presence part ofI were their for explanation in itics the , , Dondamine of La Humboldt accounts world The . the propagandizes good but equally illustrious other less and provide the reason . The romance of travel , exploration , and discovery, triggered by European expansionism during the seventeenth century, was rampant in the Europe of the Industrial Revolution , and such adventurous ob servers were far from immune to the call of the vast and unknown American continent . A glance at the titles of some then - popular books is illuminating: Billow, Aus wanderung นrid Colonisation in Interesse des deutschen Aussenhandels ( 1849); Bard , Waika; on Adventures on the Mosquito Shore ( 1855); Scherzer, Wanderungen durch die mittelamerikanischen Freistaaten ( 1857 ); Marr, Reise nach Zentral Amerika ( 1863); Boyle , A Ride across a Continent: Personal Narrative of Wanderings through Nic aragua and Costa Rica ( 1868 ); Pirn , Dottings on the Roadsides ctf Panama Nicaragua . . . ( 1868 ); Wagner, Natunx'issenschqftliche Reise in Tropical America ( 1870 ); and many others (cf . Fernandez 1972 ). We cannot boast of personalities of the stature of Alex ander von Humboldt , who scarcely saw the high volcanoes of Central America from his ship bound for Mexico but , unlike that meteor of the Parisian salons , those who came usually stayed , sowing the seed of interest in natu ral history. To this period belong several importanl names: the already -mentioned Moritz Wagner, who in association with Karl Scherzer published in 1886 Die Repuhlik Costa Rica: the Danish Anders Sandoe Orsted, who visited Costa Rica from 1846 to 1848 , publishing his

famous opus LAmerique Centrale: Recherches sur sa , printed in Co penhagen under the auspices of the Costa Rican govern ment . A somewhat unknown figure who deserves credit for his active part in advancing local natural history is William More Gabb, whose endeavors in the geology, paleontology, and zoology of the lower Talamancas ( Ferrero 1978) were crowned by later publications by E. D. Cope, L . Pilsbry, and J . A . Allen , among others. This is the period of F. Duncane Godman and Osbert Salvin , initiators of the single most comprehensive work on Central American biology, aptly entitled Biologia Centrali- Americana: it appeared between 1879 and 1915 in many volumes , with the approval of the leading au thorities on the diverse groups of fauna and flora. Karl Sapper, geologist by profession and naturalist by avocation ( Termer 1956 ) , closes the period . His collected writings on Costa Rica have been published (Sapper 1943). geographte politique , safaune etsa flora

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The Emergence of Costa Rican Biologists

Separating this period from that covered in the previous paragraphs is highly artificial , since Costa Rican field biology stems from the causes mentioned there. In 1853, scarcely ten years after the opening of the European coffee market President Mora Fernandez welcomed two

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SEARCHERS ON THAT RICH COAST

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German physicians who carried a letter of introduction from Alexander von Humboldt Carl Hoffmann ( 1833- 59) and Alexander von Frantzius ( 1821 - 77 ) . Hoffman explored the upper portions of the central volcanic ridge ( Hoffman 1856, 1857), collected plants and animals that he sent to Berlin , and fought William Walker in Nicaragua in 1856. He died in 1857 and is commemorated in the names of a dozen species. His colleague von Frantizius was also very active; his botan ical explorations made Costa Rica known to the scholarly world , and his faunal collections were used for one of the first annotated lists on the mammals and birds , both by himself and by the famous Cabanis of Berlin. Frantzius opened a drugstore in San Jose, and his apprentice , Josd c. Zeledon , became deeply interested in nature , later becoming known worldwide as an ornithologist . Fran tzius and Zeledon 'ร drugstore was a germination chamberfrom which sprang the first batch of local naturalists such as Anastasio Alfaro and J. F. Tristan ( fig. 1.1 ). Costa Rica was oriented toward England commer dally, but culturally it was French . In the natural sciencesits practitioners were truly Victorian , and soon the “drug store gang '' and their disciples were to be seen rum -maging through the countryside with nets , pillboxes , and other paraphernalia .

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I FIGURE LI Part of the drugstore gang t the Museo Nacional . , °

* at the old quarters Left George K . Cherrie , zoologist taxidermist; center, Anastasio Alfaro, first director.

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right , in front , an assistant , possibly Francisco Mendez; far right , Jos£ Gastulo Zeledon ( photo, archives of Museo Nacional de Costa Rica ).

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Under the administration of Bernardo Soto, Mauro Fernandez , the minister of public education , achieved one of the most significant feats in Costa Rican history. He developed the new pattern of public schooling totally sponsored by the government , compulsory for all citizens until the seventh grade , and he opened high schools for both men and women . Manuel Maria Peralta , ambassador to Europe , was commissioned to hire European teachers that were to staff the Liceo de Costa Rica and the Colegio Superior de Sefioritas. This precipitated the arrival of a Swiss mining engineer, Henri Francois Pittier, accompanied by others whose names are very much part of Costa Rican history: Pablo Biolley, Julian Carmiol , Gustavo Michaud , and Juan Rudin . with Pittier ( 1857- 1950) begins the golden period of Costa Rican natural history ( Conejo 1972). Determined , indefatigable , tyrannical Pittier was soon in charge of all significant scientific activity in Costa Rica. Through his enthusiasm , Pittier promoted or founded several important institutions such as the Insti tute Fisico Geografico, the Sociedad Nacional de Agri culture , and the Observatorio Nacional . He planned and led a multidisciplinary approach to field biology, inspired by his years as a disciple of the typical nineteenth -century gymnasium . Botanists owe him the national herbarium and his Primitiae Florae Costaricensis, begun in 1891. It is the first systematic flora of the country. The cloak and dagger circumstances of the Pnmitiae are summarized by G6mez ( 1978fl ,6 ). Pittier and his associates in all branches of science , notwithstanding financial and tech nical difficulties , amassed a body of information as yet unsurpassed . Revolving around him , willingly or not , were Adolphe Tonduz , Charles WferckJ £, George Cherrie , and scores of foreign researchers who at his insistence either visited the country or studied collections sent from the Instituto Geografico or the national herbarium . Until 1904, when Pittier left the country, sciences from lim nology to social anthropology flourished. It is at this time that the Museo Nacional makes its appearance. The philosophical foundation of the Museo has been briefly reviewed by Gomez ( 1973) , who attributed its origin to the Victorian craze for bric-a- brac and the French mania for “ cabinets’’ of exhibits that seized the country when it plunged into the international marketing of coffee. In this context the figure of Anastasio Alfaro is of utmost importance, since it was through his efforts and persuasiveness, as well as the usual political intrigue , that President Bernardo Soto inaugurated the national museum in 1887. Alfaro, then twenty - two years old , became its first director. The liaison of Alfaro and Zeledon was in large part responsible for opening Costa Rica to North American scientific interest . Zeledon, the disciple of Frantzius, had been sent to Washington to learn under the tutelage of Robert Ridgway, and he pre4

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pared the Smithsonian grounds for Alfaro and others. From then on the flow of United States researchers has never stopped. A glance at the publication Costa Rica en el siglo XIX ( published in 1902 ) informs US of the dozens of scholars who contributed their knowledge to Costa Rican field biology in the last twenty years of that century. Three political events are noteworthy. ( 1 ) The gov ernment of Mora asked Monsignor Llorente y La Fuente to leave the country. The immediate result was the segre gation of church and state in educational matters. This meant the introduction of krausism and fuller curricula into the system . ( 2 ) The government of Bernardo Soto enthroned positivism in the figure of Mauro Fernandez. These two events helped to form the philosophical basis of Costa Rican republicanism. The profound con sequences of these events could still be felt as late as 1921 , one hundred years after independence, in the thoughts of Ricardo Jimenez ( 1921 ). (3) The same Mauro, Fernandez , positivist par excellence and architect of the Costa Rican educational system , was moved by political reasons to close the Universidad de Sto. Tomas in 1888. The only scholastic units to remain functioning were the school of law and the school of agriculture.

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The First Half of a Bright New Century Under the tutelage of Comte and Spencer, Costa Rica started the new century as an international merchant , whose economy was based entirely on agriculture. But ,! paradoxically, it had just an incipient and meager school of agriculture that functioned only spasmodically owing to insufficient funding ( according to an editorial in La Republica in 1890; the eventual closing of this school was due to the reallocation of its funds to the building of the national theater, a project initiated and patronized by the coffee growers’ elite ). The school of law was the only other faculty at the university in these early years. It produced the professionals necessary to regulate and run the recently bom banking system , and commercial oper ations in general , and to consolidate a legalistic republi can structure. Between 1898 and 1941 , all the prominent personalities were lawyers, leading the Nicaraguan pod Dario to describe Costa Rica as “ a land of lawyers , clerks and oxen. ” On such a scene the natural sciences , regard less of the prominence of the remaining partners of the “drugstore gang , ” were considered an amenity, and field research by local or foreign naturalists was seen as pastime for the eccentric or the wealthy, a view that still persists. A review of the educational system prompted Presi dent Iglesias to suggest reopening the Universidad de Sto. Tomas ( presidential address of 1 May 1900), but this initiative had no success. It did , however, trigger the interests of the Costa Rican intelligentsia , and during the

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next two decades some of those directly influenced by Fernandez sought a diversified college education in Chile or in Europe. Important participants in this emigration were Elias Leiva , J. F. Tristan , Jose Ma . Orozco, Emel Jimenez , Lucas R . Chacon , R . Brenes Mes£ท, and Carlos Monge , all directly associated with the renovation of the high -school curriculum , educational techniques, or, eventually, the renaissance of the university. In these early years, the slow disappearance of the old Europefla , , and their direct disciples is or Europeanized teachers ' s lack of intellectual tone in all the country in reflected but fine arts and letters. In 1917 Phillip and Amelia Calvert published A Year of Costa Rican Natural History, based on a year spent studying dragonfly biology under the auspices of the Philadelphia Academy of Sciences. Their narrative, so far un excelled , is a must for all interested in field biology as well as in the social atmosphere of Costa Rica during the first decade of this century. Meanwhile the Costa Rican economy suffered a severe blow from the European war of 1914-18 , followed by the German crisis of 1923. The magic market for coffee vanished as it had appeared , on the wings of change, propelled by the flourishing of South American com petitors as well as by the banana industry (Gaspar 1979). All this motivated an orientation toward American markets and North American pragmatism . Although the 1929 Wall Street crash had serious repercussions in the Costa Rican economy, it also had beneficial effects in that the fiscal depression provoked the appearance of several non conservative political parties ( Aguilar 1978). ( Although this story of field biology may seem to be deviating into a general history of Costa Rica , our goal is to sketch the economic and political background of field biology rather than merely listing names and citing works that can be found elsewhere [Gonzalez 1976; Dobles 1927; Gomez 1975a\) The installation of other political parties meant a plu rality of ideas and , more important , of attitudes to coun-teract the narrow concepts of the coffee oligarchy. Among these changes was the reopening of the university system . In the 1920s the lack of a university was already evident , and the inauguration of the Universidad de Costa Rica ( UCR ) by the socially oriented party in power took place without adequate preparation . The sudden need for professors allowed many of the high -school docents to participate in the early development of the university, which impaired the quality of the natural sciences curric ulum from the very start . The role of the university innatural history studies and research during the first twenty years of its existence was nil . First , being overly concerned with arts and letters , it did not offer proper cheers; second , it mismanaged the Museo Nacional ( which was cared for by the university from the mid-

1940s to 1953 ). During this period the collections were badly curated , the Iibaries pillaged , and the efforts of sixty five years of straggling dedication wasted (P. c. Standley, in litt.; J . A . Echevarria, o. Jimdnez , pers . comm .). From then until the late 1960s, the Museo Nacional will be remembered by all , including some of the early participants in courses taught by the Organization for Tropical Studies (OTS ), as “the pits. ” Still , not all was darkness . In 1913 Clodomiro Picado Twight ( Picado 1964 ) returned from Paris (Sorbonne , Institute of Colonial Medicine or Institute Pasteur) as a doctor of science ( fig . 1.2), organized the first clinical laboratory of the San Juan de Dios Hospital , and dedi cated himself to studying the natural riches of the coun try. His production includes a treatise The Poisonous Snakes of Costa Rica ( 1931 , reprinted 1976), almost a hundred articles on diverse topics related to tropical med icine and microbiology, and , most important of all , his research on the ecology of the tank bromeliads that served as his doctoral dissertation and is considered a pioneer work in tropical ecology. Some of the important works of Picado have recently been reprinted (Gomez 19756 ). Picado’s activities were in full swing when the

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FIGURE 1.2. Clodomiro Picado Twight , first Costa Rican ac ademic biologist , who graduated from the Sorbonne in 1913 ( photo. L . D. G6mez collection ).

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plurality in politics took place , and he did not neglect this opportunity to participate in the country ' ร ideologies , his influence being felt more in that field than in local biol ogy. Of his 322 published articles , 131 are on scientific matters. He was a Lamarckist ( Picado and Trejos 1942), as befits a Paris trained scientist , and most of his scientific writings were published abroad , while those issued locally were of a more popular character. His almost poetic rendering of the biographies of Pasteur and Metchnikoft ( Picado 1921 ) like many of his newspaper articles, reveals a thinker of intense humanism , ardently longing for ampler horizons. Unfortunately, Picado died in 1944 , depriving the Uni versidad de Costa Rica of his enlightment in those critical years of its inception. Although for political reasons he was excluded from the planning for that institution ( R . L. Rodriguez , pers . comm . ) , had he lived longer it is quite possible that the Universidad de Costa Rica would have had an altogether different role in the development of biological sciences in Costa Rica. The Second World War played an important role in our story, since it disrupted markets and politics , but it also had a direct influence on the advancement of technology over the pure sciences. Costa Rica plunged , again with out preparation , into the atomic age. Higher education was therefore biased toward applied sciences in the same way the economy was dominated by North American capital and markets. In 1948 the revolution of the Liberal party finally decapitated the dwindling remains of conser vative ideas (Schifter 1979; Cerdas 1972) , and Costa Rica moved into a technocracy that tended to neutralize a potent electorate with bureaucracy ( Trudeau 1971 ). The establishing of a mediocre segment of the population in government positions signified the beginning of a “scientificism " at the decision- making levels that obstructed the development of true science . This massification , which could have been prevented had the politicans read Ortega y Gasset , moved inexorably toward a Jas perian Vemunft and Widen’ernuft of scientific advance-

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ment. Nevertheless , the names of two courageous idealists deserve mention: Antonio Balli was an Italian biologist brought in as part of the General Studies Program during the renovation of the Universidad de Costa Rica in the 1950s. Balli was an eclectic philosopher of science whose teachings met with skepticism and outright deri sion , and he had no impact either on the students or on the administrative groups at the higher stages of edu cation ( Balli 1974-77). Rafael Lucas Rodriguez , well known to OTS participants , earned his doctorate from the University of California , Berkeley, and was requested to create a de partment of biology for the Universidad de Costa Rica. He was responsible for the actual curriculum of that

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school , which was designed to produce able high -school teachers in the shortest time ( R . L. Rodriguez , pers. comm.). However, paradoxically, it was biased toward zoology. Rodriguez was a devoted neo- Darwinist whose influence could have been stronger had it not met with apathy from the students . Rodriguez was intellectual fa ther to many a research project and an inspiration to several younger scientists , and he played a decisive role in establishing OTS in Costa Rica. Although Rodriguez is celebrated for his multiple contributions to the com munity, particularly concerning youth , and for his un excelled talents as a botanical illustrator, the portentous depth of his thinking was little recognized among his colleagues and students. As with Picado and Balli , the scientific output and influence of Rodriguez is better known outside Costa Rica, a symptom that conditions are not yet ripe for the development of the propitious national attitude toward field and theoretical biology appropriate to a country whose natural heritage is of such magnificence. It may be that Charles Werckle , writing to Nathaniel Lord Britton to congratulate the famous botanist on the founding of the New York Botanical Garden , has proved prophetic: “ But only if a Nation has reached the highest degree of devel opment , the interest in the Natural Sciences becomes general and the fact is understood by the most advanced that these sciences , real and original , are of a higher order than the Social Sciences. *’ Thus the first half of the twen tieth century closes under the resigned looks of Spencer and Comte , the faint shadow of Lamarck , and the scornful stance of Darwin .

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Science and the Costa Rican It is difficult tc define the historical process of cultural integration in any society. It becomes particularly painful when such integration has been simultaneous with the consolidation of the national entity as a whole. Costa Rica stands in contrast to countries where one can easily examine and analyze the origins and development of the culture of an already constituted social system. We have dealt at some length with the political and economic conditions under which such a cultural mani festation the natural sciences has evolved in Costa Rican history so as to present the ontogeny of the natural sciences or attitudes toward science in the context of science as a social phenomenon . One aspect of scientific development remains to be considered: the social characteristics that enable science to be bom , to grow, or to die in a country. None of the local sociologists ( Barahona 1953; Rodriguez 1979) have touched upon the subject of science and the Costa Rican. Sancho ( 1935), with all his ingenuity and incisiveness , ignored the local attitude toward science , as has Lascaris in more recent times ( Lascaris

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1975). Nevertheless, all these authors agree , with a mix- graduate program for the Universidad de Costa Rica is

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ture of doubt and pessimism , that environmental , histor ical , and demographic elements have yielded a set of individual and collective circumstances that do not favor inquisitiveness beyond the most pragmatic reasons. Such conclusions, tinged with Lamarckism , also agree with

the thoughts expressed in Clodomiro Picado’s many arti cles in the popular press, and Picado is the landmark of . The characterization of an “ attitude toward this century " is science not an exclusively local problem but concerns the entire Latin American community and offers a fasci nating field of research (Cardoso, Pinto, and Sunkel

1970).

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Reform and Change, 1955 80 The prevailing malaise of Costa Rican field biology was to change with the reform of the Universidad de Costa Rica in 1955. Shortly thereafter a school of biology was established . The curriculum of the school was designed in consultation with Archie F. Carr, a noted North American biologist and author. Rodriguez 's role has already been mentioned . From this modest beginning has grown the well -designed and well -equipped School of Biology and the Zoological Museum , with a full time faculty of fifteen Ph. D . s, some of whom are interested in field biology. More or less simultaneously, another native Costa Rican , Alvaro Wille , a Ph D. from the University of Kansas , took over and developed the entomological section of the School of Agriculture. While the general trend of Costa Rican government agencies and institu tions has followed that of other developing countries in emphasizing pragmatic and applied science and tech nology over basic science, the Universidad de Costa Rica in its latest incarnation has had a salutary effect on this attitude toward biology. Similarly, the revitalization of the Museo Nacional in 1970 added a stimulus to both costarricenses and extranjeros to undertake modem field studies. The institution , now a semiautonomous government agency, contains the national herbarium and provides modest modem research facilities and cooperative opportunities to all field biologists . The Museo's extraordinary recovery is due in ma jor part to the botanist Luis Diego Gomez , who has elsewhere described the origins and activities of the Museo (G6mez 1975a , 1978a ,b ). There are several other agencies in Costa Rica that are contributing to the current flourishing of field studies. The Centro Agronomico Tropical de Investigacion y En sefianza ( CATIE ) was originally founded by the Or-ganization for American States as the Instituto Inter americano de Ciencias Agricolas ( IICA ) in Turrialba in1942. In 19 ^ 2 the Centro became an independent entity ul continued its applied teaching and research program 1ก agriculture , torestry, and wildlife management. A

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located there as well. CATIE has been host to many field biologists , both in Turrialba and at its lowland field sta tion at La Lola on the railroad to Limon . The Caribbean Conservation Corporation (CCC) , a group of individuals dedicated to preserving and protecting the sea turtle nesting grounds at Tortuguero, was established in 1959 by Archie F. Carr, Jr. Professor Carr, the world 's foremost authority on sea turtles , used this site to study many aspects of the biology and migration of turtles, especially green turtles. Many field biologists have worked at the field camp at Tortuguero, and it be came part of a national park in 1970. The Ministerio de Agricultura y Ganaderfa ( MAG ) , a major department of the government , is responsible for agriculture , cattle , forestry, fisheries , and wildlife and for the National Park Service. A series of the ministry ’ร agricultural experiment stations at various localities in Costa Rica , especially at Taboga and Los Diamantes , have been used by many field biologists . The recently created wildlife reserve Rafael Lucas Rodriguez Cabal lero, formerly known as Hacienda Palo Verde , is a site of many studies and will continue to be available for field research . The Tropical Science Center ( TSC) is a private consult ing firm organized in 1962 by three North Americans originally associated with the IICA at Turrialba: Leslie R . Holdridge , an internationally known tropical forester; Robert J . Hunter, an agriculturalist; and Joseph A . Tosi , forester and land-use expert . Gary Hartshorn currently is associated with two of the original partners , Holdridge and Tosi , in studies of land use and forest resources . TSC operated a field station at Rincon de Osa ( 1966-73) , and it owns and manages the Monteverde Forest Preserve ( since 1971 ). Substantial field biology studies have been carried out at both sites. In this connection we should mention Paul Slud , whose works on the birds of La Selva ( 1960) and Costa Rica ( 1964 ) are classics , and whose association with Holdridge led to these synthetic works By 1968 the cadre of professional biologists in Costa Rica had reached a point where the group organized its own professional association , the Colegio de Bidlogos , to establish standards for educational requirements and to improve the qualifications for the title ( career ) of biologo. The college holds irregular meetings with public lectures and has led a number of significant campaigns to preserve endangered natural areas , most notably the Isla del Cafio and the migratory water bird reserve at Palo Verde. Most members of the college are field biologists or are conser vation oriented . The Servicio de Parques Nacionales de Costa Rica ( SPN ) , or National Park Service , technically a part of MAG , was established in 1970 as a response to the Costa Rican people ’s demand for a park system dedicated to

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a field -oriented course in tropical biology in Costa Rica. The program was a cooperative effort by the Director of the School of Biology, Rafael L. Rodriguez , the vicedean of Science and Letters , John deAbate , who holds a Ph . D. in zoology from Tulane University, and Jay M. Savage of use, to stimulate the development of field studies in Costa Rica . The original program , sponsored by the National Science Foundation of the United States , brought groups of United States professors to Costa Rica for six- week periods of instruction in field biology by biologists resident in Costa Rica. Field trips to many major habitats were an integral part of the program . At about this same time the Universities of Miami and Kansas were developing the concept of a modest field station for tropical studies in Costa Rica; Harvard University was thinking of relocating its tropical botany offerings after Castro 's take-over of the Atkins Gardens in Cuba; and the Universities of Florida and Washington were looking for a tropical base . Finally, the University of Michigan had spent nearly ten years trying to establish a field station in southern Mexico, for purposes similar to those of the other institutions. Because of these interests and the success of the UCR USC program , a series of meetings demonstrated that the common goals and individual institutional objectives could be met through a cooperative organization . As a result , the Organization for Tropical Studies ( OTS) was established in 1963 as a small consortium of seven North American universities and the Universidad de Costa Rica (Smith 1978). The primary objective of the group was to develop a center for advanced graduate education and research in tropical sci ences centered on basic knowledge of tropical environ ments. The consortium was formed because its members recognized that no single institution could provide the material and human resources necessary for the task. Principal emphasis in the beginning of OTS was on developing a cadre of knowledgeable tropical ecologists who had course and field experience in tropical environments and whose research activities would become concentrated on tropical systems. In the intervening period OTS has grown to include twenty-seven institutions of higher education , twenty three from the United States and four from Costa Rica. It has a central office facility in San Jos£ , Costa Rica, administrative headquarters at Duke University, and a series of field stations and other sites in key env ironments throughout the Republic. OTS has an annual budget of about one million dollars to operate its facilities and course programs and to provide logistic support to groups of independent researchers. In addition , the more than fourteen hundred graduate- level students whohave partic ) ( ipated in its course programs make OTS the principal The Organization for Tropical Studies OTS of a new generation of tropical biologists whose developer the and UCR Rica ) ( Costa de In 1961 the Universidad is based on actual tropical field experience. offering knowledge began USC ( ) California University of Southern

preserving the natural areas of Costa Rica. Although several parks had been established in the past , only to be abandoned , the growing sentiment of Costa Rican con servationists , led by the remarkable a id dedicated Mario A . Boza , stimulated the government of Pepe Figueres ( in his second term as president after being out of office since 1958 ) to form the SPN in 1970. In the ensuing years , under the leadership first of Boza , then of Alvaro F. Ugalde , and most recently of Jos£ Marfa Rodriguez , and with the support of Presidents Daniel Oduber and Rod rigo Carazo, a national system of twenty parks and reserves, unrivaled by any others in Latin America , has been developed ( Boza and Bonilla 1981 ). Research con sonant with the park 'ร purposes is encouraged by the SPN . Consejo Nacional de Investigaciones Cientfficas y Tecnoldgicas (CONICIT) was established in 1972 as an arm of the planning office of the Costa Rican govern ment. It serves as a combination National Research Council and National Science Foundation , attempting to set national priorities in science and technology and providing grant support to many field -oriented research projects . Its president is Rodrigo Zeledon , a distinguished parasitologist , former dean of the School of Microbiology at UCR . Universidad Nacional Autonoma ( UNA ), a second na tional Costa Rican university, was founded in 1973 in Heredia , with emphasis on an unusual curriculum and organization as opposed to the more familiar arrangement at the Universidad de Costa Rica . A major unit at UNA is the School of Environmental Sciences , which includes forestry and marine resources programs , among others. One curious fact of Costa Rican history is the almost total absence of interest in marine biology. Very few studies were carried out even by foreign naturalists on Costa Rican marine organisms during the nineteenth and early twentieth centuries , except those coincidentally related to major expeditions that happened to collect in waters near the Republic. This situation doubtless derives from the concentration of the population , and of economic and cultural life , on the Meseta Central during most of Costa Rica 's history. Without the convivial Costa Rican hosts enjoyed by terrestrial biologists to encourage them , marine biologists were not attracted to Costa Rican shores. For these reasons the recent establishment ( 1978) of a Centro de Ciencias Marinas at the university, under the vigorous leadership of Manuel M . Murillo, a noted student of crustacean ecology, and the construction of a small marine station by CONICIT at Punta Morales are encouraging .

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Because Costa Rica borders on both the Atlantic and the Pacific and is divided by a high mountain chain that attains an elevation of 3,918 m , it has a remarkably diverse biota including approximately 800 species of birds, 350 species of amphibians and reptiles , and 8,000 species of higher plants. About 10% of the known world butterfly fauna occurs in Costa Rica. Of the 38 major tropical plant formations, 14 are represented in the Republic, ranging from lowland deciduous and rain forests to paramo on the highest peaks. Finally, the region is of great biogeographic interest because it lies on the isth mian link that has made possible the dramatic interchange of biota between the previously separated North and South American continents. OTS field sites aim to take advantage of these factors so that courses and researchers may visit many diverse environments within a short time. It is possible to visit almost any major terrestrial or marine habitat within a day 'ร ride of San Jos£ and to carry out comparative ecological studies at several sites within a week . Principal sites utilized by OTS include lowland deciduous forest at Santa Rosa National Park (0-320 m ); lowland deciduous forest and riverine swamp at Refugio Rafael Lucas Rod riguez Caballero (formerly Palo Verde, 3-183 m ); several premontane to lower montane formations at Monteverde ( 1 ,200- 1 ,800 m ); premontane rain forest at Las Cruces ( 1 ,200 m ); montane forests at Cerro de la Muerte (3,000 m ); lowland evergreen forests at La Selva ( 29-100 m ) on the Atlantic versant and Corcovado National Park on the Pacific coast (0-400 m ). Principal marine sites are at Santa Rosa , Playas del Coco ( Pacific ) , and Cahuita National Park ( Atlantic ), a coral reef area . OTS's primary field station is at La Selva. It was originally owned by Leslie R . Holdridge, whose foresight in preserving a sample of undisturbed lowland forest cannot be overstated. The property was sold to OTS by Holdridge in 1968 . This evergreen forest site has been selected by the National Research Council (United States) Committee on Research Priorities in Tropical Bi ology as one of four worldwide tropica] localities for intensive long- term ecological research . The course program of OTS, particularly its renowned "Tropical Biology: An Ecology Approach " (familiarly called the Fundamentals course) , grew out of the efforts of Daniel H . Janzen and Norman J . Scott , who in the mid -sixties increased the course length to eight weeks , emphasized a full - time intensive field experience, and centered the theme on the exciting theoretical concepts of ecology that emerged in the period 1965-75. In retrospect almost every major figure in tropical biology today m the United States has been associated with the Funda mentals course. When OTS was founded in 1963 the intent was to create a cadre of knowledgeable tropical biologists whose

educational experience would be based on a common core of field -centered biology and ecology. First , it was anticipated that the training provided by direct exposure to tropical environments would generate new approaches to thinking about tropical ecology. Second , it was hoped that students and faculty would realize that field research is possible in the tropics with only slightly more effort then elsewhere. Third , it was anticipated that the devel opment of this new generation of tropical biologists would increase basic research on tropical problems. Fourth , OTS hoped that the intellectual foment devel oped by course participation would increase the number and interest of Latin American scientists in tropical stud ies. Finally, it was anticipated that OTS graduates would become involved in planning operational aspects of renewable natural resource management. In terms of research , OTS provided the impetus for a new wave of field studies , primarily ecological . Where almost all previous field studies in Costa Rica had been taxonomic or systematic in orientation , the new ecological theories of the 1960 and 1970s were well understood , tested , and revised , and new ideas were generated by tropical field biologists interested in this exciting area . The publication of several hundred papers based on OTSassisted studies during the first fifteen years of its existence attests to the excitement generated by the opportunities in Costa Rica. Less sanguine were the attempts by OTS to mount a “big science " program of coordinated ecosystem research ( 1968-76) the necessary infrastructure simply failed to materialize , and , while individual components and individual principal investigators had successes , the idea of an integrated effort failed in practice. The concept of an ecosystem analysis of two tropical lowland forests , one deciduous ( Palo Verde ) and the other evergreen ( La Selva ) , was sound . Its implementation failed because no single scientist emerged as the coordinator and leader for the entire project. Investigators and projects within the comparative ecological study included forest commu nities and primary productivity ( Kenneth J . Turnbull ); reproductive biology of plants ( Herbert G . Baker and Gordon พ. Frankie); beetle interactions ( Jack E. Coster); early development of the forests (Gordon H . Orians ); leaf -litter herpetofauna ( Jay M . Savage and Ian R . Straughan ); reproductive cycles , biomass , and food in vertebrate consumers ( Henry ร. Fitch ); insect-plant interactions ( Lawrence E. Gilbert ); experiments with tropical forest litter and its biotic communities ( Monte Lloyd ); growth initiative in trees ( James ร. Bethel ); mineral cycling ( Dale พ. Cole ); biometeorology ( Leo Frit schen ); soils and plant nutrition (Stanley p. Gessel ); plant community interactions ( William H . Hatheway ); insect dynamics ( Daniel H . Janzen ). Subsequent studies of a more restricted and individu

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ally coordinated type , especially in population biology, have been the mode since 1976. The titles of published papers from these efforts constitute part of the bibli ographies of this book; the substance of many of them is contained in the articles that follow.

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The Future of Costa Rican Field Biology

The enormous and tragic devastation of tropical forest environments during the past decades ( Myers 1981 ) promises to continue in spite of the best efforts of all who care for natural environments. Although Costa Rica now has a cadre of biologists whose orientations have been shaped by the new theoretical ecology, the ecological movement , and the stimulus of continuing contact with United States science through OTS, can they resist this trend ? Through their efforts , Costa Rica now has a solid scientific base in its CONICIT, its universities , and the Museo Nacional . It has an awareness of ecological problems and the proper attitude to face this dilemma. The Costa Rican national park system gives some hope that the marvelously diverse communities of tropical organisms will be preserved for future generations to enjoy and for future scientists to study. But can these factors overcome the pragmatism of economic man ? Especially the pragmatism of the Latin culture? Can short- term economic gain be ignored for future long-term benefits to the nation ? Or will Costa Rica , now ideally situated to provide an example to the rest of tropical America , succumb as well ? Wfe do not know. Perhaps Costa Rica , among all the tropical countries , because of its history, its commitment to the preservation of natural areas , and its emerging scientific consciousness, can avoid the temptation of uncontrolled exploitation of its lands. If it succeeds where all others have failed it may be the beacon that provides guidance to others in the salvation of natural tropical environments . The pessimist may wish Costa Rica well yet believe there will be no room for field biology by the end of this century. Only fields of crops , a crowded and polluted Meseta Central , eroded hills , and silted rivers will remain . The optimist fears the worst yet remembers the changes in Costa Rican national attitudes in the past two decades , the dedicated young biological scientists and conservationists who have helped in developing the nation ’s environmental consciousness, and the emerging national concern for basic knowledge of the environment and its biota coupled with planning for the benefit of both man and environment . A long shot perhaps , but one that must pay off if there is to be tropical field biology in the twenty -first century. But still we remember the bright blue skies , the white clouds , the almost black forest on the slopes of the volcanoes , the driving rain , the green complexity of the forest canopy viewed from a mountain slope , and our

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own tininess within the forest s grasp . We remember the wonder and awe when a startled danta leaps up from underfoot , the morpho butterflies flit by, and the bellbird sounds from the forest top. On the trail of discovery followed by all the long line of our predecessors in the forests and on the cerros of Costa Rica , can we truly believe that man is so foolish as to completely destroy this special world ? Wfe cannot let it be so! For once gone, something special and basic about ourselves will be gone too and afterward man himself will not survive. Viva Costa Rica! All success in this noble undertaking! ’

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Aguilar B . , o. 1978 . Costa Rica y los hechospoliticos de 1948: Problematica de una decada . San Jose: Editorial Costa Rica Balli , A . 1974-77. Ideas biologicas . Department of Bi ology, University of Costa Rica. Mimeographed . Barahona J , L . 1953 , El gran incognito: Vision interna del campesino costarricense . Serie Ensayos 3. San Jose: Editorial Universidad de Costa Rica. Boza , M., and Bonilla , A. 1981. The National Parks of Costa Rica . Madrid: INCAFO. Cardoso, F. H .; Pinto, A .; and Sunkel , o. \910 . America Latina: Ciencia y tecnologia en el desarrollo de la sociedad . Coleccion Tiempo Latinoamericano. San tiago de Chile: Editorial Universitaria. Cerdas , R . 1972 . La crisis de la democracia liberal en Costa Rica . Series 6. San Jos£: EDUCA . Colon , c. 1972 . Diario de Colon . 2d ed . Madrid: Edi torial Cultura Hispanica. ConejoG . , A . 1972 . Materiales para una bio-bibliografia costarricense del Dr. Henri Pittier Dormond . Tesis de grado, Universidad de Costa Rica. Dobles ร., L. 1927. Indice bibliogrdfico de Costa Rica . Vbls. 1-3. San Jose: Lehmann . Estrada M . , L. 1965. La Costa Rica de Don Tomas de Acosta . San Jos£: Editorial Costa Rica. Fernandez G ., R. 1972. Costa Rica en el siglo XIX . 3d ed . San Jos£: EDUCA . Ferrero A . , L. 1978. พ. M. Gabb. Talamanca: El espacio y los hombres . Serie Nos Ven 7. San Jose: Ministerio de Cultura , Juventud y Deporte . Folkman , D. I . 1970. The Nicaragua route . Salt Lake City: University of Utah Press. Gaspar, J . c. 1979. Limon 1880 -1940: บท estodio de la industria bananera en Costa Rica . San Jose: Editorial GosUi Rica Gomez p , L . D. 1973. El Museo Nacional de Costa Rica. Museum ( UNESCO) 25:182-84 . . 1975«. Bibliografia geologica y paleontologica de Centroamerica y el Caribe . San Jos£: Museo Na cional de Costa Rica . , ed . 19756, Biology of Bromeliaceae. In Historia natural de Costa Rica , vol. 1 . San Jos£: Museo Nacional de Costa Rica . . 1978«. Contribuciones a la pteridologia cos tarricense. XL Hermann Christ , รน vida , obra e

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influencia 79en. la botanica nacional. Brenesia Picado T., c. 1921. Pasteur y Metchnikoff . Repertorio

12-13:25. 19786. Contribuciones a la pteridologia cosXII . Carlos Werckle. Brenesia tarricense. 14-15:361-93. Gonzalez F. , L. F. 1976. Historia de la influencia extranjera en el desenvolvimiento educacional y cientifico de Costa Rica . 2d ed . Biblioteca Patria . San Jose: Edi torial Costa Rica. . 1978. Evolucion de la institucion publica en Costa Rica . 2d ed . Biblioteca Patria. San Jose: Edi torial Costa Rica. Gonzalez V. , c. 1977. Historia flnanciera de Costa Rica . 2d ed. San Jose: Editorial Costa Rica. Hall , c. 1978. El cafe y el desarrollo historico- geografico de Costa Rica . San Jos£: Editorial Costa Rica-EUNA . Hoffmann , Carl . 1856. Excursion nach dem Volcdn de Cartago in Central America. Bonplandia , no. 6. . 1957. Eine Excursion nach dem Barba- Vulkan in Costa Rica. Bonplandia , no. 16. Houwald , G . von . 1975. Los Alemanes en Nicaragua . Serie Historica 2. Managua: Banco de America . Jimenez, R . 1921 . El Colegio de Cartago. Repertorio Americano (San Jose). Lascaris c. , c. 1964 . Desarrollo de las ideas Filosoficas en Costa Rica . San Jos£: Editorial Costa Rica. . 1975. El costarricense . San Jose: Editorial Uni versidad Centroamericana ( EDUCA ). Myers , N . 1981. The hamburger connection: How Cen tral America’ร forests became North America’s ham burgers. Ambio 10:3-8. Picado Ch ., M . 1964 . Vida y obra del Doctor Clodomiro Picado . Biblioteca Autores Costarricenses. San Jose: Editorial Costa Rica.

Americano (San Jose ). Picado T., c., and Trejos พ. , A . 1942 . Biologia hema tologica comparada . San Jos£: Editorial Universidad de Costa Rica. Rodriguez , V. E. 1979. Apuntes para una sociolog ia costarricense . 3d ed . San Jos£: Editorial Universidad Estatal a Distancia. Sancho, M . 1935. Cosla Rica, Suiza Centroamericana . San Jos6. Sapper, c. 1943. Viajes a varias partes de la Republica de Costa Rica 1899 -1924 , ed . J. F. Trejos. San Jose: Universal. Schifter, J. 1979. La fase oculta de la guerra civil en Costa Rica . San Jos£: Editorial Universitaria Centroamericana ( EDUCA ). Slud , p. 1960. The birds of Finca “ La Selva , ” a tropical wet forest locality. Bull . Am . Mus . Nat . Hist . 121:49-148. . 1964 . The birds of Costa Rica: Distribution and ecology. Bull . Am . Mus . Nat . Hist . 128: 1- 430. Smith , c. M. 1978. The impact of O.T.S. on the ecology of Costa Rica. Texas J . Sci . 30:283-89. Soley G., T. 1975. Compendio de historia economica y hacendaria de Costa Rica . 2d ed . Biblioteca Patria. San Jose: Editorial Costa Rica. Termer, F. 1956. Carlos Sapper. Explorador de CentroAmerica ( 1866-1954 ). An . Soc . Geog . Hist . Gua temala 29:55-101. Trudeau , R . H . 1971. Costa Rican voting: Its socio economic correlates. Ph . D. diss., University of North Carolina .

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2 THE CENTRAL AMERICAN niQPPDQAI

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BIOTIC HISTORY AND JZ Y

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The Paleogeography of Central America during the Phanerozoic: The Past 600 Million Years The narrow strip of land that now connects the major terrestrial biogeographic realms of Nearctica ( North America ) and Neotropica (South America ) includes Costa Rica . Today this area is usually included within the Neotropical realm , but it certainly has a distinct Nearctic flavor, and it serves as a filter bridge ( see Simpson 1965; Marshall et al. 1982) across which some organisms can 12

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p. V. Rich and T. H . Rich

The narrow strip of land that today joins the Neotropics of South America and the Nearctic of North America , two major biogeographic realms, has not always been as it is. At present Central America forms a continuous land connection between the north and the south , serving both as a corridor for terrestrial forms ( Marshall et al . 1982 ) and as a barrier to marine forms attempting an east - west movement between the Caribbean-Gulf of Mexico and the Pacific Ocean. In times past this area has been both continuous terrestrial corridor and insular stepping stones for land dwellers as well as both barrier and cor ridor for marine life. Thus it has a varied history that cannot be deduced from its present configuration . To understand the biotic associations present today in Costa Rica , as well as in the rest of Central America , it is useful to know something of the history of the area . Thus it is appropriate that this book include sections on geologic history (see chap. 4) and on plant and animal fossils. เท this chapter we deal with three major aspects of Central American history. First , we estimate what the area has looked like , both paleogeo raphically and paleoenvironmentally, during the Phanerozoic ( the past 600 million years or so). Second , we present an overview of biotic history, that is, a summary of what is known of fossil organisms, both marine and nonmarine, emphasizing the past 60 million years , the time most relevant to the development of modem faunas and floras in this area. Third , we evaluate the role this area has played in the biotic interchange between different biogeographic realms and consider how it has achieved its present unique character.

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pass. Additionally, it now serves as a barrier to marine forms in both the Caribbean and the Pacific; but , as paleontological studies have demonstrated , its role has changed many times. Let US first try to reconstruct just how Central America looked geographically and ascertain its relationship to both North and South America through time. To this end , we will present a quick resume of several current theories , none of which has met with general acceptance. No more than twenty years ago, a majority of biogeographers would have said that Central America has always been much as it is today. The only major changes suggested would have been occasional rises and falls in sea level and raising and lowering of the mountains that form the backbone of the area . But as for bits and pieces of Central America moving through several degrees of latitude or longitude, the prevailing opinion was “ not possible!" Continents , and parts thereof , were stable perhaps not vertically, but definitely with regard to “ hoand certainly rizonal " position upon the earth 'ร surface " “ , then , hypothesis stablist relative to one another. This rest the and Rica is one way of viewing the past of Costa of Central America.

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FIGURE 2.2. Cross section of a part of the upper 700 km of the earth showing major divisions between crustal plates. a modified after Sykes ( 1972 ). b , from Wyllie ( 1976).

RIDGE

II

,

volcanoes are common , as in the Lesser Antilles, an extreme example being the tremendous explosion of Mount Pele early in this century. Transform faults are a third boundary type , where two crustal plates move past one another laterally, generally

I

Plate Tectonics and Central America More recently, a number of theories with a “ mobilist" flavor have been put forward , contending that parts of Central America have moved markedly and that the present configuration cannot be assumed to extend very far into the past . All these theories assume the cor rectness of a major synthetic theory, plate tectonics ( see any recent geology text for a thorough explanation of this , such as Wyllie 1976 or Tarling and Tarling 1971 ), and the existence of sea- floor spreading. Simply stated , this theory suggests that the earth 's cmst ( the outer few hundred kilometers of rock ) is composed of a series of J plates ( see fig . 2.1 ) that meet along three types of bound 1 aries: ridges, trenches , and transform faults ( see figs. 2.21 and 2.3). Where plates meet along ridges , new molten rock is added from below, causing the plates to grow and move in opposite directions ( ridges , such as the mid-

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FIGURE 2.1. Major divisions ( lithospheric plates ) in the earth ร crust that have moved relative to one another. The major boundaries are of three types: ridges , where new cmstal material is added and plates grow; trenches, where cmstal material slides beneath other crustal material: and transform faults, where cmstal material slides past other crustal material ( e.g., the San Andreas fault in California ). In this diagram , I represents ridges; trenches; I transform faults. From Middlemiss, Rawson , and Newall ( 1971 ). Atlantic ridge, most often occur in ocean basins, but they

can have terrestrial counterparts , as in central Iceland ). Shallow earthquakes and volcanic activity often occur along such boundaries. Trenches are boundaries where one cmstal plate (lithospheric plate ) dives under another, and the lower plate is consumed or remobilized . Cmstal shortening occurs in these areas , and deep earthquakes are common along the plane of the descending plate. The oceanic trench off the west coast of South America is a familiar example, often associated with devastating earthquakes in Chile and other western South American countries. Not only are shallow, moderate , or deep earthquakes associated with such trenches, but so is volcanism. Behind trenches , above the area where one plate descends beneath another

.

resulting in shallow earthquakes. Most of these occur in ocean basins, but occasionally they are manifest on land such , as is supposedly the case with the San Andreas fault in California. Detailed studies of ocean floor magnetism , volcanism , seismicity, general geology, and several other aspects of geology all suggest that plate tectonics is the best theory currently available. One major problem remaining is just what mechanism drives these plates , although many suggestions have been made (see fig. 2.4). When the plate tectonics model is applied to the history of the area between North and South America , the Caribbean , and the Gulf of Mexico, several theories result . Most workers have dealt only with the past 100 to 200 million years of history ( Dietz and Holden 1970; Fox , Schreiber, and Heezen 1971; Freeland and Dietz 1971; Malfait and Dinkelman 1972, for example), but a few have attempted reconstructions spanning 600 million years (พ&lper and Rowett 1972), beginning with the Paleozoic era ( see figs. 2.5 and 2.6). There certainly are Paleozoic rocks in parts of Central America , all north of central Nicaragua , some dating back to least 400 million years, into the Silurian period ( Naim and Stehli 1975). But there is a lack of understanding of just what paleogeographic relationship these

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105° FIGURE 2.3. Important geographic features of terrestrial and marine environments in Central America and the Caribbean ( after Ladd 1976 ).

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Central American rocks had to each other and to those outside this area . For example , there is no general agree ment on whether the Gulf of Mexico first formed during the Mesozoic or is a much older Paleozoic or PreCambrian feature . Some workers suggest that the first and only Gulf of Mexico began in the Mesozoic and gradu ally enlarged as the North and South Atlantic developed over the past 200 million years. Others suggest that this was only the final history of a much older ocean that was

150

present during the Paleozoic and may have partially or completely closed during the late Paleozoic or early Mesozoic, only to reopen again in the Mesozoic to present. According to this latter theory, there would have been a proto-Gulf as well as the modem Gulf of Mexico ( see Cebull and Sherbet 1980, for a good summary' of this controversy ). Evidence is gradually accumulating to sug gest that the proto-Gulf /Gulf hypothesis is the more ten able ( Cebull and Sherbet 1980; Ross 1979 ). More applicable to studies on the living biota of Cen -

tral America are paleogeographic reconstructions for the late Mesozoic and Cenozoic, a time when mammals and birds as well as the flowering angiosperms radiated widely. Although there is still disagreement among mobi list geologists about the correct way to assemble and disassemble Central America , there is much more paleomagnetic data and information from general geology and fossil studies , allowing more constraints on theories pro posed for the past 200 million years than on theories about earlier times.

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FIGURE 2.4 . Proposed mechanism for movement of lithospheric plates ( after Rich 1976).

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FIGURE 2.5. Geological time scale giving both names and absolute dates for the major time divisions from the begin ning of the earth to the present ( from McAlester 1976 ).

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Formotion of Earth's cruft about 4,600 million years ogo

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until sometime in the Pliocene ( 5-6 million years ago; Raven and Axelrod 1975), no terrestrial connection exis ted between North and South America via Central Amer ica ( see our section on paleontology below, and Bandy and Casey 1973; Case et al . 1971 , among others) , al though there could well have been a connection via Eu rope and Africa .

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zoic. Most of the Mesozoic sequence , exemplified by the Nicoya complex , well exposed on the Nicoya Peninsula ( see chap. 4 ) represents deepwater marine sediments probably associated with a trench , the volcanic rocks resulting from remobilization of oceanic crust on the descending plate behind the trench ( see fig . 2.2 ). Rocks from the Cenozoic ( including the older Tertiary period and the younger Quaternary period ) include both volcanic (as well as plutonic; see chap. 4 ) and terrestrial and marine sedimentary fractions. The terrestrial sedi ments represent volcanic debris, riverine and lake deposits flowing off the rising backbone of Central America , which continues to grow through volcanic activity. According to Malfait and Dinkelman 's ( 1972 ) hypothesis , these mountains ( see figs. 2.2 and 2.8-10) have resulted from upwelling of remobilized , molten material created along a descending crustal plate behind the Middle American Trench . Only in the late Pliocene did they approach their current height. Most of the terrestrial sediments deposited as these mountains rose have very small areas of outcrop ( often only along streams and in roadcuts ); they are otherwise covered by vegetation or may have been deeply weathered , and thus fossil localities within them are extremely rare. Throughout most of the Cenozoic the area from Nic aragua to northern Colombia would probably have been reminiscent of the Lesser Antilles today — volcanically active, topographically and environmentally diverse , but certainly not a continuous terrestrial connection. This area was a perfect example of Simpson ’s ( 1965) sweep stakes route between North and South America , allowing only an occasional terrestrial form to pass , purely by chance , not by design .

B

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FIGURE 2.6. Inferred Paleozoic history of Central America , from 600 to 225 million years ago ( after Walper and Rowett

1972).

Figure 2.7 illustrates several hypotheses on continental configurations at the beginning of the Mesozoic. Al though they indicate the breadth of divergent opinions , all suggest that the separation of North and South Amer ica as well as the separation of both from Africa at this time was small , and that terrestrial forms thus probably would have had few barriers to dispersal . Pacific and Atlantic marine forms would have been affected in a variety of ways depending on the reconstruction chosen . During the Mesozoic and Cenozoic, the North and South Atlantic as well as the Gulf of Mexico and the Caribbean Sea gradually developed and increased in size (see figs. 2.8-10). Malfait and Dinkelman ( 1972 ) have suggested that during this expansion of these ocean basins the Cen tral American area was affected by the incursion of part of the Pacific crustal plate , which determined the paleo geography of this entire area . Others workers (e. g., Gose and Scott 1979; Gose and Swartz 1977; Anderson 1978; and Schmidt 1978 ) do not entirely agree with this model , preferring to define a number of small crustal plates in the eastern Pacific and Caribbean area that rotate relative to one another, eventually leading to the late Cenozoic ag gregation that we know today as terrestrial Central America . Even though it is not clear which of the multiple the ories are correct , it is clear that , from the latest Mesozoic

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The geologic history of Costa Rica , unlike that of “ nu clear Central America ' ( to the north of central Nic aragua ) spans only the middle Mesozoic to Recent times. There are no known Paleozoic rocks from northern Nic aragua to Colombia ( Naim and Stehli 1975). During this time terrestrial geologic history has been tied to volca nism , as have even the marine sediments of the Meso-

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E

FIGURE 2.7. Five hypotheses concerning the arrangement of continents just before the last stage of major drifting , during the Triassic ( early Mesozoic ), about 200 million years ago. a . " Bullard fit " after Sherbet and Cebull 1975. b , “ Bullard

fit " after Freeland and Dietz 1971. c, After Walper and Rowett 1972. d After Morel and Irving 1980. e After Freeland and Dietz 1971 .

for more than 80 million years. In Central America at present , the angiosperms — in fact the entire flora — seem most similar to those of South America . In reviewing this similarity Raven and Axelrod ( 1975) suggest that , ever since the beginning of angiosperms sometime during the Cretaceous, Africa and South America , and perhaps even some of the fragments of Central America yet uncon solidated , have constituted a site of evolutionary im portance. Much of the world’s lowland tropics were con centrated in these areas . As Africa and South America separated , from the late Cretaceous onward , and as South

America began to converge on some parts of Central America , the West Indies , and southern North America , sweepstakes dispersal occurred and South American forms moved into these areas. Raven and Axelrod ( 1975 ) further note that the reverse dispersal , from temperate North America to South America , was not so striking . “Such genera as fir ( Abies ) , alder ( Alnus ) , sweetgum ( Liquidambar ) , beech ( Fagus ), walnut ( Juglans ) and elm ( Ulms ) had already reached the mountains of southern Mexico 16 million years ago” ( p. 428). Some of these forms extended through Central

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History of the Flora of Central America ะ The Flowering Plants

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Today the angiosperms, or flowering plants , are the dom inant terrestrial plants, having retained this dominance

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FIGURE 2.8. The world during mid - Mesozoic times, 135 million years ago. Hatched lines indicate oceanic trenches where crust is being consumed ; heavy lines , spreading

ridges; light lines , transform faults. Arrows indicate motion of individual crustal plates relative to one another. White areas are terrestrial environments ( after Rich 1975).

America into South America , with walnuts reaching northern South America by at least 8 million years ago, alders by 700 ,000 years ago, and oaks (Quercus) by about 150 ,000 years ago. By the late Pliocene , some 2-3 million years ago, the mountains were high enough to provide a reliable corridor for montane -adapted plants. During the Pleistocene , cool climates first appear in tropical latitudes ( Raven and Axelrod 1975) , and this further promoted dispersal of Montane species. Such forms as the gooseberries and currents ( Ribes ), the locoweeds ( Astragalus ) , willows ( Salix ) and evening prim roses ( Oenothera ) probably dispersed south from North America then sometime during the past million years. Additionally, during the Pleistocene there were periods of greater aridity than in modem times ( Raven and Axel rod 1974 ). Grasslands expanded , as did thorn scrub , and this environmental alteration favored the exchange both

north and south of more arid -adapted animals and of grazing forms such as glypotodonts , horses , ground sloths, and camels (see below ).

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FIGURE 2.9. The history of Central America during the late Mesozoic and Cenozoic era 80 million years ago to the present (after Malfait and Dinkelman 1972 ).

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Rock Sequences in Central America 9 with Emphasis on Marine Sediments and Invertebrate Fossils: Paleozoic to Recent Woodring ( 1964 ) summarizes well the outcrop patterns of marine rocks in Central America , which are dated pri marily according to the invertebrate fossils occurring in them . Most of these fossils are foraminifers and radio larians or mollusks ( primarily bivalves and gastropods) (see fig . 2.11 ). The oldest rocks in Central America occur in the deformed mountain ranges of northern Honduras , central Guatemala , and northern Nicaragua . These are definitely

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FIGURE 2.10. The world during mid-Cenozoic times ( Oligocene ) , 35 million years ago. See figure 2.8 for legend .

older then early Permian and in fact may be Precambrian in age. Early Permian marine rocks directly overlie these older igneous and metamorphic rocks. No Triassic rocks appear to be present in Central America , and there is a record of deformation and intrusion by granitic, and some more basic , rocks . After this detormation , nonmarine Jurassic rocks were deposited north and south of these folded ranges. These rocks contain plants of early and middle Jurassic age. Except for an undocumented report of late Jurassic sediments in Hon duras, no marine Jurassic rocks are known from Central America . Early Cretaceous marine rocks are known in northern Central America , containing rhudist corals (see fig . 2.11 ) and the microscopic foraminifer Orbitolina concava texana. Although they do not crop out on land , early Cretaceous deposits dredged up off the Pacific coast of Central America have contained land - plant debris. Late Cretaceous marine deposits are known in northern Guatemala that contain the strange rudist bivalve Barrettia (converging with corals in external morphology ) and from northwestern Panama near the Costa Rican border ( dated by the contained foraminifers). Cenozoic marine deposits are decidedly more abun dant in Central America than are Mesozoic and Paleozoic deposits. Paleocene and early Eocene sediments, how ever, are mostly thin and limited in extent . During the mid to late Eocene carbonate deposits ( limestones) were

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widespread , with some terrestrial sediments and volcan ics. Although early Oligocene sediments of any kind are rare , marine limestones of late Oligocene or Miocei occur in Central America , along with extensive volcani and some terrestrial sediments. Marine Pliocene deposi are generally thin ; marine coralliferous rocks are kno from Limon , Costa Rica . Thin marine Pleistocene depos its are widely distributed on coastal terraces . They hav < not been satisfactorily differentiated into early or lat( Pleistocene. Reef corals and mollusks are common ii these deposits. Some of the more important invertebrate localities in Central America are listed below, as are faunal lists ot the better- known forms from each of these areas. Figure 2.1 illustrates a few of the characteristic invertebrate fossils from some of these locales. For more detailed coverage of this topic one should consult such references as Davies ( 1971-75) , Olsson ( 1922 , 1942 , 1943 ) Jackson ( 1917 ) Coryell and Fields ( 1937 ) , and Woodring ( 1957 ) , both tor text descriptions and for good illustrations of most species . Only a very few localities have produced Cre taceous, or pre Cenozoic, invertebrates in Central America. The fauna includes foraminifers (Globotruncana and Gumbelina ), radiolarians , and rhudist corals. The first diverse marine invertebrate faunas from southern Central America are known from the mid to late Eocene of Panama and Costa Rica (e. g., Gatuncillo For

lj

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FIGURE 2.11 . A selection of invertebrates most common in Central American Phanerozoic sediments. Ages are indicated for each suite.

mation ) and have been dated primarily by the contained microscopic Foraminifera . Smaller Foraminifera include Alabamin Al / omorphina , Angulogerina , Anomalina , Anonuilinoides , Astacolus Bathysiphon Bolivina Bub imina Cassidulina Cassidulinoides , Ceratobulimina , Chilostomella , Chilostomelloides , Chrysalogonium, Cibicides , Clavulinoides , Cornuspira Cyclammina , Dental ina , Discorbis , Dorothia , Ellipsoglandulina , Entosolenis Eponides , Frondicularia Gaudryina Glandulina , Globigerina , Globigerinoides , Globulimina Globorotalia , Globulina , Gumbelina Guttulina , Gyroidinoides Hantkenina Haplophragmoides , Hastigerinella , Hoglundina , Karreriella Lagena , Lagenoglandulina , Lagenonodosauria Loxostoma Marginulirui , Margitiulinopsis odogenerina , Nodosaria , Noion , Orthomorphina , Osngularia Planularia , Planulina , Plectina Plectch 9

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frondicularia , Pleurostomella , Pseudoglandulina , Pullenia , Pyrgo , Quinqueloculina Robulus , Rotaliatina , Saracenaria Schenckiella , Sigmoilina , Sigmomorphina , Siphonina Siphonodosaria , Siphotextularia , Spiroloculina , Spiroplectammirui Textularia , Textulariella , Triloculina , Uvigerina , Vaginulinopsis , Valvulineria , Virgu /ina . Larger foraminifers include Yaberinella , Operculinoides , Nummulites Heterostegina , Fabiania , Helicostegina , Lepidocyclina , Helicolepidina , Asterotycline and Pseudophragrnina ( Woodring 1957 ). Corals include Heliopora , Astrocoenia Astreopora , Diploastrea , Goniopora Porites Favia , Colpophyllia , Ann Ilia , and Millepora . Mollusks include Velates , Hannatoma , Xenophora , Hipponix , Calyptracea , Polonices? , Nexerita? Simon , Anuiurellina? , Pachycrommium? Turritella cf . Carinato and T. cf . samanensis. 9

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THE CENTRAL AMERICAN DISPERSAL ROUTE

Oligocene faunas possibly are represented by the Bohio Formation in Panama , which contains a diversity of smaller Foraminifera. Although much of this formation is nonmarine , with silicified wood recorded , the marine fossils are what establish the age of these sedi ments. Smaller Foraminifera include Alabamina Ammospirci , Angulogerina , Astacolus , Bothysiphon , Bolivina , Bulimina , Cassidulina , Chrysalogonium , Cibicides , Cla vulinoides , Cyclammina , Dentalina , Discorbis , Eponides , Gaudryina , Glandulina , Globigerina ciperoensis Gum belina , Gutndino , Gyroidinoides , Karreriella , Lagena , Margimdina , Nodogenerina , Nodosaria , Nonion , Osangularia , Planularia , Plectina , Plectofrondicularia , Pleurostomella , Pseudoglandulina , Pullenia , Quinqueloculino , Robulus , Schenckiella , Sigmomorphina , Si phonina , Siphonodosaria , Spiroloculina , Uvigenna , Vagi nulina Vaginulinopsis , Virgulina , and Vulvulina. Larger Foraminifera include Heterostegina anti Ilea , Archaias , Lepidocyclina , and Miogypsina. Mollusks are primarily gastropods and include Solariella , Neritina , Hemisinus , Crepidula , Natica , Polinices , Sinum , Globularia , Pachycrommium , and Turritella cf . altilira ( Woodring 1957). Miocene marine faunas are markedly diverse in Foraminifera , corals, ทในแนรkร, and a few enhinoids. Some of the more important fossiliferous rock units are the Culebra , Cucaracha , Panama , and Gatun formations , mainly in Panama , especially on Barro Colorado Island and part of the Usacari Shale and the “ Puerto Limon " Formation near Limon , Costa Rica. Foraminifers include Siphogenetian transverse , Miogypsina , Lepidocyclina miraflorensis , and several others not yet described. Corals include Stylophora , Acropora , Porites , and Mon tastrea . Mollusks include a particularly wide variety in the Gatun Formation: Calliostoma , Turbo , Neritina , Teinostoma , Anticlimax , Cvclostremiscus , Solariorbis , Episcynia , Rissoina , Xenophora , Crepidula , Calyptraea , Trochita , Crucibulum , Cheilea , Natica , Stigmaulax , Tectonatica , Polinices , Neverita , Sinum , Turritella altilira and several more species. Echinoids include Clypeaster gatuni , Encope annectens , and Schizaster panamensis. Eighteen species of ostracods have also been recorded (Coryell and Fields 1937). Other areas that have produced Miocene faunas of marked diversity include the RFo Dulce (Guatemala ) and Barranca (Costa Rica ). Fish are also rather abundant in the Gatun Formation including selachians and teleosts ( Xenodermichthys and Gobius ). The Cucaracha Formation is of particular interest because it is mainly nonmarine. Later in this chapter we will discuss fossil mammals. The formation also includes plant debris (Taenioxylon multiradiatum ) and wood . A few marine and brackish - water fossils have been found in the lower part of the formation: Anadara , Crassostrea cf . Lucina , and c. cf . Tellina . On the basis of these few

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9

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^ ^ THE CENTRAL AMERICAN

DISPERSAL ROUTE fossils the age of the Cucaracha Formation cannot be the impoverishment led to also . of the Car- 1 Cucaracha fauna ( = Gaillard Cut local fauna ) change but established , but it can be delimited because both the 2 . Panama Canal Zone ibbean province ( Woodring 1966). underlying Culebra Formation and the overlying Panama 3 f0ssils from Hemingfordian ready Miocene) " Formation are considered to be of Miocene age. ch s spwime The Pliocene is represented by such rock units as the * ร* 1 651 “ “( 1955), and Davies ( 1971 ) "^ ) , Cushman and Fischer ( 1952 Chagres sandstone ( Toro limestone member ) of Panama , 6. Class Reptilia should be consulted as starting points. the Burica sandstone of Panama and Costa Rica , and the Order Chelonia Moin Formation near Limon , Costa Rica. Faunas from Order Crocodilia The Record of Cenozoic Terrestrial Mammals in these are primarily mollusks and a few echinoids: MolClass Mammalia Central America Order Perissodactyla lusks include Dentalium , Terebra , Architectonica , Cancelhistory of terrestrial vertebrates in Central America The Family Equidae laria , Hanetia , Cantharus , Phos , Fusinus , Siphonalia , is poorly documented by direct evidence from the fossils Anchiterium Turritella , Polinices , Nucula , Area , Pecten , Chione , Lu that once lived there. Table 2.1 lists the terresArchaeohippus ciploma thyasira , and Solemya. In the Burica Formation of animals mammal sites recorded in the literature , and Family Rhinocerocidae the presence of Dentalium ( Fissidentalium ) buricum and trial fossil shows where 2.12 they are Diceratherium figure . situated None of them Nucula iphigenia suggest a Pliocene age, while Phos Order Artiodactyla , in age and Mesozoic only are one is older than Miocene: gatunensis and Turritella cf . gatunensis suggest late Mio Family Protoceratidae first tour the filths of thus the history of mammals is cene affinities. Additional mollusks from the Chagres completely unknown in Central Paratoveras ท. sp. , America and all but the sandstone include Turitella altilira and Stigmaulax fup Family Merycoidodontidae last tenth is documented by only one locality ( see figs. pians. Echinoids are represented by Clypeaster. Merycochoerus By far the most richly fossiliferous of all the Cenozoic 2.13-16). Brachycrus marine rocks are those of Pleistocene age. Mollusks are TABLE 2.1 Summary of the Vertebrates from Central . 1 Hidalgo fauna by far the most diverse of any invertebrate group American Cenozoic Sediments . Mexico 2 recorded , although a thorough search of more recent lit. 3 ( mid- Miocene ) ( Information is presented in the following order: 1 , name 4. Clarendonian erature would probably yield additional data on ForMooser ( 1959 ; , ; 2 , ; aminifera. A typical assemblage of molluskan genera is of fauna country 3 age 4, literature reference; 5, 5. North America 1963) represented by one from Monte Verde Ravine, Costa continental affinities of fauna; 6, faunal list.) 6. Class Mammalia Rica , where more than 130 molluskan species are known: Order Perissodactyla Eocene-Oligocene Area , Barbartia , Noetia , Pinna , Ostrea , Pecten , Pli- 1. Guanajuato fauna Family Equidae catula Crenella , Anomia , Placuanomia , Pandora , Thra 2. Mexico Neohipparion montezumae N . otomii tio , Eucrassatella , Crassinella , Charna , Diplodonto Car- 3. Uintan or Chadronian ( mid-Eocene to early Oligocene) N . monias dium , Cyclinella , Macrocallista , Pitar, Chione , Tellina , 4. Black and Stephens ( 1973), Fries, Hibbard , and Dunkle ( 1955) Macoma , Semele , Tagelus , Solecunus , Mactra , Labiosa , M iocene- Pliocene Corbula , Panopea , Bullaria , Terebra , Conus , Polystira , 5. North America and endemic 1 . Gracias fauna Turricula , Clathrodrillia , Crassispira , Nannodrillia , Can 6. Class Mammalia 2 . Honduras Order Perissodactyla cellaria , Oliva , Olivella , Marginella , Lyria , Latirus , Ga 3. Hemphillian ( late Miocene or early Pliocene ) Superfamily Tapiroidea leodea , Hanetia , Colubraria , Cymarium , Distorsio , En 4. Olson and McGrew ( 1941 ), McGrew ( 1944 ) Order Rodentia gine , Cantharus , Triumphis , Nassa , Phos , Metula , . North America, endemic at the specific level only 5 Family Ischyromyidae Columbella , Cosmioconcha , Strombina , Murex , Phyl . 6 Class Mammalia Floresomys guanajuatoensis lonotus , Thais , Seimcassis , Fiatร , Cypraea , รtrombus , Order Carnivora Family indet. Turritella , Architectonica , Crepidula , Crucibulum , Calyp Family Canidae Guanajuatomys hibbardi traea , Natica , Polinices , Neritina , Circulus , and Dental Osteoborus cynoides turn . Many species are indistinguishable from livin; Amphicyon sp. Miocene Order Perissodactyla forms. Corals are also diverse in these faunas. 1. El Gramal fauna Family Equidae When Cenozoic marine invertebrate faunas of the Cen 2. Oaxaca , Mexico Pliohippus hondurensis tral American area are examined , particulary the mol- 3. Hemingfordian or Barstovian (early Miocene Neohipparion montezumae lusks ( Woodring 1966), it becomes clear that , although 4. Stirton ( 1954), Wilson ( 1967) Family Rhinocerotidae species were able to move between the Caribbean and the 5. North America Subfamily Teleocerinae Pacific during the early and middle part of the Cenozoic, 6. Class Mammalia Order Artiodactyla

“

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-

-

•

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. ---

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particularly during the Miocene , that during the latest Miocene and more recently this interchange was in creasingly restricted . Interchange ceased entirely during the latest Pliocene or earliest Pleistocene , the Central American land bridge distrupting an extensive biogeographic province. This not only stifled direct inter

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Order Perissodactyla Family Equidae Merychippus sp. Order Artiodactyla Family Camelidae ?Oxydactylus Family ?Protoceratidae

Family Cervidae? Family Camelidae Procamelus sp. Order Proboscidea Family Bunomastodontidae Rhynchotherium sp . , cf . blicki Order Rodentia

22

23

THE CENTRAL AMERICAN DISPERSAL ROUTE

THE CENTRAL AMERICAN DISPERSAL ROUTE

Family Glyptodontidae

TABLE 2.1 Continued

Glyptodon Order Perissodactyla Family Equidae Equus sp. Order Artiodactyla Family Tayassuidae Family Cervidae Odocoileus sp.

Pleistocene

1. Comayagna Valley fauna 2. Honduras 3. Early Pleistocene 4 Webb ( 1972 ) 5. North and South America 6. Class Mammalia Order Proboscidea Order Notoungulata *

Family

Toxodontidae

1. Copan fauna 2. Honduras 3. Late Pleistocene 4 . Stirton and Gealey (1949) 5. North and South America 6. Class Mammalia Order Edentata Family Megatheriidae Megatherium Family Glyptodontidae Order Carnivora Family Felidae Felis concolor Order Perissodactyla Family Equidae Equus Order Artiodactyla Family Camelidae CameIops? Family Cervidae

Odocoileus? Order Proboscidea Order Notoungulata Family Toxodontidae

ft

-

K

-

.

K ,

1 . El Hatillo fauna 2. Pese, Panama 3. Late Pleistocene 4 Gazin ( 1957) 5. North and South America 6. Class Reptilia Order Chelonia Family Testudinidae Pseudemys sp. Class Aves Order Anseriformes Family Anatidae Cairina moschata Class Mammalia Order Edentata Family Megatheriidae Eremotherium rusconi Family Mylodontidae Cf . Glossotherium tropicorum Scelidoth erium?

1. Hormiguero 2. El Salvador 7

^

I

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lfA

Plpicfrvpnp

4 Stirton and Gealey ( 1949) 5. North and South America 6. Class Mammalia Order Edentata Family Megatherium Megatherium Order Carnivora Family Felidae

FIGURE 2.12. Fossil vertebrate localities in Central America , 1, Guanajuato; 2, Hidalgo; 3 El Gramal; 4 , Copan ( Coban ); 5, Gracias; 6, Comayagna valley; 7 , Hormiguero; 8, El Hatillo and La Coca; 9, Cucaracha .

.

Smilodon Order Artiodactyla Family Camelidae Family Bovidae

Quaternary faunas in Central America reflect an intermixing of elements from North and South America . This interchange began after a land connection was estabBison lished about 3 million years ago across the Bolivar Order Proboscidea Family Gomphotheriidae Trench , a seaway between the Atlantic and Pacific oceans Cuvieronius that cut through northwestern Colombia and eastern Family Elephantidae Panama ( Marshall et al. 1979). Mammuthus cf . jeffersonii By contrast , as far south as Panama the Central AmerTertiary ( pre- Pleistocene ) mammal faunas show no ican 1. La Coca fauna whatever with South America . Even the early affinity 2. Ocu , Panama Gaillard Cut local fauna ( = Cucaracha fauna ) Miocene 3. Late Pleistocene ), collected on the bank of the Panama Canal , ( fig . 2.13 ) ( 1957 4. Gazin consists of either five or six taxa , all congeneric with 5. North and South America North American forms. From just 400 km to the east in 6. Class Mammalia Order Edentata Colombia , across the Bolivar Trench , a late Miocene Family Megatheriidae fauna has been collected that consists of genera with Eremotherium rusconi exclusively South American affinities. Except for one Order Notoungulata site , this pattern that all Central American Tertiary terresFamily Tbxodontidae trial mammals are congeneric with North American speToxodon cies is consistent. The exception is the late Eocene or early Oligocene Guanajuato fuana of southern Mexico ) Besides those sites listed Stirton and Gealey ( 1949 which also happens to be the oldest known terrestrial bui mention nineteen Pleistocene sites in El Salvador vertebrate locality in Central America . ) give no further information . Rodriguez ( 1942 recordt The fauna ( fig . 2.14 ) is exceptional beGuanajuato hav < six sites in Costa Rica and one in Nicaragua that cause the two genera recognized are not known anywhere when yielded proboscidians , plus a number of others ®lse They also happen to be small mammals ( rodents ) , , isolated toxodont or horse remains have been found ! n contrast to every other Central American fossil mam the information he gives about these Pleistocene occur al ye* reported in the literature ( all are medium -sized to rences IS extremely scanty. Doubtless , if an effort were e forms). This suggests that the pi cm re of a strong made to search out the information , numerous other orth American affinity for the Central American Ter( of Pleistocene age could be found in Central America iary mammalian fauna may be an artifact of the proof site Janzen and Martin 1982 ) . One nonmammalian the! >n js more representative of ocean conditions than of day hours of sunshine at national parks (table 3.3). conditions.

Gramzow and Henry ( 1971 ) identified , through a pel tad analysis of rainfall data , the beginning, end , ai duration of the rainy season in various areas of Cent! America. According to them , the rainy season begi earlier on the Atlantic coast and later on the Pacific coas An exception is the northern coast of Honduras , whei the rainy season begins in July. In Costa Rica the rain season starts in late April on the Atlantic side and by th end of May on the Peninsula of Nicoya . The end of the rainy season ( generally in October) comes earlier in the interior portions of southern Honduras and western Nicaragua than in other parts ot Central America . In Cost Rica the rainy season ends in November on the Pacifi coastal plain and in January on the Atlantic side. Th period in which veranillos occur (July-August ) is poorl defined from the Pacific coast of Panama to the Atlanti inland coast of Nicaragua . The highlands of Costa Rica al If we make 5 = 5 , then ( ร IS ) = 1 , a value we could have a poorly defined veranillo. obtain only on a cloudless day such as during the Guana caste dry season. Here , a plus b will not necessarily be Some Peculiarities of Costa Rican Climate to one , as Fritz ( 1951 ) and other authors proposed , equal SOLAR RADIATION Q is not the value proposed by Angstrom because The amount of solar radiation received at any point on the Smithsonian 1963, p. 440); Qa is the amount of radiation ( surface of the earth depends on factors such as latitude, that would reach the surface of the sea in the absence of time, day and year, aspect and slope of the ground , el- an atmosphere , and Q is the average solar radiation that ges evation , proximity of oceans and high mountain ran| the site s reaches horizontal on the average for all cloud thickness and height , and atmospheric compo days. Thus, a plus b will besurface equal to the transmission tion . This means that an Angstrom equation of the foi coefficient , better called “atmospheric transmissivity ” QIQo = + b ( s /S ) ( Huschke 1959, pp. 589-90) , at the site in question . For reasons b must change with the elevation of the site. is a model for a local site and the constants a and b mu these In 3.2 we can see values of a and b for eight table be determined from local data. In equation ( 1 ), Q i stations at different for the year 1973. The deobserved solar radiation , Q0 is Angot’ร value or the the pendence of b on thealtitudes elevation oretical amount of radiation that would reach the surfaci with the following equation: H ( in km ) is expressed of the earth in the absence of an atmosphere (Chan) 1968 , pp. 8-9) , 5 is the observed duration of sunshine b = 0.36 + 0.056 // , and 5 is the maximum possible duration of sunshine with a coefficient of determination (r of 0.77 , excluding In places like Costa Rica , where air pollution is not ye Port Limon . Port Limon is not appropriate the model; severe outside San Jose except during dry -season pastured it has the latest value for b at sea levelfor . เท contrast , burning , the average values may be useful. For eighl Cobal, several kilometers inland at sea level on the Atstations in the country' ( table 3.2 ) , the averages , with lantic side, has the lowest value for b , which suggests their corresponding standard deviations of the constant a that clouds that develop on the Atlantic coast are pushed and b , are inland during the day by the winds. This phenomenon 0.04 occurs when the alisios ( reinforced trade winds) are K = 0.27 ss..dd.. = 0.09 strengthened by the sea breeze, producing fewer clouds = . xh = 0.42 the day than during the night on the Atlantic coast . If we make 5 = 0 in equation ( 1), then Xa is the Iran 1during )ก the Pacific coast the sea oppose the alisios, mission coefficient of an overcast day in Costa Rica. Th and cloudy skies occur morebreezes frequently is in agreement with the value of 0.25 given by Haurwi than during the night. This agrees with during the day the small value for ( 1948 ) for stratus clouds and an optical air mass of 1.1 b at the Port of Puntarenas . a The optical air mass at latitude 10° N at noon is 1.06 , the “ Angstrom equation ” for Costa Rica , except an average for the year. The value of a = 0.27 is also i forThus the edge of the Atlantic coast , takes the follow agreement with the valued = 0.29 cos d), where d> is th ing formvery : latitude (Chang 1968 , pp. 8-9).

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0

^

1.1

TEMPERATURES

Because Costa Rica is in the equatorial zone , the average temperature of the warmest month does not exceed the average temperature of the coolest month by more than 5 ° c at a given site. Figures 3.4 and 3.5 represent the vari ation of maximum and minimum temperatures through the year at different elevations. The data in table 3.4 were used to draw the isotherms. It is interesting to note that , as elevation increases , the difference between the mean maximum and mean min imum monthly temperatures decreases. At sea level the average difference is approximately 10° c on both slopes, *

Maximum differences occur on the Pacific slope during the dry season (cloudless nights and leafless vegetation allow maximum radiation of heat from the ground , and during the day the same factors lead to maximum heating of the substrate ). In all parts of the country, the coolest month may be November, December, or January; the warmest month is in March , April , or May. The small variation in the average temperature throughout the year is in striking contrast to the large diurnal variation. This variation decreases with increasing elevation ( figs. 3.4, 3.5). The largest changes of temperature during the day occur in some valleys where the cold winds draining off the mountains lower the minimum daily temperatures, Examples of this phenomenon are the low values for the minimum temperatures observed in Cachf and Naranjo ( table 3.5), which do not correspond to the temperatures expected for these elevations (fig . 3.4). The same thing may be happening at Volcan Buenos Aires and Cobal , at elevations of 450 m and 55 m , respectively, where min imum values are exceptionally low. Each station is at the foot of a large mountain mass. Average temperatures for the monthly maximum are higher on the Pacific coast than on the Atlantic coast at the same elevations (fig . 3.5). For the Pacific, the mean

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39

CLIMATE CLIMATE

TABLE 3.3

Num1

ber

Estimated Daily Hours of Sunshine in the National Parks of Costa Rica

Location P. N. Santa Rosa M.N. Guayabo P. N. Rincon de la Vieja P. N. Volean Poas P. N. Chirripo R . B. Islas del Golfo R.N. Cabo Blanco R . N. Manuel Antonio P. N. Tortuguero P. N. Corcovado P. N. Cahuita P. N. Barra Honda

Maximum possible hours of

Hb 0.08 1.10 1.50 2.๓

Jan. Feb. Mar. 8.7 9.7 9.5 4.0 4.9 5.0 7.6 8.5 8.5 6.0 7.0 6.6

Apr. May June July 9.3 6.7 5.5 5.4 4.4 4.1 3.0 3.1 8.0 5.7 4.0 4.4 6.4 4.6 4.4 3.9 4.5 5.4 4.0 3.0 2.4

2.50 4.3 5.0 0.01 8.5 9.6 8.8 0.01 8.5 8.4 9.0 0.๓ 6.6 8.1 7.1 0.๓ 4.4 4.5 5.4 0.02 6.5 8.1 6.1 0.๓ 4.7 6.2 6.1 0.๓ 8.4 9.5 8.8

sunshine at 10° N on the

9.2 6.7 5.1 5.2 9.2 6.8 4.0 5.3 7.3 6.6 2.0 3.3 5.7 4.4 3.9 2.5 6.2 5.9 3.0 4.2 5.8 5.4 4.7 4.0 8.7 6.2 5.0 5.4

Aug. 5.7 3.8 4.9

Sep. 5.0 3.2

4.4

Average for Oct. Nov. Dec. Year 5.2 6.5 8.2 7.1 3.5 3.9 4.2 3.9 4.0 5.6 6.8 6.0

3.9 3.4 3.8 3.1 3.0 3.0 6.1 5.0 5.1 6.2 4.8 5.0 3.0 4.1 5.0 5.0 4.6 4.4 4.0 4.2 4.0 4.4 4.3 5.0 5.8 4.7 5.0

3.9 5.4 4.0 4.0 6.1 7.6 5.9 7.3 4.9 6.1 4.0 4.4 4.4 5.3 4.5 4.6 6.0 8.0

4.9 3.8 6.9 6.7 5.3 4.4 5.2 5.0 6.8

11.6 11.8 12.0 12.4 12.6 12.7 12.7 12.5 12.2 11.9 11.7 11.5 Unpublished maps of sunshine for Costa Rica by E. Coen 1977. a See figure 3.3 b Average elevation above mean sea level , in km .

15th day of each month

SOURCE:

COSTA FICA

National Parks and National Reservations

III o

Sunshine Stations

.

Solar Radiation Recorders

E

.

Coen

.

PACIFIC SLOPE

.

ATLANTIC SLOPE

.

1978

FIGURE 3.3. Location of sunshine and solar radiation recording stations in Costa Rica. value at sea level is about 32.6° c, but on the Atlantic the mean is 29.9° . These observations agree with the hy pothesis that there are fewer clouds on the Pacific side during the dry season , allowing more solar radiation to reach the ground. On the Atlantic side the dry season is less pronounced , and there are more cloudy days during the year. These cloudy days depress the average monthly maximum temperatures . Mean annual temperatures decrease with elevation at a for for the maximum and at 5.2° rate of 6.5๐ the minimum temperatures . Consequently, in Costa Rica the minimum and maximum average annual temperatures for the highest peak ( Chirripo) are about 2.2 ° c and 7.2 ° respectively ( 3,819 m elevation ). With these tem peratures frost is frequent, but there is no permanent ice in the soil or on ponds; nevertheless , forklore has it that snow has been seen once on Chirripo and Cerro de la

c

c/km

c

Muerte .

c/km

Terrestrial radiation during the night depends to a lai degree on local conditions. For a given site the minimi temperatures are more uniform through the year than the maximums ( fig . 3.4). However, lack of cloud cove during the dry season also results in cooler nights th during the rainy season . Below elevations of 800 m on the Pacific side of G Rica , the mean monthly maximum temperatures are highest of the year during February, March , and A Between 800 m and 1 ,000 m the decrease in the m imum temperatures is very large ( Coen 1970, 66-77). This phenomenon , found on the Pacific slope 10 ° N in Costa Rica , occurs on all the Pacific slopes the mountains of Central America. It may in part caused by the papa gayo winds or nortes described earli These winds bring a cold , humid air mass over the mi tains and later bring a descending dry' wind that warms uf the site.

7000

1000

FIGURE 3.4. Isotherms of mean monthly minimum temperatures for parallel 10°N through

Costa Rica ( 1970- 73).

40 41

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and dugongs. J.Mamm.16:216-20. group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft .( Berlin ) 35:247-325. .Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthe mana behavior and Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of )inFlorida ( .Special Publicamanatus Trichechus tee tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ).ร (พ( Fish . Wildl .Res .Rept .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, Vampyrum spectrum screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) )do not fan their seems a distinct possibility. It is clear that these bats have ,and Diphylla ,Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

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two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull .Am.Mus . Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J.Mamm.49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vampyrum spectrum , the false vampire bat , in Panama. Can .J.Zool .47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla alba and Vampyrumof the Neotropicalthecarnivorous bat , Vampyrum spectrum ,in Costa Rica. J.Mamm spec .44:186-89. trum .J.Mamm . 58:469-78.

-

-

-

-

**

-

-

-

iN

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but 8 March. problem continues. BaiTett , o. 1935. Notes concerning manatees and 16:216-20. dugongs. Frantzius, A . von. 1869. Die saugethiere Costaricas. ( Berlin ) 35:247-325. Hartman , D. 1979. Special Publication no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee

ร (พ( 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%.

;

Vampyrum spectrum ( Vampiro Falso, False Vampire Bat)

is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate mix larger in and forms such as sects with small vertebrates such as lizards, and the large favors substantial vertebrates such as birds , are strik rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the largest and most fierce- looking species must be “ the vampire. One reported that individuals of fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated structures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vamdo not fan their and pires victims and lap blood instead of suck it. roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat species avoid these trees , with the exception of which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling **

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats can somefeed every night , that a foraging times fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. for fear that some small museum will feel the need bats of Trinidad and Tobago. M. 1% 1. A review . complete its skin collection at the expense of the remnantto of the 122 301 . populations. Fieldworkers are encouraged to observe and Greenhall , A: 191 . M. 1%8. on the behavior of the enjoy these large bats, but unnecessary disturbance and false vampire bat. Notes 49 337-40. collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1:% 9. Notes on the false vampire bat , in Panama . 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging phyllostomid bats , and avian prey and of the Neotropical carnivorousbehavior , bat in Costa Rica. 44:186-89. 58:469 78.

-

-

-

**

501

CLIMATE

-

.

de las horas merits in tropical life zones: A pilot study. Oxford : Per . 1977. Mapas del promedio mensual manuscript gamon Press. de brillo solar en Costa Rica. Unpublished Huschke, R . E. 1959. Glossary of meteorology. Boston: available from E. Coen . 1 - American Meterological Society. Daubenmire , R . 1972. Phenology and other character higher are เท passes Why mountain . 7๙ % 1 . , . H D Janzen north in forest deciduous semi istics of tropical . 49 : . 233 Nat 101 . Am . -of sexual reproduction the tropics western Costa Rica . J . Ecol. 60:147-70. . 1967/?.theSynchronization Dunn . G . E. , and Miller, B. 1. 1960. Atlantic hurricanes . of trees with dry season in Central America. EvoBaton Rouge: Louisiana State University Press. :620-37. Escoto, J . A. V. 1964 . Weather and climate of Mexico lution. 21 1973. Sweep samples of tropical foliage insects: and Central America . In Handbook of Middle Ameri Effects of seasons , vegetation types , elevation , time of can Indians , vol . 1 , Namral environment and early , and insularity. Ecology 54:687-708. day : Uni Austin . 215 , 187 . pp cultures , ed . R . c. West . 1976๙. The microclimate differences between versity of Texas Press. forest and adjacent riparian forest in deciduous Frankie , G . พ. 1975. Tropical forest phenology and pol - a , Costa Rica. Brenesia 8:29-33. Province Guanacaste animals of linator plant coevolution . In Coexolution samples of tropical deciduous p Sweep ? / . 1976 . , pp . Raven . . H and E . Gilbert and plants , ed . L . insects : Seasonal changes and inhabiting forest foliage . 192-209. Austin: University of Texas Press p in differences field inter Frankie, G. พ.; Baker, H . G .; and Opler, . A . 1974๙ . Biol.- Trop. 24:149-61. adult bugs and beetles. Rev. Comparative phenological studies of trees in tropical in . T. พ. 1968. Differences wet and dry forests in the lowlands of Costa Rica. J . Janzen , D. H . , and Schoener wetter and between diversity and insect abundance Ecol. 62:881-919. drier sites during a tropical dry season . Ecology . 1974/?. Tropical plant phenology; Applications for studies in community ecology. In Phenology and 49:%-110, .M . 1973. Distribut ion de la precipitat ion seasonality modeling , ed . H . Lieth , pp . 287-%. Ber- Mendizabal con la altura . Ciudad Universitaria: Escuela de Fisica lin: Springer- Verlag. Universidad de Costa Rica . Fritz, ร. 1951 . Solar radiation energy and its modifica- Opler , p. A .; Frankie , G . พ.; and Baker, H . G. 19 6 Compendium . In atmosphere tion by the earth and its Rainfall as a factor in the release , timing , and syuJ cf meteorology, pp . 13-33. Boston: American Meteochronization of anthesis by tropical trees and shrubs. rological Society. . 3:231 -36. Biogeog in exchange Garstang, M . 1 %8. The role of momentum . Comparative phenological studies of 1980 . State Florida : Tallahassee . flow over a heated island shrubs and treelets in wet and dry forests in the lowUniversity. Rica. J . Ecol. 68:167-88. Gentry, A . H . 1974. Flowering phenology and diversity lands, ofJ. Costa en el Atlantico . Sistemas sinopticos: Escuela 1973 . M Placido . 68 : in tropical Bignoniaceae. Btotropica 6 64de Fis Universitaria Ciudad . tropical None rainy The . 1971 พ . , . K Henry , Gramzow, R . H. and . Rica de Costa ica , Universidad seasons cf Central America. College Station: Texas Portig, พ. H . 1%5. Central American rainfall. Geogr A and M University. :68-90. Hastenrath , ร. 1%6. On general circulation and energy Rev., 55 p . B. , and Borchert , R . 1982. Phenology an( Reich . seas American Central the of area budgets in the ecophysiology of the tropical tree , Tabebuia neo J . Atmos. Sci . 23:694 -711 . ( Bignoniaceae ). Ecology 63:294-99. chrysantha in Cen . 1%7. Rainfall distribution and regime , -. Smithsonian Institution . 1 %3. Meterological tables . 6tn ser . Bioklim . Geophys . tral America. ArchiV . Meteorol , ed. Washington D.c.ะ Smithsonian Institution. B , 15:201 -41. . Ecology, flowering phenology an L _ . Transmision of solar radiation Stiles , F. G . 1975 Haurwitz, B. 1948 Costa Rican Hat pollination hummingbird ; re : . 110 5 - conia species. Ecology 56of:285some through clouds (overcast ). J . Meteorol . 310 , Washing tables meteorological printed in Smithsonian : The flowering competition Coadapted . 1977 . . Institution Smithsonian , . . : C ton D in a tropical plants pollinated hummingbird of seasons - 78. Heithaus, E. R . 1979. Community structure of Neo1177 : 198 Science . forest and Diversity : wasps tropical flower visiting bees and . 1978. Temporal organization of flowering phenology. Ecology 60:190-202. foodplants of a tropical wel Holdridge , L . R .; Grenke , พ. c.; Hathaway, พ. H .; among the hummingbird 194 : 10 Biotropica . forest -210. Liang , T.; and Tosi , J . A . , Jr. 1971. Forest environ^

46

4 GEOLOGY

R . Castillo- Munoz

...

-

The appraisal of Costa Rican geology made here is re marilv

-

arse ’ medium and fine-grained sandstone, ’, °calcareous logical map (fig. 4.1) and a table of stratigraphic re- siltstone shales , and limestone intercalated „ lationships of the principal formations and units that ha VP ,;,น .,„ rOCk y oeeur ogether in a de torn defined to date (fig 4.2a,โ), } and have been assigned 11 scnption of each of the principal formations and its o aceous (Campaman -Maestrichtian ) whereabouts , and describes the tectonic structure and p 28)‘ 8 evolution experienced by the territory of Costa Rica as This undifferentiated sequence is represented by forevidenced by the known geological characteristics. mations such as the Sabana Grande and Rivas formations that outcrop in Guanacaste and on the Nicoya Peninsula , 31, 1 die Golfito and Changuinola formations that are ex _ .. posed in the southeast part of the country (figs. 4 V 4.2 ).Geolo_ jmd. Stratigraphy units constituting the territory ;of The Sabana Grande Formation , of Santonian age range ag_e Ju! Sic tOQuatemary < fig (Galli and Schmidt 1977 ) consists principally of cherts .น e y g n H i p asjfollows by bdiology and and siliceous limestone , and the overlain Rivas For* sedimentary rocks (Jurassic mation of Santonian P. volcanic and age:;Mesozoic - Maestrichtian age (Galli and Cretaceous); Tertiary sedimentary and volcanic rocks; Schmidt 1977) is principally made up of layers of sand Cretaceous and Tertiary plutonic rocks; Quaternary sedi- stone , shale , and some limestone beds. The two formentary and volcanic rocks. mations are separated from each other by an uncon formity. MESOZOIC VOLCANIC AND SEDIMENTARY ROCKS Outcrops of the Sabana Grande Formation are seen at (JURASSIC-CRETACEOUS) the site bearing that name on the Nicoya Peninsula , at Volcanic Rocks ( Mzvs ) Guatil , along the road between Nicoya Santa Ana to Mesozoic volcanic rocks are spilitic basalts, agglomer- the northwest of Santa Cruz, and in theand Cerros de Morates, basalt , and radiolarite breccia together with sedi - acia; in the Nicoya Gulf , on Berrugate island; near Haci mentary rocks of dark gray graywacke, cherts, cherty enda El Viejo, in the Cerros Peor es Nada and near Sanshales, and aphanitic siliceous limestone. They are Lazaro ( Dengo 1%2/? ); outcrops are also found in the strongly folded together and poorly weathered . This se eastern part of the Cerro de Jesus south of Vigia ( Macquence is penetrated by intrusions of gabbro, diabase-, Donald 1920). and diorite ( Dengo 1 %2/? ). The Rivas Formation has outcrops in the northern part This lithological association forms the Nicoya Com - of the Santa Elena Peninsula , in contact with the Santa plex , which outcrops extensively over the Nicoya Pen - Elena Peridotite, and in various parts of the road that insula, in a small part of the Santa Elena Peninsula, in connects Murcielago with Playa Blanca; it is also seen in Punta Herradura and sites immediately to the east , in the Santa Elena Bay, in the road Santa Ana and Osa Peninsula , and in some parts of Punta Burica (figs. Corralillo, in the central part ofbetween the Nicoya Peninsula , 41 » 4.2). and in the valley of the Rio Tempisque near Matina and The rocks of the Nicoya Complex vary in age Rio Blanco, and in Punta Quesera ( Dengo 1 %2/? ). Jurassic ( Middle Titonian; Galli 1977 ) to Upper from Cre- Dengo ( 1 %8 ) used the term Golfito Formation to destaceous (Campanian - Maestrichtian ), according to Hen - ignate a formation that irregularly overlies the Nicoya mngsen and Weyl ( 1 %7). More recently, an Upper San . This formation consists, from bottom to top, tonian age has been considered for these rocks (Galli and- ofComplex greenish gray siliceous limestone , conglomerates , Schmidt 1977 ) , with an isotope date of 72.5 million years shales, and siltstone , with occasional beds of stratified ( Barr and Escalante 1 %9) . gray brownish sandstone with beds of siliceous lime stone. This formation appears in the area of Golfito Bay,'ๆ Mesozoic Undifferentiated Sedimentary Rocks ( Mzsd เท the hills between Golfito and the railway station at El The Mesozoic undifferentiated sedimentary rocks are pri- Alto A , and in Punta Banco on the Burica Peninsula. There

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47

GEOLOGY

GEOLOGY

good section between the exit of Golfito Bay and Punta Gallardo. , as named and described Xhe Changuinola Formation ( 1965) , is represented by limePessagno and by Fisher , grayish to clear greenish , and clear stone, usually white 1

g

, with top layers of interstratified anreenishlavayellow and dacites and clastic sediments. These

, desitic conspicuous very are rocks

at the type locality in the

Changuinola River and its principal tributary, the Rio

Pvmntuto do

Pena Blanca in the northwestern part of Panama . They are also believed to occur in southeastern Costa Rica (Dengo 1968 , p. 28). However, Fisher and Pessagno ( 1965) have found rocks similar to the Changuinola Formation in the Rio Lari and think they are assignable to Tertiary sedimentary rocks, specifically Paleocene and Middle Eocene. The Upper Cretaceous sedimentary rocks in the Nicoya Peninsula are due to volcanic detritus derived primarily from the erosion of the Nicoya Complex; how ever, these sediments also contain contemporary volcanic detritus and rocks , which indicates concomitant volcanic activity. In the same manner, the Changuinola Formation contains tuffaceous sandstone and intercalated andesitic lava and dacite.

พ' coy a

OCEANO PACIFICO

TERTIARY SEDIMENTARY AND VOLCANIC ROCKS

Tertiary Sedimentary Rocks ( Ts ) GEOLOGICAL MAP OF COSTA RICA

QUATERNARY

—

SEDIMENTARY AND VOLCANIC ROCKS

Rocont alluvial ond marina tedimentt along I ho coaola mudflomt Terracet ( pottibty Plaltlocana ) including

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TERTIARY AND CRETACEOUS PLUTONIC ROCKS Tertiary intrutiva recto , mainly dianla , granodiorita and granite mainly diarita and diobote Croiacoout mtruoivo rockk

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TERTIARY AND MES 020tC SEDIMENTARY I ( upper Miocene Pliocenei Ppiot au ;0"' C Tor I ary volcanic rock igiMmPrlit volcanic activity ) : 0 0 (โท 1 t u f f t , tedimont * lerretlrial and lacuttrine volcanic

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Dengo et al . 1969). ( FIGURE 4.1 . Geological map ot Costa Rica from

The Tertiary sedimentary rocks are represented prin cipally by marine sediments and some continental sedi ments with intercalated volcanic rocks. These rocks , which vary in age from the Paleocene to Pliocene , appear all over the Costa Rican territory and especially in the central part of the country, such as the Pacific and Caribbean slopes (fig . 4.1 ). Figure 4.2 gives the names that have been assigned to the diverse units of these rocks in different parts of the country. Each of these formations has diverse lithological characteristics that are generally associated with rudites , sandstones, and claystones, in diverse proportions and variably interstratified , and containing a major proportion of calcareous and volcanic materials. The rocks formed in marine depositional environ ments, which are the predominant ones , are sandstones , siltstones , shales , and claystones , some tuffaceous in character, and limestone , all of which display diverse degrees of intercalation and outcrop in various parts of the country. The intercalated volcanic rocks , also formed in a marine habitat , were produced by various degrees of volcanic activity that influenced the sedimentary rocks as well as the continental ones . The continental Tertiary sedimentary rocks are primarily conglomerates devel ped on the margins of the Cordillera de Talamanca. The volcanic rocks are those formed during the Lower Tertiary and appear in diverse parts of the country. In

°

association with sedimentary rocks of the Eocene and some of the Oligocene, there have been found tuffs , agglomerates , andesite and basalt flows , and some di abase sills ( Dengo 1962a , pp. 142-43). In this respect , the Tuis Formation on the Caribbean side contains numerous intercalations of andesite, basalt , and tuffaceous beds ( Dengo 1968 , p. 28). The Barra Honda Formation is made up of stratified limestones and is found among the Paleocene sedimen tary rocks on the Pacific side with extensions to the Upper Cretaceous ( Maestrichtian ). This unit , according to Dengo ( 1962b ) outcrops in Guanacaste in the Cerros de Quebrada Honda , Corralillo, Quebrada Honda , Copal , Caballito, Corral de Piedra and La Cueva on the west side of the Rio Tempisque , and in some small hills on the east side of the river. The lower contact of this formation is unconformable with the Rivas Formation . Las Palmas Formation occurs in the extreme southeast portion of the Nicoya Peninsula and is Upper Paleocene in age , or possibly Lower Eocene. It is made up of thin , hard layers of siltstone and shale , well stratified , and of lenticular masses of coral reef limestone . Important ex amples of this formation occur 1 km from Playa Naranjo on the road to Bajo Negro; it also occurs on the Rio Nacaome , to the west of Puerto Letras in the Punta Morales ( Dengo 1962b ) The upper contact of this formation is possibly concordant with the Brito Formation , and it is not concordant with the Barra Honda Formation ( Dengo 1962b 1 p. 27) It is agreed that the Eocene sedimentary rocks on the Pacific side of Costa Rica , represented primarily by the Brito Formation , are primarily clastic. However, they also contain extensive coral reef members , assigned to the Upper Eocene , which extend from Guanacaste Prov ince, with the name Colorado Member ( MacDonald 1920; Dengo 1962b ) to the southeastern part of the country along with the limestone of the Fila de Cal . This formation is known in Panama as the David Formation ( Dengo 1968 , p. 28). The Parritilla Limestone ( Caliza de Parritilla ) from the Upper Eocene is an organic limestone , dark gray to blackish and at times very fine grained , 60 m thick , with lai e foraminifers and calcareous algae ( Malavassi 1966 ) . It outcrops on the south and west flanks of the Cerro Caraigres , situated on the south bank of the Rio Candelaria , Parritilla de Acosta ( type locality ) , Tiquires , Escuadra , and Cangrejal . According to my observations this limestone is stratigraphically below the purple facies of the Pacacua Formation in the Cerro Caraigres, but the type of stratigraphy is unknown . Recently, Castillo ( 1981 ) has observed various limestone outcrops between Cangrejal and Escuadra that suggest that existence of more than one limestone layer intercalated with rather clastic units of unknown ages.

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49 48

GEOLOGY

GEOLOGY

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CUAORO OE CORRELACION GEOCRONOLOGICA Y

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Eocene sedimentary rocks on the Caribbean side of the country are represented by the Tuis Formation and the Las Animas Limestone (Caliza de las Animas ) . The Tuis Formation is represented in the Turrialba area by a sequence of sandstones and tuffaceous siltstones and claystones intercalated with basalt lava flows , conglom cr3tcs and volcanic breccias and reaching more than 3 ,000 m in thickness ( Riviere 1973 , p. 50) . Although it outcrops conspicuously in Tuis of Turrialba , the type locality, the extension , and contacts with other formations are not known .

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ALUVIONES, LAMARES , SEDIMENTOS FLUVIOL ACUSTRES DEL VALLE OE CARTAGO Y ROCAS VOLCANICAS ^ ^ BARBA S L NO DIFERENCIADAS ^ ^rCERVANTEs [ AQUIARES 1 1 IRAZU LAVAS •* •* TIRIBI IRAZU • • • 3 v NT PO 7 COLIMA - -* * PARAlso UJARRAZ

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( VALLE Y CORDILLERA CENTRAL , CERROS OE ESCA 2U Y BUSTAMANTE ) PUCNTC : DONOOLI V TORRES , 1954 ; ESCALANTE , 1* 6 «; CASTILLO , I t s ; KRUSHENSKY, 1 72 ; * NACIONES UNIOAS , 1975 ; RIVIER , 19 *0. *

*

.

TERCIARIO

PARTE CENTRAL DE COSTA RICA

RELACIONES ESTRATIGRAFICAS COSTA RICA VERTIENTE DEL PACIFICO Y SUR REGION DE GUANACASTE ( DesdeCENTRAL Puntarenos a Burica, VERTIENTE DEL CARIBE y Guanacoste de Prov ( incluyendo Herraduro , Pe Y DEL RIO SAN JUAN SERIE Peninsula de Nicoya ) กเ*กรน la de Osa , Golfiio y ALUVlONCS SEQIMCNTOS

CUATERNARIO

CORRELACION ESTRATIGRAFICA Y GEOCRONOLOGICA

••

4

Y

ORIENTAL

-

Stgun Nacionci Unitfas S« nos ( 1975) ร* gun Oondoli y Torres ( 1954 ) ร*gun Krulhtniky ( 1972 )

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of the southwest and east of Santa Ana In the central part of the country sedimentary r all Ciudad Colon to east f. ESCaZ.d/ 11,6 loWer Contact ๙ ° outcrops that have been assigned to the Eocene . i s n Ljiblt ๒'s Part ๙ the country. grouped under the names Pacacua Formation ( R* e 80 , p l 27 ) ei a Lower Mio' conglomeratesof 1969) and Caliza de Parritilla ( Malavassi 1966 ) . ใเ the Pacacua Formation . up The Pacacua Formation is a sequence made A " ใ 01ใ le itho gic including the 9 coi uniL finer ' facies of " interstratifications of breccia conglomerates and 11 . . 1 , . ใ ใ ’ was assigned fo the Upper Mio* glomentic sandstones , sandstones , siltstones , and shal« cen ° ใ1 a"ubasshale s of paleontologic evidence . Castillo ' , all tuffaceous; in some places they have a characteristl 1 " ใdiatthe erroneous Eocenic age of ml purple color, and the thickness is as great as 1 , 148 the p. 1 Po at,’apparently co d he accentuated by the litho'initially defined ° “ the type locality on the north flank of the Cerro Pacac loei unit " 1 ใ by Castillo in 1 %5 as “ Rfo e f In general this formation ranges from the south and 1

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Cafias Unit" (considered in Castillo 1969 as the Terraha Formation) , which is represented by a sequence of black shales , siltstones , and fine sandstone beds outcropping on the Rfo Cafias , southeast of Aserrf and is stratigraphically underlain by the San Miguel Limestone of Lower to Middle Miocene age; this unit was subsequently cor related in age with the Oligocene-Lower Miocene Ter-raba Formation only on the basis of lithologic consid erations ( Dengo , pers . comm . ). Therefore Castillo ( 1981 ) considers it likely that a major part of the sequence of black shales , siltstone , and fine sandstone beds out51

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GEOLOGY

formation ha* cropping in various parts of the area studied by Castillo which locally grades into limestone, .pThis a section of I in ) . 2 ( 1967 Malavassi by been reported in 1965 is the lithologic equivalent of the “ Rio Cartas the . It is Peralta and Animas Las , in Reventazon Rio well be considered as the Rio Cartas ”

GEOLOGY

may also be observed in villages such as Cdbano San volcanic rocks of the Aguacate Formation and by Malpais. Its lower contact is unconform - Quaternary volcanic and sedimentary rocks. Isidro, androcks of the Nicoya Complex , and no upper The sedimentary Miocene and Pliocene rocks on the with able , which could บแน่ , because there are no younger formations Caribbean side are represented by the Uscari ( described , breed sandstones exists sandstone calcareous ๙ contact. lithologic character- made up Formation because of Its with it limestones organic above), Gatlin , Venado, and Suretka formations. The , and over limestones isdcs , Formation named to by Dengo possible , ( Curre 19626 been ) is of not Gatlin has . Formation , described by Olsson ( 1922) ( Hoff It The shales , and The Las Animas Limestone was recognized by Sjogren algae age Costa Miocene Upper in and made outcrops RicaJ to up primarily , of formation stetter Dengo, and Weyl I 960, pp. 254-56), is Middle this where Middle sandstones, medium grained , stratified , (Hill 1898; Hoffstetter, Dengo, and Weyl 1960 pp. precisely are I formations other with with not mi in age. In Costa Rica it is made up primarily of Miocene contacts its of nature mottled 258-60); its type locality IS Flacienda Las Animas in The . of conglomerates and shales . These intercalations sandstones and some layers of lignite and conglomerates , nor known Tumalba in the Rio Reventazdn valley. It is a sequence The บscari Formation , recognized by Olsson ( 1922M rocks reach a thickness of 830 m and outcrop in typical It also has coral facies made up of coraline limestone and made up of two layers of whitish limestone separated by , Terraba below Ri Miocend the Paso Lower the Real to in Oligocene Formation Upper fossiliferous . It is encountered locally on the coast , marl from form age in layers of sandstone and conglomerate . The limestone is extends ) Valle below between Curre the in and de I Creek Escuadra . Uscari the at especially is in described Puerto locality ( Limon (Olsson 1922). It extends the type thin and fossil-bearing and also outcrops in the area of Azul , ot Charco Formation shales soft described of up by Terry darldl The made along some is primarily parts . of It the Talamanca , Caribbean coast to the east of the Azul and on the Chitana River ( Ddndoli and Torres 1954 ) in Panama , is Pliocene in age. On the Costa Rican Cordillera de Talamanca , and it seems to extend up to the states ) 64 , . 363 pp 1923 1956 ( , ( Redfield . However tones pp 41 -42) . side it is made up primarily of greenish gray shales with Valle del Rio Reventazon ( Branson 1928) เท the area of , beside shales , contains intercalations of On the central and southern parts of the Pacific side of the formation intercalations (Madrigal 1979, pers . comm . ). Tumalba Rio Bonilla , and Las Lomas. Some authors majfl thickness . Its sandstones sandstone calcareous and limestone , Costa Rica there are Oligocene to Lower Miocene rocks is typically develops! on It crops out in Punta Burica and in the Osa Peninsula think that it unconformably overlies the Uscari Formation represented by black shales of the Terraba Formation attain 1 ,500 m . The formation ! along the Rio Tigre in the extreme southern portion of the (Olsson 1922; Woodring 1928, p. 69), while others, such de Talamanca . appeal Cordillera the of east and north the ( Dengo 1961 , p. 48 ) , which suggests a semieuxinic envi ( of the Rio Reventazdn Hofl peninsula and on its eastern coastal margins. It lies un- as Dengo, consider it concordant with this last formation ronment owing to the lack of external communication of ing to attain ,the valley (Hoffstetter, Dengo, and Weyl 1960, p. 254 ). Weyl 1960 , p. 293) and in the area of I conformably over rocks of the Nicoya Complex . the sedimentation basin. These rocks appear extensively stetter, Dengo and, Tres ( Mai . A Sedimentary rocks เท the Pavones central , part and of the Equis country The Miocene Venado Formation , described by Mal , Chitaria Peralta , and in the Cordillera Costerta , in the Valle de El General iguel ® 1 Li s ( or พ Caliza de 3 ร 1 Mi avassi and Madrigal ( 1970, pp. 5-6), is made up of a . ) , . 34 p 1967 . In * " ; * " the western foothills of the Cordillera de Talamanca 11 ) and the Turrucares and Cons formations. The San sequence of limestone , calcareous sandstones , shales guel by recognized , Formation Machuca The the the northwest portion of the country, in Guanacaste up of shales and I Miguel Limestone , named by Romanes ( 1912a ), is a siltstones and sandstones , and layers of lignite. It outOligocene rocks (Upper Oligocene ) are represented by ( 1899), is Oligocene in age and is made are green , silicifie* gray bioclastic limestone, 5 to 15 m thick and poorly crops between Venado, Delicias, and Jicarito de San Carthe Masachapa Formation , which is constituted of fine sandstones. These ,sandstones hard , and piritized and they lack fossils ( MalavasB stratified . It is dens generally hard with irregular joint- los. In the vicinity of Venado it contains caves produced coarse- to fine-grained sandstones, some calcareous gray veryMadrigal 1970, p. 6). The formation outcrops in the ing and contains small fractures filled with calcite and by kartzic erosion. The formation is unconformably oversiltstones , calcareous shales , shaly limestones , and sand - and between El Castillo Viejo and abundant fossils This limestone is found primarily in the lain by the Cote Formation and possibly by basalts asstones some fossiliferous. According to Dengo ( 19626 ) , basin of the Rio San Juan , In east and south of^ Patarra and in Loma Salitral and Loma signed to the Aguacate Formation ( Malavassi and Mad . Castillo Viejo it is)overlaidl these rocks are found on the west side of the Nicoya the Rio Crucita del Norte ls limestone has been dated as Late Lower to rigal 1970, p. 5). The lower contact is unknown , , butfl 6 , . and Madrigal 1970 p Peninsula in two places: one in the lower part of the by basalt ( Malavassi ( Carballo and Fischer 1978 ) and overMiddle Miocene Canbbeafl the of The Pliocene Suretka Formation , described by Sapper' section from the Rio Nosara to Punta Peladas , and the relationships with the other formations TSrraba Formation . Locally is it . conformably ( ) ( Hoffstetter, Dengo, and Weyl 1960 pp 1905 unknown yet as are slope de Pefion to other from themouth of the Rio Manzanillo พ' by the Coris Formation and also Pliocene and unconformably , 288 ), derives its name from an abandoned village 90 Miocene side Pacific the On they because Ano. The type of contacts are unknown rlam in other areas by , Montezuma the proper Coris Carballo Punta Formation the , on the north bank of the Rio Sixaola. It by called Suretka represented are rocks . mentioned have not been examined in the places ( sh*ns > เ p Carba eomm ) . Punta The . > formations is formed Azul by a Charco , conglomerate made up of diverse-sized and 6 Cin sedimentary outcrop In the central part of the country e Iumjcares Formation named by Castillo ( 1969 ), fragments of andesite , basalt , and quartz to , Middle is ) 1961 ( Dengo by , named Formation , Miocene diorite distnbLower rocks that have been dated Oligocenea sequence made up of conglomeratic sandstone and uted in a siliceous sandy matrix with some calcareou of up made is and age in Miocene , pp 1969 lenses ( Castillo such as the TSrraba Formation 1 I conglomerates. It is approximately 200 m thick claystone, sandstone , and lignitic shale interbedded of grajl , greenish stratified , poorly sandstones grained fine of layers with interstratified 11-13). They are made up of and s exposed onI theeast flank of the Cerros de Tur- conglomerate ( Redfield 192 , p. 366). It Car Punta at outcrop They . parts in fossiliferous , and shale calcareous ' outcrops gray in van dark to sandstone , siltstone , black sequence is calcareous and very fossil - ous parts of the Caribbean coast. It is exposed betweenPuntarenas . Dengo ( 19626 ) assign « and some limestone. The rocks of this unit are exposed ballo, southeast ofrocks of the same lithology exposed I* «®erous. t has been assigned in age to the Lower Mio- the basins of the Rio La Estrella and the Rio Sixaola and extensively เท the southern part of the Valle Central in the to this formation « owing to the fauna identified by Woodring and in the Nicoya Gulf and in extends for a long distance below the Duruy and Bitey Fila Diamante , in the Cerros de Escazu , in Tablazo, in the the Isla de San Lucas avassi Peninsula s Nicoya forma ion of the un rests side ( tributaries of the Rio La Estrella ) ( MacDonald west on the Barrigona , the in Tabarcia Rio the of valley northern margin of the on orma y on the T&raba Formation and probably is rivers CompB Nicoya on the unconformably , . 142). It also outcrops along the length of the Rio 1920 p rests formation , and this of west the to and Caraigres north flank of Cerro vered by the Coris Formation ( Castillo 1969). . ) 1961 Dengo ( Formation Nanei ( ) and in the higher Nanabri Brito the on and in the Fila Guaitil . fe Coris Formation , named by Castillo ( 1969), is a as far as Bonifacio and to the lands to the northeast the of rocks าเ on resting it observed ) 1970 ( Madrigal sedimentary northwest as far as Ban Oligocene , the side On the Caribbean c, SL*9 us'nLe of quartzitic sandstone , with thin layers of anito. It also occurs in the tunnels . Formation raba part and of the railroad to Formation Senosri the by rocks are represented 3 some local lenses of lignite. It is clearly visible Limon ( Malavassi up , of ) 19626 Dengo ( Formation , Montezuma 1967 The p . . 5 ) the of The slopes formation the rests unin formations and Machuca of the ร type locality at the Alto Coris and extends to the ' horizontal of up , antj made is east age conformably over Pliocene the to Gatlin Miocene . Formation Rio San Juan. e Valle de Cartago to south to Cerros del of conglomerate, sandsti The Senosri Formation was recognized by MacDonald poorly consolidated layers typically Thi Mazo , Loma Salitral and’ Loma Asern , thence to the Tertiary Volcanic Rocks ( Tv ) villag the near outcrops . It siltstone . and , pp 1960 Weyl and , , Dengo , ; ( 1919 also Hoffstetter LOCal 0U t r 000 near Jaris in The Tertiary volcanic rocks constitute a continental sein the extreme south of the Nicoya 287-88); the type locality is the Rio Senosri , a tributary Montezuma Pun s i formation is Middle to Upper Miocene quence made up principally of lava flows , agglomerates. good a cover formation this of . The outcrops of the Rio Sixaola on the Panama side. It is made up of ninsulasouthern in ge (Fischer and Franco 1979) and is dl and , Peninsula Nicoya overlain by and breccias that are generally andesitic and basaltic in the of part agglomerate beds cemented by calcareous material , of the

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(Castillo 1969). The Monterrey Intrusive, to the south of Orosi in the Cordillera de Guanacaste) constitute the vol -

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GEOLOGY

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by composition , with some tuffs and ignimbrites intrudedalso ๙ ๐r° ina’ VaJ e Central and Guanacaste San Ignacio de Acosta , on the southern margin of the Rio basalt bodies . Fluviolacustrine sedimentary rocks , between Limonal and Las Mesas, has h^n โ Candelaria gnimbrites of rtiyoli’ * * * appear along with these rocks. ( Malavassi as 1 by a 196๑ and ver tic. considered i n * ast volcanic held . ’, The sequence is regarded as Miocene- Pliocene in age , , Monterrey (Type de in Asern , exposed San The well received and some of its principal lithological units have PLUTONIC ROCKS diverse names such as the Aguacate, Rio Pey, Paso Real). Plutonic Cretaceous Rocks 1966 ( Dengo 1962/? ), and Doan formations ( Escalante the Santa Elena Periodite ( II ) and th rocks These Tilaran g the Rocks of these units outcrop in areas such as degree in bodies ( TKi ) ofaregabbro intruding th diorite , and , diabase ๓ de Cordillera Guanacaste lesser and Aguacate mountain ranges and to a , and in the sequence of volcanic and sedimentary rocks belonging t The Pleistocene Bagaces Formation is made up of ig๙ theR 'o Macho Rio Humo, and tributaries T e the Talamanca and Costena mountain ranges ng Santa Elen 9 p a 2 ) The . 6 4 wh 6 ch earlier vanes in com mentioned Complex ๒ nimbrites of dacitic and quartz - latitic composition in Nicoya the ’ valley of the Rio Tfcrraba (fig . 4.1). granodiorite of from half th to Weyl quartz northern the ( monzonite in position outcropping ( Dengo 1962tf , p . 59 ). This formation is some , ) sections II ( Periodite by , named The rocks of the Aguacate Formation ) , is found immediately south of Santa Marfa well exposed along the Inter American Highway , * ser 29 partly . body , p igneous , an 1957 is Peninsula Elena Santa , ag flows between Dengo ( 1962/? ), are andesite and basalt lava Divisidn The It ( Dengo I 962a , p . 146 ) Cartas and Bagaces, and to the northwest of Liberia in the . . is Intrusive diorite Dota and de gabro of intrusions , with pentinized in exposed glomerates, breccias , and tuffs. They are well is made up primarily of granodiorite and granite, accord - Santa Rosa National Park . Cretaceous in age. the Cerros del Aguacate , the type locality, in the Cor- considered penetn that ing to Weyl ( 1957 , pp . 31 -35) , and is found on the hill ) ( TKi rocks The majority of the intrusive Tal dillera de Tilaran , and in parts of the Cordillera dethese , sills bodies f hipoabisales eabbro the Nicoya Complex are small ๙ ° amanca ( fig. 4.1 ). Later, Madrigal ( 1970) grouped 1 ' โ di and gabbro of stocks small perhaps and dikes and . Group rocks within the stratigraphic term Aguacate Dengo abase. Webber ( 1942, p. 2) discovered various olivi is well exposed in the Rio Liberia and in Liberia City, The Rio Pey Formation is Pliocene in age ( ( Dengo 1962a , p. 146 ) , which makes which reason it has been called the " white city," for Intrusive p Chirripo southern the in gabbros and Seco Rio the in diabases , particularly slope and to 1962a ) and is found on the Caribbean highest peak the the in up Talamanca Cordillera tf ) encountered gabb] 1962 ( IS Dengo the . north of Culebra Liberia in the of Bahia ) Blanco and and 1928 Colorado Branson ( . rivers in the valley of the Rio Reventazon granodiorite of ( Weyl 1957 , p 36). The formed Peninsu Nicoya the , of parts The various in Quaternary Cote diabases and , Formation this of which is rocks also volcanic part in The in the Cerere and Telire rivers . Ujum and Kamuk intrusives are less well known How Pliocene ( Malavassi and Madrigal 1970, p. 4), has its such as at Cabo Velas , near Sardinal , near Santa Cr formation are found in these rivers in the lower part of the Canjel Dengo ( 1962a , p. 146) found granodiorite on the type locality at Lago de Cote ( north of Tilaran on the ever . , Montezuma and , Rio Blanco Suretka Formation and consist primarily of agglomerates According side of these intrusions in Ujairas , Rio Ceibo, and slopes of the Rio San Juan ). It is represented by a se Pacific similar , intrusions ) tf 1962 ( Dengo to and of basaltic breccias with minor intercalations of ba- those in the Nicoya . Gabb ( 1874 ) reported granite and sienite in quence of pyroclastic rocks made up of ash , tuffs, sands,Cabagra Rio Q in gabbro as occur Peninsula salt lava flows and raffs ( Dengo 1962tf ). . and at times lapillis. It is also exposed in the land sur Honda de Turrubares and in the Osa Peninsula a!) Kamuk The Upper Miocene Pliocene Paso Real Formation brada ( Dengo , 1962 p . a 146 ) also thai reported possible is it , various and \ intTUM Rincon Rio e of the rounding Laguna and to the north of the Rio (Dengo 1962tf ), found on the Pacific slope, is made up the headwatersfound in other parts of the peninsula . bodies on the Caribbean slope of the Cordillera de Tal- Arenal , covering aArenal be may they goodly portion of the headwaters of associated , with water in of pyroclastic rocks deposited , generally of alkaline affinity, intruding into Oli the tributaries of the Rio Frio amanca to near San Rafael and exposed well is . It flows lava , agglomerates , breccias and H gocene and upper Eocene sediments and probably some T 1 r & ( Jicarito Plutonic Malavassi and Tertiary It Madrigal . 1970, geological map ). Escuadra the to Cuan at Rio Terraba , from Quebrada of the most alkaline members of the Talamanca Co- This formation unconformably overlies described been have rocks intrusive giranitoi of Stocks Sabalito and Tertiary sedi Gordas Cartas is poorly exposed between magmatic Series. Near Pico Aguila and the tributaries of mentary rocks of the Venado Formationthe * Tal de Cordillera leng the along . . ) tf 1962 Dengo ( along the border with Panama toe Coen , La Estrella , and Telire rivers, there are horn - The remaining volcanic rocks that make up the active . amanca as a res Q *a ( 1 (8 7 4 )าexplorations Iiamj The volcanic Doan Formation , of Pliocene age ( Esca rocks blendic diorite dikes that overlap the group of Victoria volcanoes in the Cordillera de Guanacaste are an un were grouped y \ eneo 196 /? ) under the of the! dikes dated lante 1966 , pp. 64-65), is made up of agglomerates and Talamanca as Miocene-late Pliocene. To the north of differentiated sequence of breccias, lava flows , mudflowmembers en , hard raffs that outcrop characteristically in Cerro Doan associationComagmatic Monte Matama there is also a granite stock in the upper deposit ( lahars), and diverse pyroclastic materials that were probably initially derived from a c< along the southern margin of the Rio Reventazon , and in course ol the Rio Banano. have not been completely differentiated and named , . magma other places such as Cerro Alto Araya , Copal , Cerro mon According to Dengo ( ? , , . 1962 / gabbrow 147 p , ) the eradodiorites Talamanca nrincioallv The Tivives Formation and the Orotina Ignimbrites are La- These roc Cruces , Cerro Congo, Urasca , Pertas Blancas, Cerro Comagmatic Series mon was , emplaced erites during a ofrnan period of rnrooortions ^ the , W 1 primary lithological units of the Orotina volcanic m ), and granites jas , Alto Velo de Novia, Rio Quiri ( 1 ,300-1 ,475 strong ) deformation at . 39 the p beginning of 1957 kyi the . Miocene The field . The Plio-PIeistocene Tivives Formation ( Madrigal , dong the road between Tapanti and Rio Tuas ( 1 ,600 m ), zooites diont intrusive activity produced as a sedimentary series of ary , mineralizations esequenc , p. 29) is a mudflow containing laval fragments, 1970 into and in the Naranjo river to Cerro Doan . Later and intruderock fte country in tot| priroarily metallic, that characterize the areas it mainly basalt , of various sizes (0.10 to 2.0 m ) distributed ทุ่ Krushensky ( 1972 , pp. 12 13) assigned a sedimentary influenced , to T . been has « interpretation « this , but » formation origin to this 1๙ w ด ctayol aci on ^ stocks the e าาไ รน . I also who , ) po 38 37 , . ed pp ๒ 1977 ( Berrange rejected by f tue northweSI QUATERNARY VOLCANIC AND SEDIMENTARY ROCKS Tivives cliffs, in the*Rio Jesus Maria delta , at*Perton de mapped the Tapanti quadrangle and found it to be vol dillera rnaryVoUxmic Rocks ( 1< Bajamar at Punta Carrizal at Punta in the Rio canic in nature just as Escalante had proposed in 1966. to the southeast: e in SD rocks are primarily lava , ( Berm flows Macho andesitic 0 volcanic , Surubres and near the sedi Tertiary confluence of over the Rio unconformably Machuca lies This formation 'rccc as’ tuffs , and ignimbrites that grade from andesitic with the Rio Jesus Maria . , Ujum , Durika 6 a , L ) , Dota 1977 for This Aguacate and formation Pacacua rests ( uncon rocks volcanic and mentary 8ca u Intrusive ° diyolitic composition , mudflow deposits , and accumu £ Dengo formably on ( Tertiary Kamuk P sedimentary ( rocks Punta Carballo . ) Ial , respectively mations ons of unconsolidated pyroclastic materials such as Formation ) and is overlain by the Orotina cdiately to the south situated in e In relation to these rocks, it is of particular importance San and lapilli. These rock types and the active ( Orotina Formation ) ( Madrigal 1970, p. 30Ingnimbrites , rocks ndMllVOfdioritic , IS ma euppnnc Jose that surface peneplain wide a be to appears there that olcanoes (Turrialba, Irazu , Poas in the central Cor- The Pleistocene Orotina Ignimbrites were).first recog 'dfllera , monzonites , and gabbros where the gabl developed over the Tertiary rocks in the Cordillera de dioritesappear ; Tenorio, Miravalles, Rinc6n de la Vieja, and nized of the in periphery the at principally and described by Dengo ( 1961, p. 53) and were rocks foot associated and Aguacate del Montes the , in Tilaran

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later named the Orotina Formation by Madrigal ( 1970). They are made up of ignimbrites of prismatic columnar structure, possibly derived from a quartz - latitic magma , similar to the magma that gave origin to the volcanic field of the Valle Central ( Dengo 1961 , p. 56). According to Castillo ( 1969) and Madrigal ( 1970), this formation outcrops at Hacienda Vieja and south of Orotina along the Rio Grande de Tircoles , to the south , southwest and west of Coyolar between Rio Cuarros , Quebrada Huaca , and Quebrada Pozon , near the delta of the Rio Jesus Maria on the southeast margin near the Laguna Sapo, and on the northwest side of the same river closer to the coast. This, formation rests unconformably on the Tivives Formation and over Tertiary volcanic (Aguacate Formation) and sedimentary rocks ( Punta Carballo Formation ). It is overlain by recent Quaternary sedimentary rocks , mainly near the coast. The Valle Central volcanic field is made up of two sequences of rocks that outcrop in the western and eastern portions of the Valle Central ( Valle Central Occi dental , Valle Central Oriental ), separated by the con tinental divide at the Alto de Ochomogo, between Tres Rios and Cartago. The volcanic rocks of the Valle Central Occidental are made up of andesitic and basaltic lava flows, tuff , and ignimbrites of latitic and andesitic- basaltic composition ( Williams 1952 , p. 155) , mudflows derived from the highlands to the north and northeast , and pyroclastics recently erupted by the volcanoes of the Cordillera Cen tral . Among this sequence of rocks some lithological units have been defined and named by a variety of authors (Colima , Tiribi, and Barba formations and the Irazu La vas) ( United Nations 1975). Other rocks of volcanic ori gin or nature that have not yet been named ought to be considered within the group of undifferentiated volcanic rocks of the Valle Central Occidental. The Pleistocene Colima Formation , whose type local ity is the lajo of Colima at the bridge over the Rio Virilla between San Juan de Tibas and San Domingo de Heredia , is a unit made up of piroxenic andesitic lava flows with layers of tuff and ash . It outcrops on the bottom of the canyon of the Rio Virilla for more than 30 km between the delta of the Rio Circuelas and the northern part of the city of' San Jose. These lavas were called “ intracanyon lavas by Williams ( 1952 , pp . 150-52), who considered them to be derived from Poas and Barba volcanoes. This formation lies unconformably on some Plio-Pleistocene fluviolacustrine sediments , and on Tertiary sedimentary and volcanic rocks , in several places, on the bottom of the Rio Virilla canyon . The Pleistocene Tiribi Formation has its type locality at the Rio Tiribi, before it joines the Rio Virilla at Elec triona . It is made up primarily of tuff , poorly welded

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to welded ignimbrites of latitic and andesitic basaltic composition ( Williams 1952, p. 155). Some have a co

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lumnar prismatic structure , derived from cones or frac tures related to the volcanic system of Po£s, Barba and Irazu . This formation originated the flat and slighdy un dulating topography, with a slight inclination toward the south and southwest , of the Valle Central Occidental. It is distributed from the northwest of Santo Domingo to the southwest in the direction of the Rio Torres and Rio Tiribi in the form of a narrow belt. It is always next to the canyon of the Rio Virilla until it reaches the top of the mountain mass of the Alto de Las Palomas. Also from here it is very well exposed along the canyon of the Rio virilla until it joins the Rio Grande. From here it continues yet farther to the west on both sides of the Rio Grande de TSrcoles , in the form of isolated patches until it blends into the Orotina Ignimbrites. This unit , described by Williams ( 1952, p. 53) as “glowing avalanche deposits,” rests unconformably on the Colima Formation , The Irazu Lavas together with the pyroclastic material were originated by Volcan Irazu . They are primarily an desite and outcrop to the north and east of San Isidro de Coronado to the lands adjacent to the east and south of Rancho Redondo. The stratigraphic correlation of this unit is not well established , but it could momentarily be considered contemporary with the Colima , Tiribi and Barba formations, The Recent Barba Formation , whose type locality is the Quebrada Barba to the southeast of the village of Barba , consists of various dense , well-crystalized , m sive , and fractured lava flows. They have scoriae^ and breccia facies separated by mudflows , ash , and fossi soil deposits. They extend from the southern flank of thi Cordillera Central to the valley of the Rio Virilla , alonj the length of this from Electriona to where it joins the Ril Virilla , along the length of this from Electriona to whf it joins the Rio Ciruelas and the Rio Grande to the end of this formati the Valle Central Occidental . The rocks were called as “ postavalanche lavas ' by Williams ( 1952 , pp. 162-63) and were originated from Barba and Poas volcanoes. This formation unconformably overlies the Tiribi Formation and in general is covered by ash layers of varying thickness, Within the group of undifferentiated volcanic rocks in the Valle Central Occidental , there are mudflow deposits or “ lavina ," as described by Drindoli ( Dondoli and Torres 1954 ), and recent ash deposits. The mudflows cover the central and northwest portions of the central valley c E sponding with San Jose , Zapote, Curridabat , San Pedro Guadalupe, and Moravia. They mainly consist of lafl fragments of various sizes and proportions, and of vol canic gravel and sand distributed in a clayed matrix of dark brown to yellowish brown . The deposits of ash from

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emptions reach very thick depths in some places Unarras Formation , which is described below, and the recentcover large areas of the formations described above , Parafso Lava Flow. arid Formation , the Irazu Lavas, the Barba The Aquiares Lava Flow is Recent in age and is an the such as , Tiribi mudflow deposits or lahars. the and andesite, similar to the Cervantes Formation Flow. It extends sequence of the Valle Central Orien - from the northern part of Aquiares Lava rock volcanic The to the southeast in the tai is made up of lava flows, breccias, tuffs, mudflow direction of Santa Rosa and the city of Turrialba, to the deposits, and recent pyroclastic material . These have other side of the Rio Reventazon at the place called t>een. in some cases, differentiated by stratigraphic Angostura. The unit is dissected by the Rio Turrialba and for example, the Parafso Lava Flow, the Cer- its tributary, the Rio Aquiares. The Aquiares Lava Flow names:Lava Flow, the Aquiares Lava Flow and the Irazu rests on a conglomerate made up of fragments of andesite vantes Group. and basalt , intruded by basalt dikes , which can be asThe lava flows mentioned above were named by Ddn - signed to Tertiary volcanic rocks ( Aguacate Formation ). 1954). The Paraiso Lava Flow Later, Krushensky ( 1972 ) introduced the term Irazu doli ( Dondoli and Torres ' n Irazu and is a fissured and frag Volca from derived Group to classify the group of lava flows and andesitic is mented andesite , massive and compact with columnar and basaltic breccias, tuffs and mudflows derived from jointing and scoriaceous on the surface. It extends over the Irazu and Turrialba volcanos. The Irazu Group covers the lands around Cartago and to the village of Paraiso all the flanks of Volcan Irazu and includes four formations , the way to the Rio Reventazon and very near to Turrialba . from oldest to most recent: Reventado, Sapper, Birris , It is exposed along the road between Paraiso and Orosi, and Cervantes , the last described earlier. approximately 1 km before arriving at the bridge over the The Reventado Formation ( Krushensky 1972, pp. Rio Navarro, and in the road from Paraiso to Cachi. 18-23) is made up of andesitic- basaltic lava , ash , and The first flows of the Paraiso Lava Flow apparently mudflow deposits. It is mainly visible in the canyon of closed off the flow of the Rio Reventazon and formed a the Rio Reventado and includes three members: a lower lake where were deposited the lacustrine sediments of the member that corresponds to the Paraiso Lava deUjarras Formation (described below ) which were then scribed earlier; an intermediate ash member thatFlow covers later covered by the more recent flows of the same Pa- the lands around Paraiso to the east and to Caballo ralso Lava Flow. The Paraiso Lava Flow lies uncon and Concavas and most resembles a lateritic soilBlanca proformably over the Tertiary sedimentary and volcanic duced by the laterization of the Paraiso Lava Flow ( Ber rocks. Dondoli and Torres ( 1954 ) dated the Paraiso Lava range 1977, pp. 45-46); and an upper member made upFlow tentatively to the Pliocene, but Escalante ( 1966) of lavas and breccias of andesitic basaltic composition placed it doubtfully in the Pleistocene. and mudflow deposits , distributed as a con The Cervantes Lava Flow is Recent in age; it consti- tinuous surface layer tointercalated the northwest , north , and north tutes a compact andesite internally but is scoriaceous on east of Cartago, where are found villages such as San the surface. It is derived from the southern flank of Cerro Rafael, Rancho Redondo, Tierra Blanca Juan de PasquI, a parasite cone of Volcan Irazu . It extends from Chicua, Potrero Cerrado, Cot, Pacayas,, San Capellades the lands around the village of Cervantes and reaches the Santa Teresa , and others ( Krushensky 1972 ). The Re, left margin of the Rio Reventazon in the vicinity of the ventado Formation is late Pleistocene in age and restsRio Parruas. It is well exposed in the canyon of the Rio over the Ujarras Formation . Reventazon between Fajardo bridge and Santiago and in unconformably The Sapper Formation ( Krushensky 1972 , pp. 23 26) the canyon of the Rio Birris near Birris electric plant consists of andesitic basaltic lava flows, ash , and number 2, south of Birris ( Krushensky 1972). Between mudflows. The is very well exposed at the Yas and Santiago the formation shows numerous circular Cerro Sapper, 1.7formation km southwest of the active crater of the depressions and collapsed lava canals. According to Irazu volcano; in the headwaters of the Rio Reventado it Krushensky ( 1972), the collapsed tubes generally do not lies unconformably over the Reventado . The exceed 1 km in length and a few dozen meters in width , type locality of the Sapper Formation is inFormation upper the part Exceptional collapsed tubes 2 km long and 200 m wide of the canyon of the Rio Reventado, about 250 m below can be observed from the Finca Leda on to the Camino the confluence of the Quebrada Pavas and Rio Reventado del Cerro, south ot Boqueron . Another collapsed to the crest of the Vfolcan Irazu -Cerro Pica Piedra. Also 3.4 km long extends from the Fuentes road to the westtube of there are excellent exposures in the Rio Retes, in the Arrabara along the western limit of the For headwaters of the Sucio and Blanco rivers to the north nation , to the Camino del Pedregal near Cervantes Yas . This tube of Cerro Retes, on the north face of Cerro Cabeza de ร about 200 m wide. The Cervantes 'uncontormably Lava Flow lies Vaca , and in the headwaters of the Durazno and Caj6n over Tertiary volcanic rocks and over the rivers.

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57

GEOLOGY

The Bims Formation is made up mainly of andesiticbasalitic lavas and outcrops in characteristic form in the canyon of the Rio Birris , from the Lecheria Bims to an unnamed quebrada that borders the Cerro Noche Nueva to the east . Good exposures are also visible in Gonzalez , Roscavdn, Central and Laguna Tapada quebradas ( Krushensky 1972 , p. 26 ). This Holocene formation rests unconformably over the Sapper Formation. Other volcanic rocks have been defined and described by Krushensky ( 1972 , p. 13) , such as the Toba de Avalancha de Piroclastos de San Jeronimo, in the Rio Re ventazon valley, and the Toba de Avalancha de Pirocldsticos del Rio Aguacaliente , which can also be seen in the Congo and in the valley of the Rio Reventazon. However, Kmshensky considers these units of uncertain stratigraphic position. Quaternary Sedimentary Rocks ( Qa and Qt ) These Pleistocene rocks are made up primarily of alluvial materials (Qa), coluvial -alluvial materials (Qt ) and fluvial lacustnne deposits. The coluvial alluvial materi als appear in the form of very clear elevated terraces such as in die north of the country on the slope of the Rio San Juan and in the south in the El General valley. In the include " northern part of the country these materials . mudflows . In the northern part of the country on the side of the drainage basin of the Nicaragua Lake and the Rio San Juan die coluvial -alluvial matenals and mudflows of the Plio Pleistocene have been grouped and named by Maiavassi and Madrigal ( 1970, p . 34) under the stratigraphic name Buenavista Formation . They include materials ot great heterogeneity in size and nature, pieces of andesitic and basaltic lava boulders distributed in a sandy and clayed matrix and intermediate fractions. Among the Quaternary sediimentary rocks on the Pacific slope in northern Guanacaste, there is the Pierstocene Caflas Dulces Diatomite ( Segura 1945; Madngal I 960) that appears in the area of Las Brisas-Cafias Dulces and in the Camastro hill ( Loma Camastro) (Sal azar 1977). The diatomite is white to grayish white to yellow and violet . It was formed in a lacustrine environ ment developed over volcanic Quaternary rocks, such as tuffs, breccias , and lavas. It contains some impurities such as clay, quartz , volcanic ash , and metallic oxides and occurs intercalated with thin layers of lacustrine con tinental origin and covered by volcanic rocks of more recent Quaternary origin , as occurs at Loma Camastro. Deposits similar to diatomite have been reported near Tilaran (Salazar 1977 , pers. comm . ). On the Pacific slope , specifically on the Osa Peninsula , there are rocks of the Pleistocene Armuelles Formation , Mentioned by Hoffstetter, Dengo, and Weyl ( 1960), the type locality is Puerto Armuelles in Panama. According

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erates of a clayed matrix that is stratified with greenish to Madrigal ( 1978 , p. 162), it is made up of conglom

GEOLOGY

gray, coarse-grained sandstones. These rocks are not as well stratified as those of the Charco Azul Formation, and they rest unconformably over it. TCrraba TrOUgh • and to Limon Bas,ท . In the central part of the country, the sedimentary roc Aguacaliente the niff fonnably of the Quaternary are principally alluvial , mudflows , and The fluviolacustrine sediments of the Cartago valley or Outer Arc fluviolacustrine deposits. These intercalate with or ท Valle del Guarco developed as a result of an old lake that This area is on the Pacific side of the orogen and correprincipally on Quaternary volcanic rocks. Less frewas formed by the stopping up of the valley of the Rio sponds to the Jurasic-Cretaceous rocks of volcanic , sedi quently they rest on Tertiary sedimentary and volcanic ( Dondoli and Tones 1954 ). This was done mentary, and Aguacaliente intrusive origin of the Nicoya Complex and rocks. Within this group are the Esparta and Ujarr$s the of lava emission coming from some Vblcdn Irazu by the the of Car Santa Elena Periodotite. sediments formations, the fluvial lacustrine produced the Lava Paraiso Flow later . the At end of that of ) the mudflows ( lahares , tago Valley or El Guarco the the Quaternary, the Rio Aguacaliente cut through the Ifiner Arc Valle Central , and the alluvial and uplifted terraces of the , permitting the draining of the lake and the ex obstacle I Rio Reventazon . The inner arc is the principal structure of the orogen and the plain that is the valley of Cartago. of posure The Pleistocene Esparta Formation ( Madrigal 1970, p. During the Pleistocene and Recent in the Valle Central , extends to the northeast of the external arc on its concave 34) was called Terraza de Esparta by Dondoli ( 1958 ) and . The border between the two arcs is defined by a there were some mudflows or flows of those coluvial - side is made up of a mudflow consisting primanly of phJ alluvial series of faults running from northwest to the southeast , materials called “ lavina" by Dondoli ( Dondoli neritic basalt fragments up to 0.40 m in size, subroundel and Torres 1954), which were probably previously described along Within of lateral displacement ( Dengo 1962tf , p. 136). and subangular, distributed in a reddish yellow sandi with other volcanic rocks of the the inner arc are recognized various structural Valle Central. clay matrix and deeply laterized on the surface This unit, : the Cordillera de Talamanca and its footsubdivisions Finally, during Recent times a series of alluvial terwhich is a characteristic flat plain is very evident at thd , hills up made of sedimentary rocks (Ts), volraces was developed in the eastern Valle Central , prin - canic rocks (Tvprimarily type locality to the south of the park in Esparza ( Esparta , ) plutonic and rocks (Ti ) of Tertiary age; along the valle of the Rio Reventazon on the lands on the road to Chumical ( Artieda ) , exactly where it de- cipally entra บ made up principally of Tertiary volcanear Orosf and Cachf , between Santiago and Turcurrique , scends to the Rio Esparta. Also it is very evident nic rocks and Quaternary volcanic rocks ( Qv ) , as and in the valley of Turrialba. well as some Tertiary sedimentary rocks Ts ( ); the volcaEl General valley there is a Pleistocene unit u ปี In1k the between Esparza andI the- entranceะ to Punta M ral . | to to ccmillleras the , northwest made of up Tertiary (Tv ) of coluvial -alluvial matenals or fanelomerates * 11pprimarily formation discordantly overlays the Punta Cartlallo or and Qv ( ) atemary volcanic rocks . from the Cordillera de Talamanca. This R » unithasdeveloped a set ๙ flat terraces that are dissected un formably OT e TivivK Orotinafc • io by epirogenic uplifts with a slight inclination principally The Terraba Trough In the area o amares an' to the southwest . However, some recent movements have "Hie Krraba Trough , southwest of the Cordillera Tast ^ c „ 6 ไใ ท11! modified in some places the original slope of these ter- lamarK* extends from San Isidro de El General and the < * 7 e 'the past existence ofa lake « WIth frasrncnts ๙ท primarily ๐ ๙ ' ^ , ternary in . is Panama It ' * agec which reveads made up igneous primarily , of of nature Tertiary sedimen various ~ 1- \ rvnrral เท ( , tary ) Ts rocks produced diabase ' , dikes during and the last glacial volcanic rocks of retreat that /ivtnHrki; | fuprp arp QC old Lago de WmwxWnaM 1951 ) the e a m affected Upper the Tertiary age ( ) Cordillera Tv de in the . Talamanca central On this part of the trough . formation rca develop yellowish to reddish brown laterities that ought The boundary between the trough and the Cordillera s a 1 °nl f 10 be aluminum ore. This formation rests unconformably de Talamanca is marked by subparallel faults that are ry' v°lcanic’ and intnisive rGcks ๙ 'fcrttoy It is mad# on ( 966, p 62 ) and assigned to die M consol I |n various pans ๗ the country there are alluvial and Irough and the outer arc is the Faialla Fault , which is ckar yellowish gray 0 clear brown and rflocri brtcc R«c » sedimenri atcov» fteMts racks oriented -s^dteasi . ๒ . and near the „R ,o Umon Stone and with claystone. Near the Rio Cnin and เฒ sjve COVPrj10 ^ r. ^ Basin Bimsito. close to Loajza , and in the small ^ area f PU S 71 6 Llm6n Basin is stnjchj๗ feature extends to the Joyas , the exposures are mainly very poorly Espar/ a; c . 1, ° .ntarejna, L '5 { the inner arc on hs concave side , and northeast o * f011 conglomerates , and conglomeratic sandstone or โ . ท Cordillera It displays a geosyncline of rapid development , with m° gravel. T e rocky fragments are " ' c rta e 0 6 Panamanian border and the , Osa Peninsula tense volcanic activity, where there were thick ° young) Tile lithic ragnie ( commonly peripheral accumulations of primarily Tertiary sediments (Ts) with are andesitic , commonly porphyritic and fresh , andfl TECTONIC STRiim RE an addition of rudaceous continental sediments (Suretka semble rocks of the Aguacate and Doan formations, bull „ c Central American Oro Formation C ) that occurred at the time of the final main and , differ from those of the overlying Sapper, Birris I 11 jn . เ lc‘* /( Dengo 1962๘ ) follow an arced orien uplift of the Cordillera Talamanca . The faulting and contin Cervantes formations. The stratigraphic position of, bull 1a1 0n j ryiLn0rt VVeSl 1๐ soudieast However, they are temporary folding affected rocks of late Upper Terin Ujarras Formation has not been clearly established p ed by an^approximately east - west tectonic trend tiary ( Pliocene ) in also the same basin .

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GEOLOGY

of the area into two deposition basins separated byof the emerging Cordillera Talamanca and the volcanoes the inner arc: the Limon Basin to the east and the Terrabu Trough to the west . During this time the Caribbean Sea and the Pacific Ocean remained connected across various canals that linked the minor sedimentation basins thai may have impeded continuous terrestrial travel through out the isthmus. The climax of the orogenic phase ( Miocene) was ac companied by an intrusive activity (Ti ) represented pri marily by granodiorite, gabbro , and granite along with axis of the orogen . In the late stages ot the orogenic phase, at the end of the Miocene and the Pliocene epochs, there was strong volcanic activity (Tv) characterized by andesitic and basaltic lavas , agglomerates , breccias, and tuffs , intruded by basalt dikes.

Tectonic History

According to Dengo ( 1962tf ) , the tectonic history of Costa Rica can be visualized considering the following

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igneous tectonic sequence: a prototectonic phase , an oro genic phase, and a postorogenic phase .

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Prototectonic Phase

The prototectonic phase began in the Jurassic with the formation of an insular volcanic arc with the concave side toward the Caribbean. It was the product of the elevation of the ocean floor crust in this area . This initial tectonic event produced a combination of subaerial and marine volcanic activity, erosion of the resulting rocks, and sedimentation of the detritus as well as carbonates and sil , icaceous materials originated directly from the ocean These processed produced the ophiolitic association called the Nicoya Complex . This association was partially metamorphized by tectonic forces during the Upper Cretaceous ( Dengo 1968, p . 27). The intrusive episode that took place in this phase and that characterizes the rocks of the Nicoya Complex also produced the Santa Elena Periodotite. According to Dengo ( 1962Z> ), this last unit represents an outcrop of the upper mantle, in other words, rocks that normally are found below the Mo horovicic discontinuity that came to the surface along the length of a fault zone aligned with the Clipperton fracture of the Pacific Ocean . This emplacement of the peridotite, took place during the early stages of the deformation before the deposition of the undifferentiated upper Cretaceous sedimentary' rocks ( Dengo 1962tf ).

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Postorogenic Phase

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According to Dengo ( 1968, p. 28), the separation between the prototectonic and orogenic phases is largely artificial. However, the first term has been used for con venience to describe the initial events in the history of the area and the second to define the tectonic episodes that it is possible to identify with strong certainty. Both phases are in reality part of a continuous process , and the limits , established with difficulty, are arbitrary. Dengo ( 1968 , p. 28) considers that the orogenic phase really began with the tectonic disturbances during the last part of the Cretaceous , reaching its climax during the Eocene contemporaneous with the Laramidic orogenesis. This phase began during the Upper Cretaceous with an uplift of the axis of the orogen and with a displacement to the east of the principal area of sedimentation . This formed a deposition basin on the concave part of the insular volcanic arc influenced by intermittent volcanic activity and is where the sedimentary rocks of Upper Cretaceous and Tertiary age ( Paleocene to Miocene) orig inated . The tectonic movements and the displacement of the volcanic axis to the northeast produced a fragmentation

Orogenic Phase

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The postorogenic phase extends from the Upper Plioc to the Recent ( Dengo 1962tf , p. 133) and refers to the events that occurred after the orogenic phase ot mountain building and deals with diverse events such as normal taphrogenic faulting and extensive volcanism . This vol canism (Qv ) involved emission ot lava flows, ash -flow tuff and ignimbrite avalanches , and air-fall pyroclastic eruptions. This phase is characterized more by detor mations owing to tensions in the crust than to com pression . During this phase the Guanacaste and Centraพ cordilleras were formed and produced the volcanoes now know. The terrestrial connection between North and South America took place in the Miocene or more probablyto during the Pliocene ( Dengo 1968, p. 30). According Woodring ( 1966) , the small areas ot Pliocene deposi indicate that the area of Central America correspondii to Costa Rica would have emerged almost in totali during this period ,

GEOLOGY

1981. Costa Rica. geografico de Costa Rica . San Jose: Revista Costa Rica. Dengo. G . 1961 . Notas sobre la geologia de la parte Madrigal , R . 1960 . Algunas localidades de diatonuta de central del litoral del Pacifico de Costa Rica: Inst. Costa Rica . Informes Tfccnicos y Notas Geologicas, Geogr. de Costa Rica. Inf . Semest , Inst . Geograph . no. 5. San Jose: Ministeriode Agricultura e Industrias. Sac . , July-December, pp . 43-63. . 1970. Geologia del mapa bdsico Barranca 1962a Tectonic-igneous sequence in Costa . Buddington Costa Rica Informes TScnicos y Notas Geoldgicas , Vol . , Geol . Soc . America pp. Rica. no. 37. San Jose: Ministerio de Industria y Comercio. . b. Estudio 133 611962 . 1977 . Evidencias geomorficas de movimientos geologico de la region de tectonicos recientes en el Valle de El General. Cienc . Guanacaste Costa Rica . San Jose: Instituto GeoTec . 1 , no. 1; 97-108. graph ico. . 1978. Terrazas marinas y tectonismo en la Peninsula de Osa , Costa Rica . Instituto Panameri ' ' mmfologiade America . Re Re cano de Geografia e Historia , Rev . Geograf 86-87: gionalde Ayuda Tgcnica (AID) 161-66. 969 MaPa Metalogenetico de Amer- Malavassi, E. 1966. Geologia de la hoja Caraigres . San B ica Central. aJPublA . Geol . ICAITI Jos£: Ministerio de Industria y Comercio. Restricted Dondoli , c. 1951. Zona de Palmares , estudio report. agromomico . Technical bulletin no. 5, 16. San Jose: . 1967 Resefia geologica de la zona de Turrialba. Ministerio de Agricultura e Industrias. Informes Tecnicos y Notas Geologicas no. 27. San 1958. Breve resefia geologica . In Estudio semi- Jose: Ministerio . de Industria y Comercio. detallado de los suelos de la region comprendida entra Malavassi , E. , and Madrigal , R . 1970 . Reconocimiento los rios Barranca y Lagarto . San Jose: Ministerio de geologico de de la zona norte de Costa Rica. Informes Agricultura e Industrias. Tecnicos y Notas Geologicas no . 38 . San Jose: MinisDondoli , c., and Torres , J. A . 1954. Estudio geo- terio de Industria y Comercio. agronomico de la region oriental de la Meseta Central . , A. A. 1922. The Miocene of northern Costa Technical bulletin no. 32. San Jose: Ministerio de Ag- Olsson . Rica Bull . Am . Paleontol 9, no. 39:9-20. ricultura e Industrias. Redfield , A . H . 1923. The petroleum possibilities of Escalante, G . 1966. Geologia de la cuenca superior del Costa Rica . Econ . Geol 18 , no. 3:81 101. Rio Reventazon, Costa Rica. Publ Geol . ICAITI 1 Riviere, F. 1973 . Contribucion estratigrafica sobre la (Guatemala ):59-70. geologia de la de Limon , zona de Turrialba , Fischer, R . , and Franco, J. c. 1979. La Formacion Coris Costa Rica. PublCuenca Geol . ICATTl 4:144 -59. ( Mioceno; Valle Central , Costa Rica ). Inf Semest . . 1980. Geologia del area norte de los Cerros Inst Geograph . Nac . , January -June. de , Cordillera de Talamanca , Costa Rica . Inf . Escazu Fisher, ร. p , and Pessagno, E. A. 1965. Upper Cre- Semest . , Inst . Geograph Nac . January June. taceous strata of northwestern Panama. Am . Assoc . Romanes, J. 1912 tf . Geology of a part of Costa Rica. Petrol . Geol . 49:433 44. . London Quart J . 68:105-23. Gabb, พ. M . 1874. On-the geology of Costa Rica: Am . Geol .. Soc 1912 b . Geological notes on the Peninsula of J Sci . 7:438-39; 8:388 90; 9: 198-204. , Costa Rica. Geol Mag . 9:258 -65. Nicoya Galli , c. 1977. Edad de emplazamiento y periodo de Salazar, A . 1977. Geologia de los depositos de diatomita acumulacion de la ofiolita de Costa Rica. Cienc Tec , de la Brisas-Cafias Dulces y la Loma Camastro , Li , Guanacaste y รน evaluacion preliminar.Galli , c. , and Schmidt- Effing, R . 1977. Estratigrafia de [beria Corporation Costamcense de Desarrolo] Bol . Geol . la cubierta sedimentaria supra ofiolitica cretacica de Recurs . Mineral . 1:296-319. Costa Rica. Cienc . Tec . vol. 1, no. 1. Segura , . 1945. Rapidos apuntes sobre los marmoles de A Hayes , c. พ. 1899. Physiography and geology of region Guanacaste aspectos geologicos. Rev. Inst . Deadjacent to the Nicaragua canal route. Geol . Soc . Am . fensa Cafe . y15otros : 337 47. Bull. 10: -348. , Terry R . . A . 1956 A geological reconnaissance of Henningsen285 , D. , and Weyl , R . 1967. Ozeanishe Kruste Panama . Occasional Paper no. 23. San Francisco: Cal im Nicoya- Komplex von Costa Rica ( Mittelamerika ). ifornia Academy of Sciences. Geol Rdsch . 57:33-47. Nations-Senas. 1975. Investigaciones de aguas Hill, R. T. 1898. The geological history of the isthmus of United subterrdneas Panama and portions of Costa Rica. Mus . Comp . Zool . 65-502/ 1. en zonas seleccionadas . DP/UN/COS Hanard Coll . Bull 28:151-285. Webber, B . M. 1942. Manganese deposits in Costa Rica, Hoffstetter, R .; Dengo, G.; and Weyl , R . 1960. Lexico Central America . Am . Inst . Min . Met . Tech Pub estratigrafica de America Central . San Jos£. 1445 : 339 . 45 ^ Kriishenfr er^ ky, R . 1972. Geology of the Istaru Quadrangle , Weyl , R . 1957. Contribucion a la geologia de la Corsla Rica. U S . Geol . Sun . Bull .., no. 1358. ° dillera de Talamanca de Costa Rica . San Jose: Instituto McDonald , D. F. 1920. Informe final geologico y Geografico de Costa Rica. Sinopsis sobre la geologia de . Uni' - Costa Rica , in press.

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al Barr, K . พ. , and Escalante, G . 1969del. Contribucion de Complejo esclarecimiento del problema Nicoya , Costa Rica . Publ Geol . ICAITI 2:43-47.Ta Berrange , J . p 1977. Reconnaissance geology of the -. panti Quadrangle Talamanca Cordillera , Costa Rica . Institute of Geological Sciences , Overseas Division geog the on observations Some . Branson , E. B . 1928 raphy and geology of middle-eastern Costa Rica. Unix. Missouri Stud 3:30-72. Carballo, M . A . , and Fischer, R . 1978. .La Formation San Miguel ( Mioceno, Costa Rica ). Inf Semest . Inst . Geograph Nac , January -June . Abray Castillo, R . 1969. Geologia de los mapas bdsicos yI Tecnicos Informes . Rica parte de Rio Grande Costa n de Ministerio : Jose San . . no 33 Geologicas Notas nomi'a , Industria y Comercio.

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1

61

land bridge as a sea banier. Panama The . 1966 . Cen . Meseta the \blcanic history of California Publ. Am . Philos. Soc. Proc. 110, no. 6:425-33 Wiliams, H . 1952,. Costa forMiocene . . 1961 . E . Univ Woodring , พ. , and Malavassi ,barnacle tral Occidental, no. 4:145Rica Valle the from . 80 a - Imolluscs from Bowden , aminifera , mollusks and. Paleontol 35, no. 3:489-97, Geol . Sci . 29 Woodnng , พ. 1928. Miocene Central , Costa Rica. J Carnegie : . C . D , Washington . 385 Part 2. Publication

5 SOILS

GEOLOGY

Institution.

A . Vasquez Morera

A -5 Marshy soils , occurring frequently near the various Costa Rican soil types The distribution of be the map in general seen oceans and often under mangroves (Aqu soil of can here described study compiled was from the This ) . ). ents 5.1 . ( fig Rica Costa Drenaje y Riego Direccidn del the de Very A in light- textured soils formed mainly as 6 base informationde Agriculture y Ganaderia (Section of Irri belts along littoral beaches (Psamelongated Ministerio Drainage of the Ministry of Agriculture and . ) ments and gation b) Soils of fluviolacustrine origin: ) Dondoli and Torres ( 1954 ), Vai as and Torres Livestock , ) . Nuhn 1966 et ( ) al A -7 Flat , dark , very heavy- textured soils , very . ( Centro 1977 al et , Sander ) 1958 ( ( 1968 ), Perez , Alvarado, and Ra poorly permeable to water, lumpy when dry Cientifico Tropical and sticky when wet (shrinking and swelling mirez ( 1978) and Morge ( 1978). clays); fertile but difficult to farm . The exterIn spite of the different levels of study that have pronal drainage is moderate to poor and the vided information , the map’ร level of cartographic generslopes are less than 2% ( Usterts , Uderts). alization is not uniform for the entire country In isolated cases , and in areas of very difficult access, the summaries c) Soils of organic origin A -8 Soils formed from organic materials in an are very approximate, such as occurs in some extreme advanced state of decomposition , poorly northwest zones and in the central region of the southdrained , flooded (Saprist , Hemist ). eastern part of the country. The soils are separated into four large categories, 2. Soils on undulating relief B- l Soils of coluvial -alluvial origin , on slightly based on relief . These are then subdivided according to , among , depth other local characterundulating terrain , with slopes of 2-8%, well drainage texture and drained , deep , medium to moderately heavy istics. The soils are classified to the level of suborders in texture , moderately fertile , brownish and in accordance with the soil taxonomy of the United Agriculture . of Department (Tropepts ). States B-2 Soils developed from volcanic ash deposits on gently undulating terrain , with slopes of 3 15% ; well drained , deep , dark colors , rich Classification and Description -organic The classification and description of the principal categomatter , of medium texture , friable , in ries of Costa Rican soils, with their map symbols, is as moderately fertile ( Andepts). follows: B-3 Soils developed from volcanic tuffs , on al1 . Soils on flat relief most flat to gently undulating terrain , with a ) Soils of alluvial origin slopes of 3- 15%, moderate to shallow in A - l Well -drained , deep , dark , fertile, friable, depth , brownish , medium to moderately light medium - texture soils , rich in organic matter, in texture , well to excessively drained , with with slopes of 0-2% (Udolls , Ustolls , Trolow fertility (Tropepts ). pepts ). B-4 Soils developed from old coluvial -alluvial A -2 Moderately drained , brown , clayey , modmaterials, on gently undulating relief , erately fertile soils, with slopes less than 2%, formed in areas of longitudinally dissected susceptible to occasional flooding (Tropepts , piedmonts, with slopes of 3- 15%, highly Aquepts). weathered , well drained , deep , red , clayeyA -3 Poorly drained , light-colored , clayey , hydrotextured , with low fertility ( Humults, Tromorphic soils, with weak morphogenetic depepts ) . velopment , and susceptible to flooding, with B-5 Soils developed over ancient eroded terraces slopes less than 1 % ( Aquepts). and forming a softly undulating terrain of low A -4 Very poorly drained , dark-colored , heavy hills , with slopes of 3-15%. They are deep , textured , hydromorphic soils, without morsomewhat excessively drained , reddish , phogenetic development and with slopes less heavy-textured , and have low fertility ( Hu than 1 % ( Aquents). mults , Udults).

^

62

63

H.; Perez , ร.; and others. 1977. Estudio book no. 436. Washington , D.C.: บ.ร. Government regional zona Atlantica Norte de Costa geografico Jose Printing Office. : Instituto de Tierras y Colonizacion . Vargas Vaglio, o., and Torres , J. A . 1958. Estudio preSan . Rica , A.; and Ramirez , E. 1978. Asoแทนทar de suelos de la region occidental de la Meseta Perez, ร.; Alvarado de suelos de Costa Rica ( mapa Central. Boletm ciacion de )subgrupos TScnico no. 22. San Jose: Ministerio preliminar San Jose: OPSA/MAG Instituto Geo- de Agricultura e Industria. grafico Nacional . Vdsquez M , A . , and Lara c., L. D. 1977. Mapa de , H .; et al. 1966. Estudio geografico Sander, G .;deNuhn reconocimiento de : Cuenca Baja del Rio Tamregional la zona none de Costa Rica. San Jose: pisque. San Jose: suelos Seccion de Suelos, Direccidn de Instituto de Tierras y Colonizacion . Riego y Drenaje, Ministerio de Agricultura y Gaof Agriculture , Soil Conser- naderia. United States Department vation Service. 1975. Soil taxonomy. Agriculture Hand -

SOILS

Nuhn ,

SOILS

NIC AR AGUA

-

IV

V

-

0 1

i

,

4

.

GENERAL SOIL MAP OF COSTA RICA

MINISTERIO DE AGRICULTURA Y GANADERIA Unidod d * รน* lo * Compiled 0»

Irq Aqr Al*m vdsquez Morero

SAN JOSE

- COSTA RICA

1979

Mloo

SOIL

!ED ED ED EE

LEGEND AND SYMBOLOGY

,'

dromod OHur Ol toil! (Udollt , บ* tolls Tropopts )

Ml

Uodorotoly dromod ollumol soil* ( Tropopts , Aquopts )

Poortf dromod allurial toils

T

ร!
wet forest areas are used both for seasonal pastures, pecially when there is a drought in Guanacaste , and fattening calves. Thus the old pattern of rotating pasn use has been expanded from a regional to a nation* scale. A particular subject of controversy in recent yets has been the effect of year- round use of wet lowlitf pastures on soil structure , nutrient status, and พ corrhizal populations , and the implications of this long term yields .

-

^

*

-

SPECIEร ACCOUNTS

as Research Sites Without doubt the agro ecosyste

Food and Agriculture Organization ( FAO). 1974. m is fast becoming the Monthly Bulletin of Agricultural Economic Statistics predominant system in the world. For this reason alone it 23 , 1 ( January ):23 deserves much more intensive study than it currently 1975fl . Monthly Bulletin of Agricultural Ecoreceives. But far more important is that by its very nature nomic. Statisti cs 23, 1 (January ):23. it provides the community ecologist with an important . \ 915 b. Monthly Bulletin of Agricult experimental tool. Agro-ecosystems are manipulate and nomic Statistics 24, 7/8 (July/August):17. ural Eco. 1978. Production Yearbook , / 977. Vol. 31. controllable and can be used for serious scientific study, ' enabling the ecologist to perform controlled Rome . ents Hall , c:. FAO 1976 . El Cafe y el desarrollo historico Owing to the recent rapid expansion of thisexperim habitat , ทนgeografi co de experim Costa Rica . San Jos6: Editorial Costaental situations are either already avail merous able or easily set up. Agro-ecosystems thus provide the- Harris , D. R . 1972. The origins of agriculture in the opportunity for detailed studies of ecologic al inter tropics . Am . 60:181- 93. , studies that are often much more difficult to carry Igbozurike, M..Sci actions V. 1971. Ecological balance in tropical out in the more “ pristine" environments commonly cho- agriculture. Geogr . Rev\ 61:519-27. sen as objects of study. Kass’ D C; L 1978. Poly culture cropping systems: ReAdditio , although our unders~ tanding is far from ’ nally . ' ” 7 1 ? . The changin.g nature- of New World®; complete , in comparison with other ecosyste a 7“ Parsons , J. J1“ consid- tropical . 1975 erable: amount; .is known about how agroms forests, since Europeani colonization . In_ The XV ms function. Besides the standard application -ecosyste 0f ecological guidelines for develop of ment in the western American humid scientific technique to agriculture, there exists a rich tropics , pp. 28-38. IUCN Publica col lection of associated folk knowledge in many areas of the tions, new series no. 31 . Morges, Switzerland. IUCN.. 1976. Forest to pasture: Development or detropics. structio n? Simposio intemacional sobre la ecologia de Finally, one should note that it is in the study of agro la conserv ecosystems that the science of human ecology is brought americano ation y el desarrollo en el Istmo Centro. Biol. Trop., vol. 24, suppl. 1. into clear focus. If one is to understand a functioning Perrin , R . M.. Rev 1977 . Pest management in multiple cropagro ecosystem , one must include questions of factors ping systems. Agroec osystem -118. and relations of production . Homo sapiens enters into Plath , c. V., and van der Sluis,s A3:. 931968 the deliberations at a very early point. And in a world char- de la tierra del Istmo Centroamericano. Uso potencial : FAO. acterized by a variety of imbalances among people and Risch , ร. J . 1979. Effect of plant diversity. Rome on the between people and the rest of the natural world , lation dynamics of several beetle pests in monocupopu ecolo gists must assume a certain responsibility. Homo and polycultures of com , beans, and squash in ltures sapiens Rica . Ph D. thesis, University of Michigan , Costa is influenced by and influences ecological processes Ann in Arbor . vitally important ways. It would be naive and short Sauer, c. 1966. The early Spanish Main. Berkeley: Uni sighted to initiate ecological ations that ignore- versity of California Press. such an important component investig of the ecosystem. Stone, D. 1972. Pre-Columb ian man finds ica: The archeological bridge. Cambri Central Amerdge, Mass.: Peabody Museum Press. Araya Pochet , c. 1976. Historia economica de Stone, ร. 1976. La dinastia de los conquistadores . 2d ed . Costa Rica , 1950-1970. 2d ed. San San Jose: Editorial Universitaria Centroamericana. Jose : Editorial Fer- Trenbat nandez Arce. h , B. R. 1974. Biomass productivity of mixtures. Blutstein , H . I .; Anderson , L. c.; Betters, E. c.; Dom Adv. Agron . 26: 177-210. browski , J H .; and Townsend , c. 1970. Area hand book for Costa Rica. Washington , D.C.: American- SPECIES ACCOUNTS . Bosemp, E. 1965. The conditio . . ns of,agricult ural growth. . African Oil Palm (Palma de Aceite) Chicago: Aldine. ermeer Cromartie , พ. J. 1981. The environmental control of J Vand .1 , _ -1 . insects using crop . . ,1 . . . manageZra, ed D. Pimentel 1:233-51. C..RIn.CPest l ok Senes in one ๙ toe most abundant trees in Costa Rica. Its< rapid Agriculture. Boca Raton , Fla.: C. .Handbo R.C. Press. sPread is due to a particularly effective dispersal agent, Direction General de Estadistica y Censos . 1973. Censo Homo sapiens , organize d in a particularly effective way Agropecuario, 1973. San Jose: Direction General de for such activity. The Standar Estadistica y Censos. Company and es pecially the United Fruit CompadnyFruit are rapidly expandingAgro ecosystems

——

’

1

"

1

‘

-

"

*

I

^

ร ร !ร ่_ L I

~

J

I

-

-

-

.

TT.

. diveAity

1

.

! t

bf:”5*

72

73

- - — —-

SPECIES ACCOUNTS

AGRICULTURE

dium is produced . When the inflorescence reaches 6- 12

๒* * the sameh“as those*that attack coconuts1 . are frequently

*•

aborts, depending on environmental concm it frequently , J . พ. 1972 . Tropical crops: Monocotyledons . ditions and the health ofuntilthe tree. The inflorescence is Purseglove \ ๖1 . 1 New York: John Wiley. enclosed in a sheath about one month after it

emerges from the petiole base (6 weeks before anthesis ). erma,e or female son; swinging machetes , bending backs, sweat mixed making efforts to develop disease-resistant varieties. with chaff in 99% humidity has caused more than one North American OTS staff member to gain respect for the Sugarcane ( Cana) rice she/he ate every day, after experiencing for a few E . D . McCoy hours the life of a small acreage rice farmer on the Osa Sugarcane (Sacchamm cfficinarum L. ) ( fig . 6.19) is 2 Peninsula . bugs perennial grass (Gramineae: Andropogoneae ) of Many pest species attack o. sativa . Pentatomid ( Solubea poecila ) sucking on leaves and stems can retard World origin , most probably originating in New Gui the growth of panicles and damage grains. Noctuid moth The genus Sacchamm includes six species: four of which larvae can attack leaves and cause serious damage , es- are cultigens , unable to survive long in a wild state (5 pecially after heavy rains followed by extensive drought. officinarum , ร. barberi Jeswiet , ร. sinense Roxb. , and รOne of the worst diseases of rice is caused by a vims. edule Hassk . ). ร . spontaneum L . is widely distribute

I mm* Sf i?r

IรV




I

3

FIGURE

6.19. Sacchamm officinamm. a Sugarcane fields ne^ Alajuela . , Mature plants with . b juela; person is near Ala reaching to a height ofinflorescences about 2 m. c, Tradi -

tional

_

manner

of taking harvested cane to ., Heredia, Costa Rica d, A large sugarcanea small mill , rnear mill in lowland Veracruz, Mexico ( photos, D. H. Janzen ). ,

Fig . 6.19 c-d continued next page .

L

SPECIES ACCOUNTS

are often applied ) in areas with high cies for many crops , indicating that the region ร agriculsoils (fertilizers , abundant sunlight , and heavy rainfall ( the tural pests may be poorly collected. Little work has been

AGRICULTL’ RE

temperatures plants require at least 1 ,525 mm of rain per year unless ). grown with irrigation The principal sugarcane growing regions are Australia and southern Asia , the West Indies, South America , and the lower part of North America ( Mexico and the United States ). Central America has never been a major รน carcane-growing region , and the present total area in is less than half that in Mexico. cultivation there Costa Rica and other Central American countries have substantially increased the production of sugarcane, particularly within the past ten years , and the crop has bein Costa Rica ร economy. Cane come relatively important in the country increased steadily but sugar production s , when a decline in pro1950 early slowly until the . decline was This halted in the late 1950s , duction began has production accelerated . For instance , and since then 96,000 metric tons; in about 1962 was 63 output in , metric tons. 168 000 was about it 1970-71 of low sugarcane in Costa production relatively The factors attributed several to has been the past Rica in ) of high : cane1960 the much of the altitude ( McPherson , delays which for to 15 18 maturity region growing , ; irregular of the the topography region which months does not allow efficient cultivation; transportation distances; and lack of capital . The decline in production in the early 1950s resulted from the shifting of cultivation from cane to coffee on the small fincas of the Meseta Central . Subsequent increased production in the Pacific provinces , the principal area of sugarcane farming , and a rapid proliferation of production in the Atlantic provinces , particularly San Josd , Heredia , and Alajuela , accounts for the recent increasing trend in cane sugar output in Costa Rica. A large suite of diseases and pests of sugarcane are recorded . The major diseases and many of the nematode pests are widespread , but most insect pests are localized ( Strong et al . 1977; Williams et al . 1969 ). Detailed information on cane-associated diseases is available in Hughes et al . ( 1964) and on cane-associated nematodes in Williams et al . ( 1969). Box ( 1953) records only six species of insects as feeding on sugarcane in Costa Rica: *

—

Brassolis isthmia ( Lepidoptera :

and the Pacific; ร. mbustum through to New Guinea and Brandes and Jeswiet is confined ! the commercial cultivars neighboring islands . Almost ฝinterspecific . of sugarcane now grown are 6 m), and allhybrids a have solid , Most cultivars are tall ( 2.5Africa , Asia ,

108

, and color of the joints van jointed culm . The size , shape environmental factorsand. Sugar with the cultivar and with variety ' soil of climates cane is grown in a° wide 35 °s . However , highest yie« N and 35 mostly between ferti , are obtained in heavy but well-drained highly

Brassolidae); Diatraea guatamalella and D . tabemella ( Lepidoptera: Pyralidae ); Saccharicoccus sacchari ( Homoptera: Pseudococcidae ); Phera obtusifrons ( Homoptera: Cicadellidae ); and ( ? ) Ectecephala tripunctata ( Diptera: Chloropidae ). D . lineoluta and Schistocerca paranensis ( Orthoptera: Acrididae ) probably also feed on sugarcane in Costa Rica ( Williams 1969). It may well be that the actual number of * pest fUtecct species is larger; about thirty species are own from all of there Central America. Central America is Pek*lvely depauperate in recorded numbers of pest spe-

done on the biological control of Costa Rican sugarcane pests , but Trichogramma fasciatum ( Hymenoptera: Trichogrammatidae ) and Paratheresia claripalpus ( Diptera: Tachinidae ) are known to parasitize Diatraea spp. there. Rodents are serious pests of sugarcane in many parts of the world . Costa Rica is no exception , and a rich mam malian pest fauna has been recorded for the country. เท addition to the ubiquitous Rattus rattus and R . norvegicus ( Rodentia: Muridae ) , native species of Zygodontomys ( Rodentia: Cricetidae ) , and Heterogeomys and Macrogeomys ( Rodentia . Geomyidae ) have been observed de stroying cane.

-

-

Barnes , A . c. 1974. The sugar cane. New York: Halsted Press. Box , H . E. 1953. List of sugar cane insects. London: Commonwealth Institute of Entomology. Fernandez , o. J. E. 1906. Estudio de los taladradores de la cana de azucar del genero Diatraea ( Pyralidae: Lepidoptera ) y รน importancia econdmica en Costa Rica. Thesis , Univsidad de Costa Rica. Hughes , G . c., et al . , eds. 1964 . Sugar cane diseases of the world . Amsterdam: Elsevier. Humbert , R . p. 1968. The growing of sugar cane. Amsterdam: Elsevier. McPherson , พ. K . 1960 . Informe sobre el cultivo de la cana de azucar y el desarrollo de la industria azjucarera en Costa Rica . San Jos6 : Ministro de Agriculture y Industrias. Strong , D. R . , et al. 1977. Time and the number of herbivore species: The pests of sugar cane. Ecology 58: 167-75. Williams , J . R . , et al . , eds . 1969. Pests of sugar cane. Amsterdam: Elsevier. Vegetables ( Legumbres ) M . Holle Vegetables, defined as perishable plant products that par-

ticularly contribute minerals and vitamins to the diet , are found throughout Costa Rica in four characteristic situ ations: I . The homestead garden. Such gardens are apparently disorganized arrays of plants next to homes, includ ing fruit trees , ornamentals , vegetables , and medicinal and miscellaneous plants. Careful description and anal ysis of such plantings are lacking for Costa Rica , but they exist for Guatemala ( Anderson 1950) , Java ( Soemarwoto et ฝ. 1975) , and the Philippines ( Sommers 1978 ). Casual observation here shows that the important vegetables are chayote (Sechium edule ), hot peppers ( mainly Capsicum sinense and c. frutescens in the lower altitudes and

-

-

109

AGRICULTURE

c. pubescens above 1 ,500 ทา ) , cherry tomatoes

-

( Lyco

persicon esculentum var. cerasiforme ) , squash (Cucurbita spp . ) , herbs like “ณda ” ( Ruta graveolens ) and roots like cassava ( Manihot esculenta ), sweet potato ( Ipomoea batata ) , and some weed forms like Solarium nigrum ,

whose shoots and flowers are occasionally consumed as picadillo ( a mixture of chopped vegetables with or with out scrambled eggs or meat ). Most of these plants are managed as perennial crops and can produce throughout the year. 2 . The market -oriented vegetable production system ( truck farm ) . The producer cultivates more than five species at any one time and intends the produce for the local market , mainly larger cities of the Central Valley area ( San Jos£ , Cartago, Alajuela , Heredia ). These operations are scattered , but they are found most often in the areas of : Cartago: Cervantes ( potatoes , tomatoes ); Para ISO ( leaf crops); Tejar (cabbage , green com , squashes , snap beans, etc. ); Volcdn Irazu ( potatoes , root crops , cabbage ); Guayabo (cabbage , tomatoes , snap beans ). San Jose: Santa Ana de Escazu (onions); San Antonio de Belen (sweet potatoes, miscellaneous). Heredia: Santo Domingo (garlic). Alajuela: Zarcero (a variety of species ); La Garita de Alajuela ( tomatoes ). Limon: Guapiles/Siquirres ( tropical root crops , hot peppers ). In most of these areas production is concentrated in the rainy season ( April through December ) unless water is available for irrigation . Landholdings are rarely above 15 ha . Tables 6.4 and 6.5 summarize varieties used , agronomic practices , and some of the main disease and insect problems. 3. The mixed farm . Normally agronomic crops are alternated with intermittent vegetable crops. This usually

-

constitutes a gamble and provides an opportunity to learn the management of a different and more intensive operation . These units can be found anywhere in the country. Plots rarely are more than 0.5 ha . Crops and character istics are the same as described above for truck farms. 4. The industry contract plot . The farmer arranges with a processor to grow vegetables such as tomatoes, broccoli , hot peppers , cucumbers , and peas for commer cial canning , pickling , or freezing. Such operations can sometimes be seen in the Meseta Central and in Gua nacaste. There is interest in this type of production , but at this time it is not regular or concentrated , although it will probably continue to develop , possibly in the Gua nacaste area ( drier than the rest of the country ) in con nection with irrigation projects.

-

-

-

-

-

*

Anderson , E. 1950. An Indian garden at Santa Lucia , Guatemala . Ceiba ( Honduras ) 1:97- 103. Folquer, F. 1976. Clasificacion botanica y varietal de las hortalizas. Miscellanea no. 62. Tucuman , Argentina: Facultad de Agronomia y Zootecnia , Universidad National de Tucuman. Smith , p. G ., and Welch , J . E. 1964. Nomenclature of vegetables and condiment herbs grown in the United States. Proc. Am . Soc. Hort . Sci. 84:535- 48. Soemarwoto, o., et al . 1975. The Javanese homegarden as an integrated agroecosystem . Paper presented at the International Congress of Scientists on the Human En vironment , 16- 26 November 1975 , Kioto, Japan . Sommers , p. 1978. Description and analysis of the home garden in four areas of the Philippines . M . s . thesis University of the Philippines at Los Banos , Institute of Human Ecology ( tentative title ). Terrell , E. E. 1977. A checklist of names for 3 ,000 \ ttscw-j lar plants of economic importance. Agriculture Hand book no. 505. Washington , D. C.: Agricultural Re search Service , USDA. '

-

.

--

TABLE 6.4 Common (Spanish and English ) and Scientific Names of Plants Grown as vegetables in Costa Ri Spanish Name

Scientific Name

Vegetable Crops

Beta vulgaris L. Capsicum spp *

Acelga Aji\ pimiento, chile Ajo Apio

Allium sativum Apium graveolens

Arveja Arracacha Ayote. zapallo Berenjena Broccoli

Pi sum sativum Arracacia xanthoriza Cucurbita spp . h Solanum melongena Brassica oleracea .

Camote, batata

Ipomoea batatas

no

.

L.

( dulce group)

L.

( italica group ) ( Poir )

L.

L

L.

English Name

Family

Swiss chard Pepper, chili pepper, red pepper Garlic Celery

Chenopodia< Solanaceae

Pea , garden pea

Liliaceae Umbelliferae

Squash Eggplant Broccoli

Leguminos; Umbellifen Cucurbitac< Solanaceae Cruciferae

Sweet potato

Convolvul

—

SPECIES ACCOUNTS

TABLE 6.4 conL

Name Caupi * rabiza , frijol de costa Spanish

Cebolla

Scientific Name Vigna sinensis

(Stickm . ) Savi

Allium cepa

English Name Cowpea, southern pea Onion

Hassk .

L.

(common onion group )

Chayote

Sechium edule

Chayote

Coliflor

Brassica oleracea ( botrytis group ) Momordica charantia

Cauliflower

Condeamor Jicama

Lechuga

Mafz (dulce, choclo. elote) Melon Okra , quimbombo Papa

Pepinillo, pepino Pepino ( mata serrano) Porro, puerro Rabano Remolacha , betarraga Repollo (col )

Repollo. (col china , pe-sai ) Sandia . patilla

( Jacq .) รพ.

Lactuca sativa Zea mass L

.

L.

Lettuce

Sweet com

.

Cucurnis melo L ( reticulatus group ) Hibiscus esculentus Solarium tuberosum

white potato Cucumber

L.

Cucurnis sativus

Solarium muricatum

Ait .

Allium ampelloprassum

( leek

Melon, muskmelon , cantaloupe Okra , gumbo Potato, Irish potato,

L.

group )

Pepino, melon shrub

L.

Leek

Radish Garden beet , table beet

Raphanus sativus Beta vulgaris L.

Brassica oleracea L. ( capitata group ) Brassica campestris L. ( pekinensis group ) Citrullus lanatus ( Thunb.) ( Mansf /C vulgaris ) Lycopersicon esculentum

Vainita,

Phaseolus vulgaris

Zanahoria

Daucus carota

Mill.

Cabbage Chinese cabbage , pe-tsai Watermelon Tomato Snap bean , green bean , string bean Carrot

. .

Tomate ejote

Balsam pear, bitter melon Jicama , yam bean

L.

Pachyrrhizus erosus

( Urban )

L.

L.

L.

Family Leguminosae

Liliaceae Cucurbitaceae Cruciferae Cucurbitaceae Leguminosae

Compositae

Graminae Cucurbitaceae Malvaceae Solanaceae Cucurbitaceae Solanaceae Liliaceae Cruciferae

Chenopodiaceae

Cruciferae Cruci ferae

Cucurbitaceae Solanaceae Leguminosae

Umbelliferae

Herbs

Albahaca

Bern) de agua Culantro. cilantro Perejil Ruda

Ocimun bailicum

L.

Nasturtium officinale

Brown

( partial list )

R.

Coriandrum sativum L . Petroselinum crispurn ( Mill ) Ruta graveolens L.

.

Basil

Labiatae

Watercress

Cm ciferae

Coriander Parsley

Umbeli ferae

Umbelliferae

me Rutaceae NOTE: Useful supplements to this checklist are Terrell 1977; Folquer 1976Common ; and Smith and Welch . 1964 The species of this genus found cultivated in Costa Rica are Capsicum annuum .; L c. frutensens L.; c. baccatum L. var. pendulum ( wild ) Eshbaugh; . pubescens. Ruiz and Pavon. The first two are most common . The species of this genus cfound cultivated in Costa moschata ( Duch . ) Duch. ex Poir; c . pepo L.; c. nfolia Bouche ( chiverre); c. maxima Duch .; c mixtaRicaPangare. Cucurbita The first two species are the most common. .

111

SPECIEร ACCOUNTS

AGRICULTURE TABLE 6.5

Vegetable ( Spanish Name) Acelga Ajo

Apio Arracacna Arveja Ayote

Descriptive Data for Costa Rican Vegetable Crops

Areas Where Commonly Seen Cartago

Cultivar

Commonly

Seen Lucullus

Vblcdn Irazu Parafso

Local (criollo) Tall Utah 52 7 OR ,

-

Washington self blanching Local Reserve early

perfection , sprite Local , sello de oro

Berenjena

Black beauty

Broccoli

Gem

Camote

C- 15

Caupi ( rabiza)

Centa 105 Yellow granex , tropicana

Cebolla Chayote

Coliflor Col de bruselas Cubaces Lechuga

Mai'z (elote) Melon

Okra Papa Pepino

Pepino Paerro Rabano Remolacha

112

.

9

DF DF, N

Direct Direct

1

Cooked Soup , cooked ,

Information limi DS IF

Direct ( roots ) Direct

3, 4

Vblcan Irazu , Cartago Ujarras ,

200

300

3

240 130-65 60-80

0.2

0.2 I

90 150

250 225

2 2

2-3

2 3

-

1๓

2๓

I

1.0-1.2

0.6-0.8

n.d. 3

n .d .

n . d.

0.75 0.5

0.3

125

3๓

2

0.2

100- 140

40

507

1

0.5

0.1

90

2

2๓

3๓:

3๓ 6๓

3

Months

0.10-0.15 Home garden

90

80- 110

0.4

0.6

168

4๓

2

0.5

1.0

2๓

3๓

Home garden 75-90

3

Home garden 0.25-0.3

Home garden 0.25-0.3

1๓

1๓

2

Direct

100- 140

1.0

0.5

90

90

2

Direct

120-80

2-3

n.d.

IS, N

Transplant

1.2

Same as snap

Ana

Direct

2

Cooked

spp., DF DF, DS

Transplant

1, 4

Cooked , pickled

DS , IS, DF N , V

Direct

2

Cooked

IF IF

Transplant

30

Transplant

30

Information limil

Direct

DF, V, IF If

Transplant

2, 4

Meseta Central

2, 3

Green seeds cooked Salad

Generalized

2, 3

Cooked , tamales

Dry Pacific

3, 4

Fruit

2

Rare

1.2

Vblcan Irazu Meseta Central

3

Local

Rare Rare

I

Detroit dark red

0.4-0.5

Cooked

Meseta Central Above 1 ,2๓ m

Meseta Central (Cervantes ) Meseta Central (Cervantes )

1.0-1.2

45-60

2

( limited ) Frozen ( rarely used )

2, 3

2 2

2

.

Cooked Salad , pickled ( limited ) Fruit Cooked ( limited ) Salad , fresh Cooked ( limited )

salad

.

Same as melon

IL DF, IL IS , N IS , N

Direct

Direct Direct

Transp|afl (

90-120

90- 120 90- 180

Transplant

IS , IF, N , DS, V DF IF DF, DS , IF, V

Number of Applications

100- 150

120

Direct

.

Fertilization ( kg/ha )

Home garden

Transplant

bean

Harvest

Distance within

0.7

2, 4

2

Distance between Rows ( m )

Days to

Row ( m ) 0.3

Practice

.

Days to Transplant 30

DF, IF, V, N 1L IF

Rare

Cherry belle

.

Sowing

Soup, dessert cooked Cooked ( limited ) Frozen ( limited )

Clemson spineless Atzimba Poinsett American flag

sauce

canning

Meseta Central ( rarely seen ) San Antonio de Belen Rarely seen

Vblcan Pois

White bouton Great Lakes Local white and yellow com varieties

2 2

Condiment Soup , salad ( limited )

2, 1 , 2,

1 , 2, 3

generalized

Snowball , X, Y Jade cross

Transplant

Type of Farm System

Cartago/Tejar, generalized Cartago ( rarely

seen )

Local

Local

Cartago

Problems with Other Biological Organisms8 IF, DE, IF, V, N

Uses in Local Dishes Cooked , salad , cooked hot

30

60

30

120- 80 60 90

90- 120

90- 140

120-80 90- 120 60-90

^^splant

Home garden 90- 150

Direct

30

Direct

60-80

0.15

1.5-2 0.7-1.0 1.2

0.15 0.3 0.25

N

0.08

0.10-0.15

Home garden

4

p

0.5-1.0

n .d .

n .d . n.d .

0.2 0.2

260 150

5๓

2

0.15

n . d.

n .d.

n .d .

0.1

n .d .

n.d .

n .d .

0.03

1๓

506

2

0.5 1.0

640

n .d . n .d . I

113

AGRICULTURE

SPECIES ACCOUNTS

—

TABLE 6.5 corn.

Vegetable (Spanish Name ) Repollo

Sandia Tomato Vainita

Zanahoria

Cultivar Commonly Seen

Golden acre, Copenhagen market Charleston gray Tropic floradel , manapal tripigro,

.

tropared

Tendergreen , harvester,

Guaria , extender, contender

Chantanay

Local

Problems with Other Biological Organisms*

Type of Farm System 2, 3

Uses in Local Dishes Cooked , salad

Guanacaste Meseta Central , Tilaran .

2, 3 2, 3

Fruit Cooked , salads ,

DF DD, IF, V, N

Direct Direct

Meseta Central ,

2.3

Cooked

DF, DS, IF, V

Direct

Cooked soup

IS , DF IL

Direct Direct

Areas Where Commonly Seen

Meseta Central , Zarcero

Bijagua

.

Tejar Guayabo,

Vblcdn Irazu Vblcan Irazu

2 1

Generalized

processed

.

Sowing

Practice

IF DF

Transplant

.

V, IF, DF, N

Hierbas de sabor Arguedas and c . Gonzalez Villalobos, Recomendaciones( para la SOURCE: Most of this information was adapted from o. P6rez comments from: พ. Canessa and J . Hernandez Estacion siembra de hortalizas (San Jos£: MAG - UCR , 1978. ) Additional Rica , 1980). , Costa , Alajuela Experimental Fabio Baudrit , Universidad de Costa Rica 1 . Beds 0.9- 1.0 m wide. 2. If grown for storage , add 1๓ kg K:0/ha . 3. N.d. = no data . M . Hansen

freIn the tropics throughout the world , root tubers the . In carbohydrates of source quently are an important is crop root important most the New World and Africa family Eu , the of ) utilissima . M ( Manihot esculenta = or phorbiaceae , commonly known as yuca , cassavafor, so cultivation under been has ) . It manioc ( fig. 6.20 long that it can no longer be found in the wild . The genus is strictly American , although M . esculenta is cultivated: throughout the tropics , with two centers of specialization northeastern Brazil and western and southern Mexico. M ., esculenta is believed to have originated in the former and more since a greater variety of cultivars or clones most species the since , and there exist plant uses of the ) 1968 6 n Le ( there found also are it to related closely drought extremely is , it climates tropical all in Grown resistant and does well in areas where there is a long dry season or rains are sporadic and where the soil is of moderate to poor quality. Although primarily a lowland to 2 ,000 tropical crop , it can be grown at elevations upvery poor on m in the Andes. Because it can be grown soils that are unsuitable for other crops , it is often the last crop planted before a fallow cycle begins or is often the only crop grown. It is cultivated throughout Costa Rica and appears to be most abundant on the west coast.

.

*

-

.

114

is a short-lived shrub 1-5 m in height , with large palmate leaves. It is extremely variable in all its characters. The growth form is highly variable de pending on clonal variety and method of propagation . Plants grown from seed tend to have a single stout trunk with perhaps a small amount of branching at the ape whereas those grown vegetatively tend to have a trunia that branches once or twice , with these branches then, dividing further into an umbrella-shaped growth form The latter form is more common , since most propagation by humans is vegetative. Whether the trunk bifurcates close to the ground or higher up , the angle of the branches to the main stem and the angle of leaves to the branches are varietal characters, determining whether the plant has an erect or a spreading growth form , The leaves are extremely variable in size , shape , color, number of lobes , shape of lobes , and so forth , depending the on clonal variety. In general they are near the apex of are branches and are spirally arranged . In form they large , glabrous , and palmate with from three to eleven lobes , five to seven being average . Shape of the lobes can vary from strictly linear-lanceolate to obovate-lanceolate in some varieties there is even a distinct bulge than mato> occur anywhere along the length of the lobe from base4 to apex ( Rogers 1963). The size of a lobe ranges from 20 cm . Both position on the mink and environmental conditions affect the number of lobes per leaf . As oof M . esculenta

Yuca ( Yuca , Cassava)

-

Days to Transplant 30 22

Days to Harvest 90- 120 120-50 90- 150

60-90

75-90 Home garden

Fertilization

Distance between Rows ( m ) 0.25

Distance within Row ( m ) 0.25

N n .d .

2-4 1.2

0.5

-

3005

1 2

p n .d .

Number of Applications n .d .

600

3

1๓

1๓

2

1๓

1๓

2

70- 150

0.1-0.2

0.3

( kg/ha )

0.3

2๓

4. Specialized production practices in Ujarras/Cartago. 5. Additional foliar applications of Poliboro (5๓ g per 2๓ I water ) and molybdenum ( 10 g per 2๓ 1 water). 6. If less than 50 ppm of p in soil analysis. 7. Add 1๓ kg K:0/ha. 8. IF = foliar insects; IS = soil insects; V = vims; DF = foliar diseases; DS = soil diseases; N = nematodes.

advances up the trunk , the number of lobes per leaf increases; it is not uncommon to find cultivars in which the basal leaves are entire. In addition , the number of lobes is generally smaller during the rainy season than during the dry season . This phenomenon is noticeable within a clone, since the average life of a leaf is only 1 2 months . The leaves are bicolored , with the top ranging from a bright green to green with a tinge of red to green and yellow variegation , while the underside glau cous , ranging from gray to bluish . Color isof more the midrib and the veins is also variable , ranging from green to yellow to red . In some clones there may be variation in the vein color, with a different colorintraleaf each side ot the midrib ( Le6n 1968). The petiolar coloronvaries from purple to red to green and from uniform to splotched depending on the variety. The petiole longer than the lamina , being from 20 to 40 cm islongusually . At the base พ a pair of lateral stipules that are long or short , have smooth or dentate edges, and usually have three to five lanceolate lobes each . The leaf scars are very noticeable nd variable in color, ranging from gray greenish yellow to dark green , red to dark brown , ortostreaked with Pu plc

-

branched inflorescence with stalks that are 5 15 cm long and bear approximately fifty to sixty small unisexual flowers, the majority being male , with the female ones situated at the base of the panicle. Hermaphroditic flowers occur in some clones, as do panicles with only staminate flowers . All floral types lack petals; the sepals look like petals. There are five glabrous sepals that vary from greenish to yellow; in some clones they may be tinged with red or purple, either in the center or around the edges. Sepals also contain latex glands , as do most of the other parts of the plant. Staminate flowers are smaller than pistillate flowers and have a short , straight pedicel 0.5- 1.0 cm long and a calyx 6 8 mm long . In the center of the flower is a fleshy orange disk with nectaries; around this are two whorls of stamens, altemately short and long . Pistillate flowers have a long curved pedicel 1 2.5 cm long and a 1 cm or more in length . In the center of the flower iscalyx a large yellowish or reddish ten lobed disk , on which sits an elliptical six sided ovary 3-4 mm long that contains three carpels, each with a single ovule. The pistil con tains a short style that is divided into three finely lobed stigmata . The primary pollinators , thought to be dipter *Tbe ans , visit the flowers , which open for 2 3 h aroundcommonly are in the axils of the for 8-10 consecutive days. Female flowers open leaves, inflorescences though they may also be near the ends of the 6midday 8 days - , before the male flowers. Very few flowers set ches. They are panicles, or a loose, several times seed a process that takes 5 months. The fruit is a green

-

^

^^

-

-

-

-

-

-

-

- -

115

ends on table ) in market . Guanacaste Province , c Rica ( photos , D. H . Janzen ).

FIGURE 6.20. Manihot utilissinui . a View under canopy of mature crop. b. Tubers ( on ground ) and sectioned tubers

( white

1

oval capsule 1.5 cm in length and contains three seeds. Upon maturation , the hard capsule explodes and ejects the mottled grayish seeds , which are flat and elliptical , 1 cm long , and have large caruncles attached to them. When grown from a seed , M . esculenta นรนฝly has a single main root; tubers develop as swellings on some of the secondary or adventitious roots. When the plant is grown vegetatively, pieces of the stem 20-30 cm long are planted with a few inches sticking out of the top of the soil; adventitious roots , some of which develop into tu bers, form from the leaf scars. Extensive variability ex istร in tuber size, number, shape, color, flavor, and amount of secondary compounds. The average number of tubers per plant is five to ten , each being 5- 100 cm long and either cylindrical , branched , or tapering. The tuber consists of two parts: the cortex , or rind , which is vari able in color, ranging from white to dark brown or pink to red , and containing the phloem; and the xylem , which is the edible portion of the tuber and composes approxi mately 80% of total tuber weight . The xylem contains few latex glands . The carbohydrate content , however, is very high; it can supply more calories per area planted than com , rice, or other tubers . Protein content , on the other hand , is low, averaging 2% of fresh weight . The

-

-

*'rir,ะ?:;r : . ระ SSEST S ร;E5EH5

dent of the distribution of HCN . Ybca may be prepared for eating in a number of ways. Sweet yuca can be peeled and eaten raw, whereas bitter yuca must be cooked and repeatedly washed to remove die HCN . Sweet or bitter forms may be cut into strips or sections and boiled , cooked , or fried . Flour can be made from sun -dried slices. Starch is extracted by grinding washed , peeled tubers and then squeezing them in repea ๗ changes of water. I is en cooked or fned Mon eating . Yuca is also used as a broth for making a tvoe of iUs hea vegetable stew. In i to it agglutinate causes into small balls , forming which tapioca . The major nations importing tapioca are the United States and countries of Europe, where it IS used as a dessert and in the manufacture of various confectionery products . The major producer and exporter of tapioca before World War II was Indonesia. Two vimses are the major serious pests of M . esculenta . In many areas the mosaic vims can be a serious problem , causing up to 95% mortality on susceptible cultivars. Vectors for the vims are whiteflies of the genus Bemisia. It can also be spread by the use of infected cuttings for propagation . The vims attacks only young leaves, with symptoms occurring in 2-3 weeks if the

,

116

^

rSSESr^r

AGRICULTURE

.InTnesia

*

^

SPECIEร ACCOUNTS

into the stem and the root. Infected leaves are usually deformed and small . Brown streak vims is also a serious pest , especially in the coastal regions of Africa. It attacks older leaves , turning them yellow, but no deformation of the leaf blade occurs. Brown streaks appear in the corti cal stem tissue and congeal into patches , followed byshrinkage and death of intemodal tissue and discolored areas in the roots. The vims also makes the plants sensi 1 s e are rea p ® of 8 p ' a nt ?to . i cl1 1 1,. ,.,๗, ร.,, M HY ,ร /๓ ' because of its resistance to locusts. There are a number of programs for breeding M . esculenta with other species in the same genus in an attempt to produce strains with resistance to the mosaic and black streak vimses as well as to increase the protein content of the leaves.

^

“ บรพ Jon pX " “- "' "^ * '“'"

._

1

,

" . pSly

Leon, J. 1968. Fundamentos botanicos de lOS cultivos tropieales . San Jos6: Instituto Interamericano de Cien cias Agricolas de la OEA . Purseglove, J. พ. 1968. Tropical crops: Dicotyledons. VblT 1. New York: John Wiley. Rogers, D. J. 1963. Studies of Manihot esculenta Grantz and related species. Bull. Torrey Bot . Club 90:43-54.

leaves may contain up to 18% protein and are eaten parts of Africa in addition to the tubers. Leaves also make excellent cattle fodder owing to their high protein content . The tubers themselves are rich in vitamin B , iron. and phosphorous but are low in calcium ( Leon 1968 ). For a long time people thought there were two forms of yuca one with soft , white, sweet xylem , the other with a tougher, more yellow, bitter xylem but now it IS realized that there is a gradient from the bitter form to the sweet form . The bitterness is due in part to the presence of hydrogen cyanide ( HCN) and varies with environ mental conditions. In the sweet forms HCN is usuallyis restricted to the rind or cortex , but in the bitter form it also usually found in the xylem , which must be processed to remove it. The bitter form is found predominantly ii1 areas with very poor soil conditions and can be grow where other crops will not grow. It is long -lived , rc quiring a year or more to produce tubers. Tuber pffr# duction may last 3 to 4 years without deterioration in tf tuber quality. The sweet form , in contrast , is short - lived faster growing , and occurs primarily in areas with bettef soil conditions. Tubers may be harvested after as little * 6 months, but after 9-11 months the tuber quality start also can clone or variety a of to deteriorate. The flavor

—

—

117

INTRODUCTION

7 PLANTS

INTRODUCTION

G . ร . Hartshorn

the reader to the This chapter is meant to introduce emphasis is given to vegetation of Costa Rica. Majorunderstanding , or lack the description and ecological thereof , of the vegetation of the seven field sites on which OTS courses have focused tropical forest vegetation can General descriptions of coverage by Richards ( 1952) be found in the pantropical treatments by Whit and Baur ( 1968) or in the regional and Jenflc ( 1974). Though no more ( 1975) and Longman vegetation exists, here regional treatment of Neotropical I shall attempt to provide a cursory hemispheric perspecfive.• ecological underA significant proportion of ourRica is a direct con standing of the vegetation of Costa past fifteen years by sequence of studies done over the researchers. Studies ori OTS students , professors, andand ecology ented toward natural history orevolutionary . research have dominated OTS sponsored affiliated of study pilot Except for the Holdridge et al. ( 1971 )known of forest is nothing practically , zones tropical life of processes ecosystem functions or the ecophysiological . al et Mooney ( tropical plants in their natural habitats

-

พ

-

necessity, this chapter takes an 1980). By both design andapproach . evolutionary ecological courses and emThe seven sites frequented by OTS National Parkj Rosa are: Santa recently phasized in this chapter ren ( Refuge Pdo Verde National Wildlife La , Selva ) Caballero Rodriguez Refugio Rafael Lucas , ALLPark National Biological Reserve, c orcovado Cruces Field Static , Las * h verde Cloud Forest Reserve erro - Mu the , and Garden Botanical Iropical and vernacular title erte ("Hill of Death ). The shortened M \\ he Selva La example shown here in italics for

.

A

\

11

! ;

11

ะะ

Botanical Exploration century brought m The middle and late nineteenth Eu interesting , often eccentric, but very competent i first The . ) 1957 pean naturalists to Costa Ricaof(Pittier Costa of vegetation the naturalist to publish an account Andres Oersted ( 1871 ), botanist Danish the Rica was collections ๙ who in 1846- 48 made the first substantial botanists foreign Costa Rica 's flora . Other notable the Polish gard include collected in Costa Rica physician Carl Hoffmann , the German , the Warscewicz

the swamp between the mangroves Monospecific stand of Pterocarpus officinalis trees in Park , Osa Peninsula , Costa Rica National , and the mixed forest at Llorona Corcovado ) . ( photo, D . H . Janzen

118

1-

(

t s to used throughout the chapters title, unless spec tieally represent the correct and longer

o n egeta

stated otherwise,

gardeners Hermann Wendland and Jules Car0! and the German professor Helmut Polakowsky. Moritz Wkgner and Carl Scherzer The German naturalists Rica during 1853-54 , offering a ( 1856) explored Costa fascinating account of their arduous journey from Puerto Viejo de Sarapiqui to the Meseta Central . A new era began in 1887 with the arrival of the Swiss educator, botanist , and naturalist Henri Pittier, the found ing of the Museo Nacional , and the official' involvement of Costa Rican botanists. Anastasio Alfaro s 1888 listing of higher plants and ferns known from Costa Rica totaled 1 ,218 species . Standley ( 1937-38) suggests that three Alfaro listed were discovered by fourths of the species ’ laudatory writings indicate a strong and ร Pittier Oersted. with Anastasio Alfaro that not collaboration productive knowledge increased the of the Costa Rican only greatly Costa Rican bota stimulus to profound was a flora but , Adolfo Tonduz , Juan as such Cooper naturalists nists and , , Carlos Wferckle Brenes. Pitand Alberto Biolley Pablo years in Costa culminated sixteen Rica with a ) tier ( 1957 on essay plants the useful of Rica ( pubCosta remarkable ) of Pittier The and extraordinary impact . 1908 in lished can Rican from be botany gleaned Stand Costa on Alfaro ley’s comments that the Costa Rican National Herbarium in 1903 was unequaled south of the Rio Grande del Norte ( Mexico) and increased the known flora of Costa Rica to about 5,000 species. After Pittier 'ร departure in 1903, Alberto Brenes made the first systematic collections for Central America in the mountains around San Ramon . Brenes’s fine work is evident in the substantial number of endemic species based on his collections and in the fact that some of the new species collected by him in the 1900s have not been collected again. Other Costa Rican botanical contributors of the early 1900s were Oton Jimenez , Ruben Torres Rojas, Juvenal Valerio Rodriguez, and Manuel Valerio. The 1920s brought the prolific botanist Paul Standley to Costa Rica during the Chicago winters of 1923-24 and 1925-26. Standley ' ร prodigious collecting of 15,000 plants led to the 1937-38 publication of his Flora of Costa Rica , an annotated listing of 5,815 species of higher plants native to Costa Rica. Present estimates of the Costa Rican higher plant flora are about 8 ,000 species ov. Buiger, pers. comm .). Such an extraordinarily rich flora in so small a country (51 ,100 km 2) is truly remark able and no doubt is still an important attraction to botathsts. The Chicago Field Museum of Natural History has recently undertaken to publish a new Flora Costaricensis under the direction of William Burger. Two issues cov ering fifteen families have been published to date ( Burger19? 1 , 1977 ). The first attempts to describe the phytogeography of osta Rica differentiated a few floristic regions , primarfly on the basis of altitude . Pittier ( 1957 ) recognized three German

ret

ะ^

^

altitudinal belts: ( 1 ) a basal zone from sea level to 1 ,000 m with mean annual temperatures between 280 and 21๐ C; ( 2) an intermediate or mountainous zone from 1 ,000 to 2 ,600 m with mean annual temperatures between 21 and 14° C; and (3) a superior or Andean zone above 2 ,600 m and mean annual temperatures between 15° and 5 ° c . In an enlightened essay on the phytogeographic regions of Costa Rica , Werckle ( 1909) describes four re gions. ( 1 ) Atlantic or Caribbean region from sea level to 800 m; ( 2) Pacific region from sea level to 800 m; (3) temperate region from 800 to 1 ,500 m; and (4) cold region above 1 ,500 m. Werckle gives 2 ,000 m as the lower limit of frost. He also says there are no paramos in Costa Rica a statement no doubt generated by his ex perience in Colombia and his unfamiliarity with the high est parts of the Talamancas. Standley ( 1937-38 ) follows Werckle 's phytogeographic divisions but states that the cold region must be subdivided into lower and upper belts. Standley is the first to point out the difficulty of assigning a single alti tudinal limit to a particular type of vegetation.

-

—

-

Holdridge Life Zones

L. R . Holdridge ร dissatisfaction with existing vegetation classification systems applied to the mountainous regions of Haiti led to the development of a new classification system based on simple climatic data ( Holdridge 1947). In Latin America , Holdridge 's classification system has been extensively used to prepare ecological maps of twelve countries and as a basis for detailed studies of land -use capability, natural resources management , and environmental impact assessment. Preliminary ecologi cal mapping and trial applications of the Holdridge system have also been successfully conducted in Nigeria , Thailand , and East Timor. Because of the objectivity, applicability, and generality of the Holdridge classifica tion system , it is used here as a framework for describing the vegetation of Costa Rica. Holdridge's classification system gives first importance to temperature and rainfall and the seasonal vari ation and distribution of these two climatic parameters as the primary determinants of the world vegetation . The bioclimatically defined units are called Life Zones. Al though a Holdridge Life Zone may be somewhat anal ogous to the plant geographer ’s “formation ' or the ecol ogist 's “ biome," Holdridge avoids the botanical bias of the former term . Each Holdridge Life Zone has a distinctive vegetation physiognomy and structure that occurs wherever similar bioclimatic conditions exist. Since Holdridge’s Life Zone system is independent of floristic relationships , the same Life Zone may occur on opposite sides of the world for example, the tropical moist forest Life Zone in Latin America , Africa , and Southeast '

*

Asia.

—

119

INTRODUCTION

PLANTS

Holdridge ( 1967) states that associations ofarenaturally and animals recog evolved communities of plants an association as a nizable in the tropics, and he defines has distinctive unique ecosystem within a Life Zone that plants and ani associated and environmental conditions Holdridge in the mals. Associations form a lower level the scale in difference The . classification system association level is sobetween that great Life Zone level and the an “associ Tosi has added an intermediate leveltotermed the “catena ” seation grouping ” that is analogous “association grouping ” level quence in soil science. Themedium -scale ecological mapis particularly useful for ping (Tosi 1976; Hartshorn 1977). , and association The Life Zone, association grouping are all consystem levels in Holdridge’s hierarchical primary vegetation , , or , undisturbed natural cemed with disruptive influence on natMan’s pervasive and usuallythe classification ural vegetation enters into Holdridge types of Altered . system as a fourth -order component successecondary and vegetation such as pasture or crops naturally the to in relation sional stages are described . replace they associations occurring Holdridge’s Life Zone system is probably known best

-

(fig. 7.1) depicting for the deceptively simple diagramActually , the diagram . Zones Life some thirty hexagonal , for it is a appears it than complex is considerably more 116 Life approximately the of three-dimensional model parameters three of consists diagram Zones on earth . The isogonally. Mean an scaled logarithmically and arranged is defined as the Celsius degrees in nual biotemperature , with zero substituted mean of unit- period temperatures 0 ° c and above 30° c. below values for all unit -period is used as the millimeters in Mean annual precipitation . The third parameter in the second independent parameterevapotranspiration ratio , is Life Zone diagram , potential is ; it the parameters dependent on the two independent ( PET ) ratio of mean annual potential evapotranspiration on dependent is PET . Since precipitation to mean annual the constant temperature , Holdridge et al. ( 1971) derived annual biotemperature to 58.93 to be multiplied by mean The PET ratio give PET in millimeters of precipitation.moisture avail is a biologically meaningful index of occurs when the ability; for example, a PET ratio of 1.0 PET, and it also to the mean annual precipitation is equal ) and subhumid (PET 1.0 ratio < PET ( humid separates ratio > 1.0) humidity provinces.

-

OR PLANT FORMATIONS THE CLASSIFICATION OF WORLD LIFE ZONES

DIAGRAM FOR

by L . R Holdridge

ALTITUDINAL LATITUDINAL REGIONS IWM
nds , has produced a remarkable array of altitudinally compressed life zones . The protected Pacific valleys and slopes are in a dry season rain shadow nearly as severe as ln the Guanacaste lowlands . Only near the diyide do the moisture- laden winds spill over tocontinental bathe the uPpermost Pacific slopes in a nourishing mist that lnc°ngrous with the bright , sunny weather. Thoseseems Moisture-laden winds forced up the Atlantic valleyssame and

slopes have produced a beautifully sculptured and streamlined elfin forest on the exposed upper Atlantic slopes . The ecological map of Costa Rica (Tosi 1969) shows the following Life Zones in the Monteverde area: ( 1 ) Tropical Premontane Moist Forest in the upper San Luis valley and along the cliff edge; ( 2) Tropical Premontane Wfet Forest encompasses most of the extended community and farms; ( 3 ) Tropical Lower Montane Wet Forest on the upper Pacific slopes such as in the vicinity of the TSC field station and much of the Sendero ( trail ) Bosque Nuboso; ( 4) Tropical Lower Montane Rain Forest along the divide and mountains generally above 1 ,5๓ m , ineluding the Elfin Forest; and ( 5) Tropical Premontane Rain Forest in the upper Pertas Blancas valley. The first three Life Zones in the Monteverde area , as well as the elfin forest , should be considered atmospheric associ ations owing to the strong influence of the dry -season trade winds . The relatively level , swampy area along the Sendero Pantanoso is a hydric association . Most of the continental divide and upper Pacific slopes have had substantial inputs of andesitic volcanic ash , suggesting that they are largely edaphic associations . Perhaps only the Tropical Premontane Rain and the more easterly Lower Montane Rain Life Zones are close to the respective climatic associations . Practically the entire Monteverde Reserve can be considered cloud forest because of the prevalence of moisture-bearing clouds throughout the year. During the April - November rainy season the trade winds lessen in velocity but still contribute substantial moisture to the Monteverde area . Convectional thunderstorms bring al most daily rain during May to October with a slight veranillo in July. The physiographic complexity of the 141

\

RESERVA BOSQUE NUBOSO L EY E

MONTEVERDE (LEGEND N D A

)

—

L

INTRODUCTION

accentuates the dominant role of wind Tilardn Cordillera determining types of forest in the Mon in and clouds . The incredibletheabundance diversity of teverde area and bryophytic epiphytes andsome growing

—

both vascular on top of each other amid dense carpets of soft mosses 10 cm thick imparts a characteristic aeriel-draped cloud forest where it is sometimes impossible to locate the leaves of the host tree. Of the major cloud forests in tropical America produced by the northeast trade winds ( Rancho Grande in Venezuela , the Blue Mountains of Jamaica , and the Luquillo Mountains in Puerto Rico), cloud forest is most luxuriantly developed in Monteverde. Within the Monteverde Reserve , Lawton and Dryer ( 1980) describe the vegetation of six forest types: cove,

—

Strangler Ficus tuerckheimii , Meliosma verrucosa , Quararibea platyphylla , Sapium oligoneuron , and pachy stachys, as well as several members of , Myr

ร. Lauraceae taceae , and Sapotaceae. Of the six forest types , the cove forest has the least-developed epiphytic load; bryophytes cover less than 30% of trunk surfaces , while canopy epiphytes are mostly drought- resistent , shrubby forms. Leeward cloud forest is more exposed to the trade wind spillover than is the cove forest and occurs in a broad , irregular arc from Cerro Sin Nombre to the Brillante Gap. This forest ( fig . 7.11a ) has a more open can opy, is shorter ( 25-30 m ), and has heavier epiphyte loads than cove forest. Byrophytes cover 50-70% of the tree tranks , reflecting a much less stressful dry season . The authors claim the leeward cloud forest is transitional from tropical Lower Montane Wet to Lower Montane Rain Life Zone. Typical tree species include Calocarpum vir-

leeward cloud , oak ridge, windward cloud , elfin , and swamp forests. The following summaries are based on the Lawton and Dryer descriptions. Cove forest within the Monteverde Reserve is restricted to the protected ide , Citharexylum macradenium , Daphnopsis americana , western base of Cerro Roble and the upper, west-facing the strangler Ficus crassiuscula , Guarea tuisana , Melislope of Quebrada Alondra . The forest is well developed , osma vernicosa . Persea americana , Quararibea platy30-40 m tall , and evergreen despite the dry-season loss phylla , Sapium pachystachys , Sloanea medusula, and of moisture . Characteristic tree species include the huge several members of the Lauraceae and Myrtaceae.

*•°

CURE 7.11. Profile of upper Pertas Blancas valley in Monteverde; the very steep, uppermost

FIGURE

142

7.10. Ma|

s pes are covered with elfin forest ( photo, G . ร. Hartshorn ).

PLANTS

Oak ridge forest occurs on the exposed ridges of Cerros Sin Nombre , Amigo, and Roble. The dominant oaks, Quercus corrugata and Q . seemannii , may attain 1 m dbh and 25 m in height on the less windswept ridges. Epi phyte cover is practically complete on all surfaces , in cluding the ground: 75-100% for bryophytes and 30-80% for angiosperms. Other tree species include Billia hippocastanum , Brunellia costaricensis , the strangler Ficus crassiuscula , Guatteria consanguinea , Hieronyma poasana , Oreopanax xalapensis , Persea ameri cana , p. schiedeana , the stilt-rooted Tovomita nicaraguensis , Weinmannia pinnata , พ. werc klei , Zanthoxylum melanostichum , and members of the Lauraceae and Myrtaceae . Windward cloud forest is the most extensive type within the lower montane rain life zone in the Monteverde Reserve. It is an extremely saturated habitat with a very discontinuous canopy, seldom more than 20 m tall , and a dense understory. Many of the tree species are the same as those in the leeward cloud forest; typical tree species include Alchomea latifolia , Ardisia palmana , Citharexylum macradenium , Daphnopsis americana , Dendropanax gonotopodus , the strangler Ficus crassiu scula , Guarea tuisana , Guatteria consanguinea , Hieronyma poasana , Inga longispica , Meliosma sp., Persea schiedeana , Sapium pachystachys , Weinmannia wercklei , Zanthoxylum melanostichum , and several members of the Lauraceae. Windward cloud forest has the most luxuriant load of epiphytes , including beautiful Begonia estrel lensis and Hydrangea peruviana , plus carnivorous Utricularia spp . Elfin forest occurs along the crest of the Brillante Gap as well as on other very exposed ridges. It is a short , gnarled , dense , sodden forest whose canopy appears sculptured and streamlined by the relentless wind . The smooth profile of the elfin forest viewed (fig. 7. Mb ) from the overlook on the old road cut to Pertas Blancas belies the tangled mass of roots , branches , and fallen dwarfs that is nearly impenetrable. The plants that make up the elfin forest are the same ones as in the windward cloud forest , except for those species such as Cecropia polyphlebia and Heliocarpus popayensis that are unable to tolerate the wind stress. Stilt-rooted Clusia alata trees may protrude above the low canopy as true emergents , especially on the ridge crest above elfin forested slope. The Araliaceae , Didymopanax pittieri , Oreopanax nubi genum , and o. sanderianus are common trees in the elfin forest . Swamp forest occupies the poorly drained plateau between Cerro Roble and Cerro Sin Nombre. The authors note that the western drainage is a blackwater tributary to the Rio Guacimdl . The forest is a mosaic of scattered patches of trees to 25 m tall among lower thickets of small trees. Characteristic tree species include Alchomea 144

latifolia , Clusia alata , Conostegia spp. , Dendropanax arboreus , D. gonatopodus , Guatteria consanguinea , Hieronmya poasana . Magnolia sp. , Persea schiedeana , Podocarpus oleifolius , Quercus corrugata , Q . seenuinnii, Sapiumpachystachys , Tetrorchidium sp . , Tovomita nicara guensis , and Weinmannia wercklei . Lawton and Dryer ( 1980) do not include descriptions of the Tropical Pre montane Rain Life Zone in the upper Pertas Blancas valley or the Tropical Premontane Wet Life Zone around the Monteverde community. The Pertas Blancas valley is so poorly known botanically and ecologically that it is presumptuous to try to characterize the forests there. Suffice it to say that the forest is taller and that huge strangler figs are much less frequent than in other Monteverde forest types. Only remnant patches of forest re main around the Monteverde community, but they clearly indicate that it was a magnificent forest , extremely rich in species of Lauraceae and Sapotaceae, as well as huge strangling Ficus . The tree species in the vicinity of the Monteverde community are reasonably well documented in the checklist of trees found later in this chapter.

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LAS CRUCES

The tropical premontane rain forest that covered the upper Coto Brus valley has been virtually exterminated in the past fifteen years. One of the few remaining remnants occupies 100 ha of Las Cruces. It is a tall forest ( 30 35 m ) with abundant Quercus spp. and Lauraceae. The epiphyte load is appreciably less than at Monteverde , pre sumably because of the slight dry season and absence ๙ wind-driven mist at Las Cruces. L. J. Poveda is pre paring a report on the Las Cruces forest .

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TABLE 7.14 Province

INTRODUCTION Stand Characteristics of a 0.4 Ha Plot at Km 97 of the Inter-American Highway on the Cerro, Cartago Density

Species Quercus costaricensis Sficonia bipulifera Vaccinium consanguineum Weinmannia pinnata Oidymopanax pittieri

Sublocal: lop 5 species Subtotal: other 15 species

Frequency

Basal Area

Stems (% )

(% )

101 (41.2) 39 (15.9) 25 ( 10.2) 16 (6.5) 10 (4.1)

m 1 (% )

20.43 15.05 11.82 8.60 6.45

10.93 (61 ) 1.78 (9.9) 0.71 (3 9) 1.75 (9.8) 0.85 (4.8)

40.89 13.63

191 (78 ) 34 (22)

62.35 37.65

16.02 (89) 1.90 ( 11 )

76.58 23.42

17.92 ( 1๓)

1๓.๓

245 ( 1๓) Total: 20 species SOURCE. Data from Holdridge et al. 1971 , site 6

1๓.๓

Physiognomic Features and Patterns

Distinctive tropical forest features such as buttresses , crown shapes , drip tips , epiphytes , and lianas have long attracted naturalists , phytogeographers , and plant

ecologists (Schimper 1903; Richards 1952), leading to considerable speculation on the functional significance or evolutionary advantage of particular structures . Unfortunately, our understanding of some tropical forest features has not progressed much beyond the speculative stage . Notwithstanding the dearth of testable hypotheses such features as buttresses , drip tips, and epiphyte distri bution often attract OTS students. In this section the more prominent physiognomic features are briefly reviewed to highlight hypotheses , point out ecological patterns , and serve as entries to the pertinent literature .

.

BUTTRESSES

AND

STILT ROOTS

Of all the distinctive or interesting physiognomic features CERRO of tropical , none has elicited more conjecture and Pension La Georgina is close to the present transition speculationforests than the question , “Why do some tropical from oak forest to paramo. The Holdridge et al. ( 1971 ) trees have buttresses ? ” Buttressing ( fig . 7.12) is largely Villa Mills study site just south of the pensidn illustrates restricted to the tropical basal belt in Costa Rica and is ( table 7.14 ) the complete canopy dominance of Quercus best developed in tropical Moist and Wet Life Zones. costaricensis . Most of the oak forests along the Inter- Buttress height is well correlated with trunk diameter American Highway have been severely degraded by log ( Holdridge et al . 1971 ) , but not all canopy trees are but ging and charcoal - making . The scandent bamboo, Swul tressed . Buttressing is more prevalent in poorly drained lenochloa sp., aggressively dominates the shrub layer of areas , but some exceptionally well buttressed trees , for be the degraded and open oak stands and appears to , example Dussia macroprophyllata and Huberodendron inhibiting the regeneration of oaks . potinoi , occur on well-drained sites. The height , thick Pdramo vegetation on the Cerro is often dominated by °ess, form , and degree of bifurcation of buttresses are Hypericum shrubs 1-1.5 m tall with occasional emerg generally very useful in species identification. Buttress ( to 5-8 m ) trees. The drier, rocky hillsides are cov growth is strongly ( Richards 1952 ) , that is , the by clump grasses with occasional forbs. Bogs have all anatomical center ofepinastic the buttress near ground level interesting assemblage of plants such as Cirsium sp * and growth is strongly skewed tois attheorupper edge of the Cir Puya dasylirioides , and the Andean fern . Jamesonia . tottress . of ’ cumstantial evidence suggests that man s frequent use The various explanations of buttress formation are fire on the Cerro is effectively lowering the tree linfi grouped into four theories , discussed Richards ( 1952): Regeneration after fire is extremely slow ( Janzen 1973)* fl ) adaptive responses to wind orin gravity stresses;

-

5

-

Importance Value (% )

8.66

8.30 5.10

( 2) (4 )

negative geotropism ; ( 3) conduction shortcut; and mechanical stimulation of strains caused by winds. Smith ( 1972 ) updates and reviews the buttressing theories. An analysis of buttresses using engineering models indicates buttresses are excellent support structures ( Hen wood 1973 ) . Smith ( 1979 ) found a strong negative correlation between buttressing and bark thickness , sug gesting why thick- barked temperate trees are not buttressed. It is a common generalization ( Comer 1940; Richards 1952; Spruce 1908; Whitmore 1975) that taprooted trees do not form buttresses , and vice versa . A common mis conception holds that buttresses are attached to the cen tral trunk: “In felling tropical timber trees the cut is necessarily made above the buttresses , so the amount of waste is considerable '' ( Richards 1952, p. 74 ). A recent paper by Black and Harper ( 1979) indicates that the misconception is still alive and well. Most well - buttressed trees completely lack a central bole. Stilt roots differ from buttresses in that the former are adventitious roots . Some stilt roots elongate in a vertical plane such that they may appear as raised buttresses. The best known example of stilt roots occurs on the red man grove, Rhizophora mangle , sometimes originating more than 5 m above “ground ' level and making passage extremely difficult . Stilt roots occur in other dicot trees such as Bravaisia integerrima , Protium spp., Symphonia globulifera , Tovomita pittieri , Virola surinamensis , and nonepiphytic species of Clusia . Virola surinamensis and Protium spp. usually have vertically elongated stilt buttresses. Numerous palms also have stilt roots , for exampie, Chamaedorea spp . , Cryosophila albida , Euterpe macrospadix , Iriartea gigantea , and Socratea durissima . In the latter, the original stem below the stilt roots actu ally dies, so that all support and translocation is through the adventitious stilt roots As with buttresses, there is considerable speculation on why stilt roots have evolved. In palms at least the

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*

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145

PLANTS

Terminalia oblonga ) . Trees with exthin sheets foliating bark consistently have fewer epiphytes and climbing vines on the trunk and branches than do trees without exfoliating bark . Some bark surfaces bear spines , (eg . , Bombacopsis quinatum , Hura crepitans, Pith ecellobium dulce , and Zanthoxylum spp. ), while other species have conspicuous spines as juveniles but exceed ingly few or none as mature trees (e .g . , Ceibapentandra , Jacaratia costaricensis , Lacmellea panamensis , and Zanthoxylum mayaทนทใ ). A few genera ( e.g . , Cryosophila spp. and Xylosma spp .) have multiple- branched spines , (e g. ,

LEAVES It is well documented that the overwhelming majority of tropical tree2 leaves are mesophylls that is , between 20 and 182 cm in upper surface area ( Beard 1945; Richards 1952; Greig-Smith 1952). Many compound leaves have leaflets of mesophyll size. Compound leaves are generally more common in tropical lowlands than in tropical montane forests ( Cain etal. 1956; Grubb et al. 1963), but Tasaico ( 1959) found that simple and compound leaves occur in the same proportion in tropical montane areas as in the lowlands. Compound leaves occur commonly not only in areas with a dry season , but on species that depend on rapid increase in height (Givnish 1978). With increasing altitude , average leaf length decreases at a rate of about 0.5 cm per 100 m (Tasaico 1959). Entire leaf margins, drip tips, and thin leaves are most frequent in wet lowland forests. The adaptive significance of drip tips has long been associated with more rapid runoff and drying of the leaf surface (Richards 1952). An experiment with Machaerium arboreum ( Fabaceae ) leaflets confirms the role of drip tips in speeding water runoff and drying of the leaf surface ( Dean and Smith 1978). The occurrence of tropical forest understory or ground cover leaves with red spots or red undersurface has interested ecologists. Lee, Lowry, and Stone ( 1979) have recently shown that the abaxial anthocyanin layer en hances photosynthesis by functioning as a reflective sur-face that bounces light back through the mesophyll .

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7.12. Tree line lowered by fire at Cerro de la Muerte ( photo, G. ร. Hartshorn ). -specific that sec- and sap, are sufficiently varied yet species answer seems obvious, for monocots cannot makemany identifi species for characters diagnosti c useful they are ondary xylem; for example , Welfia georgii take is it , how true were generaliz ation quoted the If . cation years to make a large trunk below ground before the trunk same those use competen tly so finders tree local that emerges like a telephone pole. The stilt- rooted palms are characters for species identification ? an on fast grow must and intoleran t much more shade Rica bark colors are predominantly dark , Costa เท inverted cone to take advantage of canopy openings. Theis ranging from black to various hues of red , brown , and s (com evolution of adventitious stilt roots in monocotbuilding species have light-colored bark , although Very . gray a good example) allows fast growth without Holdridgea white, red ,few or blue crustose lichens may camouflage the final stem diameter at or below ground level . . Bark thickness is generally between 5 and 10 mm, et al . ( 1971) report that in the Tropical Premontane Rain bark with some species having thinner' bark , but very few have Forest Life Zone about 10 of the tree species have stilt bark than 10 mm . Smith ร ( 1979) observation that roots , but Tropical Dry, Premontane Moist , and Lower thick thicker trees do not have buttresses appears to be Montane and Montane Forest Life Zones do not have valid-barked for Costa Rican species. stilt rooted trees. Bark texture is probably more varied than are color and.. thickness; some examples are: coarsely fissured. (eg BARK , finely fissured ( e.g Cal“The bark of rain -forest trees is usually remarkably thin Lecythis costaricensis ) ophyllum brasiliense ), shaggy (e.g. , Lxsilonui di and is generally smooth and light-coloured " ( Richards ), coarsely lenticulate (e.g . , Hernandia icata 1952, p. 58 ). That gross generalization is still widely lenticulate (e.g . , most Moraceae anil dynuintha ) repeated (see Walter 1973 for a recent restatement even Mimosac ) , finely eae ) , minutely exfoliating (e.g., Bursera )sinto though it is incorrect. Isolated trees do have light , รนท , e*' , but the ruba , Inga coruscanst Pithcellobium pedicellareCartfM bleached bark that appears deceptively smooth . . ( . g e light foliating เท scalloped or concentric patterns same species in the forest generally do not have exfoliating in hu£* , and ) caribaea , Albizzia guianen sis odor with , and thickness , along ,

FIGURE

fl

-

^

-

. diveA -

-

bark . Bark color texture

146

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CROWN SHAPE According to Richards ), canopy or emergent tree crowns are wider than( 1952 they are deep and often are umbrella-shaped; subcanopy tree crowns are deeper than

wide , or are equal ; and understory tree crowns are coni cal, much deeper than wide. In their of useful physiognomic and structural features ,analyses Holdridge et ฟ. ( 1971 ) report a rough positive correlatio of n branching gle with trunk shape ( trunk shape defined as clear bole length divided by trunk diameter), a slight correlation of crown diameter with trunk shapenegative a stronger negative correlation of crown volume with, and trunk

^

INTRODUCTION

shape . They found that the largest crown volumes occur in Tropical Moist and in poorly drained alluvial associ ations in Tropical Wet Life Zones. The smallest crown volumes occur in Tropical Lower Montane Rain and Premontane Rain Life Zones. Tropical Wet ( well -drained associations) and Premontane Wet are intermediate in crown volume , The generalization that a tree changes crown shape to conform with the characteristic shape in each stratum as it grows to the canopy ( Richards 1952 ) appears to com pletely ignore gap- phase regeneration ( Hartshorn 1980). The recognition of the ecological differences between monolayer and multilayer crowns ( Horn 1971 ) also con tradicts Richards ' s generalization . Monolayer crowns are characteristic of shade- tolerant “ climax " species and shade- intolerant pioneer species , whereas multilayer crowns are common in shade- intolerant gap species. The ecological rationale is that in the low-light conditions of the forest understory it pays to maximize the coverage of the photosynthetic surface area , that is , in a monolayer, while in the better light regime in gaps it pays to stack one or more additional layers of leaves below the uppermost layer. The latter rationale would intuitively hold for pio neer species, except that they are often in such a strongcompetitive milieu that only the uppermost layer of leaves is important for photosynthesis.

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STRATIFICATION The presence or absence of strata ( layers, stories , and tiers are used synonymously with strata ) of trees , shrubs , and herbs has engendered considerable controversy. Richards ( 1952 ) states that three tree strata occur in most humid tropical forests, whereas two tree strata is the ณle for temperate forests. Shrubs and herbs are considered to form one stratum each ; hence they are ignored in virtu ally all discussions of forest stratification. Richards ( 1952, p. 23) defines a stratum as “a layer of trees whose crowns vary in height between certain limits" and says that “ in a several - layered forest each stratum will have a distinctive floristic composition." However, he admits that the presence of juveniles can considerably obscure the well -defined strata . Other authors such as Mildbraed ( 1922, cited in Richards 1952) state that strata have no objective basis. Strata are most easily recognized in species- poor secondary forest , but they become in creasingly cryptic (if they exist) in heterogeneous, ma-ture forest. Because it is so difficult for a ground observer to see tree strata in the forest (an interesting twist to not seeing the trees for the forest ) Davis and Richards ( 1933, 1934 ) constructed profile diagrams depicting all the trees on a 7.6 X 61 m strip of forest as an aid to recognizing tree strata. Major problems with profile diagrams are the im possibility of replicating even two sample strips within 147

PLANTS

the same heterogeneous forest and that dynamic phases are either ignore! or avoided Holdridge et al . ( 1971 ) avoided the problems of actual profile diagrams by using only mature individuals to prepare an i(kalized profile diagram . "Sketches of mature individuals of the most “ important species are placed together in a profile to represent species richness , basal area , density, and frequency on actual 10 X 100 m sam pie strips. The idealize! profile diagram is used to define the height limits of tree strata. However, Sawyer and Lindsey ( 1971), using the same tree height and crown depth data , often give different height limits for strata and , in a few instances, disagree on the number of tree strata. The polemic on the presence of tree strata has avoided the more fundamental question of their importance if they do indeed exist. What are the ecophysiological and competitive consequences of numerous tree crowns in the same stratum? Is reproductive biology hindered in a dense stratum of tree crowns? Is herbivory affected by the density of tree crowns in a stratum? Despite the difficulty with observing tree strata , the multistratal concept has produced a terminology for the upper height limit attained by mature individualร within a forest. The differentiation of a forest into abstract can opy (and/or emergent ), subcanopy, and understory strata , without necessarily defining height limits , is an ecologically meaningful positioning of the hundreds of tree species in a tropical forest. Canopy, subcanopy, and un derstory tree species may have different modes of regen eration , reproductive syndromes, root systems, and so on , that are discussed in other sections of this chapter. It is in this spirit that subjective assignments to the canopy, subcanopy, or understory are made in the next section

.

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,

EPIPHYTES

The abundance and richness of the epiphytic flora is much more pronounced in the Neotropics than in the Paleotropics ( Brieger 1969). Epiphytic Bromeliaceae are strictly Neotropical , and only the epiphytic cactus Rhipsalis occurs in Africa and Sri Lanka. A preliminary com parison ( table 7.15) with tropical West Africa and Java clearly demonstrates the richness of dicot epiphytes in Costa Rica.

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Dicot families with prominent epiphytic members in j the Neotropics include the Araliaceae, Begoniaceae, Cactaceae , Ericaceae, Gesneriaceae , Melastomataceae , Piperaceae , and Solanaceae. Monocot epiphytes are most II rich in the Araceae, Bromeliaceae , and Orchidaceae, with about 1 ,200 Costa Rican species in the latter family. Costa Rica also has an incredibly rich array of epiphytic ferns and mosses , Epiphytes are most conspicuous in biomass and spe cies richness in Premontane and Lower Montane Rain Life Zones , probably reflecting a correlation with avail able moisture. Cloud cover and wind -driven mist are more important to epiphytism than is rainfall (Grubb et al . 1963; Grubb and Whitmore 1966; Sudgen and Robins 1979). Epiphytic loads appear heavier in La Selva than in Corcovado, even though total rainfall is approximately the same. The nearly year-round presence of conden sation drip at La Selva and the three month dry season at Corcovado may account for the difference in epiphyte abundance. Epiphyte abundance and richness are dras tically less in the Dry and Moist Life Zones , although a very few species of Bromeliaceae , Cactaceae , and Orchidaceae may be fairly abundant . Large tank bromeliads holding up to 1 ,000 ml of water are most conspicuous in the Montane Rain Life Zone . Epiphytic shrubs such as Ericaceae and Araliaceae are largely restricted to Lower Montane and Montane Wet and Rain Life Zones The interesting observations of Perry ( 1978) suggest that arboreal mammals , primarily monkeys , may play an important role in preventing epiphyte dominance on upper branch surfaces used as canopy highways, OTS courses have generated numerous attempts to de termine the degree of host specificity between epiphytes and trees , but I am not aware of any publications report ing detailed studies of epiphyte distribution and abundance in the Neotropics. Hazen ( 1966) found a species ๙ Guzmania to have a random distribution of four branches of an isolated tree on CADE grounds near Turrialba. On a larger scale , there does appear to be a remarkable fidelity of most epiphytic species with Life Zones (J. Utley, pers. comm.). Epiphylls are tiny epiphytes occurring on leaf surfa and by-and -large are restricted to lower plants such as mosses, liverworts , lichens , and algae. Epiphylls are

II

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,

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TABLE 7.15. Preliminary Comparison of Dicot Epiphyte Floras of Costa Rica , Java , and Tropical West Afiri ( km 2)

Total area Total dicot families Epiphytic families Epiphytic genera Epiphytic species SOURCE: Burger 1980.

148

Costa Rica 51 ,1๓ 155 19 51 247

Java 130,987 153

12 22

107

Tropical West Africa ca. 1.000 000 154 9 16 58

.

INTRODUCTION

most prominent in the perhumid and superhumid life

Desmoncus costaricensis present at La Selva and Corcovado. เท contrast to the erect , emergent stature of some montane altitudinal belt. of the Old World rattins Neotropical canopy lianas , in Understory trees and shrubs have the heaviest loads of eluding Desmoncus , are scandent , draping themselvesepiphylls, not only because of the shaded conditions, but over tree crowns. also because they are slow -growing hosts whose leaves Lianas tend to be more frequent in the dry forest of the appear to be held longer than those of canopy members. tropical lowlands, yet the thickest “stems " ( -40 cm in Epiphylls appear especially abundant on shade-tolerant diameter ) are found in Tropical Wet Forests30 . Because members of the Rubiaceae and Palmae.^^^^^^^^^^ their very slow diameter increment, the presence of thickof The classic literature on epiphylls (as cited in Richards lianas is considered an excellent indicator of undisturbed 1952 ) generally viewed them as detrimental to the host forest ( Budowski 1965). leaves. In the extremely low light conditions of the forest In contrast to shade intolerant lianas , the herbaceous understory, an epiphyllous covering must seriously hin climbers are often shade-tolerant and restricted to the der photosynthesis in the host leaf . Richards ( 1952) sum - forest understory. Some understory climbers locate marized by stating that most epiphylls are at least par- able hosts by growing toward dark areas (Strong and Ray tially parasitic on the host leaf . Drip tips are purportedly 1975). Most climbers have the remarkable ability to adaptive in that they enhance surface runoff of rainwater, greatly vary intemode length . เท dense shade , intemode which in turn reduces the availability of moist substrate length is long and leaf size reduced ; in sunny patches , for epiphyllous colonization. Edmisten ( 1970) found that intemodes are drastically shortened and leaf size is inblue green algae among the epiphylls fix atmospheric creased ( see Givnish and Venmeij 1976). Herbaceous nitrogen , suggesting a beneficial input to the host leaf . climbers in Monstera ( Araceae ) have appressed , disklike Barbara Bentley and Amos Bien are at present in - juvenile leaves that effectively cover the terminal meri vestigating the role of nitrogen -fixing epiphylls in the La stem . In apparently more favorable light regimes 5- 10 m Selva understory. above the ground , Monstera produces more typical aroid Canopy lichens also contribute significant quantities of leaves perpendicular to the climbing stem . fixed nitrogen to tropical ecosystems. Foreman ( 1975) found that canopy lichens in a Colombian forest fixed STRANGLERS about as much atmospheric nitrogen (roughly 5 kg/ha/yr) Some species of Ficus ( Moraceae ) and Clusia (Guttias enters the ecosystem through rainfall inputs. Canopy ferae ) start out as epiphytes but send down woody, clasplichens occur mostly on twigs and branches in the 3 12 ing roots that anastomose around the host trunk host cm diameter range and generally do not occur in the tree often dies , and after it rots away, a hollow. The shell of understory. anastomosed roots supports an independent tree . Although these are genetically termed “strangler figs," the CLIMBERS actual cause of death of the host tree has not been eluciHerbaceous and woody climbers the latter also called dated . Constriction of seems unlikely; root vines , lianas, and bush ropes are conspicuous and inte competition is possible;thebut trunk I think crown competition gral components of tropical forests. The distinction be- may be a more likely cause. Stranglers able to tween herbaceous and woody is diffuse , more often re- easily overtop the host crown with a dense seem monolayer of lated to size than to taxonomic affinity, yet some delicate eveigreen leaves vines are remarkably strong . Climbers occur in more Stranglers are most frequent in Tropical Moist and families than do epiphytes, with Apocynaceae , Araceae , Premontane Wet Zones, where they may be among Bignoniaceae, Combretaceae, Convolvulaceae, Dille the tallest trees inLife the forest . The greater abundance of niaceae, Hippocrateaceae , Leguminosae, Malpighia stranglers in Corcovado than in La Selva suggests that an ceae, Passifloraceae , Sapindaceae , Smilacaceae , Ulma effective dry season may be ideal for seedling establish ceae, and Vitaceae being well represented among the ment as epiphytes. climbers. Bignoniaceae usually has the greatest number f climbing species (about twenty in any tropical forest; PARASITES AND SAPROPHYTES °Gentry 1976). Parasitic shrubs in several genera of the Loranthaceae are The gymnosperm Gnetum is a canopy liana in La Selva well represented the wetter Life Zones of Costa Rica Corcovado. Rourea glabra ( Connaraceae ), Com- from the Tropicalin basal to Lower Montane. Gai Return farinosum ( Combretaceae ), and Davilla kunthii adodendron poasense is abeltparasitic tree with beautiful Thlleniaceae ) are common lianas in the Tropical Dry yellow flowers that grows in Lower Montane and Mon . Climbing palms the rattins of the tropical Far tane Rain Life Zones. All the other Costa Rican Lo ast are very depauperate in the Neotropics , with only ranthaceae are crown parasites , often forming dense-

zones without an effective dry season , but they are not nearly as common in the cooler

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sum-

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cr^ ests

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149

PLANTS

shrubby clusters. Loranthaceae are much more prevalent in isolated trees left standing in pastures and such , than in undisturbed forest . I am unaware of any evidence for host specificity. Among Costa Rican members of the Olacaceae , root parasitism has been documented for Schoepfia schreberi ( Werth , Baird , and Musselman 1979 ) and Ximenia americana ( DeFillips 1969) , but no evidence of host specificity has been found for these two genera. Achlorophyllous saprophytic plants are rare in Costa Rica . I have seen Leiphaitnos ( Gentianaceae ) only in virgin forest above Punta Llorona in Corcovado. An obligate association with mycorrhiza may restrict the occurrence of Leiphainws to small areas ( D. Janos, pers.

comm . ).

Community Ecology

PRIMARY PRODUCTIVITY AND NUTRIENT CYCLING

No studies of primary productivity of natural forests have been done in Costa Rica , nor are data on plant biomass ( standing crop ) available. Scarcity of data , of course , has not hindered generalizations for tropical forests (e.g., Rodin and Bazilevich 1967; Kira and Ogawa 1971; Rodin , Bazilevich , and Rozov 1975; Golley 1975 ). An nual gross primary productivity for humid tropical forests appears to range between 75 and 150 tons/ha/yr, yielding net primary productivity of 25-50 tons/ha/yr. Dry forests appear to have productivity values roughly half those for moist and wetter forests. Total plant biomass appears to range between 600 and 1 ,200 tons/ha . Estimates available are crude at best and tell us practically nothing about the variability of primary productivity and total plant biomass in different Life Zones , or about the ecology of the site . Productivity studies in El Verde , Puerto Rico ( Odum and Pigeon 1970) , Darien , Panama (Golley et al . 1975) , Amazonian Brazil near Manaus ( Klinge et al . 1975) , and the Rio Negro region of Amazonian Venezuela ( Herrera et al . 1978 ) should not be considered representative of Costa Rican forests . All four study areas are on soils much poorer in nutrients than most Costa Rican soils. The hypothesis of direct nutrient cycling ( Went and Stark 1968 ) and the realization that luxuriant tropical forests may occur on quite infertile soils have stimulated considerable investigation of nutrient cycling in tropical forests. Most of the productivity studies mentioned above also include information on nutrient ( or mineral ) cycling . Jordan and Kline ( 1972) review the state of knowledge about nutrient cycling in tropical forests. My earlier ex pression of caution about generalizing from productivity studies to Cost Rican forests is even more appropriate to nutrient cycling . The infertile Ultisols near Manaus and the mosaic of practically sterile white sands ( Spodosols )

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and lateritic ( Oxisols ) soils in the Rio Negro region of Venezuela and Brazil are very different from the relatively fertile soils of Costa Rica ( Harris, Neumann , and Stouse 1971). In an analysis of nutrient status on volcanic (andesitic) soils in San Carlos , Costa Rica , Krebs ( 1975) found that permanent agriculture did not cause serious deterioration of soil nutrients. On a fairly fertile soil near Tumalba , Costa Rica , Harcombe ( 1973) found that early secondary succession accumulated nutrients in approximately equal proportions from decomposition of organic debris and from external inputs such as rain , dust , and soil mineral ization. , As pan of the OTS comparative ecosystem project ’ s Col Washington of University the investigators from lege of Forest Resources collected considerable quantities of data on nutrient cycling , primarily at La Selva and secondarily at Palo Verde. The few publications that have come out of that project are based on rather short data runs lasting a few days to a few weeks. Substantial quantities of nutrients are leached out of the canopy by intense rainfall and moved by throughfall and stemflow into the soil and to a lesser degree on through the watershed by groundwater ( McColl 1970). Carbonic acid is postulated as the primary cation leaching agent in non calcareous tropical soils by Johnson , Cole , and Gessel 1975). These authors report that litter decomposition accounts for only 25-50% of the CO; produced in the soil; the remainder is attributed to root respiration and micro bial activity. CO; entrapment and buildup occurs because of poor diffusivity in the soil . Passage of a wetting front creates appreciable quantities of carbonic acid that leaches cations through the soil profile. The authors report that C02 evolution is an order of magnitude greater in La Selva soil than in a Douglas fir forest soil in western Washington state. Carbonic acid leaching of tropical forest soils may be an overlooked flux in the current polemic over whether tropical forests are a source or sink for the global carbon cycle.

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PHENOLOGY

Phenological studies of the periodicity of leaf production and senescence, flowering , and fruiting of’ tropical trees in Costa Rica were stimulated by Janzen ร ( 1967) paper on the synchronization of sexual reproduction with the strong dry season in Guanacaste. Despite the inimical, physical conditions caused by severe annual drought numerous tree species flower during the dry season. Jan zen ( 1967 ) interpreted dry season flowering as in partiman optimum time for pollinator activity, but , more reportant , he pointed out the advantage of being

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INTRODUCTION

productive at a time when adjacent tree crowns are not ally report unusual behavior. The dry forest understory engaging in vegetative competition because they are qui tree Jacquinia pungens ( Theophrastaceae ) is deciduous escent and leafless. during the rainy season but produces leaves , flowers , and Knowledge of tropical tree and forest phenology has fruit during the dry season; Jacquinia has a very long been greatly advanced by the considerable efforts of Gor taproot , which enables it to be in full leaf below the don Frankie , Herbert Baker, and Paul Opler ( 1974๗,/? ), deciduous canopy ( Janzen 1970๗ ). Being dormant durwhoin 1968 initiated detailed phenological monitoring of ing the rainy season , even in full shade , costs the tree a majority of tree species in a tropical dry forest area ( the about a third of its starch reserves ( Janzen and Wilson front part of Comelco ranch , near Bagaces ) and in a 1974). tropical wet forest ( La Selva ). Only 8% ( N = 331 ) of the Some trees regularly flower and fruit once in two years tree species in La Selva are deciduous, and virtually all ( e . g ., Andira inermis and Hymenolobium pulcherrimum , of them are canopy species; hence the canopy remains both Fabaceae ); others do irregularly once in three to evergreen with scattered deciduous crowns during the six years (e. g., Licania sp.so, Chrysobalanaceae , Ouratea of

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driest time year. The dry forest canopy is almost en tirely deciduous during the long dry season , while the understory is partially deciduous. Periodicity of leaf fall coincides well with the onset of the primary dry season at each site , even though February- March is seldom effectively dry at La Selva . In the wet forest leaf flushing peaks in January, a generally wet month , but in the dry forest it occurs just before and after the rainy season . In the wet forest , Frankie, Baker, and Opler ( 1974๗ ) differentiate seasonally flowering ( X = 6.5 weeks ) from extended- flowering species ( x = 23 weeks ) and report that the length of flowering is very similar for overstory and understory tree species. Seasonally flowering species have a slight peak during February March and gradually taper off through the rest of the year. Extended- flowering species have a major peak during May-August. A definite fruiting peak occurs in September-October. Dry forest flowering has two peaks a broad , mid dry -season period and a sharp peak immediately after the rainy season starts . Sporadic light rain showers in the few weeks before the heavy rains start are the proximal cue to synchronization of flowering of many tree species at the end of the dry' season (Opler, Frankie, and Baker 1976). The peak of fruit maturation isi spread through the dry

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season . The most complete phenological study of all flowering plants was done on Barro Colorado Island , Panama ( Fos ter 1973; Croat 1975). Barro Colorado has a peak ot flowering activity one in the dry seasondouble and one in the wet season — that corresponds well with its inter-

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mediate situation ( tropical moist forest ) between dry and that more species flower and fruit in the rainy season than in the dry season , although the greater season syn chrony produces a higher peak . Foster ’sdry ( 1973) detailed study demonstrates that an unusually wet “dry season ” in 1970 prevented many species from flowering and fruit Ing , with drastic consequences for frugivore populationsDetailed phenological studies of indigenous species *** very few ( e .g . , Mori and Kallunki 1976) and gener-

wet sites studied by Frankie ’s team. Croat ( 1975) reports

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Ochnaceae ). However, almost no Neotropical intervals like the tropical Far Eastern Dipterocarpaceae ( see Janzen 1974 ). Perennial plants that flower once before dying , such as bamboos and some palms, have long attracted the atten lion of biologists ( Janzen 1976๗ ). Among the few dicot trees that are monocarpic , that is, die after reproducing once, Tachigalia versicolor (Caesalpiniaceae) cohorts appear to be four years offset from each other ( Foster 1977). Some individuals in a population flower, then die as the large fruit crop matures. In some trees only a branch or part of the crown is reproductive and then dies , while the nonreproductive branches remain healthy, Phenological patterns may be important components of community organization , such as with nectivorous guilds of birds ( Feinsinger 1976; Stiles 1978) , or the high species richness of sympatric species (Gentry 1974 , 1976). lucens ,

tree species synchronously produce mast fruiting at long

REPRODUCTIVE BIOLOGY

Research on the reproductive biology of tropical plants has focused on three aspects: breeding systems , polli -

nation , and seed dispersal and predation . The generally held interpretation that individuals of a species are hyperdispersed in species-rich tropical forests (e.g ., Richards 1952 ) gave rise to the assumption that most tropical tree species must be self -compatible (e.g ., Baker 1959; Fedorov 1966). The pioneering studies of Kamal Bawa on tropical tree breeding systems indicates that self incompatibility mechanisms are more prevalent in tropi cal dry forest than in temperate forests and that dioecy issurprisingly frequent for biotically pollinated species ( Bawa 1974; Bawa and Opler 1975 ). Recent studies dem onstrate that bees are effective long -distance pollinators (Janzen 1971๗ ; Frankie, Opler, and Bawa 1976). Evidence is also accumulating that tropical tree species are not spatially hyperdispersed but are generally regularly or randomly distributed , or even occasionally clumped ( Hubbell 1979), but nevertheless are much farther apart

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than it they grew in proportion to the density of their seed shadows (Janzen 1971๙ ). The predominance of biotic pollination in lowland tropical forests has led to spectacularly coevolved polli nation systems involving very complex floral attractions and deceptions. Some of these systems , such as for orare chids and bees , have received much attention; others wasps not nearly as well known , such as for figs and ( Janzen 1979). It is impossible to adequately review the fascinating field of pollination here, but some recent papers on tropical pollination systems should provide the necessary entrde to the abundant literature (Cmden et al;. 1976; Essig 1971; Gentry 1978; Gilbert and Raven 1975 Janzen 1968; Linhart 1973; Opler, Baker, and Frankie 1975; Toledo 1977; Uhl and Moore 1977). The relationship of seed predation and spacing of tropical trees has received considerable attention , particularly owing to Janzen ’ร prolific research on insect seed predation in tropical dry forests. Host-specific predators devastate seed crops unless seeds escape in space or time or both via dispersal or toxicity or satiating the predators ( Janzen 1969, 1971/?,c , e , 1972 , 1975๘ , /?; Janzen and Wilson 1977; Silander 1978). The high degree of seed and seedling predation precludes establishment of regeneration near the parent tree, hence herbivores may have an important effect on species packing in tropical forests (Janzen 1970/?, 1971๙ ). Much less is known about vertebrate predation on seeds (e.g., Higgins 1979). Vertebrates are extremely important vectors for seed dispersal in tropical forests . Wind dispersal of tree seeds accounts for 31% of dry forest trees ( Frankie, Baker, and Opler 1974๘ ) and only 8% for La Selva trees ( Hartshorn 1978). Including wind dispersed canopy lianas and epiphytes would be expected to raise the percentage a few points , but the floras are still too poorly known to permit a more complete analysis of dispersal modes. For La Selva trees ( N = 320 species) Hartshorn ( 1978) estiby mates that 50% have diaspores of the type dispersed birds and 13% have those of the type dispersed by bats The remainder are dispersed by arboreal and terrestrial mammals or by wind . Only the riparian Ficus glabrata, ( Moraceae) appears to be fish -dispersed . This contrasts not unexpectedly, with the important role Amazonian fish play in dispersal of varzea forest seeds (Gottsberger 1978). A few detailed studies of seed dispersal by birds have been done in Costa Rica on Casearia corymbosa in the; Flacourtiaceae ( Howe and Primack 1975; Howe 1977

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Howe and Vande Kerckhove 1979) and on Stemmadenia donnell - smithii in the Apocynaceae ( McDiarmid , Rick lefs , and Foster 1977; Cant 1979). Bats are effective dispersal agents of aggregate fruits like Ficus and Piper as well as of large single-seeded fniits such as Dipteryx and Andira (e.g . , Morrison 1978; Vasquez- Yanes et al. 152 ,

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ysis should Stimulate some exciting new areas of research on tropical trees and forestร. Because of our considerable ignorance about the root systems of tropical trees ( but see Longman and Jenik 1974; Jenik 1978), architecture models only vaguely address root systems. In Neotropical forests, buttresses and taproots appear to be mutually exclusive. Nonbuttressed canopy trees for example, Bertholletia excelso Lecythis costaricensis ( both Lecythidaceae) , Sterculia recordiana ( Sterculiaceae ) , Chaunochiton kappleri

to night 1975). The propensity of bats to carry diasporesshadows feeding roosts often produces discernible seed ( Janzenetal . 1976; Janzen 1978๘, /? ). I have also encoun tered substantial concentrations of Brosimum alicastrum) ( Moraceae ) and Calophyllum brasiliense ( Guttiferae seeds under trees of different species obviously used as feeding roosts by bats. Terrestrial mammals can be important seed dispersal agents , as in the scatter-hoarding of seeds by agoutis (Smythe 1970) or the diaspores that easily attach to pas sersby ( Bullock and Primack 1977). Species characteristic of swamps and coastal habitats often have diaspores that float (e.g., Carapa guianensis, Meliaceae; Prioria copaifera , Caesalpiniaceae). Man grove species have dispersal properties that are well. adapted to their habitat preferences ( Rabinowitz 1978)

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I of tropical

characterization trees by architectural models has recently appeared in an expanded English version ( Halid , Oldeman , and Tomlinson 1978) that is a veritable gold mine of information on tropical trees . For the thousands of tropical tree spe- 1 cies , only twenty-three architectural models have been found . Each architectural model is based on a set of morphological characters that includes the life-span of meristems and the degree and type of differentiation of vegetative meristems; the latter involves sexual versus) axes vegetative differentiation, plagiotropy ( horizontal ) episodic ( , rhythmic ) axes versus orthotropy ( vertical de branch of chronology , and versus continuous growth velopment . Some examples of well- represented architectural mod-, els are Comer ร model (Carica papaya Cocos nuciferas Welfia georgii ); Tomlinson ’s model ( banana ); Koriba model ( Hura crepitans ); Nozeran ร model (Theobroma cacao ); Aubreville s model (Terminalia catappa ) ; Mas'

and Sacoglottis trichogyna ( Humiriaceae ) well a have -developed system of feeder roots near the . In surface contrast , nonbuttressed subcanopy trees soil large conical a taproot , for example, Naucleopsis with naga ( Moraceae ) and Griasfendleri ( Lecythidaceae) , do not have extensive surface feeder roots. Slow-growing , shade-tolerant subcanopy trees may be obtaining nutri ents from the subsoil rather than competing for nutrients from litter decomposition Even less is known about the root systems of understory trees, but I have seen no evidence of taproots on shade-tolerant species . Fast growing , shade-intolerant understory trees and treelets appear to have superficial root systems. Janzen ( 1976/? ) suggests that hollow tninks may be adaptive by providing a roosting site for bats , whose guano could then be tapped by the “host ” tree ’s roots There is some evidence that a hollow tree ’s roots exploit the nutrient resources beneath the hollow trunk ( Dickenson and Tanner 1978). ( Olacaceae),

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TREE ARCHITECTURE Halid and Oldeman ’s ( 1970)

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sart ’s model ( Araucaria heterophylla , Ceiba pentandra ); and Cook ’s model (Castilla elastica ) . Architectural models are most easily determined arein young , free-growing trees. Mature trees in the forestplas more difficult to categorize because of vegetative; how ticity and responses to competition and damage ever, reiteration of the basic architectural model is evident in vegetative sprouting , models An intriguing potential use of tree architecturemanage and dynamics forest of investigation is in the ment. Do pioneer, gap, and climax tree species represent different models? Are shade-tolerant and shade-intolerant tree species mutually exclusive models? Are there opti mum mixtures of architectural models for photosynthetK efficiency? Are forest strata occupied by trees with different architectural models? The use of architectural anal

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DOMINANCE

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FOREST SUCCESSION

DIVERSITY

Single-species dominance is most pronounced in tropical montane rain forest on the Cerro ( table 7.14) , where Quercus costaricensis ( Fagaceae ) has an important value of 41% , precisely triple the value of the second most important species, Miconia bipulifera ( Melastomata ceae ). The only lowland tropical oak , Quercus oleoides , dominates the dry forest on pumice and rhyolitic ash soils with an importance value of 37% ( table 7.2), more than three and a half times the value of Byrsonima crassifolia ( Malpighiaceae) , the second most important species. Pentaclethra macroloba ( Mimosaceae ) dominates the virgin forests of La Selva (tables 7.9- 12) (Hartshorn 1972), with importance values between % and 23% , more than double the values of the second 18most important species, except in the swamp ( table 7.10) , where Carapa guianensis ( Meliaceae ) has an importance value of 11% . In the Palo Verde forest , Calycophyllum candidissium ) is nearly twice as important ( 11% ) as the Rubiaceae ond-ranked species ( table 7.3). Of the four sites with adequate plot data , Corcovado’ร forests show the weakest dominance. Dominant species evident where restrictive site factors substantially

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INTRODUCTION

reduce species diversity ( tables 7.7 and 7.8; also Jan zen 1978c). It is clear ( tables 7.4 and 7.5) that domi -nance does not occur in the forests of the Corcovado uplands. Species diversity is greatest in the upland forests of Corcovado undoubtedly the richest forest in Central America . Dominance of the La Selva forests by Pentaclethra macroloba greatly reduces the alpha diversity of trees in the La Selva forest. Even though the hetero geneous forests of Palo Verde and Santa Rosa lack astrongly dominant species , the alpha diversity of these forests is less than at La Selva because of lower species richness in the dry forests. It is hoped that more plot data will become available for Monteverde , Corcovado, and Santa Rosa that will then permit much more detailed analyses of patterns in Costa Rican tree species diversity Analyzing the wealth of data on tree species diversity and soil nutrients from the Holdridge et al . ( 1971 ) study sites in Costa Rica , Huston ( 1980) found a strong nega five correlation between tree species diversity and soil fertility. This pattern of lower species diversity on more fertile soils appears equally valid within sites (e.g., Cor covado) and between sites and typifies Amazonian for ests as well . Even the species-rich forests of Corcovado ( 100- 120 species/ha ) pale in comparison with Ama zonian forests, where the number of species 10 cm and larger in diameter ranges between 150 and 200 per hectare AND

DYNAMICS

Several papers summarizing important aspects of Neotropical secondary succession appeared in a supplemental issue of Biotropica ( Ewel 1980), hence it would be presumptuous to attempt to review here this vast field . Much less attention has been given to successional and dynamic processes in late secondary and mature forests. In the same Biotropica supplement , I review our present understanding of Neotropical forest dynamics. ( Hartshorn 1980). Based on my travels to many forests in tropical Amer ica , I believe La Selva is at the fast end of the range ofNeotropical forest dynamics; that is , tree falls are more frequent in La Selva than in any other mature forest I have visited in tropical America. That of course excludes obvious successional forests such as the Euterpe palm breaks on the windward ridges of Puerto Rico maintained by hurricanes. In Corcovado, tree falls do not appear to be as prevalent as in La Selva . Drier forests (e.g., Palo Verde , Santa Rosa ) and midelevation forests (e.g., Mon teverde ) do not appear to be as dynamic as La Selva; however, long-term monitoring will be necessary to determine if gaps are any less important to species regen eration elsewhere than at La Selva.

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. 1977 . Flora costaricensis . Fieldiana , Bi . \914 b . Tropical plant phenology: Applications Forest dynamic processes in western Amazonia ( west- 40:291 ed population dynamics of Pentaclethra macroloba , a . for studies in community ecology. In Phenology and tropical wet forest dominant em Brazil , northern Bolivia , eastern Pern and Ecuador, Cain , ร. A .; Oliveira Castro , G . M . de; Murca Pires , J .; and Stryphnodendron exseasonality modeling , ed . H . Lieth , pp . 287-96. New celsum , an occasional associate and southern Colombia ) appear to be similar to those and Tomas de Silva , N . 1956 . Application o ! IJC . Ph D diss . . UniverYork : Springer- Verlag p. of Washington . sity found to be important in Costa Rican forests ( see Harts- phytosocioiog cal techniques to Brazilian rain foreal , G . พ. ; Opler . A . ; and Bawa , K . ร . 1976 . Frankie . 1977 . Criterios para la clasificacion de bosques horn 1978 , 1980 ) . However, in central and northeastern , Foraging behaviour of solitary : bees Implications for y , la determinacion del uso potencidl de tierras en Para. 1979 . Dispersal ๙ ร emmadenia donned outcrossing of a Neotropical forest tree species . . J . guay .strikinglyadifferent . Near San Carlos de R o Negro Ven- C a n t , J1,,Cby. Hbirds t Informe 8 . Asuncidn: FAO/FO: DP/PAR / £ cnico . 122 11 : Biotropica . monkeys and . 64: 1049- 57 . Bed / 72 001 . most . 2 Malaya ) noticed vols of I Spodosols ( trees . Wayside soils . sand 1940 white . ezuela , on Corner E J H Gentry, A . H . 1974. Flowering phenology and diversity . 1978 . Tree falls and tropical forest dynamics . In Singapore: Government Printing Office. trees die while still standing rather than being uprooted or Bignoniaceae tropical . Biotropica 6 : in 64 68 . Tropical trees as living systems , ed . p. B Tomlinson and habit of behavior and having the entire crown snap off . The gradual crown Croat , T. B . 1975 . Phenological 1976 . . Bignoniaceae of southern Central Amer . M H . Zimmermann , pp . 617-38 . London: Cam ( } Panama Island Ca Colorado classes on Barro attrition of dead branches followed by the fall of a habitat : Distribution and ecological specificity. Biotropica ica bridge University Press . . 77 270 7 : Biotropica ) . Zone nal branchless bole does not open up a substantial canopy Cruden 8: 117-31 . . 1980 . Neotropical , D; forest dynamics . Biotropica . E , . R ; Stockhouse , . ร พ Kinsman , : . R 1978 . Anti -pollinators for mass-flowering gap , nor does it cause high mortality of nearby trees . A Linhart , Y. B . 1976 . Pollination , fecundity, and the ( suppl . ): 23- 30 . 12 small , one-crown canopy gap is filled by competing pole- distribution of moth -flowered plants . Biotropica plants? Biotropica 10( l ):p68-69 . Hazen , พ. E . 1966 . Analysis Gilbert , L. E. , and Raven , . H . , eds . 1975 . Coe\olution phytes . Ecology 47:634- 35. of spatial pattern in epi sized trees that survived the death of the canopy tree . 8 - 201 10 , „ m cf animals and plants . Austin: University of Texas Heithaus , E. R . 1979. Community structure of Neothat most tree ^Daubenmire , R 1972 . „Ecology ofr. Hyparrhenia suggests This type of canopy !replacement rufy ‘- V 7 Press . tropical flower visiting bees and wasps: Diversity and species must be shade- tolerant , which is quite different ( Nees) in derived savanna in north - western c , T. J . 1978 . On the adaptive significance of Givnish phenology . Ecology 60: 190- 202. from the abundance of shade- intolerant species in the La Rica . J . Appl . Ecol . 9: 11 -23 . compound leaves , with particular reference to tropical Henwood , K . 1973 . A structural model of forces in but p can tolerant veg The . shade of . พ 1933 . , proportion Richards , w greater . . A and forest A T Selva Davis trees . In Tropical trees as living systems , ed . p. B . tropical rain forest trees . Biotropica 5:83-93.tressed : An ecological opy species would indicate more deterministic forest dy- tion of Moraballi Creek , British Guianaforest Tomlinson and M . H . Zimmermann , pp . 351-80. Lon- Herrera , R .; Jordan , c. F.; Klinge , H .; and Medina , E. . Part I . J namics than occurs in the highly stochastic forests of study of a limited area of tropical rain : Cambridge University Press. don 1978 . Amazon ecosystems: ไาใeir structure and funcEcol . 21:350-84 . Central America and western Amazonia . , . . T , J and , Vermeij Givnish G J . . 1976 . Sizes and shapes tioning with particular emphasis on nutrients . Inter. 1934 . The vegetation of Moraballi Creek , Britliane leaves . . Nat . 110:743- 78 . of Am 3 ciencia :223- 31 . area limited of ish Guiana: An ecological study of a , F. B . 1975 . Productivity and mineral cycling in Herwitz , ร. 1979. The Golley regeneration of selected tropical . 155 106 : . 22 Ecol . J . tropical rain forest . Part 2 p Allen , p. H . 1956 . The rain forests of Golfo Dulce . Dean tropical forests . In of Productivity world , ecosystems wet forest tree species in Corcovado National Park , anc Behavioral . 1978 . . , A Smith and , . J , . M Gainesville: University of Florida Press. pp 106- 15 . Washington , D . C.: NAS . Costa Rica . . A . thesis , University of California , M to high plant tropical a of adaptations morphological in Anderson , R . , and Mori , ร. 1967 . A preliminary Golley, F. B .; McGinnis , J . T. ; Clements , R . G .; Child , Berkeley. 10: 152- 54 . vestigation of Raphia palm swamps , Puerto Viejo , rainfall,. Biotropica .; Duever, M . J .; plus Duke , J .; Ewel , J .; and Gist , Higgins , M . L . 1979. Intensity of seed predation on DeFillips R . 1969 . Parasitism in Ximenia . Rhodorcl cG.. I1975 Costa Rica . Turrialba 17:221-24 . . Mineral cycling in a tropical moist forest Brosimum utile by mealy parrots ( Amazona farinosa ) . Baker, H . G . 1959 . Reproductive methods as factors in 71:439-,43 ecosystem . Athens: University of Georgia Press. Biotropica 11 :80 . I Exploitation . V . 1978 . J , . E Tanner , and . . A T Dickinson Harbor Spring speciation in flowering plants . Cold , G . 1978 . Seed dispersal by fish in the in - Holdridge , L. R . 1947 . Gottsberger Biotropiccl Determination of world plant . of hollow trunks by tropical trees Symp . Quant . Biol . 24: 177- 91 . undated regions of , Amazonia . Biotropica Humaita formations from simple climatic data . Science 105: 10:231-33 . Bauar, G . N . 1968 . The ecological basis cf rainforest Edmisten 10: 170-83. 367 68 . nitroge the of studies Preliminary . J 1970 . , management . Sydney: Forest Commission of New -Smith , p. 1952 . Ecological observations on de. 1967 . Life zone ecology. Rev. ed . San Josd budget of a tropical rain forest . In A tropical rait Greig graded South Wales , Australia . and secondary forest in Trinidad , British West Tropical Science Center. . H 211 chap , Pigeon . F . R , ed . H . T. Odum and Indies Bawa , K . ร. 1974 . Breeding systems of tree species .of a forest . I . General features of the vegetation . J . Ecol . Holdndge . L . R . ; Grenke , พ c.; Hatheway, พ. H ; . USAEC C : . . , . D Washington 215 . 92 40:283-315. lowland tropical community. Evolution 28:85in-tropical , ; Liang . T and , J Tosi . , . A Jr . 1971 Bac . in Forest pollination on environ Observations . . 1971 . , F B Essig Gmbb , p. J .; Lloyd , J . R .; Pennington , T. D .; and Whit- ments in tropical life zones: A pilot study. PergamonBawa , K . ร . , and Opler, PA . 1975. Dioecism : . 24 20 15 Principes . tris 79 . more , T. c. 1963 . A comparison of montane and low- Press. forest trees . Evolution 29: 167. Ewel , J . 1980 . Special issue on tropical succession . Bioland rain forest in Ecuador. 1 . The forest structure , Horn , H . ร. 1971 . The adaptive geometry cf trees . PrinceBeard , J . ร . 1945 . The natural vegetation cf Trinidad tropica 12 (suppl . ): l . Oxford Forest Memoir no . 20. Oxford: Oxford Univer- Fedorov physiognomy and floristics . . Ecol . 51 : 567-601 . ton: Princeton University Press . , A . A . 1966 . The structure of the tropical rair Gmbb , p. J . , and Whitmore , T.J c sity Press. . . 1966 A comparison , H . F. 1977 . Bird activity and seed dispersal of a Howe Ecd . J . tropics forest and speciation in the humid of montane and lowland rain forest in Ecuador. 2. The tropical Black , H . L . , and Harper. K . T. 1979. The adaptive forest tree . Ecology 58:539- 50. value of buttresses to tropical trees: Additional hypoth - 54: 1 - 11,. p. 1976 . Organization of a tropical guild Cf climate and its effects on the distribution and phys- Howe , H . Fwet , . and Primack , R . B . 1975 . Differential seed Feinsinger eses . Biotropica 11 : 240. iognomy of the forests . J . Ecol . 54:303-33 . dispersal by birds of the tree Casearia nitida ( Flacour. 91 257 46 : . Monogr . Ecol . birds nectarivorous na parques Halte , F. , and Oldeman , R . A . A . 1970. Essai sur Boza , M . A . , and Bonilla , A . 1978 . Los tiaceae ) . Biotropica grce blue -83 . with Foreman , R T. T. 1975. Canopy lichens / architecture et la dynamique de croissance des arbres Howe , H . F. , and Vande7:278 cionales de Costa Rica . Madrid: INCAFO . ! fore , G . ร . 1979 . Fecun rain Kerckhove Colombian a in source nitrogen :A * tropicaux . Paris: Masson . Brieger, F. G . 1969 . Patterns of evolutionary and geo- algae dity and seed dispersal of a tropical tree . Ecology 84 . 1176 56 : Ecology . graphical distribution in Neotropical orchids . Biol . J Foster, R . B . 1973 . Seasonality of fruit production aiy Malle , F. ; Oldeman , R . A . A .; and Tomlison , p. B . 1978 . 60: 180-89. Tropical Linn . Soc . 1 : 197-217 . and forests: An architectural analysis . Hubbell , ร . p. 1979 . Tree dispersion , abundance , and fall in a tropical forest ecosystem in Panama New Yorktrees Budowski , G . 1965 . Distribution of tropical American seed : Springer - Verlag . diversity in a tropical dry forest . Science 203: . , Duke University. Harcombe , p. A . 1973 . Nutrient cycling in secondary rain forest species in the light of successional pro- Ph . D.. diss Ntf 1299- 1309. suicidal is a versicolor Tachigalia . 1977 plant succession in a humid tropical forest region ( Tiir- Huston cesses . Turrialba 15:40- 42. , M . 1980 . Soil nutrients and community structure tropical tree . Nature 268:624- 26 . , Costa Rica ) . Ph . D. diss . , Yale University. nalba Bullock , ร. H . , and Primack , R B . 1977 . Comparative p in Costa 1974 Rican forests . J . Biogeogr. 7: 147- 57 . . , A . ; Opler and . , . , G . พ. ; Baker H G * . A .; Neumann , A . M . ; and Stouse , p. A . , Jr. Janzen ร experimental study of seed dispersal on animals . Ecol - Frankie , tropitf . D 1967 . Synchronization of sexual rein of trees studies phenological Comparative L The major soil zones of Costa Rica . Soil ร๗. production Hof. trees ogy 58 :681 -86 . .J Rica Costa of within the dry season in Central lowlands the in forests dry and wet , Fieldiana 122:439- 47 . Bulger, พ. , ed . 1971 . Flora costaricensis . America . Evolution :620- 37 . 21 . 919 881 : 62 . Ecol Mart s h ๐!ท , G . ร . 1972 . The ecological life history and Bot . 35: 1 - 227 . . 1968 . Reproductive

INTRODUCTION

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aJ

.

*

.

1

.

.

'

’

.

.

154

^

J

behavior in the Passi -

155

PLANTS

floraceae and some of its pollinators in Central America . Behaviour 32:33-48. versus seed size , number, . 1969. Seed-eaters toxicity and dispersal. Evolution 23:1-27. . 1970a . Jacquiniapungens , a heliophile from the understory of tropical deciduous forest. Biotropica 2:112-19. . 1970/7. Herbivores and the number of tree species in tropical forests. Am . Nat. 104:501-28. . 1971๙ . Euglossine bees as long-distance polli nators of tropical plants. Science 171:203-5. . 197 1/7. Escape of Cassia grands L. beans from predators in time and space . Ecology 52:964-79. . 1971c. The fate of Scheelea rostrata fruits beneath the parent tree: Predispersal attack by bruchids. Principes 15:89- 101 .

_ ___ _

-

. 1971๔. Seed predation by animals. Ann . Rev. Ecol. Syst . 2:465-92. of juvenile Dioclea megacarpa . 1971c. )Escape predators in a deciduous from vines ( Leguminosae tropical forest. Am. Nat. 105:97- 112. by Sterculia apetala seeds . 1972. Escape in spacefasciatus in a Costa Rican

—

from the bug Dysdercus deciduous forest . Ecology 53:350-61 . . 1973 Rate of regeneration after a tropical high elevation fire . Biotropica 5:117 22. rivers , animals, and . 1974. Tropicaltheblackwater . Biotropica Dipterocarpaceae fruiting by mast 6:69-103. variations in Gua. 1975a. Intra- and interhabitat predation by Am) seed zuma ulmifolia ( Sterculiaceae Rica. Ecology ) Costa in Bruchidae ( cistelinus blycerus 56:1009- 13 . 1975/7 . Behavior of Hymenaea courbaril when its predispersal seed predator is absent . Science 189: 145-47. . 1976a. Why bamboos wait so long to flower. Ann . Rev. Ecol . Syst . 7:347-91. . 1976/7 . Why tropical trees have rotten cores . Biotropica 8:110. . 1978a. The size of a local peak in a seed shadow. Biotropica 10:78 . 1978/7 . A bat -generated fig seed shadow in rainforest . Biotropica 10: 121. . 1978c. Description of a Pterocarpus officinalis ( Leguminosae ) monoculture in Corcovado National Park , Costa Rica . Brenesia 14 15:305-9. . 1979. How to be a fig . Ann . Rev. Ecol . Syst . 10:13-51. -Jones, J .; Pond , Janzen , D. H .; Miller, G . A .; Hackforth c M .; Hooper, K .; and Janos , D. p. 1976. Two Costa Rican bat generated seed shadows in Andira inermis ( Leguminosae). Ecology 57:1068-75. Janzen , D. H and Wilson , D. E. 1974 . The cost of being dormant in the tropics . Biotropica 6:260-62. . 1977. Natural history of seed predation by Rosella sickingiae Whitehead (Curculionidae ) on Sick ingia maxonii ( Rubiaceae ) in Costa Rican rainforest . Coleopt . Bull . 31:19-23.

.

-

.

.

-

-

.

-

156

•*

Jenik , J . 1978. Roots and root systems in tropical trees: aspects. เท Tropical trees as Morphologic and ecologic p . Tomlinson and M . H. . B . , ed systems living Zimmermann , pp. 323-49. London: Cambridge University Press. Johnson , D.; Cole , D. พ.; and Gessel , ร. p. 1975. Processes of nutrient transfer in a tropical rain forest. Biotropica 7: 208- 15. Jordan , c F. , and Kline, J. R . 1972. Mineral cycling: Some basic concepts and their application in a tropical rain forest. Ann. Rev. Ecol. Syst. 3:33-50. Kira , T. , and Ogawa , H . 1971. Assessment of primary production in tropical and equatorial forests. Proc. UNESCO Brussels Symp ., pp. 309-21. Klinge , H .; Rodrigues, พ. A .; Brunig, E.; and Fitthau , E. J . 1975. Biomass and structure in a central Amazonian rain forest. In Tropical ecological systems: Trends in terrestrial and aquatic research , ed . F. B. Golley and E. Medina , pp. 115- 22. New Yi Springer- Verlag . Krebs , J . E. 1975. A comparison of soils under agriculture and forests in San Carlos , Costa Rica . In Tropical ecological systems: Trends in terrestrial and aquatic research , ed . F. B. Golley and E. Medina , pp . 381- 90. New York: Springer- Verlag . Lawton , R . , and Dryer, V. 1980. The vegetation of the Monteverde Cloud Forest Reserve. Brenesia 18: Lee , D. พ.; Lowry, J . B.; and Stone, B . c. 1979. Abaxial anthocyanin layer in leaves of tropical rain forest plants: Enhancer of light capture in deep shade. Biotropica 11:70-77 . Lewis , B. E. 1982. Land-use and conservation on the Osa Peninsula , Costa Rica . Ph D. diss. , University of California , Berkeley. Linhart , Y. B. 1973. Ecological and behavioral determinants of pollen dispersal in hummingbird-pollinated Heliconia . Am. Nat . 107:511-23. Longman , K . A. , and Jenfk, J . 1974. Tropical forest and its environment . London: Longman . McColl , J . G . 1970. Properties of some natural waters in a tropical wet forest of Costa Rica . Bioscience 20:1096-1100. McDiarmid , R . พ.; Ricklefs R . E.; and Foster, M. ร 1977. Dispersal of Stemmadenia donnell-smithii ( Apocynaceae ) by birds. Biotropica 9:9-25. Montoya Maqurn , J . M . 1966. Notas fitogeografica" sobre el Quercus oleoides Cham , y Schlecht . Turrialbo 16:57-66. Mooney, H . A .; BjGrkman , o ; Hall , A . E.; Medina , E ; and Tomlinson , p. B. 1980. The study of the phys iological ecology of tropical plants current static and needs . Bioscience 30:22-26. Mori , ร. A . , and Kallunki , J . A . 1976. Phenology aiKii floral biology of Gustax ia superba ( Lecythidaceae) * central Panama . Biotropica 8: 184-92 . Morrison , D. พ. 1978. Foraging ecology and energetics of the frugivorous bat Artibeus jamaicensis . Ecology 59:716-23.

Standley, p. c. 1937-38. Flora of Costa Rica . Chicago: Field Museum of Natural History. . . Stiles. F. G. 1978. Temporal organization of flowering Energy Commission . among the hummingbird foodplants of a tropical wet , A . ร. 1871. L’ Amerique centrale , recherches sur Oersted forest . Biotropica 10: 194- 210. sa flore et sa geographiephysique: Resultats dun voy- Strong , D. R ., Jr.; and Ray, T. ร. , Jr. 1975. Host tree age dans les Etats de Costa Rica et de Nicaragua , location behavior of a tropical vine Monstera gigantea execute pendant les annees 1846 -1848 . Copenhagen . by skototropism. Science 190:804-6. opler, p. A .; Baker, H . G .; and Frankie. G . พ. 1975. Sugden , A M ., and Robins , R . J . 1979 . Aspects of the Reproductive biology of some Costa Rican Cordia spe- ecology of vascular epiphytes in Colombian cloud farcies ( Boraginaceae ). Biotropica 7:234-47. ร. I. The distribution of the epiphytic flora . Bioest Opler, p. A .; Frankie, G. พ.; and Baker, H . G . 1976. tropica 11:173-88. Rainfall as a factor in the synchronization, release, and Tasaico, H . 1959. La fisionomi'a de las hojas de arboles timing of anthesis by tropical trees and shrubs. J . Bioen algunas formacidnes tropicales . M . Agr. thesis, geogr. 3:231-36. IICA , Turrialba . Perry D. R . 1978 . Factors influencing arboreal epiphytic Toledo, V. M . 1977. Pollination of some rain forest phytosociology in Central America . Biotropica 10: plants by non - hovering birds in Veracruz , Mexico. 235-37. Biotropica 9:262 67. de ordenacidn del bosque de la Tosi , J. A., Jr. 1969-. Mapa ecologico, Republica de Costa Petriceks ,“ J . 1956. Plan Finca La Selva. " M. Agr. thesis, Instituto Inter- Rica: Segun la clasificacion de zonas de vida del mundo americano de Ciencias Agricolas , Turrialba . de L . R . Holdridge . San Josd: Centro Cientffico TropiPittier, H . 1957. Ensayo sobre plantas usuales de Costa cal . Rica . 2d ed . San Josd: University of Costa Rica. Orig . 1973. The Corcovado basin on the Peninsula de inally published 1908 . Osa . San Josd: Tropical Science Center. Pool , D. J.; Snedaker, ร. c.; and Lugo, A . E. 1977. . 1976. La zonificacion ecologica de la Structure of mangrove forests in Florida , Puerto Rico , region de Darihi en la Republica de preliminar Colombia. San Mexico and Costa Rica. Biotropica 9:195-212. Jos£: Centro Tropical . Rabinowitz , D. 1978. Dispersal properties of mangrove Uhl , N. พ. , andCientffico , H . E. , Jr. 1977. Correlations of Moore propagules . Biotropica 10:47-57. inflorescence, structure , and floral anatomy Richards , p. พ. 1952. The tropical rainforest: An eco- with pollinationflower in some palms. Biotropica 9: 170-90. logical study. Cambridge: Cambridge University Vasquez- Yanes , c.; Orozco, A .; Francois , G .; and Trejo, Press. L . 1975. Observations on seed dispersal by bats in a Rodin , L. E., and Bazilevich , N . I. 1967 . Production and tropical humid region in Veracruz , Mexico. Biotropica mineral cycling in terrestrial vegetation . Edinburgh : 7:73-76. Oliver and Boyd . Vaughan , c . Plan maestro para el manejo y deRodin , L. E.; Bazilevich , N. I.; and Rozov, N. N. 1975. sarrollo del. 1979 Parque Corcovado, Peninsula de Productivity of the world ’s main ecosystems. In Pro- Osa , Costa Rica. MNacional . s . thesis , CATTE , Turrialba . ductivity of world ecosystems, pp. 13-26, Washing- Wagner, M., and Scherzer , c. 1856 La Republica ton , D.C.: NAS. Costa Rica en Centro America ( 1974 transl. ) Sawyer, J. o , and Lindsey, A. A . 1971. Vegetation of de Vol . 1. San Josd: Ministera de Cultura , Juventud y the life zones of Costa Rica. Indiana Acad. Sci. Deportes. Monogr. no. 2. , H . 1973. Vegetation of the earth in relation to Schimper, A . R . พ. 1903. Plant geography upon a phys- Walter climate and the ecophysiological conditions . New York: iological basis . Oxford: Clarendon Press Springer- Verlag . Silander, J. A., Jr. 1978. Density-dependent. control of Wfeber , H . 1959. Los pdramos de Costa Rica y รน conca reproductive success in Cassia biflora . Biotropica 10: tenacidn fitogeografica con los Andes Suramericanos. 292-%. San Jos : £ Instituto Geograflco Smith , A. p. 1972. Buttressing of tropical trees: A de- Wfent , F. พ. , and Stark , N. 1968 . .Mycorrhiza . Bioscience scriptive model and new hypotheses. Am . Nat . 18:1035-39. 106:32-46. , c . 1909. La subregion fitogeografica co. 1979. Buttressing of tropical trees in relation to Wferckia . San Jos£: Sociedad Nacional de Agronostarricense bark thickness in Dominica , B. W. I . Biotropica 11: 159 60. , c. R .; Baird , พ. V; and Musselman , L. J. 1979. Smythe-, N. 1970. Relationships between fruiting sea- Wferth Root in Schoepfia Schreb. ( Olacaceae ) . Biosons and seed dispersal methods in a Neotropical for- tropicaparasitism 140 11 : 43. est . Am. Nat . 104:25-35. , Whitmore . c . , and Burnham , c . p. 1975. Tropical rain T Spruce , R . 1908 . Notes of a botanist on the Amazon and forests of the Far East . London: Oxford University Andes . 2 vols. London . Press. 1970. A tropical rain forest: A study of irradiation and ecology at El Verde , Puerto Rico. Washington , D C.: บ ร Atomic

.

.

.

INTRODUCTION

Odum , H . T., and Pigeon , R . F. , eds .

.

.

.

—

I

I

157

PLANTS

The information above is presented in the checklist as a three-part code: category 1 by an upper-case letter, category 2 by a lower-case letter, and category 3 by one or more upper-case letters. As an example , PentaclethrQ macroloba ( Mimosaceae) at La Selva has the code CcWLI , which means mature individuals ( 1 ) are canopy trees , ( 2) are common , and (3) occur in swamps and on slopes and ridges. In some instances we are un certain about the abundance or habitat preferences of a species; hence we have simply left blank those parts of categories and their classes are: 1. Typical physiognomic position attained by mature the code. Let us stress that the checklist is not complete for any individuals single site. A site such as La Selva is much better known c = canopy ( or emergent ) floristically than is Corcovado or Monteverde . Such difร = subcanopy ferences are readily apparent when one looks at the บ = understory checklist entries for plant families with numerous species T = treelet ( < 10 cm dbh or 5 m tall ) of small trees and treelets , such as the Rubiaceae , Piper2. Abundance of mature individuals aceae , or Melastomataceae. c = common (> 10/ha) Aereal extent and habitat diversity also affect the spef = frequent ( 1-10/ha ) cies richness at a site: La Selva is small , but the trees art o = occasional (0.1-1 /ha ) reasonably well known; Corcovado is huge and sh r = rare (0.1-0.01/ha or between 1 /10 ha and have at least five hundred species of trees; Montev 1/1๓ ha) includes three life zones and should also have at least V = very rare ( < 0.01/ha or < 1/ 1๓ ha ) five hundred species of trees; Las Cruces is very small 3. Habitats and not very well known. It should be evident that it is R = riparian ( streamside) not valid to use the presence data in our checklist to A = alluvial ( floodable, but good drainage) make any detailed comparisons of the tree flora be พ = swamp ( floodable, but poor drainage ) P = plateau or flatlands (slopes < 5% , moderate tween sites . We thank Luis Diego Gomez for making available to drainage) L = slope (slopes > 5% , good drainage ) us his informative list of tree ferns for the seven sites. We also thank Valerie Dryer and Bill Haber for valuable help I ridge ( well drained to excessive drainage) o == open country, cultivated areas, pastures , etc. with Monteverde trees. Finally, we welcome correcti G = secondary forest and additions to this checklist. CHECKLIST OF TREES G . ร . Hartshorn and L . J . Poveda พร attempt to provide a reasonably up-to-date checklist of trees known at the seven major sites described in the preceding section . Rather than limiting our effort to a simple presence list , we have incorporated information on canopy stature and abundance of mature individuals , plus major habitats where the species occur. These three

-

Cerro de

Santa Rosa Palo Verde Corcovado La Selva Monteverde Las Cruces La Muerte

Acanthaceae Aphelandra deppeana

Schlecht. & Cham .

TfA

TfPL

Bravaisia integerrima

(Spreng.) Standi .

Justicia aurea Schlecht Razisea spicata Oerst

.

CoA

.

Actinidiaceae Saurauia veraguasensis

CfA TrAG ToPL

Donn . Smith Spondias sp. Annonaceae Anaxagorea Anaxagorea

crassipetala

( Bert . & Balb. ) Skeels

Jacq

. Astronium graveolens Mauria heterophylla HBK Spondias mombin Spondias purpurea .

.

CfL

CfRA CfPLI

. & Sessd Annona reticulata Annona sp.

Moc

Crematospermum

L.

sp.

( Donn .

.

( Rob ) Safford

Safford

Desmopsis maxonii Desmopsis microcarpa Fries Guatteria aeruginosa Guatteria amplifolia Triana & . Guatteria consanguinea

CfPLO UoPLO

CfPLO CfA UoPLOAP UrO

CrA

SfPL SoPL

Klotzsch.

UrA TrA

( HBK . ) Standi .

Fries Safford . Wms.

Standi

ToLG TcPLG

Standi .

ท Sm .) Woodson !e *madenia donnell- smithii ( ) Woodson

_ Rose t

yudenia glabra Benth.

^

UfPL

CoAL SoPL

CoAP; fG

UoL

SoWPL UoO ToLP

UoOL

UoAW

SoPL

CrLI

SrAPL CrPL

CfPL

CvP

CvLI CrPL

SoPLI

SoLI

CoPL SoPL ToL

UoA

SoAP

nadenia macrantha

Stand].

UoP

UrPL

Arg.

. kernellea panamensis OMxxls.) Markgr. Plumeria rubra L. var. acutifolia ( Ait7) Woods.

UrA TrAPL UrAWP SoPL

Planch

Sapranthus palanga L nonopsis pittieri Xylopia sericophylla .& L

UoLI

SrL

Guatteria costaricensis Fries Guatteria inuncta Fries Guatteria tonduzii Diels Guatteria verrucosa Fries Rollinia microsepala Standi . Sapranthus campechianus

Muell.

SoOGI

SoPL SoPL

UfPLI UfLI

SrPLI

Standi.

Aspidosperma myristicifolium (Markgr ) Woodson

CfRA

UrA

Smith ) Fries

Cymbopetalum sp. Desmopsis bibracteata

CoPL

UfPL

.

Cymbopetalum costaricense

'fDon

CrLI

HemsI.

phaeocarpa Mart Annona holosericea Safford Annona purpurea

vnolfia purpurascens U mmadenia alfari

Anacardium excelsum

Crl SrR

Standi.

Tapirira brenesii

Apocynaceae Aspidosperma megalocarpon

Anacardiaceae

L. L

Spondias radlkoferi

TvO

UfLIO

Seem.

Santa Rosa Palo Verde Corcovado La Selva Monteverde Las Cruces Cerro de La Muerte CfRA CoA

Taxa

-

ะ

Taxa

CHECKLIST OF TREES

UvPLOG

Tvp

158

159

PLANTS

CHECKLIST OF TREES

Taxa

Santa Rosa

Palo Verde

UoPL

UfPLRO

Corcovado

La Selva

Monteverde

Las Cruces

Cerro de La Muert(

Santa Rosa

Palo Verde

UoOG

UoO

CoroV ado

La Selva

CoA CvPLI CfA ; cG CoAW

CrAL

CoL

CoWA CrWA

CoL

Monteverde

Las Cnices

Bixaceae

Stemmadenia obovata ( Hook . & Am .) Schum.

Tabernaemontana arborea Rose Tabernaemontana chrysocarpa Blake Tabernaemontana sp. (Dryer 1349) Thevetia o\>ata (Cav. ) A. DC.

Bixa Orellana L.

(Jacq . ) Dugand

ToPR

ร ToPL

TrPL

Araliaceae Dendropanax albertii- smithii Nevl. Dendropanax arboreus (L. ) Dene. & Planch . Dendropanax gonatopodus ( D. Sm . ) A . c. Smith Dendropanax praestans Standi. Dendropanax stenodontus (Standi.) A. c. Smith Didymopanax pittieri March . Oreopanax nubigenum Standi . Oreopanax oerstedianum March . Oreopanax xalapensis ( HBK . ) Dene. & Pkanch . Sciadodendron excelsum Griseb .

CrPL

SoPL

CoL

SoPL Sri

SoPL

CfLI SoLI

CfLI

UoPL

UoPL

SrPL

CcLIG

SrRG

CfWO UfA

UvPL SoLI

CrPL

CrL

CfPL CfPL

CfPL CfAPL

Dugand

CfPL

CfPL

CvW; fl

CfWA

SfPL UvR TvR

SoLPG

Bourreria costaricensis (Standi.) A. Gentry Bourreria quirosii Standi. Cordia alliodora (R . & p. ) Oken Cordia bicolor DC. Cordia collococca L. Cordia cymosa ( D. Sm. ) Standi . Cordia dwyeri Nowicke Cordia gerascanthus L. Cordia lucidula I . M. Johnston Cordia nitida Vahl . Cordia panamensis Riley Ehretia austin- smithii Standi. Toumefortia glabra L. Toumefortia ramonensis Standi.

SrPL

CvL

SfPL

SoPL

CoP

SoPL

CcA CoPL

SoPL

SoPL

SrPL; cOG

SfPL

UfPL

SfPL

SoPLG

CvL

CrA

SrPL ToPL

UoPL

SoPO ToPLG

ToLO

TrO

SOIL

ScPL

SfPLR

Burseraceae

Bursera °graveolens (HBK ) Triana

CrL

Bursera simaruba ( L.) Sarg . Bursera permollis Standi. & Steyerm .

Bursera tomentosa CrW

SoPL

Brunelliaceae

UoA

CfPL

CoAP

Brune Ilia costaricensis Standi. Brune Ilia standleyana Cuatr.

SoPI

CoP

CoAP

Boraginaceae

SrLI

UoA

Huberodendron patinoi Ochroma lagopus Swartz

Quararibea platyphylla Pitt . & D. Sm.

UrR

Betulaceae Alnus acuminata HBK .

Bignoniaceae Crescentia alata HBK . Enallagma latifolia (Mill . ) Small Enallagma sp. ( Dryer # 1160 ) Godmania aesculifolia ( HBK . ) Standi . Jacaranda copaia ( Aubl . ) D . Don Parmentiera valerii Standi . Tabebuia guayacan (Seem.) Hemsl. Tabebuia impetiginosa ( Mart. ) Standi . Tabebuia ochracea ( Cham . ) Standi . Tabebuia rosea ( Vertol . ) DC.

SoPLI

CoLl

CvA CrAL

Quararibea asterolepis Pittier Quararibea bracteolosa (Ducke) Cuatr. Quararibea sp. Quararibea panifolia Standi.

SrLl SfPL

CfPL

(Jacq .)

SrP

SoPL

CfPL

Ceiba aesculifolia (HBK . ) Bntt . & Baker Ceiba pentandra ( L.) Gaertn .

Pachira aquatica Aubl. Pseudobombax septenatum

SoPL

.

Cerro de La Muerte

Bombacaceae Bombacopsis quinatum

CrP

Aquifoliaceae Hex lamprophylla Standi . Ilex skutchii Edwin Ilex vulcanicola Standi

160

Taxa

CrW

CrPL

(Jacq . ) Triana & Planch.

TrI CfLI

Sri CfPLI

Ufl

บก

CrA; fl

CrA

SoPL

161

CHECKLIST OF TREES

PLANTS

Cerro de Muerte La Cruces Las Monteverde La Selva Santa Rosa Palo Verde Corcovado

Taxa Protium costaricense

( Rose )

Protium g labrum

( Rose) Engler

. M . Johnston

Protium panamense

( Rose ) I

Capparidaceae Capparis baducca Capparis indica

UrPL SrPL

Engler

SoAPL SoPL

Protium pittieri ( Rose ) Engler Protium Protium Tetragastris panamensis

sp. sp.

D. M. Porter (Standi . ) Swart .

. & Rendle .

SfPL SfPL UrAR CoLI

UoA forchhammeria pallida Liebm . CoA TrRA Morisoma americana L . Caprifoliaceae

Sambucus mexicana

( Oersted. ) Hemsl

UfA

UoA

Sow

UfPL

SrP

SfP

Presl.

UoR unG

Caricaceae UoLl

Britt . & Rose

. .

Jacaratio dolichaula ( D. Sm ) I M . Johnston Jacaratia costaricensis

Uol

I . M . Johnston

Caesalpiniaceae

-

A teleia herben smithii

Pittier

L

(Jacq .) Willd.

Caesalpinia eriostachys Benth Caesalpinia exostemma DC Cassia emarginata . Cassia fruticosa Mill . Cassia grandis Lf Cassia papillosa ( B . & ) Standi . Cassia reticulata Willd . Copaifea aromatica Dwyer Cynometra hemitomophylla

.

L

Can ocaraceae

ScP UfPLO

Bauhinia ungulata . Caesalpinia coriaria

.

.

R.

SfP; rP

Anthodiscus

Haematoxylon brasiletto Karst Hymeruiea courbaril L . Macrolobium costaricense

TrPL

CoL

CrAP

UrRPL

UoRPL

. SoAI

CfPL

Sri

พ. Burger

CcA

L

( Veil . )

(Swartz ) Spreng .

UrPL

Tachigalia versicolor

CoPLI

Standi . & L. Wms .

CvAPL

CrL

( Loes ) Lundell

UrO

SrL

UrP

Euonymous costaricensis

Standi .

TrP

Maytenus woodsonii Lundell Maytenus sp. ( Hartshorn 1470) Maytenus sp. ( Dryer # 1125 ) Perrottetia longistylis Rose Zinowiewia integerrima Turcz

SoLO CoPLG SrPL CoPL

.

TrLI CoL

Chloranthaceae

SoL

CrL

Swartzia simplex

SrAL

p.

( Miq . ) A . c. Smith

CrPL

UrI

UoPL

CfAPLI

Cuervea kappleriana

CoPL

CcRA

R

Alzatea verticellata . & Crossopetalum tonduzii

UoAP

UfRAO CoL

CoG

Blake Sclerolobium sp.

& Wils.) Standi.

UoA CvL

CoL

Mora oleifera (Triana ) Ducke Parkinsonia aculeata . Peltogyne purpurea Pittier Schizolobium parahybum

Swartzia cubensis

Celastraceae

CfPL

TfP ToPL

UoL

CrPL

Donn. Smith

( D. Sm .) Britt . & Rose ( Aubl. ) Steud .

sp.

Caryoca r costaricense

Dialium guianense

162

.

UoA UfAW

Viburnum costaricanum

CrAL

Lemaireocereus aragonii

( Britt .

UoAP UoAW

.

Cactaceae ( Weber )

TrG

(L.) Fawc

CrLI

Trattinnickia aspera

L.

Capparis odoratissima Jacq . Capparis pittieri Standi Capparis pseudocacao Schum . Crataeva tapia L

SfPL

( Engl . ) o. Kuntze

Tetragastris tomentosa

Cerro de Santa Rosa Palo Verde Corcovado La Selva Monteverde Las Cruces La Muerte

Taxa

Hedy osmum costaricense c. E. Wood Hedyosmum montanum

UoLI

พ. Burger Standi . Chrysobalanaceae

UoLI

Hedyosmum scaberrimum

CrL CrPL

Chrysobalanus icaco L. Couepia polyandra

UfPL

Hirtella lemsii

UoPL ToA

( Kunth .)

Rose L. Wms. & Prance

Hirtella racemosa

Lam .

CvPL

UrPL

UoPL

UoPL

163

PLANTS

CHECKLIST OF TREES

Taxa

Licania arborea Seem. Licania hypoleuca Benth . Licania platypus ( Hemsl . ) Fritsch

Santa Rosa

Palo Verde

Corcovado

CfAL

CcAP

CrL

CoL

SoPL; fO

SoA

SoRA

Gaertn .

SfA

SfRA

SfA

.

Compositae Clibadium leiocarpum Steetz Critonia daleoides DC. Eupatorium angulare Rob. Eupatorium hebabotryum ( DC. ) Hemsl . Eupatorium hylonomum Robinson Eupatorium morifolium Mill. Eupatorium pittieri Klatt Lasianthaea fruticosa ( L. ) K . Becker Senecio arborescens Senecio cooperi Greenm. Verbesina oerstediana Benth .

CoRA

CrR

CrL

CoL

CoL

CrAL

CvAL

CoA

CfA

Santa Rosa

Palo Verde

Cxathea holdridgeana Cyathea multiflora

SoPL

Metaxya rostrata Sephelea cuspidata Sephelea erinacea Sephelea mexicana Sephelea polystichoides Sphaeropteris brunei Sphaeropteris elongata Sphaeropteris horrida Trichipteris costaricensis Trichipteris microdonta Trichipteris schiedeana

CoLI

Trichipteris stipularis Trichipteris trichiata Trichipteris ursina Trichipteris wendlandii Trichipteris williamsii

Corcovado La Selva

Cerro de Monteverde Las Cruces La Muerte

T

T

T

T

T

T

T UfL

Dilleniaceae Curatella americana L.

UrLP

T

T

T UrR

UoL

T T UoPL T

T T

T T T

T

T T

T

T T T T T T

T

Dichapetalaceae Dichapetalum axillare Wnodson Dichapetalum brenesii Standi . Dichapetalum nenatum Cuatr.

TrLOG UrPL ToPLIOG

ToWAP UfPL TrL

T

T

T T

T

T Tr

UrL

CfPO

UoLP

Ebenaceae Diospyros nicaraguensis Standi. Diospyros sp. ( Dryer # 1178)

TrR

ToP

TfLPG

ToO

ToPG

ToOG

SrP

SrP

UrGO

UrL TfOG TrOGI

Elaeocarpaceae Muntingia calabura L.

TfOG

CrPL

Cunoniaceae

CoPL

CoPLGO CoPLGO

Weinmannia pinnata L. Weinmannia wercklei Standi.

164

Taxa

Cxathea suprastrigosa Dicksonia gigantea Oicksonia lobulata

Comaceae Cornus disciflora DC.

Cyatheaceae ( sensu lato ) Cnemidaria choricarpa Cnemidaria cochleana Cnemidaria grandifolia Cnemidaria horrida Cnemidaria mutica Culcita coniifolia Cyathea divergens Cyathea fulva

Ccrro de La Muerte

SoP; fO

SoA

Terminalia amazonia (J . F. Gmel . ) Exell Terminalia bucidioides Standi . & L. Wms. Terminalia oblonga ( R . & p.) Steud

Las Cruces

CoA

Combretaceae Conocarpus erecta L. Laguncularia racemosa ( L.)

Monteverde

SrL

Clethraceae Clethra lanata Mart . & Gal . Cochlospermaceae Cochlospermum vitifolium ( Willd . ) Spreng

La Selva

T T

T

T

Sloanea terniflora (Moc . & Sess£) Standi. Sloanea zuliaensis Pittier

T

T

T T

T

T T

T

T

SrPL; fO

SrP; fO

SoR SoAP CrL

CvL

CvP

CrPL CrPL CoPL

CfP

SvL

Ericaceae Vaccinium consanguineum

T T

Sloanea faginea Standi. Sloanea fragrans Rusby Sloanea laurifolia ( Benth .) Benth Sloanea medusula Schum . & Pitt. Sloanea picapica Standi.

T

Klotzch

T

Erythroxylaceae Erythroxylon amplum Benth .

CfL; Cel

UrAP

UoPL

165

PLANTS

CHECKLIST OF TREES

Taxa

.

Erythroxylon fimbriatum Peyr Erythroxylon havanense Jacq . Erythroxylon lucidum HBK Erythroxylon rotundifolium

.

Lunan.

Cerro de La Cruces Las Monteverde Muerte Selva La Santa Rosa Palo Verde Corcovado UrPL TcL ToP TrAPI

SrL

Taxa fftrorchidium sp. (Dryer # 1071 ) Veconcibea pleiostemona (D. Sm.) Pax. & Hoffm.

de Santa Rosa Palo Verde Corcovado La Selva Monteverde Las Cruces LaCerro Muerte

SoILW CoL

CoPL

SoWA

UrAPL

CrPL

Fabaceae j{

Euphorbiaceae

Acalypha diversifolia Jacq . Acalypha macrostachya Jacq Acalypha schiedeana Schlecht . Adelia triloba

.

( Muell. Arg.)

TfG UoAG

T

.

Boiss

Garcia nutans Rohr Gymnanthes lucida Swartz Hieronyma guatemalensis

Donn . Smith

Hieronyma oblonga

.

var.

.

.

Mill .

Jatropha aconitifolia Jatropha curcas

L.

Mabea occidentalis Benth Manihot carthagenesis

.

.

L

.

(Jacq . ) Muell Arg.

Margaritaria nobilis

(L. f .) Muell. Arg

.

&

Uf?L

TrPI UoA UoPL

บก

Muell.

. . Schum . Schum . ( HBK .) Muell . Arg.

Sapium oligoneuron . & Pitt Sapium pachystachys . & Pitt Sapium thelocarpum & Pitt Stillingia zelayensis

CoA CrAL ToA ToP

CoRA CrL

SoAL

SoA

Tetrorchidium euryphyllum

CoP

CrPL

SoPL CoPL

SoAP CoLI

CvL

CoPL

Dipteryx panamensis

( Pitt . ) Record Dussia cuscatlanica (Standi .)

CfP

Standi. & Steyerm . ( D. Sm.) Harms Erythrina cochleata Standi . Erythrina fusca Lour.

CrPL

Dussia macroprophyllata

SfA SrRL

erw

Erythrina lanceolata

Standley

CoPL CrA CrA ToL

CoPL

CoLI

SoPLO

Lonchocarpus costaricensis ( D. รทใ . ) Pittier Lonchocarpus minimiflorus

CoP

Cf?

Sf?0

SoP

( Schlecht. ) Sousa

ToL

TrWG SrPL CoAP

CoLPO

CoPL CoP

TrL SrPL

I Micheli I Huellera sp.

CoPL CoA

CoPLI

UoPLRO

TrL

SoPO

Ur

SoRA UoPL

UoP SrRA TrPI

Benth .

Machaerium biovulatum

Cf?L CrW

Lonchocarpus eriocarinalis Micheli Lonchocarpus rugosus Benth . Lonchocarpus velutinus

Cf?; oL SoA UrW

Gliricidia sepium ( Jacq . ) Steud . Lonchocarpus acuminatus

Donn Smith Hermann Lonchocarpus orotinus Pittier Lonchocarpus pentaphyllus (Poir. ) DC

CrA

SoA

CoAL CoPL

SoL SoL

Benth .

Lonchocarpus oliganthus

Schum

166

SvL

CoPL SfO

UcAL

.

( & .) Pax Hoffm Sapium jamaicense Swartz Sapium macrocarpum

Standi .

Donn. Smith

Diphysa robinioides

ToP

Phyllanthus valerii Standi . Richeria cf racemosa

P. E

UfLR

SoA

Hemsl.

Dalbergia retusa Dalbergia rucurrensis

CrIL

benthamii (Tul ) Muell Arg . Hieronyma poasana Standi Hippomane mancinella . Hura crepitans L

.

SoRL

ร

Alchornea latifolia waltz Croton gossypiifolius Vahl . Croton killipianus Croisat Croton reflexifolius HBK Croton schiedeanus Schlecht Drypetes standleyi Webster Euphorbia schlechtendalii

Andira inermis (Swartz ) HBK .

UfAW

Hemsl .

Alchornea costaricensis

Pax & Hoffm.

TfG UfAG

cosrmum panamense ( Bentb . ) Yakoul .

CvA SoPL

SoP

I v>rospermum frutescens I Ormosia velutina Rudd Jacq . UoP I Ormosia sp. I Ormosila sp. I ftscidia carthagenensis I Hatymiscium pinnatum Jacq . SoL (Jacq.) Dugand I ^Qtymiscium pleiostachyum D°nn. Smith CrPL

UrP

SvL

CoPL

UoP

CrL

CoPL

CoAL

CrPL 167

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). . . Res . Rept .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear 8ด As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am . Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool . 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrumof the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). ร (พ( FishWildl . . Res. Rept .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

นโ

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). ร (พ( FishWildl . Res. Rept. .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear หว As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus ). . separated from their group , for example , for foraging , . Res. Rept .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; Reynolds , J. E. 1979. The semisocial manatee. Reviewof mouths much like that found in wolves . A rapid high June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most re. is the largest bat in the New World ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro , intermediate Most of these prey are either fairly smelly or roost in fruit plus or insects insects nycteris feed on at night . Some of the species taken match the bats in groups larger mix Trachops Chrotopterus and as forms such , . This same study suggested that not all bats and in large weight as such lizards the sects with small vertebrates can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp 1977. Observations on the foraging behavior phyllostomid bats , Ectophylla and avian prey albaand Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

SPECIES ACCOUNTS ,and adult nearly ,butpup one male ,one half performs and eats Snooks .Baby -grown .1973 Anonymous sized two sites in which Vampyrum roosts are now kept secret ,G. Goodwin G.,and , 8March . Tribune .Tampa continues the bats roost problem .Inthe bred had not yet ,A M 1%1.A review form female that Greenhall a fear that for some small museum will feel the of n eed the to bats of andcontact T rinidad manatees concerning ,o .Notes .. - 1935 and BaiTett ,and T obago .Bull. Am. Mus. social tobehighly appear .They cluster Nat. complete its skin collection at the expense Hist 122 ofthe :191301 remnant . . : 20 . 16 216 .J. Mamm dugongs long periods for stable mayremain group composition . . populations Fieldworkers are ,A.M.1%8.Notes encouraged to . When observe Costaricas .Die saugethiere ,A.von .1869 andGreenhall Frantzius onthe behavior ,the asoft enjoy group emits whole ofthe roost inthe disturbed these large bats ,but false vampire u nnecessary .J. Mamm bat . :247 )35 325 . (Berlin . Naturgesch Archiv . 49 disturbance : and 337 40 . of is the noise c haracteristic sound This . spe chittering collecting should ,R.L.,and Peterson beavoided ofthemana ,D. .Ecology andbehavior whenever -1979 ,p.1%9.Notes — Kirmse Hartman possible . onVamfor the p resence marker a good and c onstitutes cies of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama tree groups Captive . hollow in given a Vampyrum have Can.J. Zool. 47 : . ofMammalogists 14015 Society .5.American . no tion . signals of social variety a wide demonstrated Animals ,— Vehrencamp -w (Trichechus ,F.G. Stiles Indian manatee ,— .The Wfest ;and -L.1977 Husar ,R.— ,J.1977 Bradbury Casebeer -w ,B.— Linsky . -w and ,c N elson . 1 % 3 . ,for ,for example The group their foraging , from Observations on ). the . Res. Rept. separated .t Wildl . Serv FishWildl manatus foraging behavior phyllostomid and , a vian bats prey Ectophylla alba and Vampyrum ofthe roost byaninterlocking the to ontheir return are greeted Neotropical 7: 22. 1carnivorous ,Vampyrum bat spectrum , in Costa spec Rica .J. Mamm . 44 : 186 . 89 trum. J. Mamm ; ofmouths .A rapid . 58 inwolves found high .Review like : manatee that semisocial much ,J.E.1979 .The Reynolds 46978 . ,pp %. wings .93June closed ofthe is byvibration accompanied piping a the toagitate seems behavior that contagious whole often emit aloud tofeed .Bats a roost leaving group ( Vampiro Falso, times it while several asthey screech and depart repeat in , since ofthis sound is unclear the .The function flight bats False Vampire Bat) singly . to appear forage ,these .Most are bats carnivorous re above Asnoted agree ,feces ,or analyses litter roost . ports NewWorld ofstomach bat inthe largest is the that Vampyrum spectrum ,rodents ,and major are adult ofupto with other bats the 200birds weights animposing animal Itis constituents maybetaken .Some is theofthe diet fruit andinsects 80cm.The species on awingspan approaching gand ,but .Onelong a occasion bats share this toberare appears that ofphyllostominine -term inaseries extreme . study aroost prey larger for back to bias brought ofprey onthe remains and adietary in physiognomy similar ,and Guanacaste ,canidlike nonpasserine muzzles bias for birds showed aheavy pinnae complete large have All ,and ,parakeets ,motmots ,doves asMicro trogons - such . asanis forms such .Smaller leaves nose elongate orroost fairly smelly ,intermediate are ofthese either prey in Most fruit plus orinsects nycteris oninsects feed taken match the species ofthe bats at in groups night .Some mixlarger Trachops Chrotopterus and as forms such bats not that all ,and suggested study .This same inweight large aslizards such the with small vertebrates sects Vampyrum , feed ,that can some every night asbirds such aforaging favors substantial vertebrates Vampyrum leaving after are strik its stomach within the anhour fill jaws ,and .The ofVampyrum times other bats rodents at forage great necessarily ,and donot ofcar the bats roost that arow with canine teeth and endowed ingly large the although showed that ;the .Italso from the roost skull distances apronounced nassiallike molars bears shearing behind inthe wasleft group study pup inthe .Weresuckling muscles jaw powerful anchor the crest to sagittal orsubadult adult “guarded byone ,it easily always ,this was could betaken bat leaf have roost not to anose it by ofprovisioning evidence ofthe .Some group form member . inhead dog for asmall , inTrinidad was observed .Early pair explorers ofacaptive name isa misnomer the adult male Thescientific .It species theand behavior bats assumed that maybeacommon inthis simply tales ofvampire this hearing ** remains ofprey for presence certainly be“the would the must account species largest fierce andmost -looking floor ofthe insuf ofVampyruminthe :these individuals roost that .Onereported vampire are quantitatively in bats ,and ficient ofall the energy needs with their for the account to large wings sleep fanned their victims to some enough substantial but towarrant are nose associated the and struc group leaf another the that suggested to back prey brought are blood not eaten .Since all .Bothexplanation used toremove devices were suction tures sitter (and ,provisioning baby ,since vam the ofyoungsters the true roost speculation clearly were suggestions - ) - the have ) these bats slear that (Desmodus .Itic their donot adistinct fan , and , Diaemus Diphylla possibility seems pires with many parallels mayshow family lives that complex . victims and blood instead ofsuck it lap ofupto five roosts Vampyrum ingroups intheother . spectrum ascanids carnivores such mammalian American ofthe awide range occurs over ofhollow bat other .Most upper spe Vampyrum dark reaches trees ,with is but is afunction presumably .This exception tropics never ofSaccopteryx cies avoid these the dense trees dispersion sparse , which .This carnivore asalarge status mayroost opening .The ofits near bilineata the tree skin museum years by of has consist to appear inrecent captured exacerbated have been been that groups few who biologists , , field c areless and group .Onesuch their fanciers young in collectors pair one adult and recent pet are Rica ,one Costa and .Trinidad suckling adults done the bats two ofthe consisted will leave not Guanacaste

..

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-

-

-

-

-

-

**

-

-

-

501

MAMMALS

SPECIES ACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the manatus ) . ป๙ ๐๙Fish Wildl . Serv.t Wildl . Res . Rept . separated foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear ห , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). ร (พ( FishWildl . Res. Rept. .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup, one half-grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett, o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remainstable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbedin the roost, the whole groupemits a soft . ( Berlin) 35:247-325. . Naturgesch Archiv is the spe - chitteringsound. This noise characteristicof ofthemana andbehavior Hartman, D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrateda wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee(Trichechus from their group, for example, for foraging, separated ). บ .ร . FishWildl . . Res . Rept .t Wildl . Serv manatus are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) appearto forage singly. As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost litteranalyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds - such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in groupsat night. Some of the species takenmatchthe bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can some favors substantialvertebratessuch as birds, feed every night, that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains in the floor of the roost: these are quantitativelyinsuf vampire. One reported that individuals of Vampyrum fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. Vampyrum spectrum Vampyrum occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spe tropics but is never dense. This is presumablya function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

-

two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull. Am . Mus . Nat. Hist122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat, in . Panama Can . J. Zool . 47:140 15. , Stiles, F. G.; and Bradbury, J. 1977. Casebeer, R. Linsky, B . and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian prey phyllostomid bats, Ectophylla albaand Vampyrumof the Neotropical carnivorousbat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

’

’ , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group V • screech times itseveral while astheydepart andrepeat in , since ofthissound isunclear thebats .Thefunction flight [ singly . toforage appear J . Bradbury , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

7

MAMMALS

SPECIES ACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich V ampyrum roosts are n ow kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. A review female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . social to be highly . Theyappear cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group composition . . Fieldworkers populations areencouraged , A. M. 1%8. Notesonthebehavior toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius , thewhole asoft group emits of the the roost in d isturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the noise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . hollow in given a Vampyrum have Can . J . Zool . 47:14015. . of Mammalogists Society tionno. 5. American . signals ofsocial variety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical byaninterlocking spectrum , inCosta totheroost ontheirreturn aregreeted 7:1-22. bat, Vampyrum specRica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapid high .Review; ofmouths manatee muchlikethatfoundinwolves , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, timeswhile itseveral astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear 8ด , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andother ofupto200 birds withadult batsarethemajor weights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term a occasion batsthatshare to be rare of phyllostominine inaseries extreme . studyontheremains prey backtoa roostin biasforlarger of preybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles birds showed a heavybiasfornonpasserine pinnae complete Allhavelarge , andtrogons , parakeets , motmots , doves . formssuchasMicro- suchasanis . Smaller noseleaves elongate orroostin fairlysmelly , intermediate Mostof thesepreyareeither plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors withinanhourafterleaving the , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily roost withlarge canine teethanda rowofcar inglyendowed the thatalthough ;theskullbears .Italsoshowed fromtheroost distances apronounced nassiallike molars shearing in the wasleftbehind pupin thestudygroup . Were suckling jawmuscles thepowerful cresttoanchor sagittal orsubadult “guarded byoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . forasmalldogin headform , inTrinidad . Early pairwasobserved explorerstheadult ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient of allthebatsin needs withtheirlarge fortheenergy toaccount wings fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , provisioning , since ofyoungsters thetruevam- theroost speculation wereclearly suggestions -sitter) ) batshave thatthese ( Desmodus , Diaemus , andDiphylla donotfantheir seems . Itisclear adistinct possibility pires with family livesthatmayshowmanyparallels complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe other Vampyrum spectrum roosts ingroups . carnivores suchascanids mammalian of theAmerican overa widerange of hollow batspe Vampyrum occurs . Mostother darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The of itsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin yearsby museum of hasbeenexacerbated toconsist appear in recent thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, Vampyrum spectrum screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

iN

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear yf As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

^ฬ

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. - , 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร, 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร, L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ) Rept . Wildl Wildl Res . . . . Serv Fish . ( ( manatusร พ t are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . . The scientific name is a misnomer Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร,‘ Stiles , F. G .; and Bradbury, J . Casebeer, R . ร,‘ Linsky, B . ร,‘ and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). ร (พ( FishWildl . . Res. Rept .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

SPECIES ACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group Vampyrum spectrum ( Vampiro Falso, times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear J . Bradbury , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

^ฬ

501

MAMMALS

SPECIESACCOUNTS , andone adult nearly , but pup male , one half performs and eats Snooks . Baby -grown .1973 Anonymous sized twosites in which Vampyrum roosts are n ow kept secret , G.G., and Goodwin , 8March . Tribune . Tampa continues thebats problem . Intheroost had notyetbred , A M 1%1. Areview form female that Greenhall a that forfear some small museum willfeeltheneed ofthe to b ats of andcontact T rinidad manatees concerning ,o . Notes . 1935 and BaiTett , and T obago . Bull. Am. Mus social tobehighly appear . They cluster .Nat. complete itsskin collection atthe expense Hist 122 ofthe :191301 r emnant . :216 20. . 16 . J . Mamm dugongs periods forlong remain stable may group composition . . populations Fieldworkers are , A. M. 1% encouraged toobserve . When 8. Notes Costaricas . Diesaugethiere , A. von . 1869 andGreenhall Frantzius onthebehavior , the asoft group emits whole ofthe roost inthe disturbed enjoy these large bats , butunnecessary false vampire . bat J . :247 . ) 35 325 . (Berlin . Naturgesch Mamm Archiv . 49 disturbance : and 337 40 . of is the n oise characteristic sound This . s pe chittering collecting should , R. L., and Peterson beavoided ofthemana , D. 1979 . Ecology andbehavior whenever , p. 1% Kirmse Hartman possible . 9. Notes onVam for the presence marker a good and c onstitutes cies of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama tree groups Captive . hollow in given a Vampyrum have Can.J. Zool. 47:14015. . ofMammalogists Society . 5. American no tion . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles Indian manatee , L. 1977 . TheWfest Husar , R. Linsky , J. 1977 Casebeer , B. and . ,c N elson . 1 % 3 . , forforaging , for example The group their , from Observations on ). the . separated . Res. Rept .t Wildl . Serv FishWildl manatus f oraging behavior phyllostomid and , a vian bats prey Ectophylla alba and Vampyrum oftheNeotropical roost byaninterlocking spectrum tothe ontheir return are greeted 7:1-22. carnivorous , Vampyrum bat , inCosta spec Rica . J . Mamm . 44 :186 -89. trum. J . Mamm ; ofmouths . A rapid . 58 inwolves found high .Review likethat :46978. manatee much , J. E. 1979 . Thesemisocial Reynolds , pp wings .93-%. June closed ofthe isa byvibration accompanied piping thewhole toagitate behavior thatseems contagious often emit a loud tofeed . Bats a roost leaving group ( Vampiro Falso, times itseveral while asthey screech and depart repeat in ,since ofthis sound isunclear the .Thefunction flight bats False Vampire Bat) singly . toforage appear , these . Mostrearecarnivorous bats above Asnoted agree , feces , orroost analyses litter . ports World ofstomach batintheNew isthelargest that Vampyrum spectrum , rodents , and arethemajor adult ofupto200 birds with other bats weights animal It isanimposing constituents maybetaken . Some isthe ofthediet fruitandinsects 80cm .Thespecies on awingspan approaching gand , butthis . Onelong a occasion bats share toberare appears that ofphyllostominine -term inaseries extreme . study prey forlarger back toaroost bias brought ofprey ontheremains and adietary in physiognomy similar , and Guanacaste , canidlike muzzles bias fornonpasserine birds showed aheavy pinnae complete large Allhave , and , parakeets , motmots , doves asMicro trogons - such . asanis forms such . Smaller leaves nose elongate orroost fairly smelly , intermediate areeither ofthese prey in Most fruit plus orinsects nycteris oninsects feed taken match the species ofthe bats atnight in groups .Some mixlarger Chrotopterus and asTrachops forms such notallbats that ,and suggested study . This same largeinweight aslizards such the with small vertebrates sects Vampyrum , feed , that cansome every night asbirds such aforaging favors substantial vertebrates Vampyrum leaving after are strik times within the anhour fillitsstomach jaws , and . The ofVampyrum other bats rodents atgreat forage , and donotnecessarily ofcar thebats roost that arow with canine teeth and endowed ingly large the although showed that ;the .Italso from the roost skull distances apronounced nassiallike molars bears shearing inthe leftbehind was group pup inthestudy . Weresuckling jawmuscles thepowerful crest toanchor sagittal orsubadult adult “guarded byone , itwas easily always , this betakenroost batcould leaf anose itnottohave by ofprovisioning evidence ofthe . Some group form member . inhead dog forasmall , inTrinidad was observed . Early pair explorers ofacaptive name isa misnomer the adult male Thescientific . It species behavior bats assumed thatthe and may inthis simply tales ofvampire this beacommon hearing ** remains ofprey forthepresence certainly be“the would must account species largest fierce andmost -looking oftheroost insuf ofVampyruminthefloor : these thatindividuals . Onereported vampire arequantitatively in , and ficient ofallthebats needs with their fortheenergy toaccount large wings fanned their victims tosleep some enough butaresubstantial towarrant nose associated structhegroup leafand another the that suggested to back arebrought blood not eaten . Since allprey . Bothexplanation used toremove devices were suction tures sitter (and , provisioning baby ,since vam the ofyoungsters the true speculation clearly were suggestions - ) - theroost have ) these bats that (Desmodus . Itisclear do not adistinct fantheirseems , and , Diaemus Diphylla possibility pires with show many parallels may family lives that complex victims and instead ofsuck it. lapblood ofuptofive roosts Vampyrum ingroups inthe other . spectrum ascanids carnivores such mammalian oftheAmerican awide range occurs over ofhollow batspe Vampyrum other . Most upper dark reaches trees , with ispresumably butisnever afunction .This exception tropics ofSaccopteryx cies avoid these the dense trees dispersion sparse , which . This carnivore asalarge may opening . The ofitsstatus near bilineata thetree roost skin years bymuseum of has toconsist appear inrecent captured exacerbated have been been that fewgroups who b iologists , , field c areless and group . One their such young in collectors pair one adult and recent petfanciers are Rica , one Costa and . Trinidad suckling done thebats ofthetwoadults consisted willnotleave Guanacaste

. .

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-

-

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-

**

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). ร (พ( FishWildl . . Res. Rept .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

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-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ) Rept . Wildl Wildl Res . . . . Serv Fish . 8 8 manatusอ ร t are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear 8ด As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . . The scientific name is a misnomer Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , Ectophylla alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

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**

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5๓

501

MAMMALS

SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , Goodwin G. G., andGreenhall . . Tampa Tribune, 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm. 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , but false v ampire unnecessary . bat J . . ) 35:247 325 Mamm Archiv. Naturgesch. (Berlin . disturbance : 49 and 337 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana- ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , Publica the tee (Trichechus manatus ) in Florida . Special false v ampire , bat in . Panama tree groups . Captive Vampyrum in a givenhollow have Can. J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . Animals Casebeer ofsocialsignals a widevariety , Vehrencamp (Trichechus demonstrated , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 , B. andNelson . , c . 1 % 3 . , , for example The for g roup their foraging , from separated Observations on ) Rept the . Wildl Wildl Res . . . . Serv t Fish manatus . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum, inCosta Rica . J . Mamm. 44:186 -89. trum. J . Mamm. 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear J . Bradbury , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrumspectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrumarestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing the behind in left w as g roup pupin thestudy . Were suckling jawmuscles thepowerful cresttoanchor sagittal subadult or one adult “guardedby , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , Trinidad in . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific It . behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking floor insuf of Vampyrum in the oftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested to back prey brought Since a re Both blood not e aten . a ll explanation usedtoremove . devices turesweresuction sitter ( , baby , vam and the of since t he true youngsters roost provisioning speculation wereclearly suggestions - ) - the have ) t hese b ats is that It ( . their distinct clear do not a f an , , Diphylla possibility seems piresDesmodus Diaemus and with show many p arallels m ay family lives t hat complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrumspectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata, whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who , andcareless , petfanciers fieldbiologists . Onesuchgroup young in collectors oneadultpairandtheirrecent Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

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-

-

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**

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นโ

501

MAMMALS

Anonymous . 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von . 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv is the spe - chittering sound . This noise characteristicof ofthemana andbehavior Hartman, D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists . demonstrated a wide variety of social signals. Animals Husar, ร. L . 1977. The Wfest Indian manatee (Trichechus from their group, for example , for foraging, separated ). . . Res . Rept .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso , screech as they depart and repeat it several times while in flight . The function of this sound is unclear, since the bats False Vampire Bat ) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World . ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the major constituents g and a wingspan approaching 80 cm . The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae, canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves , parakeets, motmots , and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds, feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles . Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult member of the group. Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings , and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation. Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam - the roost, provisioning of youngsters ( and the baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidad and Costa Rica are

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two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES

ACCOUNTS

Goodwin , G. G. , and Greenhall, A . M . 1% 1. A review of the batsof Trinidad and Tobago. Bull. Am . Mus . Nat. Hist122:191 -301 . Greenhall, A . M . 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat, in . Panama Can . J. Zool . 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J. Casebeer, R . ร.; Linsky , B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats, Ectophylla prey alba and Vampyrumof the Neotropical carnivorous bat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

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SPECIES ACCOUNTS ,and adult nearly ,butpup one male ,one half performs and eats Snooks .Baby -grown .1973 Anonymous sized two sites in which Vampyrum roosts are now kept secret ,G. Goodwin G.,and , 8March . Tribune .Tampa continues the bats roost problem .Inthe bred had not yet ,A M 1%1.A review form female that Greenhall a fear that for some small museum will feel the of n eed the to bats of andcontact T rinidad manatees concerning ,o .Notes . 1935 and BaiTett ,and T obago .Bull. Am. Mus social tobehighly appear .They cluster .Nat. complete its skin collection at the expense Hist 122 ofthe :191301 remnant . . : 20 . 16 216 .J . Mamm dugongs long periods for stable mayremain group composition . . populations Fieldworkers are ,A.M.1%8.Notes encouraged to . When observe Costaricas .Die saugethiere ,A.von .1869 andGreenhall Frantzius onthe behavior ,the asoft enjoy group emits whole ofthe roost inthe disturbed these large bats ,but false vampire u nnecessary .J . Mamm bat . :247 )35 325 . (Berlin . Naturgesch Archiv . 49 disturbance : and 337 40 . of is the noise c haracteristic sound This . spe chittering collecting should ,R.L.,and Peterson beavoided ofthemana ,D. 1979 .Ecology andbehavior whenever ,p.1%9.Notes Kirmse Hartman possible . onVam for the p resence marker a good and c onstitutes cies of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama tree groups Captive . hollow in given a Vampyrum have Can.J. Zool. 47 : . ofMammalogists 14015 Society .5.American . no tion . signals of social variety a wide demonstrated Animals , Stiles Vehrencamp (Trichechus ,F.G. Indian manatee , L.1977 .The Wfest ;and Husar ,R. Linsky ,J.1977 Bradbury Casebeer ,B. and . ,c N elson . 1 % 3 . ,for ,for example The group their foraging , from Observations on ). ร (พ( FishWildl the . separated . Res. Rept .t Wildl . Serv manatus foraging behavior phyllostomid and , a vian bats prey Ectophylla alba and Vampyrum ofthe roost byaninterlocking the to ontheir return are greeted Neotropical 7: 22. 1carnivorous ,Vampyrum bat spectrum , in Costa spec Rica .J . Mamm . 44 : 186 . 89 trum. J . Mamm ; ofmouths .A rapid . 58 inwolves found high .Review like : manatee that semisocial much ,J.E.1979 .The Reynolds 46978 . ,pp %. wings .93June closed ofthe is byvibration accompanied piping a the toagitate seems behavior that contagious whole often emit aloud tofeed .Bats a roost leaving group ( Vampiro Falso, times it while several asthey screech and depart repeat in , since ofthis sound is unclear the .The function flight bats False Vampire Bat) singly . to appear forage J . Bradbury ,these .Most are bats carnivorous re above Asnoted agree ,feces ,or analyses litter roost . ports NewWorld ofstomach bat inthe largest is the that Vampyrum spectrum ,rodents ,and major are adult ofupto with other bats the 200birds weights animposing animal Itis constituents maybetaken .Some is theofthe diet fruit andinsects 80cm.The species on awingspan approaching gand ,but .Onelong a occasion bats share this toberare appears that ofphyllostominine -term inaseries extreme . study aroost prey larger for back to bias brought ofprey onthe remains and adietary in physiognomy similar ,and Guanacaste ,canidlike nonpasserine muzzles bias for birds showed aheavy pinnae complete large have All ,and ,parakeets ,motmots ,doves asMicro trogons - such . asanis forms such .Smaller leaves nose elongate orroost fairly smelly ,intermediate are ofthese either prey in Most fruit plus orinsects nycteris oninsects feed taken match the species ofthe bats at in groups night .Some mixlarger Trachops Chrotopterus and as forms such bats not that all ,and suggested study .This same inweight large aslizards such the with small vertebrates sects Vampyrum , feed ,that can some every night asbirds such aforaging favors substantial vertebrates Vampyrum leaving after are strik its stomach within the anhour fill jaws ,and .The ofVampyrum times other bats rodents at forage great necessarily ,and donot ofcar the bats roost that arow with canine teeth and endowed ingly large the although showed that ;the .Italso from the roost skull distances apronounced nassiallike molars bears shearing behind inthe wasleft group study pup inthe .Weresuckling muscles jaw powerful anchor the crest to sagittal orsubadult adult “guarded byone ,it easily always ,this was could betaken bat leaf have roost not to anose it by ofprovisioning evidence ofthe .Some group form member . inhead dog for asmall , inTrinidad was observed .Early pair explorers ofacaptive name isa misnomer the adult male Thescientific .It species theand behavior bats assumed that maybeacommon inthis simply tales ofvampire this hearing ** remains ofprey for presence certainly be“the would the must account species largest fierce andmost -looking floor ofthe insuf ofVampyruminthe :these individuals roost that .Onereported vampire are quantitatively in bats ,and ficient ofall the energy needs with their for the account to large wings sleep fanned their victims to some enough substantial but towarrant are nose associated the and struc group leaf another the that suggested to back prey brought are blood not eaten .Since all .Bothexplanation used toremove devices were suction tures sitter (and ,provisioning baby ,since vam the ofyoungsters the true roost speculation clearly were suggestions - ) - the have ) these bats slear that (Desmodus .Itic their donot adistinct fan , and , Diaemus Diphylla possibility seems pires with many parallels mayshow family lives that complex . victims and blood instead ofsuck it lap ofupto five roosts Vampyrum ingroups intheother . spectrum ascanids carnivores such mammalian American ofthe awide range occurs over ofhollow bat other .Most upper spe Vampyrum dark reaches trees ,with is but is afunction presumably .This exception tropics never ofSaccopteryx cies avoid these the dense trees dispersion sparse , which .This carnivore asalarge status mayroost opening .The ofits near bilineata the tree skin museum years by of has consist to appear inrecent captured exacerbated have been been that groups few who biologists , , field c areless and group .Onesuch their fanciers young in collectors pair one adult and recent pet are Rica ,one Costa and .Trinidad suckling adults done the bats two ofthe consisted will leave not Guanacaste

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, andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous -sized twositesinwhichVampyrum roostsarenowkeptsecret Goodwin , G. G., andGreenhall . . Tampa Tribune , 8 March continues Intheroostthebatsforma problem . , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. พ. 1935 . Notesconcerning and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. ร. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , ร.; Stiles Vehrencamp (Trichechus , F. G.; andBradbury manatee , ร. L. 1977 . TheWfestIndian Husar , R. ร.; Linsky , J. 1977 Casebeer , B. ร.; andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the manatus ) . อ 8ร 8 Fish Wildl . Serv.t Wildl . Res . Rept . separated foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ฟ screech Vampyrum spectrum / times itseveral while astheydepart andrepeat in , since ofthissound isunclear thebats .Thefunction flight 1 singly . toforage appear J . Bradbury , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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, andonenearly adult , but pup male , onehalf andperforms eats Snooks . Baby -grown .1973 Anonymous -sized , 8March . Tribune . Tampa continues thebats Intheroost problem . form female thathadnotyetbred a and contact manatees concerning , o. . Notes 1935 BaiTett , and social tobehighly . Theyappear cluster :216 . 16 . J . Mamm dugongs -20. forlongperiods stable mayremain group composition . . When Costaricas . Diesaugethiere , A. von . 1869 Frantzius , thewhole asoft group emits the roost in disturbed . :247 ) 35 . (Berlin . Naturgesch Archiv -325 ofthespe ischaracteristic noise sound . This - chittering ofthemana , D. 1979 . Ecology andbehavior Hartman forthe presence m arker a good a nd constitutes cies of ) inFlorida Publica . Special manatus tee(Trichechus t ree groups Captive . h ollow in given a Vampyrum have . ofMammalogists Society tionno. 5. American . Animals signals ofsocial variety a wide (Trichechusdemonstrated manatee , L. 1977 . TheWfestIndian Husar , , for example for group their foraging , from separated ). . . Res. Rept .t Wildl . Serv FishWildl manatus byaninterlocking totheroost ontheir return are greeted 7:1-22. ; ofmouths . A rapid inwolves high .Review likethatfound manatee much , J. E. 1979 . Thesemisocial Reynolds , pp wings .93-%. June oftheclosed isa byvibration accompanied piping thewhole to agitate behavior thatseems contagious emita loud tofeedoften . Bats a roost leaving group (VampiroFalso , Vampyrum spectrum times it several while asthey screech and depart repeat in ,since ofthissound isunclear thebats .Thefunction flight FalseVampireBat) singly . toforage appear 8ด , these . Mostrearecarnivorous bats above Asnoted agree , feces , orroost litteranalyses . ports World ofstomach batintheNew isthelargest that Vampyrum spectrum , rodents , and arethemajor ofupto200 birds withadult other bats weights animal It isanimposing constituents maybetaken . Some isthe of thediet fruitandinsects 80cm .Thespecies on approaching gandawingspan , butthisappears . Onelong a occasion bats toberare thatshare ofphyllostominine -term inaseries extreme . study prey forlarger back toaroost bias ofpreybrought ontheremains andadietary in physiognomy similar , and Guanacaste , canidlike muzzles bias fornonpasserine birds showed aheavy pinnae complete large Allhave , andtrogons , parakeets , motmots , doves asMicro - suchasanis . forms such . Smaller leaves nose elongate orroost fairlysmelly preyareeither in Mostofthese fruit, intermediate plus orinsects nycteris oninsects feed taken match thebats ofthespecies atnight in groups .Some mixlarger Chrotopterus and asTrachops forms such thatnotallbats ,and suggested study . Thissame aslizards such thelargein weight withsmall vertebrates sects Vampyrum , feed , thataforaging cansome every night asbirds such favors substantial vertebrates Vampyrum leaving arestrik times within the anhourafter fill itsstomach , and . Thejaws ofVampyrum other bats rodents atgreat forage , and donotnecessarily roost thatthebats withlarge canine teeth andarowofcar inglyendowed the thatalthough ; theskull .Italsoshowed from theroost distances apronounced nassiallike molars bears shearing in the wasleftbehind group pupin thestudy . Weresuckling jawmuscles thepowerful crest toanchor sagittal orsubadult adult “guarded byone , it was easily always , thisbatcould betakenroost leaf anose it nottohave by ofprovisioning evidence ofthegroup . Some form member . dogin head forasmall , inTrinidad observed . Early pairwas explorers ofacaptive name is a misnomer theadult male Thescientific . It behavior bats assumed thatthe andthismaybeacommon in thisspecies simply tales of vampire hearing ** remains ofprey forthepresence certainly be“the would must account species largest fierce andmost -looking insuf of Vampyrumin theflooroftheroost : these thatindividuals . Onereported vampire arequantitatively in , and ficient ofallthebats needs withtheir fortheenergy toaccount large wings fanned their victims tosleep some enough butaresubstantial towarrant associated struc thegroup leafand another thatthenose suggested to back arebrought blood notallprey eaten . Since . Both explanation used toremove devices were suction tures sitter (and , provisioning ,since vam thebaby ofyoungsters thetrue speculation clearly were suggestions - ) - theroost have ) bats thatthese (Desmodus . It isclear donotfantheir seems adistinct , and , Diaemus Diphylla possibility pires with show many parallels family lives thatmay complex victims andlapblood instead ofsuck it. ofuptofiveinthe other roosts Vampyrum ingroups . spectrum ascanids carnivores such mammalian oftheAmerican awide range occurs over ofhollow batspe Vampyrum other . Most darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably tropics ofSaccopteryx cies avoid these dense trees dispersion , which . Thissparse carnivore asalarge mayroost . The ofitsstatus near bilineata thetreeopening skin years bymuseum of hasbeen toconsist appear in recent captured exacerbated been thathave fewgroups who biologists , , field c areless and group . One such young in collectors pairandtheir one adult recent petfanciers are Rica , onesucklingwillnotleave Costa and . Trinidad done thebats of thetwoadults consisted Guanacaste

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SPECIESACCOUNTS

inwhich twosites Vampyrum roosts arenowkeptsecretGoodwin , G.G., and , A M 1%1. A review Greenhall forfearthatsome small museum willfeeltheneed ofthe t o bats of Trinidad and T obago . Bull. Am. Mus .Nat. complete itsskin collection atthe expense oftheremnantHist 122 :191301 . . Fieldworkers populations areencouraged , A. M. 1% toobserve 8. Notes and Greenhall onthebehavior ofthe enjoy these large bats , butunnecessary false vampire .J . Mamm bat . 49 disturbance : and 337 40 . collecting should , R. L., andKirmse Peterson beavoided whenever , p. 1% possible . 9. Notes onVam pyrumspectrum , thefalse vampire , bat in . Panama Can.J. Zool. 47:14015. , Vehrencamp , F. G.; andBradbury Stiles , R. Linsky , J. 1977 Casebeer , B. and . , . 1% N elson 3 . The Observations on the foraging behavior phyllostomid and , Ectophylla a vian bats prey alba and Vampyrum of theNeotropical carnivorous , Vampyrum bat spectrum , inCosta spec Rica . J . Mamm . 44 :186 -89. trum.

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Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

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two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

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Trachops ์ Chrotopterus

Vampyrum ฺ์

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ฺ จ ฺ์ ด จด ฺ หด สด 5! 5ด ห ๑ ์ ด . Am Bull . ด Mus Nat. ๑ Hist ฌ ฐ5 5 ด ด ๑ ฺ หด สด 5 ๑ 0 ! ด ์ ๑ ๑ ๑ ๑ฺ ๑ ๑ ด J . Mamm . ฐ“ ์๑ ฺ ด ไ วด็ ฺ ์ ฺด ด ์ 5! ด ฝ ด Vam pyrumspectrum , ๑ ๑ ๑ ฺ ด Can.J. Zool. ฐ" ด ฺ ฺ ด ±์ อ จด ฺ ฺ วด 8 ด ด ฺ ๐ด ์ ฺ ] ด 5 ! ํ ด ๑ ๑ ์ ฺ Ectophylla alba ์ Vampyrum ๑ ฺ spectrum Vampyrum , ๖ด spec J . Mamm . ฐไ ๊ด trum. J . Mamm . ฐฝด

Vampyrum

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์ Vampyrum

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ฺ ๑ ๑ ด ์ ๑ ์ ๖ 9 9 ๑ ์ ๑ ์ ไ Vampyrum ๑ ๑ ฐ ๑๑ ด๑๑ ๑ ๑ ๑ ฺ์ ๑ ์ ]๑ ์๑ ๑ ์ ด ๑ ด ้ ์๑ ไ ิ ฺ ๑ ๑ ไ ๑ฺ ) ๑ (Desmodus ด ๖ , ์ , Diaemus Diphylla ์ ์ ์ ด ๑ Vampyrum ด spectrum ๖ ๑ ๑ ด ๑ Vampyrum ๖ Saccopteryx ด ์ ๑ฺ ๑ ๑ , ๖ ๑ ด๑ ด๑ bilineata ๑ ์ ฺ์ ฺ ์ ด๑ ๑ ์ ฺ ๖ ๑ ์ ด ๑ ๑ ์๑ ๑

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). ร (พ( FishWildl . Res. Rept. .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

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two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

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SPECIES ACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich roosts are n ow kept s ecret , G. G., andGreenhall Goodwin . March ๖ ฺ8๑ . continues problem . Intheroostthebatsforma , A M 1%1. A review female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . ดด . ด social to be highly . Theyappear cluster complete itsskincollection attheexpense oftheremnant :191301. 122 20. ด 16:216 .8 ด dugongs forlongperiods stable mayremain group composition . . Fieldworkers populations areencouraged , A. M. 1%8. Notesonthebehavior toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius , thewhole asoft group emits of the the roost in d isturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. 8ด ด49:337 . ) 35:247 325 ด (Berlin ด disturbance and 40 . of i s the noise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided ๑ ไ ciesandconstitutes , D. 1979 . ์ whenever , p. 1%9. Notes Hartman possible . on ไ forthe presence marker a good of ฺ ิ ้ Publica the ด Special ๖ ๑ false vampire , in Panama bat . t ree groups Captive . hollow in given a have 8ด ด47:14015. ด . of Mammalogists Society tionno. 5. American . signals ofsocial variety a wide demonstrated Animals , Vehrencamp ้ , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on ิ the ด ด ด separated ด ด ด foraging behavior phyllostomid and , avian bats p rey and of theNeotropical byaninterlocking totheroost ontheirreturn aregreeted 7:1-22. carnivorous ฺinCosta ไ Rica .8 ด ด44:186 -89. ด 8ด ด58:46978. bat, . A rapid high . ; ofmouths manatee muchlikethatfoundinwolves , J. E. 1979 . Thesemisocial Reynolds , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso , timeswhile itseveral astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat ) singly . toforage appear J . Bradbury , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld isthelargest that , rodents , andother ofupto200 birds withadult batsarethemajor weights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term a occasion batsthatshare to be rare of phyllostominine inaseries extreme . studyontheremains prey backtoa roostin biasforlarger of preybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles birds showed a heavybiasfornonpasserine pinnae complete Allhavelarge , andtrogons , parakeets , motmots , doves ไ suchasanis . formssuchas . Smaller noseleaves elongate orroostin fairlysmelly , intermediate Mostof thesepreyareeither plusfruit orinsects feedoninsects taken match thebats atnight in groups . Someofthespecies mixlarger and forms suchas thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects , feedeverynight , thata foraging cansome suchasbirds favors substantial vertebrates arestrik timesfillitsstomach withinanhourafterleaving the , andother . Thejawsof bats rodents atgreat forage , andthatthebatsdonotnecessarily roost withlarge canine teethanda rowofcar inglyendowed the thatalthough ;theskullbears .Italsoshowed fromtheroost distances apronounced nassiallike molars shearing in the wasleftbehind pupin thestudygroup . Were suckling jawmuscles thepowerful cresttoanchor sagittal orsubadult “guarded byoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . forasmalldogin headform , inTrinidad . Early pairwasobserved explorerstheadult ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking in theflooroftheroost insuf of : thesearequantitatively thatindividuals . Onereported vampire , and ficient of allthebatsin needs withtheirlarge fortheenergy toaccount wings fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , provisioning , since ofyoungsters thetruevam- theroost speculation wereclearly suggestions -sitter) ิ batshave thatthese ้ . Itisclear donotfantheir seems adistinct ฺand ฺ possibility pires with family livesthatmayshowmanyparallels complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe other roosts ingroups . carnivores suchascanids mammalian of theAmerican occurs overa widerange of hollow batspe . Mostother darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably tropics of ciesavoid thesetrees dense dispersion ฺwhich . Thissparse . The of itsstatusasa largecarnivore ๖mayroostnearthetreeopening skin yearsby museum of hasbeenexacerbated toconsist appear in recent thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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, andonenearly adult , but pup male , onehalf andperforms eats Snooks . Baby -grown . 1973 Anonymous -sized . . Tampa Tribune , 8 March continues thebats Intheroost problem . hadnotyetbred form female that a and contact manatees , o. . Notesconcerning 1935 BaiTett , and social tobehighly . Theyappear cluster . . J . Mamm . 16:216 dugongs -20 forlongperiods stable mayremain group composition . . When Costaricas saugethiere , A. von . 1869. Die Frantzius , thewhole asoft group emits the roost in disturbed ) 35:247 Archiv. Naturgesch . (Berlin -325. ofthespe ischaracteristic noise sound . This chittering , D. 1979 . Ecology and behavior of the mana - cies Hartman for the presence marker a good and constitutes of Publica tee (Trichechus manatus ) in Florida . Special t ree groups Captive . h ollow in given a Vampyrum have . ofMammalogists Society tionno. 5. American . Animals signals ofsocial a widevariety (Trichechus demonstrated manatee , L. 1977 . TheWfestIndian Husar , , for example for group their foraging , from separated manatus ) . ร (พ( Fish Wildl . Serv.t Wildl . Res . Rept . byaninterlocking totheroost ontheir return aregreeted 7:1-22. . A rapid inwolves high .Review; ofmouths likethatfound manatee much , J. E. 1979 . Thesemisocial Reynolds , pp wings . 93-%. June oftheclosed isa byvibration accompanied piping thewhole to agitate behavior thatseems contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear J . Bradbury , these . Mostrebatsarecarnivorous above Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andother arethemajor ofupto200 birds withadult bats weights animal Itisanimposing constituents maybetaken . Somefruitandinsects isthe of thediet 80cm. The species on approaching ganda wingspan , butthisappears . Onelong a occasion batsthatshare toberare ofphyllostominine -term inaseries extreme . study prey back toa roost biasforlarger ofpreybrought ontheremains anda dietary in physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae complete large Allhave , andtrogons , parakeets , motmots , doves . forms suchasMicro- suchasanis . Smaller noseleaves elongate orroost fairly smelly , intermediate in Mostofthesepreyareeither fruit plus orinsects nycteris feedoninsects taken match thebats ofthespecies atnight in groups . Some forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge inweight suggested study . Thissame aslizards such withsmall vertebrates sects Vampyrum cansome , feedevery , thata foraging night suchasbirds substantial vertebrates Vampyrum favors leaving after within the anhour , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , and donotnecessarily roost thatthebats withlarge canine teeth anda rowofcar endowed ingly the although that ;theskull .Italsoshowed from theroost distances apronounced nassiallike molars bears shearing inthe wasleftbehind group pupinthestudy . Weresuckling jawmuscles thepowerful crest toanchor sagittal orsubadult “guarded byoneadult , itwasalways easily betakenroost a noseleaf, thisbatcould itnottohave by ofprovisioning evidence ofthegroup . Some form member . doginhead forasmall , inTrinidad wasobserved . Early pair explorers ofacaptive name is a misnomer theadult male Thescientific . It behavior batssimply assumed thatthe andthismaybea common inthisspecies talesof vampire hearing remains ofprey forthepresence certainly be“the** would must account species largest andmostfierce -looking insuf of Vampyrum intheflooroftheroost : these thatindividuals . Onereported vampire arequantitatively , and ficient ofallthebatsin needs withtheir fortheenergy toaccount large wings fanned their victims tosleep some enough butaresubstantial towarrant struc thegroup leafandassociated another thenose that suggested to back arebrought blood notallprey eaten . Since . Both explanation usedtoremove devices weresuction tures sitter (and , provisioning , since vam thebaby ofyoungsters thetrue speculation clearly were suggestions - ) - theroost have ) these bats that ( Desmodus , Diaemus , and . Itisclear adistinct Diphylla donotfantheirseems possibility pires with many parallels family livesthatmayshow complex victims andlapblood instead ofsuckit. ofuptofiveinthe other Vampyrum spectrum roosts ingroups . carnivores suchascanids mammalian oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most upper dark reaches trees , withtheexception butisnever afunction .Thisispresumably ofSaccopteryx tropics ciesavoid these dense trees dispersion . Thissparse carnivore asa large mayroostnearthetreeopening . The ofitsstatus bilineata , which skin years bymuseum of hasbeenexacerbated toconsist appear inrecent captured been thathave fewgroups who biologists , , field careless and . Onesuchgroup young in collectors pair oneadult andtheir recent petfanciers are Rica , onesucklingwillnotleave andCosta . Trinidad done thebats of thetwoadults consisted Guanacaste

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inwhich twosites Vampyrum roosts arenowkeptsecretGoodwin , G. G., and , A M 1% Greenhall 1. Areview forfearthatsome small museum willfeeltheneedto ofthebats of Trinidad and T obago . . Am. Mus.Nat . Bull complete itsskin collection attheexpense oftheremnantHist 122:191301 . . Fieldworkers populations areencouraged , A. M. 1% Greenhall toobserve 8. Notes and onthebehavior ofthe enjoy these large bats , butunnecessary falsevampire . J . Mamm . 49 bat disturbance : and 337 40 . collecting should , R. L., andKirmse Peterson beavoided whenever , p. 1% possible . 9. Notes onVampyrum spectrum , thefalsevampire , bat in . Panama Can . J . Zool . 47:14015. , Vehrencamp , F. G.; andBradbury Stiles , R. Linsky , J. 1977 Casebeer , B. and . , . 1% N elson 3 . The Observations on the f oraging behavior phyllostomid and , Ectophylla alba andVampyrum a vian bats prey of theNeotropical carnivorous , Vampyrum specbat spectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . SPECIES ACCOUNTS

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SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group (VampiroFalso , Vampyrum spectrum times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight FalseVampireBat) singly . toforage appear 8ด , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals for example , for foraging, ). ป๙ manatus ๐๙FishWildl. Serv.t Wildl. Res. Rept. separated from their groupto, the roost by an interlocking are greeted on their return 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear หว As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro , intermediate Most of these prey are either fairly smelly or roost in nycteris feed on insects or insects plus fruit mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops , sects with small vertebrates such as lizards and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum hour leaving after are strik its stomach within the an Vampyrum fill jaws , . of The times other bats rodents and at forage great necessarily , do not row car of and the bats roost that a with canine teeth and endowed ingly large nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but March. ๖ ฺ8๑ problem continues. BaiTett , o. 1935. Notes concerning manatees and ด 16:216-20. dugongs. 8ด Frantzius, A . von. 1869. Die saugethiere Costaricas. ด ( Berlin ) 35:247-325. ด ๑ ไ ์ Hartman , D. 1979. ิ ้ Publica ด Special ๖ ๑ tion no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee ้

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals ิ ด ด ด separated from their group , for example , for foraging , ด ด ด are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ไ such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds , feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufvampire. One reported that individuals of fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ิdo not fan their seems a distinct possibility. It is clear that these bats have ฺand ฺ pires ้ complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees , with the exception of ฺwhich ๖may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. 1% 1. A review for fear that some small museum will feel the need bats of Trinidad and Tobago. M. ด ด . ด complete its skin collection at the expense of the remnantto of the 122 : 191 301 . populations. Fieldworkers are encouraged to observe and Greenhall , A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. 8ด ด49:337-40. of the and collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1%9. Notes on ไ ฺthe false 8ด ด47:140 15. vampire bat , in Panama. ด , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , and of the Neotropical carnivorous ฺin Costa Rica. 8 ไ ด ด44:186-89. ด 8ด ด58:469 78. bat ,

-

-

-

**

501

MAMMALS

, andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous -sized , 8 March . Tribune . Tampa continues Intheroostthebatsform problem . female thathadnotyetbred a and contact manatees , o. . Notesconcerning 1935 BaiTett , and social to be highly . Theyappear cluster :216 . 16 . J. Mamm dugongs -20. forlongperiods stable mayremain group composition . . When Costaricas . Diesaugethiere , A. von. 1869 Frantzius , thewhole asoft group emits the roost in disturbed ) 35:247 . (Berlin . Naturgesch Archiv -325.ofthemana ofthespe ischaracteristic sound . Thisnoise - chittering , D. 1979 . Ecology andbehavior Hartman forthe presence marker a good and constitutes cies of ) inFlorida Publica . Special manatus tee(Trichechus t ree groups Captive . hollow in given a Vampyrum have . ofMammalogists Society tionno. 5. American . Animals signals ofsocial a widevariety demonstrated (Trichechus manatee , L. 1977 . TheWfestIndian Husar , , for example for g roup their foraging , from separated ). . . Res . Rept .t Wildl . Serv FishWildl manatus byaninterlocking totheroost ontheir return aregreeted 7:1-22. ; ofmouths . A rapid inwolves high .Review likethatfound manatee much , J. E. 1979 . Thesemisocial Reynolds , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear , these . Mostrebatsarecarnivorous above Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld isthelargest that Vampyrum spectrum , rodents , andother ofupto200 birds withadult batsarethemajor weights animal Itisanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong a occasion batsthatshare to be rare ofphyllostominine -term inaseries extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae complete Allhavelarge , andtrogons , parakeets , motmots , doves - suchasanis . forms suchasMicro . Smaller noseleaves elongate orroostin fairly smelly , intermediate Mostofthesepreyareeither plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies mixlarger andChrotopterus forms suchasTrachops thatnotallbats , andthelarge inweight . Thissamestudysuggested aslizards such withsmall vertebrates sects Vampyrum , feedeverynight , thata foraging cansome suchasbirds favors substantial vertebrates Vampyrum arestrik timesfillitsstomach within the anhourafterleaving , andother . ThejawsofVampyrum bats rodents atgreat forage , andthatthebatsdonotnecessarily roost withlarge canine teethanda rowofcar inglyendowed the thatalthough ;theskull .Italsoshowed from theroost distances apronounced nassiallike molars bears shearing inthe wasleftbehind pupinthestudygroup . Were suckling jawmuscles thepowerful cresttoanchor sagittal orsubadult “guarded byoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some form member . doginhead forasmall , inTrinidad . Early pairwasobserved explorers ofacaptive name is a misnomer theadult male Thescientific . It behavior batssimply assumed thatthe andthismaybea common inthisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking intheflooroftheroost insuf of Vampyrum : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheir fortheenergy toaccount large wings fanned their victims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices weresuction tures (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) batshave thatthese (Desmodus . Itisclear donotfantheir seems adistinct , and , Diaemus Diphylla possibility pires with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe other roosts Vampyrum ingroups . spectrum carnivores suchascanids mammalian oftheAmerican occurs overa widerange ofhollow batspe Vampyrum other . Most darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably tropics ofSaccopteryx ciesavoid these dense trees dispersion , which . Thissparse carnivore asa large mayroostnearthetreeopening . The ofitsstatus bilineata skin yearsby museum of hasbeenexacerbated toconsist appear inrecent thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors pairandtheir oneadult recent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

-

SPECIES ACCOUNTS

twositesinwhich Vampyrum roosts arenowkeptsecretGoodwin , G. G., andGreenhall , A M 1%1. A review forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and T obago . Bull. Am. Mus .Nat. complete itsskincollection attheexpense oftheremnant Hist122:191301 . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve and Greenhall onthebehavior ofthe enjoytheselarge bats , butunnecessary falsevampire bat. J. Mamm . 49 disturbance : and 337 40 . collecting should , R. L., andKirmse Peterson beavoided whenever , p. 1%9. Notes possible . onVam pyrum spectrum , thefalse vampire , bat in . Panama Can . J. Zool . 47 :14015. , Vehrencamp , F. G.; andBradbury Stiles , R. Linsky , J. 1977 Casebeer , B. andNelson . , . 1%3.The Observations on the foraging behavior phyllostomid and , Ectophylla a vian bats p rey alba and Vampyrumof theNeotropical carnivorous bat, Vampyrum spectrum , inCosta spec Rica . J. Mamm . 44 :186 -89. trum .

. .

-

c

-

-

J. Mamm . 58 :46978.

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. พ. 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ร. 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus manatus ) . ร ท- ท Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrum spectrum ( Vampiro Falso, False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla alba and Vampyrum prey of the Neotropical bat , Vampyrum specspectrum , in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:carnivorous 469-78.

-

**

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. พ. 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv. Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ร. 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus Fish Wildl . Serv.t Wildl . Res . Rept . manatus ) . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. ( Vampiro Falso,

False Vampire Bat)

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vampyrum spectrum , the false vampire bat , in Panama. Can. J . Zool . 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . on the foraging behavior and avian1977 phyllostomid bats , Ectophylla alba and Vampyrum ofObservations prey the Neotropical bat , Vampyrum specspectrum, in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:carnivorous 469-78.

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SPECIES ACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich V ampyrum roosts are n ow kept s ecret , G. G., andGreenhall Goodwin , 8 March . Tribune . Tampa continues problem . Intheroostthebatsforma , A M 1%1. A review female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull. Am. Mus social to be highly . Theyappear cluster .Nat. complete itsskincollection attheexpense oftheremnant Hist122:191301 . :216 20. . 16 . J. Mamm dugongs forlongperiods stable mayremain group composition . . populations Fieldworkers areencouraged , A. M. 1%8. Notesonthebehavior toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius , thewhole asoft group emits of the intheroost disturbed enjoytheselarge b ats , butunnecessary false v ampire bat. J. Mamm . ) 35:247 325 . (Berlin . Naturgesch Archiv . 49 disturbance : and 337 40 . of i s the noise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided ofthemana , D. 1979 . Ecology andbehavior whenever , p. 1%9. Notes Hartman possible . onVam forthe presence marker a good and constitutes cies of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . hollow in given a Vampyrum have Can . J. Zool. 47:14015. . of Mammalogists Society tionno. 5. American . signals ofsocial variety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on ). the . separated . Res. Rept .t Wildl . Serv FishWildl manatus foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical byaninterlocking spectrum totheroost ontheirreturn aregreeted 7:1-22. carnivorous bat, Vampyrum , inCosta spec Rica . J. Mamm . 44 :186 -89. trum . J. Mamm ; ofmouths . A rapid . 58 high .Review :46978. manatee muchlikethatfoundinwolves , J. E. 1979 . Thesemisocial Reynolds , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, timeswhile itseveral astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear 8ด , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld isthelargest that Vampyrum spectrum , rodents , andother ofupto200 birds withadult batsarethemajor weights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term a occasion batsthatshare to be rare of phyllostominine inaseries extreme . studyontheremains prey backtoa roostin biasforlarger of preybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles birds showed a heavybiasfornonpasserine pinnae complete Allhavelarge , andtrogons , parakeets , motmots , doves - suchasanis . formssuchasMicro . Smaller noseleaves elongate orroostin fairlysmelly , intermediate Mostof thesepreyareeither plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies mixlarger andChrotopterus forms suchasTrachops thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum , feedeverynight , thata foraging cansome suchasbirds favors substantial vertebrates Vampyrum arestrik timesfillitsstomach withinanhourafterleaving the , andother . ThejawsofVampyrum bats rodents atgreat forage , andthatthebatsdonotnecessarily roost withlarge canine teethanda rowofcar inglyendowed the thatalthough ;theskullbears .Italsoshowed fromtheroost distances apronounced nassiallike molars shearing in the wasleftbehind pupin thestudygroup . Were suckling jawmuscles thepowerful cresttoanchor sagittal orsubadult “guarded byoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . forasmalldogin headform , inTrinidad . Early pairwasobserved explorerstheadult ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrumin theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient of allthebatsin needs withtheirlarge fortheenergy toaccount wings fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , provisioning , since ofyoungsters thetruevam- theroost speculation wereclearly suggestions -sitter) ) batshave thatthese (Desmodus . Itisclear donotfantheir seems adistinct , andDiphylla , Diaemus possibility pires with family livesthatmayshowmanyparallels complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe other roosts Vampyrum ingroups . spectrum carnivores suchascanids mammalian of theAmerican occurs overa widerange of hollow batspe Vampyrum . Mostother darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably tropics ofSaccopteryx ciesavoid thesetrees dense dispersion , which . Thissparse mayroostnearthetreeopening . The of itsstatusasa largecarnivore bilineata skin yearsby museum of hasbeenexacerbated toconsist appear in recent thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

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two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

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SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , thefalsevampire ) ( Publica tee Trichechus manatus in Florida . Special , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, Vampyrum spectrum times itseveral while astheydepart screech andrepeat in , since ofthis s ound isunclear the . The f unction flight ) bats Bat Vampire False singly . toforage appear หว , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific It . behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , provisioning , since ofyoungsters thetruevam speculation wereclearly theroost suggestions -sitter) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla ) donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , andcareless , petfanciers field . Onesuchgroup young in collectors oneadultpairandtheirrecent Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. - , 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ร, 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร, L. 1977. The Wfest Indian manatee (Trichechus manatus ) . ร (พ( Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrum spectrum ( Vampiro Falso, False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

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pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , ร,‘ Stiles , F. G .; and Bradbury, J . Casebeer, R . ร,‘ Linsky, B . ร,‘ and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla alba and Vampyrum prey of the Neotropical bat , Vampyrum specspectrum , in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:carnivorous 469-78.

-

**

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). ร (พ( FishWildl . Res. Rept. .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). อ 8ร 8 FishWildl . Res. Rept. .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

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-

-

-

**

-

-

-

501

’

’ , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the manatus ) . ร (พ( Fish Wildl . Serv.t Wildl . Res . Rept . separated foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group V • screech times itseveral while astheydepart andrepeat in , since ofthissound isunclear thebats .Thefunction flight [ singly . toforage appear หว , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

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-

-

-

-

-

**

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-

-

7

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. พ. 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ร. 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus manatus ) . อ 8ร 8 Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrum spectrum ( Vampiro Falso, False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

. G. , and , A. M . 1 % 1 . A of the bats of TrinidadGreenhall and Tobago. Bull . Am . Musreview . Nat . Hist 122: 191 301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla alba and Vampyrum prey of the Neotropical bat , Vampyrum specspectrum , in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:carnivorous 469 78. Goodwin , G

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**

ๆะ

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. ร * 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. อ*1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, อ*L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). . . Res. Rept .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear 8ด As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool. 47:140 15. , อ*' Stiles , F. G .; and Bradbury, J . Casebeer, R . อ*' Linsky, B . อ*' and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group , Vampyrum spectrum (VampiroFalso times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight FalseVampireBat) singly . toforage appear ห , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

5๓

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group (VampiroFalso , times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight FalseVampireBat) singly . toforage appear 8ด , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. - , 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร, 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, ร, L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). . . Res . Rept .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am . Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool . 47:140 15. , ร,‘ Stiles , F. G .; and Bradbury, J . Casebeer, R . ร,‘ Linsky, B . ร,‘ and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrumof the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

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-

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but March. ๖ ฺ8๑ problem continues. BaiTett , o. 1935. Notes concerning manatees and ด 16:216-20. dugongs. 8ด Frantzius, A . von. 1869. Die saugethiere Costaricas. ด ( Berlin ) 35:247-325. ด ๑ ไ ์ Hartman , D. 1979. ิ ้ Publica ด Special ๖ ๑ tion no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee ้

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals ิ ด ด ด separated from their group , for example , for foraging , ด ด ด are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ไ such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds , feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufvampire. One reported that individuals of fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ิdo not fan their seems a distinct possibility. It is clear that these bats have ฺand ฺ pires ้ complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees , with the exception of ฺwhich ๖may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. 1% 1. A review for fear that some small museum will feel the need bats of Trinidad and Tobago. M. ด ด . ด complete its skin collection at the expense of the remnantto of the 122 : 191 301 . populations. Fieldworkers are encouraged to observe and Greenhall , A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. 8ด ด49:337-40. of the and collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1%9. Notes on ไ ฺthe false 8ด ด47:140 15. vampire bat , in Panama. ด , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , and of the Neotropical carnivorous ฺin Costa Rica. 8 ไ ด ด44:186-89. ด 8ด ด58:469 78. bat ,

-

-

-

**

iN

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. ๐11935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv. Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ป11979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ป1L. 1977. The Wfest Indian manatee (Trichechus Fish Wildl . Serv.t Wildl . Res . Rept . manatus ) . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrumspectrum( Vampiro Falso, False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vampyrum spectrum , the false vampire bat , in Panama. Can. J . Zool . 47:140 15. ,ป 18Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . ป18Linsky, B . ป18and Nelson , c. 1%3. The Vehrencamp on the foraging behavior and avian prey phyllostomid bats , Ectophylla alba and Vampyrum ofObservations the Neotropical bat , Vampyrum specspectrum, in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:carnivorous 469-78.

-

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501

MAMMALS

ดด

ฺ์ ๑ ฺ ฺ ๑ไ ์ ไ ์ , 0 ด Tampa Tribune ด ๑ ๑ ด ์ ๖ ์ ฺด ด ฺ์ ด . ฐไ ๊ด ด J. Mamm ด ๑ด ฺ หด ด ด ๑ ฺ๑ ๑ ๖ ๑ ์ “ด « . ้ ิ ฐไ . Naturgesch Archiv ๑๑ ๑ ์ ด ofthemana ฺ andbehavior Ecology ฟด ด ๑ ๖ ์ ) inFlorida . manatus tee(Trichechus ไ๖ ๑ ด ด Vampyrum๖ ด ด « ด ๖ (Trichechus ๑ ฺ็ ด ด๑ ฺ ฺ ๑ ฺ ์ ). . Res. Rept. .t Wildl . Serv FishWildl manatus ๑ ๑ ์ ด “ ฐ ไ 5 ด ห ์ ์ ด Review ๑ ± ฺ 8 ด ด ด๑ ช ฺด ! ด ๑ ์ ๖ ไ ๑ ๑ ๑ ๖ ด ๖ ( Vampiro Falso , Vampyrum spectrum ๑ ์ ์ ฺ ๑ ๑ ด๑ False Vampire Bat )

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๑ ๑ ด Trachops ์ Chrotopterus ฺ์ ๑ ๑ ์ ด Vampyrum ฺ ฺ ๖ Vampyrum ๑ ฺ์ ด ๑ Vampyrum๑ ฺ์ ๑ ๖ ์ ์ ๑ ไ ๑ ๑ด ๖ ๑ ±๑ ๑ ๑ ด ๑ ฺ ์ ๑ ฺ ์ ๖ ๑ ด ด ๖ ฺ ด ๖ ๑ ๑ ๖ ๑ ด ์ ๑ ์ ๑ ๖ 9 9 ๑ ์ ๑ ์ ไ Vampyrum ๑ ๑ ฐ ๑๑ ด๑๑ ๑ ๑ ๑ ฺ์ ๑ ์ ]๑ ์๑ ๑ ์ ด ๑ ด ้ ์๑ ไ ิ ฺ ๑ ๑ ไ ๑ฺ ) ๑ (Desmodus ด ๖ , ์ , Diaemus Diphylla ์ ์ ์ ด ๑ Vampyrum ด spectrum ๖ ๑ ๑ ด ๑ Vampyrum ๖ Saccopteryx ด ์ ๑ฺ ๑ ๑ , ๖ ๑ ด๑ ด๑ bilineata ๑ ์ ฺ์ ฺ ์ ด๑ ๑ ์ ฺ ๖ ๑ ์ ด ๑ ๑ ์๑ ๑

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Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. . - 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior -1979. Ecology Hartman , D. — for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. a wide variety of social signals. Animals -L. 1977. The Wfest Indian manatee (Trichechusdemonstrated Husar, — their group , for example , for foraging, from separated ). ร (พ( FishWildl . Res. Rept. .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. ,— -w Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . — -w Linsky, B . — -w and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. ๐11935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ป11979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ป1L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). ร (พ( FishWildl . Res. Rept. .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. ,ป 18Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . ป18Linsky, B . ป18and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , Ectophylla alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

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-

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup, one half-grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett, o. ร * 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remainstable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbedin the roost, the whole groupemits a soft . ( Berlin) 35:247-325. . Naturgesch Archiv is the spe - chitteringsound. This noise characteristicof ofthemana andbehavior Hartman, D. อ*1979. Ecology for the presence marker a good and constitutes of ) inFlorida . Special Publica- cies manatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have . tion no. 5. American Society of Mammalogists . Animals Husar, อ*L. 1977. The Wfest Indian manatee(Trichechusdemonstrateda wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus roost an by the to on their return interlocking are greeted 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats ) False Vampire Bat to forage singly. appear 8ด As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost litteranalyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds - such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in groupsat night. Some of the species takenmatchthe bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can some favors substantialvertebratessuch as birds, feed every night, that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitativelyinsuf fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. Vampyrum spectrum Vampyrum occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spe tropics but is never dense. This is presumablya function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

-

two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull. Am. Mus . Nat. Hist 122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat, in . Panama Can.J. Zool. 47:140 15. , อ*' Stiles, F. G.; and Bradbury , J. 1977. Casebeer, R. อ*' Linsky, B . อ*' and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behavior phyllostomid bats, Ectophylla and avian prey albaand Vampyrum of the Neotropical carnivorousbat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. - , 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ร, 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร, L. 1977. The Wfest Indian manatee (Trichechus manatus ) . ร (พ( Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. ( Vampiro Falso,

False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , ร,‘ Stiles , F. G .; and Bradbury, J . Casebeer, R . ร,‘ Linsky, B . ร,‘ and Nelson , c. 1%3. The Vehrencamp 1977. Observations on the foraging behavior phyllostomid bats , Ectophylla alba and Vampyrum and avian prey of the Neotropical carnivorous bat , Vampyrum spec spectrum , in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:469-78.

-

**

501

MAMMALS

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Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee (Trichechus manatus ) . บ .ร . Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. ( Vampiro Falso,

False Vampire Bat) ห

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior phyllostomid bats , Ectophylla alba and Vampyrum and avian prey of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J . Mamm . 44:186-89. spectrum . J . Mamm . 58:469-78.

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Vampyrum ฺ์

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ฺ จ ฺ์ ด จด ฺ หด สด 5! 5ด ห ๑ ์ ด . Am Bull . ด Mus Nat. ๑ Hist ฌ ฐ5 5 ด ด ๑ ฺ หด สด 5 ๑ 0 ! ด ์ ๑ ๑ ๑ ๑ฺ ๑ ๑ ด J . Mamm . ฐ“ ์๑ ฺ ด ไ วด็ ฺ ์ ฺด ด ์ 5! ด ฝ ด Vam pyrumspectrum , ๑ ๑ ๑ ฺ ด Can.J. Zool. ฐ" ด ฺ ฺ ด ±์ อ จด ฺ ฺ วด 8 ด ด ฺ ๐ด ์ ฺ ] ด 5 ! ํ ด ๑ ๑ ์ ฺ Ectophylla alba ์ Vampyrum ๑ ฺ spectrum Vampyrum , ๖ด spec J . Mamm . ฐไ ๊ด trum. J . Mamm . ฐฝด

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ฺ ๑ ๑ ด ์ ๑ ์ ๖ 9 9 ๑ ์ ๑ ์ ไ Vampyrum ๑ ๑ ฐ ๑๑ ด๑๑ ๑ ๑ ๑ ฺ์ ๑ ์ ]๑ ์๑ ๑ ์ ด ๑ ด ้ ์๑ ไ ิ ฺ ๑ ๑ ไ ๑ฺ ) ๑ (Desmodus ด ๖ , ์ , Diaemus Diphylla ์ ์ ์ ด ๑ Vampyrum ด spectrum ๖ ๑ ๑ ด ๑ Vampyrum ๖ Saccopteryx ด ์ ๑ฺ ๑ ๑ , ๖ ๑ ด๑ ด๑ bilineata ๑ ์ ฺ์ ฺ ์ ด๑ ๑ ์ ฺ ๖ ๑ ์ ด ๑ ๑ ์๑ ๑

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup, one half-grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett, o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remainstable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbedin the roost, the whole groupemits a soft . ( Berlin) 35:247-325. . Naturgesch Archiv is the spe - chitteringsound. This noise characteristicof ofthemana andbehavior Hartman, D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrateda wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee(Trichechus from their group, for example, for foraging, separated ). 18๓8FishWildl . . Res . Rept .t Wildl . Serv manatus are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) appearto forage singly. As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost litteranalyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds - such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in groupsat night. Some of the species takenmatchthe bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can some favors substantialvertebratessuch as birds, feed every night, that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains in the floor of the roost: these are quantitativelyinsuf vampire. One reported that individuals of Vampyrum fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. Vampyrum spectrum Vampyrum occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spe tropics but is never dense. This is presumablya function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

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two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull. Am . Mus . Nat. Hist122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat, in . Panama Can . J. Zool . 47:140 15. , Stiles, F. G.; and Bradbury, J. 1977. Casebeer, R. Linsky, B . and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian prey phyllostomid bats, Ectophylla albaand Vampyrumof the Neotropical carnivorousbat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

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iN

501

MAMMALS

, andonenearly adult , but pup male , onehalf andperforms eats Snooks . Baby -grown . 1973 Anonymous -sized , 8 March . Tribune . Tampa continues thebats Intheroost problem . hadnotyetbred form female that a and contact manatees , o. . Notesconcerning 1935 BaiTett , and social tobehighly . Theyappear cluster . :216 . 16 . J. Mamm dugongs -20 forlongperiods stable mayremain group composition . . When Costaricas saugethiere , A. von . 1869. Die Frantzius , thewhole asoft group emits the roost in disturbed ) 35:247 . (Berlin . Naturgesch Archiv -325.ofthemana ofthespe ischaracteristic noise sound . This - chittering , D. 1979 . Ecology behavior and Hartman for the presence marker a good and constitutes cies of ) inFlorida Publica . Special manatus tee(Trichechus t ree groups Captive . h ollow in given a Vampyrum have . ofMammalogists Society tionno. 5. American . Animals signals ofsocial a widevariety demonstrated (Trichechus manatee , L. 1977 . TheWfestIndian Husar , , for example for group their foraging , from separated ). . . Res . Rept .t Wildl . Serv FishWildl manatus byaninterlocking totheroost ontheir return aregreeted 7:1-22. ; ofmouths . A rapid inwolves high .Review likethatfound manatee much , J. E. 1979 . Thesemisocial Reynolds , pp wings . 93-%. June oftheclosed isa byvibration accompanied piping thewhole to agitate behavior thatseems contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, Vampyrum spectrum times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear ห , these . Mostrebatsarecarnivorous above Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld isthelargest that Vampyrum spectrum , rodents , andother arethemajor ofupto200 birds withadult bats weights animal Itisanimposing constituents maybetaken . Somefruitandinsects isthe of thediet 80cm. The species on approaching ganda wingspan , butthisappears . Onelong a occasion batsthatshare toberare ofphyllostominine -term inaseries extreme . study prey back toa roost biasforlarger ofpreybrought ontheremains anda dietary in physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae complete large Allhave , andtrogons , parakeets , motmots , doves - suchasanis . forms suchasMicro . Smaller noseleaves elongate orroost fairly smelly , intermediate in Mostofthesepreyareeither fruit plus orinsects nycteris feedoninsects taken match thebats ofthespecies atnight in groups . Some mixlarger andChrotopterus forms suchasTrachops thatnotallbats , andthelarge inweight suggested study . Thissame aslizards such withsmall vertebrates sects Vampyrum , feedevery , thata foraging cansome night suchasbirds favors substantial vertebrates Vampyrum leaving after arestrik times within the anhour fillitsstomach , andother . ThejawsofVampyrum bats rodents atgreat forage , and donotnecessarily roost thatthebats withlarge canine teeth anda rowofcar endowed ingly the although that ;theskull .Italsoshowed from theroost distances apronounced nassiallike molars bears shearing inthe wasleftbehind group pupinthestudy . Weresuckling jawmuscles thepowerful crest toanchor sagittal orsubadult “guarded byoneadult , itwasalways easily betakenroost a noseleaf, thisbatcould itnottohave by ofprovisioning evidence ofthegroup . Some form member . doginhead forasmall , inTrinidad wasobserved . Early pair explorers ofacaptive name is a misnomer theadult male Thescientific . It behavior batssimply assumed thatthe andthismaybea common inthisspecies talesof vampire hearing remains ofprey forthepresence certainly be“the** would must account species largest andmostfierce -looking intheflooroftheroost insuf of Vampyrum : these thatindividuals . Onereported vampire arequantitatively , and ficient ofallthebatsin needs withtheir fortheenergy toaccount large wings fanned their victims tosleep some enough butaresubstantial towarrant struc thegroup leafandassociated another thenose that suggested to back arebrought blood notallprey eaten . Since . Both explanation usedtoremove devices weresuction tures sitter (and , provisioning , since vam thebaby ofyoungsters thetrue speculation clearly were suggestions - ) - theroost have ) these bats that (Desmodus . Itisclear donotfantheirseems adistinct , and , Diaemus Diphylla possibility pires with many parallels family livesthatmayshow complex victims andlapblood instead ofsuckit. ofuptofiveinthe other roosts Vampyrum ingroups . spectrum carnivores suchascanids mammalian oftheAmerican occurs overa widerange ofhollow batspe Vampyrum other . Most upper dark reaches trees , withtheexception butisnever afunction .Thisispresumably tropics ofSaccopteryx ciesavoid these dense trees dispersion , which . Thissparse carnivore asa large mayroostnearthetreeopening . The ofitsstatus bilineata skin years bymuseum of hasbeenexacerbated toconsist appear inrecent captured been thathave fewgroups who biologists , , field careless and . Onesuchgroup young in collectors pair oneadult andtheir recent petfanciers are Rica , onesucklingwillnotleave andCosta . Trinidad done thebats of thetwoadults consisted Guanacaste

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SPECIES ACCOUNTS

inwhich twosites Vampyrum roosts arenowkeptsecretGoodwin , G. G., and , A M 1% Greenhall 1. Areview forfearthatsome small museum willfeeltheneedto ofthebats of Trinidad and T obago . . Am Bull . Mus .Nat. complete itsskin collection attheexpense oftheremnantHist122:191301 . . Fieldworkers populations areencouraged , A. M. 1% toobserve 8. Notes and Greenhall onthebehavior ofthe enjoy these large bats , butunnecessary falsevampire . J. Mamm bat . 49 disturbance : and 337 40 . collecting should , R. L., andKirmse Peterson beavoided whenever , p. 1% possible . 9. Notes onVam pyrum spectrum , thefalse vampire , bat in . Panama Can . J. Zool . 47 :14015. , Vehrencamp , F. G.; andBradbury Stiles , R. Linsky , J. 1977 Casebeer , B. and . , . 1% N elson 3 . The Observations on the f oraging behavior phyllostomid and , Ectophylla a vian bats prey alba and Vampyrumof theNeotropical carnivorous , Vampyrum bat spectrum , inCosta spec Rica . J. Mamm . 44 :186 -89. trum .

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J. Mamm . 58 :46978.

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but March. ๖ ฺ8๑ problem continues. BaiTett , o. 1935. Notes concerning manatees and ด 16:216-20. dugongs. 8ด Frantzius, A . von. 1869. Die saugethiere Costaricas. ด ( Berlin ) 35:247-325. ด ๑ ไ ์ Hartman , D. 1979. ิ ้ Publica ด Special ๖ ๑ tion no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee ้

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals ิ ด ด ด separated from their group , for example , for foraging , ด ด ด ป๙๐๙ are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, Vampyrum spectrum screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear หว As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ไ such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds , feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufvampire. One reported that individuals of fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ิdo not fan their seems a distinct possibility. It is clear that these bats have ฺand ฺ pires ้ complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees , with the exception of ฺwhich ๖may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. 1% 1. A review for fear that some small museum will feel the need bats of Trinidad and Tobago. M. ด ด . ด complete its skin collection at the expense of the remnantto of the 122 : 191 301 . populations. Fieldworkers are encouraged to observe and Greenhall , A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. 8ด ด49:337-40. of the and collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1%9. Notes on ไ ฺthe false 8ด ด47:140 15. vampire bat , in Panama. ด , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , and of the Neotropical carnivorous ฺin Costa Rica. 8 ไ ด ด44:186-89. ด 8ด ด58:469 78. bat ,

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501

MAMMALS

SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group (VampiroFalso , times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight FalseVampireBat) singly . toforage appear J . Bradbury , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrumspectrum( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but March. ๖ ฺ8๑ problem continues. BaiTett , o. 1935. Notes concerning manatees and ด 16:216-20. dugongs. 8ด Frantzius, A . von. 1869. Die saugethiere Costaricas. ด ( Berlin ) 35:247-325. ด ๑ ไ ์ Hartman , D. 1979. ิ ้ Publica ด Special ๖ ๑ tion no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee ้

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals ิ ด ด ด separated from their group , for example , for foraging , ด ด ด are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ไ such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds , feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufvampire. One reported that individuals of fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ิdo not fan their seems a distinct possibility. It is clear that these bats have ฺand ฺ pires ้ complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees , with the exception of ฺwhich ๖may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. 1% 1. A review for fear that some small museum will feel the need bats of Trinidad and Tobago. M. ด ด . ด complete its skin collection at the expense of the remnantto of the 122 : 191 301 . populations. Fieldworkers are encouraged to observe and Greenhall , A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. 8ด ด49:337-40. of the and collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1%9. Notes on ไ ฺthe false 8ด ด47:140 15. vampire bat , in Panama. ด , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , and of the Neotropical carnivorous ฺin Costa Rica. 8 ไ ด ด44:186-89. ด 8ด ด58:469 78. bat ,

-

-

-

**

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup, one half-grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a 21935. Notes concerning manatees and contact cluster. They appear to be highly social, and BaiTett, o. ๓ . 16:216-20. dugongs. J. Mamm group composition may remainstable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbedin the roost, the whole groupemits a soft . ( Berlin) 35:247-325. . Naturgesch Archiv is the spe - chitteringsound. This noise characteristicof ofthemana andbehavior Hartman, D. 121979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrateda wide variety of social signals. Animals Husar, 12L. 1977. The Wfest Indian manatee(Trichechus from their group, for example, for foraging, separated ). . . Res. Rept .t Wildl . Serv FishWildl manatus are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) to forage singly. appear หว As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost litteranalyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds - such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in groupsat night. Some of the species takenmatchthe bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can some favors substantialvertebratessuch as birds, feed every night, that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains in the floor of the roost: these are quantitativelyinsuf vampire. One reported that individuals of Vampyrum fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. Vampyrum spectrum Vampyrum occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spe tropics but is never dense. This is presumablya function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

-

two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull. Am. Mus . Nat. Hist122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat, in . Panama Can . J. Zool. 47:140 15. ,1 9Stiles, F. G.; and Bradbury, J. 1977. 2 Casebeer, R. 12 9 Linsky, B . 12 9 and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian prey phyllostomid bats, Ectophylla albaand Vampyrum of the Neotropical carnivorousbat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but ้8 March. problem continues. BaiTett, o. 1935. Notes concerning manatees and ว 16:216-20. dugongs. หว Frantzius, A . von. 1869. Die saugethiere Costaricas. ว ( Berlin) 35:247-325. ว ํ Hartman, D. 1979. : ’ วSpecial Publication no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee’

pup, one half-grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social, and group composition may remainstable for long periods. When disturbedin the roost, the whole groupemits a soft chitteringsound. This noise is characteristicof the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrateda wide variety of social signals. Animals : ว ว ว separated from their group, for example, for foraging, ว ว ว 18๓8 are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost-litteranalyses agree that It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds ํ such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in- groupsat night. Some of the species takenmatchthe bats and forms such as sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can somefavors substantialvertebratessuch as birds, feed every night, that a foraging are strik- times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains in the floor of the roost: these are quantitativelyinsufvampire. One reported that individuals of fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc- the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) :do not fan their seems a distinct possibility. It is clear that these bats have ้and ้ pires ’ complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spetropics but is never dense. This is presumablya function cies avoid these trees, with the exception of ้which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts arenow kept secret Goodwin, G. G. , and Greenhall, A. 1%1. A review for fear that some small museum will feel the need of the batsof Trinidadand Tobago. M. ว to ว . ว complete itsskin collection at the expense of the remnant :191 301 . 122 populations. Fieldworkersareencouragedto observe and Greenhall, A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessarydisturbance false vampire bat. หว ว49:337-40. of the and collecting should be avoided whenever possible. Peterson, R. L. , and Kirmse, p. 1%9. Notes on ํ ้the false หว ว47:140 15. vampire bat, in Panama. ว , Stiles, F. G.; and Bradbury, J. 1977. Casebeer, R. Linsky, B . and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian prey phyllostomid bats, and of the Neotropical carnivorous ้in Costa Rica. ห ํ ว ว44:186-89. ว หว ว58:469 78. bat,

-

-

-

**

บ'

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). . . Res. Rept .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear หว As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

ดด

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MAMMALS

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ฺ์ ๑ ฺ ฺ ๑ไ ์ ไ ์ Tampa Tribune, 0 ด ด ๑ ๑ ด ์ ๖ ์ ฺด ด ฺ์ ด ๊ด ด J . Mamm. ฐไ ด ๑ด ฺ หด ด ด ๑ ฺ๑ ๑ ๖ ๑ ์ “ด « Archiv. Naturgesch. ้ ิ ฐ ไ ๑๑ ๑ ์ ด ฺ Ecology and behavior of the manaฟด ด ๑ ๖ ์ tee (Trichechus manatus ) in Florida . ไ๖ ๑ ด ด Vampyrum ๖ ด ด « ด ๖ (Trichechus ๑ ฺ็ ด ด๑ ฺ ฺ ๑ ฺ ์ manatus ) . อ 8ร 8 Fish Wildl. Serv.t Wildl. Res. Rept. ๑ ๑ ์ ด “ ฐ ไ 5 ด ห ์ ์ ด Review ๑ ± ฺ 8 ด ด ด๑ ช ฺด ! ด ๑ ์ ๖ ไ ๑ ๑ ๑ ๖ ด ๖ ( Vampiro Falso , ๑ ์ ์ ฺ ๑ ๑ ด๑ False Vampire Bat )

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ฺ จ ฺ์ ด จด ฺ หด สด 5! 5ด ห ๑ ์ ด . Am. Musด Bull Nat . ๑ Hist ฌ ฐ5 5 ด ด ๑ ฺ หด สด 5 ๑ 0 ! ด ์ ๑ ๑ ๑ ๑ฺ ๑ ๑ ด J . Mamm. ฐ“ ์๑ ฺ ด ไ วด็ ฺ ์ ฺด ด ์ 5! ด ฝ ด Vampyrum spectrum, ๑ ๑ ๑ ฺ ด Can. J . Zool . ฐ" ด ฺ ฺ ด ±์ อ จด ฺ ฺ วด 8 ด ด ฺ ๐ด ์ ฺ ] ด 5 ! ํ ด ๑ ๑ ์ ฺ Ectophylla alba ์ Vampyrum ๑ spectrum, Vampyrum spec๖ด J . Mamm. ฐไ ๊ด trum. J . Mamm. ฐฝด ฺ

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Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

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two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

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MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup, one half-grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett, o. ร * 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remainstable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbedin the roost, the whole groupemits a soft . ( Berlin) 35:247-325. . Naturgesch Archiv is the spe - chitteringsound. This noise characteristicof ofthemana andbehavior Hartman, D. อ*1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, อ*L. 1977. The Wfest Indian manatee(Trichechusdemonstrateda wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) appearto forage singly. As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost litteranalyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds - such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in groupsat night. Some of the species takenmatchthe bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can some favors substantialvertebratessuch as birds, feed every night, that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitativelyinsuf fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. Vampyrum spectrum Vampyrum occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spe tropics but is never dense. This is presumablya function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

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two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull. Am. Mus . Nat. Hist 122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat, in . Panama Can.J. Zool. 47:140 15. , อ*' Stiles, F. G.; and Bradbury , J. 1977. Casebeer, R. อ*' Linsky, B . อ*' and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian phyllostomid bats, Ectophylla prey alba and Vampyrum of the Neotropical carnivorousbat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

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Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). อ 8ร 8 FishWildl . Res. Rept. .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, Vampyrum spectrum screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear หว As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. พ. 1935 . Notesconcerning and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. ร. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , ร.; Stiles Vehrencamp (Trichechus , F. G.; andBradbury manatee , ร. L. 1977 . TheWfestIndian Husar , R. ร.; Linsky , J. 1977 Casebeer , B. ร.; andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group Vampyrum spectrum ( Vampiro Falso, times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup, one half-grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett, o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remainstable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbedin the roost, the whole groupemits a soft . ( Berlin) 35:247-325. . Naturgesch Archiv is the spe - chitteringsound. This noise characteristicof ofthemana andbehavior Hartman, D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrateda wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee(Trichechus from their group, for example, for foraging, separated ). อ 8ร 8 FishWildl . . Res . Rept .t Wildl . Serv manatus are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost litteranalyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds - such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in groupsat night. Some of the species takenmatchthe bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can some favors substantialvertebratessuch as birds, feed every night, that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains in the floor of the roost: these are quantitativelyinsuf vampire. One reported that individuals of Vampyrum fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. Vampyrum spectrum Vampyrum occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spe tropics but is never dense. This is presumablya function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

-

two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull. Am . Mus . Nat. Hist122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat, in . Panama Can . J. Zool . 47:140 15. , Stiles, F. G.; and Bradbury, J. 1977. Casebeer, R. Linsky, B . and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian prey phyllostomid bats, Ectophylla albaand Vampyrumof the Neotropical carnivorousbat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

iN

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. ร / 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv. Naturgesch . ( Berlin ) 35:247-325. Hartman , D. บ/ 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, บ/ L. 1977. The Wfest Indian manatee (Trichechus manatus ) . ร (พ( Fish Wildl . Serv.t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrum spectrum ( Vampiro Falso, False Vampire Bat)

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vampyrum spectrum , the false vampire bat , in Panama. Can. J . Zool . 47:140 15. , บ/ี Stiles , F. G .; and Bradbury, J . Casebeer, R . บ/ี Linsky, B . บ/ี and Nelson , c. 1%3. The Vehrencamp . on the foraging behavior and avian1977 phyllostomid bats , Ectophylla alba and Vampyrum ofObservations prey the Neotropical bat , Vampyrum specspectrum, in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:carnivorous 469-78.

-

**

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but March. ๖ ฺ8๑ problem continues. BaiTett , o. 1935. Notes concerning manatees and ด 16:216-20. dugongs. 8ด Frantzius, A . von. 1869. Die saugethiere Costaricas. ด ( Berlin ) 35:247-325. ด ๑ ไ ์ Hartman , D. 1979. ิ ้ Publica ด Special ๖ ๑ tion no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee ้

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals ิ ด ด ด separated from their group , for example , for foraging , ด ด ด อ 8ร 8 are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( VampiroFalso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats FalseVampireBat) to forage singly. appear ห As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ไ such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds , feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufvampire. One reported that individuals of fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ิdo not fan their seems a distinct possibility. It is clear that these bats have ฺand ฺ pires ้ complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees , with the exception of ฺwhich ๖may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIESACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. 1% 1. A review for fear that some small museum will feel the need bats of Trinidad and Tobago. M. ด ด . ด complete its skin collection at the expense of the remnantto of the 122 : 191 301 . populations. Fieldworkers are encouraged to observe and Greenhall , A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. 8ด ด49:337-40. of the and collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1%9. Notes on ไ ฺthe false 8ด ด47:140 15. vampire bat , in Panama. ด , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , and of the Neotropical carnivorous ฺin Costa Rica. 8 ไ ด ด44:186-89. ด 8ด ด58:469 78. bat ,

-

-

-

**

501

MAMMALS

ดด

ฺ์ ๑ ฺ ฺ ๑ไ ์ ไ ์ , 0 ด Tampa Tribune ด ๑ ๑ ด ์ ๖ ์ ฺด ด ฺ์ ด . ฐไ ๊ด ด J. Mamm ด ๑ด ฺ หด ด ด ๑ ฺ๑ ๑ ๖ ๑ ์ “ด « . ้ ิ ฐไ . Naturgesch Archiv ๑๑ ๑ ์ ด ofthemana ฺ andbehavior Ecology ฟด ด ๑ ๖ ์ ) inFlorida . manatus tee(Trichechus ไ๖ ๑ ด ด Vampyrum๖ ด ด « ด ๖ (Trichechus ๑ ฺ็ ด ด๑ ฺ ฺ ๑ ฺ ์ ). . Res. Rept. .t Wildl . Serv FishWildl manatus ๑ ๑ ์ ด “ ฐ ไ 5 ด ห ์ ์ ด Review ๑ ± ฺ 8 ด ด ด๑ ช ฺด ! ด ๑ ์ ๖ ไ ๑ ๑ ๑ ๖ ด ๖ Vampyrum spectrum ( Vampiro Falso , ๑ ์ ์ ฺ ๑ ๑ ด๑ False Vampire Bat )

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Vampyrum spectrum ๑ ๖ ์

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Micro -

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ฺ จ ฺ์ ด จด ฺ หด สด 5! 5ด ห ๑ ์ ด . Am Bull . ด Mus Nat. ๑ Hist ฌ ฐ5 5 ด ด ๑ ฺ หด สด 5 ๑ 0 ! ด ์ ๑ ๑ ๑ ๑ฺ ๑ ๑ ด J. Mamm . ฐ“ ์๑ ฺ ด ไ วด็ ฺ ์ ฺด ด ์ 5! ด ฝ ด Vampyrumspectrum , ๑ ๑ ๑ ฺ ด Can.J. Zool. ฐ" ด ฺ ฺ ด ±์ อ จด ฺ ฺ วด 8 ด ด ฺ ๐ด ์ ฺ ] ด 5 ! ํ ด ๑ ๑ ์ ฺ Ectophylla alba ์ Vampyrum ๑ ฺ spectrum Vampyrum , ๖ด spec J. Mamm . ฐไ ๊ด trum. J. Mamm . ฐฝด

Vampyrum

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๑ ๑ ด Trachops ์ Chrotopterus ฺ์ ๑ ๑ ์ ด Vampyrum ฺ ฺ ๖ Vampyrum ๑ ฺ์ ด ๑ Vampyrum๑ ฺ์ ๑ ๖ ์ ์ ๑ ไ ๑ ๑ด ๖ ๑ ±๑ ๑ ๑ ด ๑ ฺ ์ ๑ ฺ ์ ๖ ๑ ด ด ๖ ฺ ด ๖ ๑ ๑ ๖ ๑ ด ์ ๑ ์ ๑ ๖ 9 9 ๑ ์ ๑ ์ ไ Vampyrum ๑ ๑ ฐ ๑๑ ด๑๑ ๑ ๑ ๑ ฺ์ ๑ ์ ]๑ ์๑ ๑ ์ ด ๑ ด ้ ์๑ ไ ิ ฺ ๑ ๑ ไ ๑ฺ ) ๑ (Desmodus ด ๖ , ์ , Diaemus Diphylla ์ ์ ์ ด ๑ Vampyrum ด spectrum ๖ ๑ ๑ ด ๑ Vampyrum ๖ Saccopteryx ด ์ ๑ฺ ๑ ๑ , ๖ ๑ ด๑ ด๑ bilineata ๑ ์ ฺ์ ฺ ์ ด๑ ๑ ์ ฺ ๖ ๑ ์ ด ๑ ๑ ์๑ ๑

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. ๐11935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ป11979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ป1L. 1977. The Wfest Indian manatee (Trichechus manatus ) . ร (พ( Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrum spectrum ( Vampiro Falso, False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. ,ป 18Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . ป18Linsky, B . ป18and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior phyllostomid bats , Ectophylla alba and Vampyrum and avian prey of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J . Mamm . 44:186-89. spectrum . J . Mamm . 58:469-78.

-

**

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. พ. 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ร. 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus manatus ) . ร ท- ท Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. ( Vampiro Falso, Vampyrum spectrum False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp 1977. Observations on the foraging behavior phyllostomid bats , Ectophylla alba and Vampyrum and avian prey of the Neotropical carnivorous bat , Vampyrum spec spectrum , in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:469-78.

-

**

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv. Naturgesch . ( Berlin ) 35:247-325. Hartman , D. 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee (Trichechus manatus ) . บ .ร . Fish Wildl . Serv.t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. ( Vampiro Falso,

False Vampire Bat)

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vampyrum spectrum , the false vampire bat , in Panama. Can. J . Zool . 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp on the foraging behavior phyllostomid bats , Ectophylla alba and Vampyrum ofObservations and avian prey the Neotropical carnivorous bat , Vampyrum spectrum, in Costa Rica. J . Mamm . 44:186-89. spectrum . J . Mamm . 58:469-78.

-

**

501

MAMMALS

SPECIES ACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich V ampyrum roosts are n ow kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. A review female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. พ. 1935 . Notesconcerning and BaiTett , and Tobago . Bull . Am. Mus.Nat . social to be highly . Theyappear cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group composition . . Fieldworkers populations areencouraged , A. M. 1%8. Notesonthebehavior toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius , thewhole asoft group emits of the the roost in d isturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the noise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. ร. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . hollow in given a Vampyrum have Can . J . Zool . 47:14015. . of Mammalogists Society tionno. 5. American . signals ofsocial variety a wide demonstrated Animals , ร.; Stiles Vehrencamp (Trichechus , F. G.; andBradbury manatee , ร. L. 1977 . TheWfestIndian Husar , R. ร.; Linsky , J. 1977 Casebeer , B. ร.; andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical byaninterlocking spectrum , inCosta totheroost ontheirreturn aregreeted 7:1-22. bat, Vampyrum specRica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapid high .Review; ofmouths manatee muchlikethatfoundinwolves , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, timeswhile itseveral astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear J . Bradbury , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andother ofupto200 birds withadult batsarethemajor weights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term a occasion batsthatshare to be rare of phyllostominine inaseries extreme . studyontheremains prey backtoa roostin biasforlarger of preybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles birds showed a heavybiasfornonpasserine pinnae complete Allhavelarge , andtrogons , parakeets , motmots , doves . formssuchasMicro- suchasanis . Smaller noseleaves elongate orroostin fairlysmelly , intermediate Mostof thesepreyareeither plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors withinanhourafterleaving the , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily roost withlarge canine teethanda rowofcar inglyendowed the thatalthough ;theskullbears .Italsoshowed fromtheroost distances apronounced nassiallike molars shearing in the wasleftbehind pupin thestudygroup . Were suckling jawmuscles thepowerful cresttoanchor sagittal orsubadult “guarded byoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . forasmalldogin headform , inTrinidad . Early pairwasobserved explorerstheadult ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient of allthebatsin needs withtheirlarge fortheenergy toaccount wings fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , provisioning , since ofyoungsters thetruevam- theroost speculation wereclearly suggestions -sitter) ) batshave thatthese ( Desmodus , Diaemus , andDiphylla donotfantheir seems . Itisclear adistinct possibility pires with family livesthatmayshowmanyparallels complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe other Vampyrum spectrum roosts ingroups . carnivores suchascanids mammalian of theAmerican overa widerange of hollow batspe Vampyrum occurs . Mostother darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The of itsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin yearsby museum of hasbeenexacerbated toconsist appear in recent thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup, one half-grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett, o. ร * 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remainstable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbedin the roost, the whole groupemits a soft . ( Berlin) 35:247-325. . Naturgesch Archiv is the spe - chitteringsound. This noise characteristicof ofthemana andbehavior Hartman, D. อ*1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, อ*L. 1977. The Wfest Indian manatee(Trichechusdemonstrateda wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, Vampyrum spectrum screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) to forage singly. appear 8ด As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost litteranalyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds - such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in groupsat night. Some of the species takenmatchthe bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can some favors substantialvertebratessuch as birds, feed every night, that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitativelyinsuf fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. Vampyrum spectrum Vampyrum occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spe tropics but is never dense. This is presumablya function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

-

two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull. Am. Mus . Nat. Hist 122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat, in . Panama Can.J. Zool. 47:140 15. , อ*' Stiles, F. G.; and Bradbury , J. 1977. Casebeer, R. อ*' Linsky, B . อ*' and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian phyllostomid bats, Ectophylla prey alba and Vampyrum of the Neotropical carnivorousbat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but March. ๖ ฺ8๑ problem continues. BaiTett , o. ๐11935. Notes concerning manatees and ด 16:216-20. dugongs. 8ด Frantzius, A . von. 1869. Die saugethiere Costaricas. ด ( Berlin ) 35:247-325. ด ๑ ไ ์ Hartman , D. ป11979. ิ ้ Publica ด Special ๖ ๑ tion no. 5. American Society of Mammalogists. Husar, ป1L. 1977. The Wfest Indian manatee ้

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals ิ ด ด ด separated from their group , for example , for foraging , ด ด ด are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ไ such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds , feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufvampire. One reported that individuals of fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ิdo not fan their seems a distinct possibility. It is clear that these bats have ฺand ฺ pires ้ complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees , with the exception of ฺwhich ๖may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. 1% 1. A review for fear that some small museum will feel the need bats of Trinidad and Tobago. M. ด ด . ด complete its skin collection at the expense of the remnantto of the 122 : 191 301 . populations. Fieldworkers are encouraged to observe and Greenhall , A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. 8ด ด49:337-40. of the and collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1%9. Notes on ไ ฺthe false 8ด ด47:140 15. vampire bat , in Panama. ด ,ป 18Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . ป18Linsky, B . ป18and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , and of the Neotropical carnivorous ฺin Costa Rica. 8 ไ ด ด44:186-89. ด 8ด ด58:469 78. bat ,

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**

501

MAMMALS

SPECIES ACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. ๐11935 . Notesconcerning and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. ป11979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals ,ป Vehrencamp 18Stiles (Trichechus , F. G.; andBradbury manatee , ป1L. 1977 . TheWfestIndian Husar , R. ป18Linsky , J. 1977 Casebeer , B. ป18andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the manatus ) . ร (พ( Fish Wildl . Serv.t Wildl . Res . Rept . separated foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but ้8 March. problem continues. BaiTett, o. 1935. Notes concerning manatees and ว 16:216-20. dugongs. หว Frantzius, A . von. 1869. Die saugethiere Costaricas. ว ( Berlin) 35:247-325. ว ํ Hartman, D. 1979. : ’ วSpecial Publication no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee’

pup, one half-grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social, and group composition may remainstable for long periods. When disturbedin the roost, the whole groupemits a soft chitteringsound. This noise is characteristicof the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrateda wide variety of social signals. Animals : ว ว ว separated from their group, for example, for foraging, ว ว ว 18๓8 are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost-litteranalyses agree that It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds ํ such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in- groupsat night. Some of the species takenmatchthe bats and forms such as sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can somefavors substantialvertebratessuch as birds, feed every night, that a foraging are strik- times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains in the floor of the roost: these are quantitativelyinsufvampire. One reported that individuals of fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc- the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) :do not fan their seems a distinct possibility. It is clear that these bats have ้and ้ pires ’ complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spetropics but is never dense. This is presumablya function cies avoid these trees, with the exception of ้which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

two sites in which

SPECIESACCOUNTS

roosts arenow kept secret Goodwin, G. G. , and Greenhall, A. 1%1. A review for fear that some small museum will feel the need of the batsof Trinidadand Tobago. M. ว to ว . ว complete itsskin collection at the expense of the remnant :191 301 . 122 populations. Fieldworkersareencouragedto observe and Greenhall, A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessarydisturbance false vampire bat. หว ว49:337-40. of the and collecting should be avoided whenever possible. Peterson, R. L. , and Kirmse, p. 1%9. Notes on ํ ้the false หว ว47:140 15. vampire bat, in Panama. ว , Stiles, F. G.; and Bradbury, J. 1977. Casebeer, R. Linsky, B . and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian prey phyllostomid bats, and of the Neotropical carnivorous ้in Costa Rica. ห ํ ว ว44:186-89. ว หว ว58:469 78. bat,

-

-

-

**

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but March. ๖ ฺ8๑ problem continues. BaiTett , o. 1935. Notes concerning manatees and ด 16:216-20. dugongs. 8ด Frantzius, A . von. 1869. Die saugethiere Costaricas. ด ( Berlin ) 35:247-325. ด ๑ ไ ์ Hartman , D. 1979. ิ ้ Publica ด Special ๖ ๑ tion no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee ้

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals ิ ด ด ด separated from their group , for example , for foraging , ด ด ด อ 8ร 8 are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ไ such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds , feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufvampire. One reported that individuals of fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ิdo not fan their seems a distinct possibility. It is clear that these bats have ฺand ฺ pires ้ complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees , with the exception of ฺwhich ๖may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. 1% 1. A review for fear that some small museum will feel the need bats of Trinidad and Tobago. M. ด ด . ด complete its skin collection at the expense of the remnantto of the 122 : 191 301 . populations. Fieldworkers are encouraged to observe and Greenhall , A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. 8ด ด49:337-40. of the and collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1%9. Notes on ไ ฺthe false 8ด ด47:140 15. vampire bat , in Panama. ด , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , and of the Neotropical carnivorous ฺin Costa Rica. 8 ไ ด ด44:186-89. ด 8ด ด58:469 78. bat ,

-

-

-

**

ๆะ

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett, o. - , 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin) 35:247-325. Hartman, D. ร, 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร, L. 1977. The Wfest Indian manatee(Trichechus manatus ) . อ 8ร 8 Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds, J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. ( Vampiro Falso,

False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae, canidlike muzzles, and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit, intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebratessuch as lizards, and the large Vampyrum favors substantialvertebratessuch as birds, rodents, and otherbats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallikeshearing molars; theskull bearsa pronounced sagittal crest to anchorthe powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form. The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “the vampire. One reported that individuals of Vampyrum fannedtheirvictims to sleep with their large wings, and anothersuggested that the nose leaf andassociatedstruc tures were suction devices used to remove blood. Both suggestions were clearly speculation, since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood insteadof suck it. Vampyrum spectrum roosts in groupsof up to five in the dark upper reachesof hollow trees. Most other bat spe cies avoid these trees, with the exception of Saccopteryx bilineata , which may roost near the tree opening. The few groups that have been capturedappearto consist of one adult pairand theirrecent young. One such group in Guanacaste consisted of the two adults, one suckling

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pup, one half-grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social, and group composition may remainstable for long periods. When disturbedin the roost, the whole groupemits a soft chitteringsound. This noise is characteristicof the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrateda wide variety of social signals. Animals separated from their group, for example, for foraging, are greeted on their returnto the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats appearto forage singly. As noted above, these bats are carnivorous. Most reportsof stomach, feces , or roost-litteranalyses agree that birds, rodents, and other bats are the majorconstituents of the diet. Some fruit and insects may be taken on occasion, but this appears to be rare. One long- term study on the remainsof prey brought back to a roost in Guanacasteshowed a heavy bias for nonpasserinebirds such as anis, doves, parakeets, motmots, and trogons. Most of these prey are either fairly smelly or roost in groupsat night. Some of the species takenmatchthe bats in weight. This same study suggested that not all bats feed every night, that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost, and that the bats do not necessarily forage at great distancesfromthe roost. It also showed that althoughthe suckling pup in the study group was left behind in the roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby theadultmale of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainlyaccountfor the presenceof prey remains in the floor of the roost: these are quantitativelyinsufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation. Since not all prey eaten are broughtback to the roost, provisioningof youngsters( andthe baby-sitter) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallelswith other mammaliancarnivoressuch as canids. Vampyrum occursover a wide range of the American tropics but is never dense. This is presumablya function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors, pet fanciers, and careless field biologists who will not leave the bats done. Trinidadand Costa Rica are

two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull . Am . Mus. Nat . Hist 122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat, in Panama. Can . J . Zool . 47:140 15. , ร,‘ Stiles, F. G.; and Bradbury , J. 1977. Casebeer, R. ร,‘ Linsky, B . ร,‘ and Nelson, c. 1%3. The Vehrencamp Observationson the foraging behavior phyllostomid bats, Ectophylla alba and Vampyrum and avian prey of the Neotropical carnivorousbat, Vampyrum spec spectrum , in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:469-78.

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**

501

MAMMALS

SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , Goodwin G. G., andGreenhall . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , but false v ampire unnecessary . bat J . . ) 35:247 325 Mamm Archiv. Naturgesch . (Berlin . disturbance : 49 and 337 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus Trichechus tee false v ampire , bat in . Panama tree groups . Captive Vampyrum in a givenhollow have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . Animals Casebeer ofsocialsignals a widevariety , Vehrencamp (Trichechus demonstrated , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 , B. andNelson . , c . 1 % 3 . , , for example The for g roup their foraging , from separated Observations on ) Rept the . Fish Wildl . Serv.t Wildl . Res . manatus . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group (VampiroFalso , times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight FalseVampireBat) singly . toforage appear 8ด , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing behind in the pupin thestudygroupwasleft . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific It . behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested back arebrought to blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction ( , , vam andthebaby ofyoungsters since thetrue - theroostprovisioning speculation wereclearly suggestions -sitter) have ) t hese b ats is that It . clear adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with show many parallels family livesthatmay complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion s parse , . The c arnivore This as status a large . ofits bilineata whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and g roup One . their s uch f anciers young in pair oneadult and recent collectors pet Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear yf As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. - , 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ร, 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร, L. 1977. The Wfest Indian manatee (Trichechus manatus ) . ป๙ ๐๙Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. ( Vampiro Falso,

False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

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-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , ร,‘ Stiles , F. G .; and Bradbury, J . Casebeer, R . ร,‘ Linsky, B . ร,‘ and Nelson , c. 1%3. The Vehrencamp 1977. Observations on the foraging behavior phyllostomid bats , Ectophylla alba and Vampyrum and avian prey of the Neotropical carnivorous bat , Vampyrum spec spectrum , in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:469-78.

-

**

501

MAMMALS

SPECIES ACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich roosts are n ow kept s ecret , G. G., andGreenhall Goodwin . 8 March . continues problem . Intheroostthebatsforma , A M 1%1. A review female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . social to be highly . Theyappear cluster . complete itsskincollection attheexpense oftheremnant :191301. 122 20. 16:216 . dugongs forlongperiods stable mayremain group composition . . populations Fieldworkers areencouraged , A. M. 1%8. Notesonthebehavior toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius , thewhole asoft group emits of the intheroost disturbed enjoytheselarge b ats , butunnecessary false v ampire bat. . ) 35:247 325 (Berlin disturbance : 49 and 337 40 . of i s the noise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . whenever , p. 1%9. Notes Hartman possible . on forthe presence marker a good and constitutes cies of Publica the Special falsevampire bat, in Panama . t ree groups Captive . hollow in given a have 47:14015. . of Mammalogists Society tionno. 5. American . signals ofsocial variety a wide demonstrated Animals , Vehrencamp , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup separated on the foraging behavior phyllostomid and , avian bats p rey and of theNeotropical byaninterlocking totheroost ontheirreturn aregreeted 7:1-22. carnivorous bat, inCosta Rica . 44:186 -89. . A rapid high . ; ofmouths manatee muchlikethatfoundinwolves 58:46978. , J. E. 1979 . Thesemisocial Reynolds , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, timeswhile itseveral astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld isthelargest that , rodents , andother ofupto200 birds withadult batsarethemajor weights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term a occasion batsthatshare to be rare of phyllostominine inaseries extreme . studyontheremains prey backtoa roostin biasforlarger of preybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles birds showed a heavybiasfornonpasserine pinnae complete Allhavelarge , andtrogons , parakeets , motmots , doves . formssuchas suchasanis . Smaller noseleaves elongate orroostin fairlysmelly , intermediate Mostof thesepreyareeither plusfruit orinsects feedoninsects taken match thebats atnight in groups . Someofthespecies mixlarger and forms suchas thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects , feedeverynight , thata foraging cansome suchasbirds favors substantial vertebrates arestrik timesfillitsstomach withinanhourafterleaving the , andother . Thejawsof bats rodents atgreat forage , andthatthebatsdonotnecessarily roost withlarge canine teethanda rowofcar inglyendowed the thatalthough ;theskullbears .Italsoshowed fromtheroost distances apronounced nassiallike molars shearing in the wasleftbehind pupin thestudygroup . Were suckling jawmuscles thepowerful cresttoanchor sagittal orsubadult “guarded byoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . forasmalldogin headform , inTrinidad . Early pairwasobserved explorerstheadult ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking in theflooroftheroost insuf of : thesearequantitatively thatindividuals . Onereported vampire , and ficient of allthebatsin needs withtheirlarge fortheenergy toaccount wings fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , provisioning , since ofyoungsters thetruevam- theroost speculation wereclearly suggestions -sitter) batshave thatthese . Itisclear donotfantheir seems adistinct possibility and pires with family livesthatmayshowmanyparallels complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe other roosts ingroups . carnivores suchascanids mammalian of theAmerican occurs overa widerange of hollow batspe . Mostother darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably tropics of ciesavoid thesetrees dense dispersion . Thissparse . The of itsstatusasa largecarnivore whichmayroostnearthetreeopening skin yearsby museum of hasbeenexacerbated toconsist appear in recent thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup, one half-grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett, o. ร * 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remainstable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbedin the roost, the whole groupemits a soft . ( Berlin) 35:247-325. . Naturgesch Archiv is the spe - chitteringsound. This noise characteristicof ofthemana andbehavior Hartman, D. อ*1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrateda wide variety of social signals. Animals Husar, อ*L. 1977. The Wfest Indian manatee(Trichechus from their group, for example, for foraging, separated ). . . Res. Rept .t Wildl . Serv FishWildl manatus are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost litteranalyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds - such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in groupsat night. Some of the species takenmatchthe bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can some favors substantialvertebratessuch as birds, feed every night, that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains in the floor of the roost: these are quantitativelyinsuf vampire. One reported that individuals of Vampyrum fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. Vampyrum spectrum Vampyrum occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spe tropics but is never dense. This is presumablya function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

-

two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull. Am. Mus . Nat. Hist122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat, in . Panama Can . J. Zool. 47:140 15. , อ*' Stiles, F. G.; and Bradbury , J. 1977. Casebeer, R. อ*' Linsky, B . อ*' and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian phyllostomid bats, Ectophylla prey alba and Vampyrum of the Neotropical carnivorousbat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

^ฬ

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. พ. 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ร. 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus Fish Wildl . Serv .t Wildl . Res . Rept . manatus ) . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrum spectrum ( Vampiro Falso, False Vampire Bat)

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp 1977. Observations on the foraging behavior phyllostomid bats , Ectophylla alba and Vampyrum and avian prey of the Neotropical carnivorous bat , Vampyrum spec spectrum , in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:469-78.

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**

501

MAMMALS

SPECIES ACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich V ampyrum roosts are n ow kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. A review female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. ร / 1935 . Notesconcerning and BaiTett , and Tobago . Bull . Am . Mus.Nat . social to be highly . Theyappear cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group composition . . Fieldworkers populations areencouraged , A. M. 1%8. Notesonthebehavior toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius , thewhole asoft group emits of the the roost in d isturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the noise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. บ/ 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . hollow in given a Vampyrum have Can. J . Zool . 47:14015. . of Mammalogists Society tionno. 5. American . signals ofsocial variety a wide demonstrated Animals , บ/ี Stiles Vehrencamp (Trichechus , F. G.; andBradbury manatee , บ/ L. 1977 . TheWfestIndian Husar , R. บ/ี Linsky , J. 1977 Casebeer , B. บ/ี andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical byaninterlocking spectrum, inCosta totheroost ontheirreturn aregreeted 7:1-22. bat, Vampyrum specRica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapid high .Review; ofmouths manatee muchlikethatfoundinwolves , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, timeswhile itseveral astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andother ofupto200 birds withadult batsarethemajor weights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term a occasion batsthatshare to be rare of phyllostominine inaseries extreme . studyontheremains prey backtoa roostin biasforlarger of preybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles birds showed a heavybiasfornonpasserine pinnae complete Allhavelarge , andtrogons , parakeets , motmots , doves . formssuchasMicro- suchasanis . Smaller noseleaves elongate orroostin fairlysmelly , intermediate Mostof thesepreyareeither plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors withinanhourafterleaving the , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily roost withlarge canine teethanda rowofcar inglyendowed the thatalthough ;theskullbears .Italsoshowed fromtheroost distances apronounced nassiallike molars shearing in the wasleftbehind pupin thestudygroup . Were suckling jawmuscles thepowerful cresttoanchor sagittal orsubadult “guarded byoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . forasmalldogin headform , inTrinidad . Early pairwasobserved explorerstheadult ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient of allthebatsin needs withtheirlarge fortheenergy toaccount wings fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , provisioning , since ofyoungsters thetruevam- theroost speculation wereclearly suggestions -sitter) ) batshave thatthese ( Desmodus , Diaemus , andDiphylla donotfantheir seems . Itisclear adistinct possibility pires with family livesthatmayshowmanyparallels complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe other Vampyrum spectrum roosts ingroups . carnivores suchascanids mammalian of theAmerican overa widerange of hollow batspe Vampyrum occurs . Mostother darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The of itsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin yearsby museum of hasbeenexacerbated toconsist appear in recent thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

-

-

-

-

-

-

-

**

-

-

-

501

MAMMALS

Vampyrum

Tampa Tribune , -, J . Mamm . Archiv . Naturgesch . Ecology and behavior of the mana ร, ) in Florida . ( manatus tee Trichechus Vampyrum (Trichechus ร, Fish Wildl . Serv . Wildl . Res . Rept . manatus ) . Review

SPECIES ACCOUNTS

Bull . Am . Mus Nat .

Hist

J . Mamm .

ร,‘

t

spectrum ,

ร,‘

Ectophylla alba J . Mamm .

pyrum spectrum , Can . J . Zool .

Vampyrum

Vam -

ร,‘

trum . J . Mamm .

Vampyrum spec-

Vampyrum spectrum ( Vampiro Falso, False Vampire Bat)

Vampyrum spectrum

nycteris

Micro-

Trachops

Vampyrum

Chrotopterus

Vampyrum

Vampyrum

Vampyrum

( Desmodus , Diaemus ,

Diphylla )

Vampyrum spectrum bilineata ,

Saccopteryx

Vampyrum

501

MAMMALS

Anonymous . 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. - , 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von . 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv is the spe - chittering sound . This noise characteristicof ofthemana andbehavior Hartman, D. ร, 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists . . Animals Husar, ร, L . 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight . The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World . ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the major constituents g and a wingspan approaching 80 cm . The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae, canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves , parakeets, motmots , and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds, feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles . Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult member of the group. Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings , and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation. Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam - the roost, provisioning of youngsters ( and the baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata of has been exacerbated in recent years by museum skin few groups that have been captured appear to consist group One in collectors, pet fanciers, and careless field biologists who one adult pair and their recent young . such , suckling one adults two will not leave the bats done. Trinidad and Costa Rica are Guanacaste consisted of the

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall, A . M . 1% 1. A review of the batsof Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122:191 -301 . Greenhall, A . M . 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat, in . Panama Can.J. Zool. 47:140 15. , ร,‘ Stiles , F. G .; and Bradbury, J. Casebeer, R . ร,‘ Linsky , B . ร,‘ and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats, Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

, andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous -sized . . Tampa Tribune, 8 March continues Intheroostthebatsform problem . female thathadnotyetbred a and contact manatees , o. ร * 1935 . Notesconcerning BaiTett , and social to be highly . Theyappear cluster . J . Mamm. 16:216 dugongs -20. forlongperiods stable mayremain group composition . . When Costaricas . Diesaugethiere , A. von. 1869 Frantzius , thewhole asoft group emits the roost in disturbed ) 35:247 Archiv. Naturgesch. (Berlin -325. ofthespe ischaracteristic sound . Thisnoise chittering , D. อ*1979 . Ecology and behavior of the mana- cies Hartman forthe presence marker a good and constitutes of Publica tee (Trichechus manatus ) in Florida . Special t ree groups Captive . hollow in given a Vampyrum have . ofMammalogists Society tionno. 5. American . Animals signals ofsocial a widevariety (Trichechus demonstrated manatee , อ*L. 1977 . TheWfestIndian Husar , forforaging , forexample , fromtheirgroup Fish Wildl. Serv.t Wildl. Res. Rept. separated manatus ) . byaninterlocking totheroost ontheir return aregreeted 7:1-22. . A rapid inwolves high .Review; ofmouths likethatfound manatee much , J. E. 1979 . Thesemisocial Reynolds , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, Vampyrum spectrum times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear ห , these . Mostrebatsarecarnivorous above Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld that Vampyrumspectrum isthelargest , rodents , andother ofupto200 birds withadult batsarethemajor weights animal Itisanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong a occasion batsthatshare to be rare ofphyllostominine -term inaseries extreme . prey larger backtoa roostin biasfor ofpreybrought ontheremains anda dietary study physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae complete Allhavelarge , andtrogons , , motmots , . forms suchasMicro- suchasanisdovesparakeets . Smaller noseleaves elongate orroostin fairly smelly , intermediate Mostofthesepreyareeither plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger n ot that a llbats , s uggested . Thissamestudy andthelarge inweight aslizards such withsmall vertebrates sects Vampyrum , , can some that foraging suchasbirds feedeverynight a substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrumarestrik timesfillitsstomach bats rodents at forage great n ecessarily , d o not row car of and the bats roost that a withlarge canine teethand inglyendowed the thatalthough ;theskull .Italsoshowed from theroost distances apronounced nassiallike molars bears shearing the behind in left was g roup study the pup in suckling Were . muscles jaw powerful anchor the crest to sagittal orsubadult “guarded byoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some form member . doginhead forasmall , inTrinidad . Early pairwasobserved explorers ofacaptive name is a misnomer theadult male Thescientific It . behavior batssimply assumed thatthe andthismaybea common inthisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum intheflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheir fortheenergy toaccount large wings fanned their victims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices weresuction tures (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning batshave thatthese ( Desmodus, Diaemus , and . Itisclear adistinct Diphylla ) donotfantheir seems possibility pires with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe other Vampyrumspectrum roosts ingroups . carnivores suchascanids mammalian oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably ofSaccopteryx tropics ciesavoid these dense trees dispersion . Thissparse carnivore asa large . The ofitsstatus bilineata, whichmayroostnearthetreeopening skin yearsby museum of hasbeenexacerbated toconsist appear inrecent thathavebeencaptured fewgroups who , andcareless , petfanciers fieldbiologists . Onesuchgroup young in collectors pairandtheir oneadult recent Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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twositesinwhich Vampyrum roosts arenowkeptsecretGoodwin , G. G., andGreenhall , A M 1%1. A review forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and T obago . Bull . Am. Mus.Nat . complete itsskincollection attheexpense oftheremnant Hist 122:191301 . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve and Greenhall onthebehavior ofthe enjoytheselarge bats , butunnecessary false v ampire bat. J . Mamm. 49:337 disturbance and 40 . collecting should , R. L., andKirmse Peterson beavoided whenever , p. 1%9. Notes possible . onVampyrum spectrum, thefalsevampire , bat in . Panama Can. J . Zool . 47:14015. , อ*' Stiles Vehrencamp , F. G.; andBradbury , R. อ*' Linsky , J. 1977 Casebeer , B. อ*' andNelson . , . 1%3.The Observations ontheforaging behavior phyllostomid and , Ectophylla alba andVampyrum a vian bats p rey of theNeotropical carnivorous bat, Vampyrum specspectrum, inCosta Rica . J . Mamm. 44:186 -89. trum. SPECIES ACCOUNTS

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SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , Goodwin G. G., andGreenhall . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , but false v ampire unnecessary . bat J . . ) 35:247 325 Mamm Archiv. Naturgesch . (Berlin . disturbance : 49 and 337 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , Publica the tee (Trichechus manatus ) in Florida . Special false v ampire , bat in . Panama tree groups . Captive Vampyrum in a givenhollow have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . Animals Casebeer ofsocialsignals a widevariety , Vehrencamp (Trichechus demonstrated , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 , B. andNelson . , c . 1 % 3 . , , for example The for g roup their foraging , from separated Observations on ) Rept the . manatus . ร (พ( Fish Wildl . Serv.t Wildl . Res . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, Vampyrum spectrum times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear J . Bradbury , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing the behind in pupin thestudygroupwasleft . Were suckling jawmuscles thepowerful cresttoanchor sagittal subadult or “guardedbyoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific It . behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested to back prey brought Since a re Both blood not e aten . a ll explanation usedtoremove . devices turesweresuction sitter ( , baby , vam and the of since t he true youngsters roost speculation wereclearly suggestions - ) - the provisioning have ) t hese b ats is that It ( . their distinct clear do not a fan possibility seems piresDesmodus , Diaemus , andDiphylla with show many p arallels m ay family lives that complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion s parse , . The c arnivore This as status a m ay opening large . ofits bilineata which roostnearthetree skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who , andcareless , petfanciers fieldbiologists . Onesuchgroup young in collectors oneadultpairandtheirrecent Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). ป๙ manatus ๐๙FishWildl. Serv.t Wildl. Res. Rept. are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

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two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

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, andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous -sized . . Tampa Tribune, 8 March continues Intheroostthebatsform problem . female thathadnotyetbred a and contact manatees , o. ร * 1935 . Notesconcerning BaiTett , and social to be highly . Theyappear cluster . J . Mamm. 16:216 dugongs -20. forlongperiods stable mayremain group composition . . When Costaricas . Diesaugethiere , A. von. 1869 Frantzius , thewhole asoft group emits the roost in disturbed ) 35:247 Archiv. Naturgesch. (Berlin -325. ofthespe ischaracteristic sound . Thisnoise chittering , D. อ*1979 . Ecology and behavior of the mana- cies Hartman forthe presence marker a good and constitutes of Publica tee (Trichechus manatus ) in Florida . Special t ree groups Captive . hollow in given a Vampyrum have . ofMammalogists Society tionno. 5. American . Animals signals ofsocial a widevariety (Trichechus demonstrated manatee , อ*L. 1977 . TheWfestIndian Husar , , for example for g roup their foraging , from separated Fish Wildl. Serv.t Wildl. Res. Rept. manatus ) . byaninterlocking totheroost ontheir return aregreeted 7:1-22. . A rapid inwolves high .Review; ofmouths likethatfound manatee much , J. E. 1979 . Thesemisocial Reynolds , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, Vampyrum spectrum times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear , these . Mostrebatsarecarnivorous above Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld that Vampyrumspectrum isthelargest , rodents , andother ofupto200 birds withadult batsarethemajor weights animal Itisanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong a occasion batsthatshare to be rare ofphyllostominine -term inaseries extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae complete Allhavelarge , andtrogons , parakeets , motmots , doves . forms suchasMicro- suchasanis . Smaller noseleaves elongate orroostin fairly smelly , intermediate Mostofthesepreyareeither plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge inweight . Thissamestudysuggested aslizards such withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrumarestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily roost withlarge canine teethanda rowofcar inglyendowed the thatalthough ;theskull .Italsoshowed from theroost distances apronounced nassiallike molars bears shearing inthe wasleftbehind pupinthestudygroup . Were suckling jawmuscles thepowerful cresttoanchor sagittal orsubadult “guarded byoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some form member . doginhead forasmall , inTrinidad . Early pairwasobserved explorers ofacaptive name is a misnomer theadult male Thescientific . It behavior batssimply assumed thatthe andthismaybea common inthisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum intheflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheir fortheenergy toaccount large wings fanned their victims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices weresuction tures (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) batshave thatthese ( Desmodus, Diaemus , and . Itisclear adistinct Diphylla donotfantheir seems possibility pires with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe other Vampyrumspectrum roosts ingroups . carnivores suchascanids mammalian oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably ofSaccopteryx tropics ciesavoid these dense trees dispersion . Thissparse carnivore asa large . The ofitsstatus bilineata, whichmayroostnearthetreeopening skin yearsby museum of hasbeenexacerbated toconsist appear inrecent thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors pairandtheir oneadult recent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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twositesinwhich Vampyrum roosts arenowkeptsecretGoodwin , G. G., andGreenhall , A M 1%1. A review forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and T obago . Bull . Am. Mus.Nat . complete itsskincollection attheexpense oftheremnant Hist 122:191301 . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve and Greenhall onthebehavior ofthe enjoytheselarge bats , butunnecessary falsevampire bat. J . Mamm. 49:337 disturbance and 40 . collecting should , R. L., andKirmse Peterson beavoided whenever , p. 1%9. Notes possible . onVampyrum spectrum, thefalsevampire , bat in . Panama Can. J . Zool . 47:14015. , อ*' Stiles Vehrencamp , F. G.; andBradbury , R. อ*' Linsky , J. 1977 Casebeer , B. อ*' andNelson . , . 1%3.The Observations on the foraging behavior phyllostomid and , Ectophylla alba andVampyrum a vian bats p rey of theNeotropical carnivorous bat, Vampyrum specspectrum, inCosta Rica . J . Mamm. 44:186 -89. trum. SPECIES ACCOUNTS

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MAMMALS

Anonymous . 1973. Baby Snooks eats and performs, but ฟ8 March. problem continues. BaiTett , o. 1935. Notes concerning manatees and ห 16:216-20. dugongs. ห Frantzius, A . von . 1869. Die saugethiere Costaricas. ห ( Berlin ) 35:247-325. ห ส Hartman, D. 1979. 1 / หSpecial Publication no. 5. American Society of Mammalogists . Husar, L . 1977. The Wfest Indian manatee /

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social, and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristicof the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals 1 ห ห ห separated from their group, for example , for foraging, ห ห ห are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight . The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear yf As noted above , these bats are carnivorous. Most reis the largest bat in the New World . ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the major constituents g and a wingspan approaching 80 cm . The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae, canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ส such as anis, doves , parakeets, motmots , and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebratessuch as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds, feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents, and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles . Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult member of the group. Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of fanned their victims to sleep with their large wings , and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation. Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam - the roost, provisioning of youngsters ( and the baby-sitter) 1do not fan their seems a distinct possibility. It is clear that these bats have ฟand ฟ pires / complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees, with the exception of ฟwhich may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall, A . 1% 1. A review for fear that some small museum will feel the need batsof Trinidad and Tobago. M . ห ห . ห complete its skin collection at the expense of the remnantto of the 122 :191 301 . populations. Fieldworkers are encouraged to observe and Greenhall, A . M . 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. ห ห49:337-40. of the and collecting should be avoided whenever possible. Peterson, R . L. , and Kirmse , p. 1%9. Notes on ส ฟthe false หห ห47:140 15. vampire bat, in Panama. , Stiles , F. G.; and Bradbury, J. 1977. Casebeer, R . Linsky , B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats, and of the Neotropical carnivorous bat, ฟin Costa Rica. ห ส ห44:186-89. หห ห58:469 78.

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501

MAMMALS

SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin , 8 March . Tribune . Tampa continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. . Notesconcerning 1935 and BaiTett , and Tobago . Bull. Am. Mus . Theyappearto be highlysocial cluster .Nat. complete itsskincollection attheexpense oftheremnant Hist122:191301 . :216 20. . 16 . J. Mamm dugongs forlongperiods stable mayremain group c omposition . . populations Fieldworkers areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe intheroost disturbed enjoytheselargebats , butunnecessary false v ampire bat. J. Mamm . ) 35:247 325 . (Berlin . Naturgesch Archiv . 49 disturbance : and 337 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided ofthemana , D. 1979 . Ecology andbehavior whenever , p. 1%9. Notes Hartman possible . onVam forthe presence marker a good andconstitutes cies of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J. Zool. 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , Vehrencamp (Trichechus , F. G.; andBradbury Stiles manatee , L. 1977 . TheWfestIndian Husar , R. Linsky , J. 1977 Casebeer , B. andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on ). อ 8ร 8 FishWildl the . separated . Res. Rept .t Wildl . Serv manatus foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. carnivorous bat, Vampyrum spectrum , inCosta spec Rica . J. Mamm . 44 :186 -89. trum . J. Mamm ; ofmouths . A rapidhigh . 58 inwolves .Review :46978. manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group , Vampyrum spectrum (VampiroFalso times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight FalseVampireBat) singly . toforage appear J . Bradbury , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld isthelargest that Vampyrum spectrum , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots - suchasanis, doves . formssuchas Micro . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies mixlarger andChrotopterus forms suchasTrachops thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum , feedeverynight , thata foraging cansome suchasbirds favors substantial vertebrates Vampyrum arestrik timesfillitsstomach within the anhourafterleaving , andother . ThejawsofVampyrum bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrumin theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave donotfantheir seems adistinct , andDiphylla , Diaemus possibility pires(Desmodus with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian roosts Vampyrum ingroups . spectrum carnivores suchascanids oftheAmerican occurs overa widerange ofhollow batspe Vampyrum other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably tropics ofSaccopteryx ciesavoid thesetrees dense dispersion , which . Thissparse mayroostnearthetreeopening . The ofitsstatusasa largecarnivore bilineata skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

. .

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-

-

-

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-

**

-

-

-

501

MAMMALS

ดด

ฺ์ ๑ ฺ ฺ ๑ไ ์ ไ ์ , 0 ด Tampa Tribune ด ๑ ๑ ด ์ ๖ ์ ฺด ด ฺ์ ด . ฐไ ๊ด ด J. Mamm ด ๑ด ฺ หด ด ด ๑ ฺ๑ ๑ ๖ ๑ ์ “ด « . ้ ิ ฐไ . Naturgesch Archiv ๑๑ ๑ ์ ด ofthemana ฺ andbehavior Ecology ฟด ด ๑ ๖ ์ ) inFlorida . manatus tee(Trichechus ไ๖ ๑ ด ด Vampyrum๖ ด ด « ด ๖ (Trichechus ๑ ฺ็ ด ด๑ ฺ ฺ ๑ ฺ ์ ). . . Res. Rept .t Wildl . Serv FishWildl manatus ๑ ๑ ์ ด “ ฐ ไ 5 ด ห ์ ์ ด Review ๑ ± ฺ 8 ด ด ด๑ ช ฺด ! ด ๑ ์ ๖ ไ ๑ ๑ ๑ ๖ ด ๖ ( Vampiro Falso , ๑ ์ ์ ฺ ๑ ๑ ด๑ False Vampire Bat )

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ฺ จ ฺ์ ด จด ฺ หด สด 5! 5ด ห ๑ ์ ด . Am Bull . ด Mus Nat. ๑ Hist ฌ ฐ5 5 ด ด ๑ ฺ หด สด 5 ๑ 0 ! ด ์ ๑ ๑ ๑ ๑ฺ ๑ ๑ ด J. Mamm . ฐ“ ์๑ ฺ ด ไ วด็ ฺ ์ ฺด ด ์ 5! ด ฝ ด Vam pyrum spectrum , ๑ ๑ ๑ ฺ ด Can . J. Zool. ฐ" ด ฺ ฺ ด ±์ อ จด ฺ ฺ วด 8 ด ด ฺ ๐ด ์ ฺ ] ด 5 ! ํ ด ๑ ๑ ์ ฺ Ectophylla alba ์ Vampyrum ๑ ฺ spectrum Vampyrum , ๖ด spec J. Mamm . ฐไ ๊ด trum . J. Mamm . ฐฝด

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๑ ๑ ด Trachops์ Chrotopterus ฺ์ ๑ ๑ ์ ด Vampyrum ฺ ฺ ๖ Vampyrum ๑ ฺ์ ด ๑ Vampyrum๑ ฺ์ ๑ ๖ ์ ์ ๑ ไ ๑ ๑ด ๖ ๑ ±๑ ๑ ๑ ด ๑ ฺ ์ ๑ ฺ ์ ๖ ๑ ด ด ๖ ฺ ด ๖ ๑ ๑ ๖ ๑ ด ์ ๑ ์ ๑ ๖ 9 9 ๑ ์ ๑ ์ ไ Vampyrum ๑ ๑ ฐ ๑๑ ด๑๑ ๑ ๑ ๑ ฺ์ ๑ ์ ]๑ ์๑ ๑ ์ ด ๑ ด ้ ์๑ ไ ิ ฺ ๑ ๑ ไ ๑ฺ ) ๑ (Desmodus ด ๖ , ์ , Diaemus Diphylla ์ ์ ์ ด ๑ Vampyrum ด spectrum ๖ ๑ ๑ ด ๑ Vampyrum ๖ Saccopteryx ด ์ ๑ฺ ๑ ๑ , ๖ ๑ ด๑ ด๑ bilineata ๑ ์ ฺ์ ฺ ์ ด๑ ๑ ์ ฺ ๖ ๑ ์ ด ๑ ๑ ์๑ ๑

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). ร (พ( FishWildl . . Res. Rept .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool. 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrum of the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

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-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. พ. 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ร. 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus £. £ Fish Wildl . Serv . t Wildl . Res . Rept . manatus ) . — 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrum spectrum ( Vampiro Falso, False Vampire Bat) หว

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp 1977. Observations on the foraging behavior phyllostomid bats , Ectophylla alba and Vampyrum and avian prey of the Neotropical carnivorous bat , Vampyrum spec spectrum , in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:469-78.

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but March. ๖ ฺ8๑ problem continues. BaiTett , o. 1935. Notes concerning manatees and ด 16:216-20. dugongs. 8ด Frantzius, A . von. 1869. Die saugethiere Costaricas. ด ( Berlin ) 35:247-325. ด ๑ ไ ์ Hartman , D. 1979. ิ ้ ดSpecial ๖ Publica๑ tion no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee ้

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals ิ ด ด ด separated from their group , for example , for foraging , ด ด ด are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ไ such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds , feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . . Early explorers name is misnomer the adult male of a captive pair was observed in Trinidad, a The scientific the bats assumed that simply vampire tales of and this may be a common behavior in this species. It hearing largest and most fierce- looking species must be “ the ** would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufvampire. One reported that individuals of fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ิdo not fan their seems a distinct possibility. It is clear that these bats have ฺand ฺ pires ้ complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees , with the exception of ฺwhich ๖may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. M. 1% 1. A review for fear that some small museum will feel the need of the to bats of Trinidad and Tobago . ด ด . ด complete its skin collection at the expense of the remnant : 191 301 . 122 populations. Fieldworkers are encouraged to observe and Greenhall , A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. 8ด ด49:337-40. of the and collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1%9. Notes on ไ ฺthe false 8ด ด47:140 15. vampire bat , in Panama. ด , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , and of the Neotropical carnivorous ฺin Costa Rica. 8 ไ ด ด44:186-89. ด 8ด ด58:469 78. bat ,

-

-

-

**

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup, one half-grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett, o. ร * 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remainstable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbedin the roost, the whole groupemits a soft . ( Berlin) 35:247-325. . Naturgesch Archiv is the spe - chitteringsound. This noise characteristicof ofthemana andbehavior Hartman, D. อ*1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, อ*L. 1977. The Wfest Indian manatee(Trichechusdemonstrateda wide variety of social signals , , for example for group their foraging, from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) appearto forage singly. As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost litteranalyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds - such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in groupsat night. Some of the species takenmatchthe bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can some favors substantialvertebratessuch as birds, feed every night, that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitativelyinsuf fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. Vampyrum spectrum Vampyrum occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spe tropics but is never dense. This is presumablya function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

-

two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull. Am. Mus . Nat. Hist 122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat, in . Panama Can.J. Zool. 47:140 15. , อ*' Stiles, F. G.; and Bradbury , J. 1977. Casebeer, R. อ*' Linsky, B . อ*' and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian phyllostomid bats, Ectophylla prey alba and Vampyrum of the Neotropical carnivorousbat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup, one half-grown male, and one nearly adult-sized , 8 March. Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett, o. ร * 1935. Notes concerning manatees and contact cluster. They appear to be highly social, and . 16:216-20. dugongs. J. Mamm group composition may remainstable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbedin the roost, the whole groupemits a soft . ( Berlin) 35:247-325. . Naturgesch Archiv is the spe - chitteringsound. This noise characteristicof ofthemana andbehavior Hartman, D. อ*1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrateda wide variety of social signals. Animals Husar, อ*L. 1977. The Wfest Indian manatee(Trichechus from their group, for example, for foraging, separated ). . . Res. Rept .t Wildl . Serv FishWildl manatus are greeted on their returnto the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds, J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompaniedby vibrationof the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they departand repeat it several times while in flight. Thefunctionof thissoundis unclear, since the bats False Vampire Bat) appearto forage singly. As noted above, these bats are carnivorous. Most reis the largest bat in the New World. portsof stomach, feces , or roost litteranalyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds, rodents, and other bats are the majorconstituents g and a wingspan approaching80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion, but this appears to be rare. One long- term similar physiognomy and a dietary bias for larger prey. study on the remainsof prey brought back to a roost in All have large complete pinnae, canidlike muzzles, and Guanacasteshowed a heavy bias for nonpasserinebirds - such as anis, doves, parakeets, motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit, intermediate Most of these prey are either fairly smelly or roost in mix larger in groupsat night. Some of the species takenmatchthe bats and Chrotopterus forms such as Trachops sects with small vertebratessuch as lizards, and the large in weight. This same study suggested that not all bats can some favors substantialvertebratessuch as birds, feed every night, that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents, and otherbats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost, and that the bats do not necessarily forage at great nassiallikeshearing molars; theskull bearsa pronounced distancesfromthe roost. It also showed that althoughthe sagittal crest to anchorthe powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost, it was always “guarded by one adult or subadult memberof the group. Some evidenceof provisioningby for a small dog in head form. The scientific name is a misnomer. Early explorers theadultmale of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “the would certainlyaccountfor the presenceof prey remains in the floor of the roost: these are quantitativelyinsuf vampire. One reported that individuals of Vampyrum fannedtheirvictims to sleep with their large wings, and ficient to account for the energy needs of all the bats in anothersuggested that the nose leaf andassociatedstruc the group but are substantial enough to warrant some tures were suction devices used to remove blood. Both explanation. Since not all prey eaten are broughtback to suggestions were clearly speculation, since the true vam- the roost, provisioningof youngsters( andthe baby-sitter) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallelswith victims and lap blood insteadof suck it. roosts in groupsof up to five in the other mammaliancarnivoressuch as canids. Vampyrum spectrum Vampyrum occursover a wide range of the American dark upper reachesof hollow trees. Most other bat spe tropics but is never dense. This is presumablya function cies avoid these trees, with the exception of Saccopteryx , which may roost near the tree opening. The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been capturedappearto consist of has been exacerbated in recent years by museum skin one adult pairand theirrecent young. One such group in collectors, pet fanciers, and careless field biologists who Guanacaste consisted of the two adults, one suckling will not leave the bats done. Trinidadand Costa Rica are

-

two sites in which Vampyrum roosts arenow kept secret for fear that some small museum will feel the need to complete itsskin collection at the expense of the remnant populations. Fieldworkersareencouragedto observe and enjoy these large bats, but unnecessarydisturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin, G. G. , and Greenhall, A. M. 1%1. A review of the batsof Trinidadand Tobago. Bull. Am. Mus . Nat. Hist122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson, R. L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat, in . Panama Can . J. Zool. 47:140 15. , อ*' Stiles, F. G.; and Bradbury , J. 1977. Casebeer, R. อ*' Linsky, B . อ*' and Nelson, c. 1%3. The Vehrencamp Observations on the foraging behaviorand avian phyllostomid bats, Ectophylla prey alba and Vampyrum of the Neotropical carnivorousbat, Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune, 8 March. BaiTett, o. ร * 1935. Notes concerning manatees and dugongs. J . Mamm. 16:216-20. Frantzius, A . von . 1869. Die saugethiere Costaricas. Archiv. Naturgesch. ( Berlin) 35:247-325. Hartman, D. อ*1979. Ecology and behavior of the manatee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, อ*L. 1977. The Wfest Indian manatee (Trichechus Fish Wildl. Serv.t Wildl. Res. Rept. manatus ) . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrum spectrum ( Vampiro Falso, False Vampire Bat)

Vampyrumspectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae, canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit, intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebratessuch as lizards, and the large Vampyrum favors substantial vertebrates such as birds, rodents, and other bats. The jaws of Vampyrumare strik ingly endowed with large canine teeth and a row of carnassiallikeshearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and anothersuggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation, since the true vampires ( Desmodus, Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrumspectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees, with the exception of Saccopteryx bilineata, which may roost near the tree opening. The few groups that have been captured appear to consist of one adult pair and their recent young. One such group in Guanacaste consisted of the two adults, one suckling

-

-

-

pup, one half-grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social, and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound. This noise is characteristicof the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group, for example, for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. Thefunctionof thissound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach, feces , or roost-litter analyses agree that birds, rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion, but this appears to be rare. One long- term study on the remains of prey brought back to a roost in Guanacasteshowed a heavy bias for nonpasserine birds such as anis, doves , parakeets, motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night. Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night, that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost, and that the bats do not necessarily forage at great distancesfrom the roost. It also showed that although the suckling pup in the study group was left behind in the roost, it was always “guarded by one adult or subadult member of the group. Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation. Since not all prey eaten are brought back to the roost, provisioning of youngsters ( and the baby-sitter) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors, pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need to complete its skin collection at the expense of the remnant populations. Fieldworkersare encouragedto observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall, A . M. 1% 1. A review of the batsof Trinidad and Tobago. Bull . Am. Mus. Nat . Hist 122:191 -301 . Greenhall, A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm. 49:337-40. Peterson, R . L. , and Kirmse , p. 1%9. Notes on Vampyrum spectrum, the false vampire bat, in Panama. Can. J . Zool . 47:140 15. , อ*' Stiles , F. G .; and Bradbury, J. Casebeer, R . อ*' Linsky, B . อ*' and Nelson, c. 1%3. The Vehrencamp . on the foraging behavior and avian1977 phyllostomid bats, Ectophylla alba and Vampyrum ofObservations prey the Neotropical bat, Vampyrum specspectrum, in Costa Rica. J . Mamm. 44:186-89. trum. J . Mamm. 58:carnivorous 469-78.

-

**

501

MAMMALS

ดด

ฺ์ ๑ ฺ ฺ ๑ไ ์ ไ ์ , 0 ด Tampa Tribune ด ๑ ๑ ด ์ ๖ ์ ฺด ด ฺ์ ด . ฐไ ๊ด ด J. Mamm ด ๑ด ฺ หด ด ด ๑ ฺ๑ ๑ ๖ ๑ ์ “ด « . ้ ิ ฐไ . Naturgesch Archiv ๑๑ ๑ ์ ด ofthemana ฺ andbehavior Ecology ฟด ด ๑ ๖ ์ ) inFlorida . manatus tee(Trichechus ไ๖ ๑ ด ด Vampyrum๖ ด ด « ด ๖ (Trichechus ๑ ฺ็ ด ด๑ ฺ ฺ ๑ ฺ ์ ). . Res. Rept. .t Wildl . Serv FishWildl manatus ๑ ๑ ์ ด “ ฐ ไ 5 ด ห ์ ์ ด Review ๑ ± ฺ 8 ด ด ด๑ ช ฺด ! ด ๑ ์ ๖ ไ ๑ ๑ ๑ ๖ ด ๖ ( Vampiro Falso , Vampyrum spectrum ๑ ์ ์ ฺ ๑ ๑ ด๑ False Vampire Bat )

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๑ ๑ ด Trachops ์ Chrotopterus ฺ์ ๑ ๑ ์ ด Vampyrum ฺ ฺ ๖ Vampyrum ๑ ฺ์ ด ๑ Vampyrum๑ ฺ์ ๑ ๖ ์ ์ ๑ ไ ๑ ๑ด ๖ ๑ ±๑ ๑ ๑ ด ๑ ฺ ์ ๑ ฺ ์ ๖ ๑ ด ด ๖ ฺ ด ๖ ๑ ๑ ๖ ๑ ด ์ ๑ ์ ๑ ๖ 9 9 ๑ ์ ๑ ์ ไ Vampyrum ๑ ๑ ฐ ๑๑ ด๑๑ ๑ ๑ ๑ ฺ์ ๑ ์ ]๑ ์๑ ๑ ์ ด ๑ ด ้ ์๑ ไ ิ ฺ ๑ ๑ ไ ๑ฺ ) ๑ (Desmodus ด ๖ , ์ , Diaemus Diphylla ์ ์ ์ ด ๑ Vampyrum ด spectrum ๖ ๑ ๑ ด ๑ Vampyrum ๖ Saccopteryx ด ์ ๑ฺ ๑ ๑ , ๖ ๑ ด๑ ด๑ bilineata ๑ ์ ฺ์ ฺ ์ ด๑ ๑ ์ ฺ ๖ ๑ ์ ด ๑ ๑ ์๑ ๑

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). . . Res. Rept .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

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two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

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SPECIESACCOUNTS , andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous sized twositesinwhich Vampyrum roosts are now kept s ecret , G. G., andGreenhall Goodwin . . Tampa Tribune , 8 March continues problem . Intheroostthebatsforma , A M 1%1. Areview female thathadnotyetbred forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and contact manatees , o. พ. 1935 . Notesconcerning and BaiTett , and Tobago . Bull . Am. Mus.Nat . . Theyappearto be highlysocial cluster complete itsskincollection attheexpense oftheremnant Hist 122:191301 . 20. . J . Mamm . 16:216 dugongs forlongperiods stable mayremain group c omposition . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve . When Costaricas . Diesaugethiere , A. von. 1869 and Greenhall Frantzius onthebehavior , thewhole a soft group emits ofthe the roost in disturbed enjoy t hese l arge b ats , butunnecessary false v ampire bat. J . Mamm . 49:337 . ) 35:247 325 Archiv. Naturgesch . (Berlin disturbance and 40 . of i s the n oise characteristic sound This . spe chittering collecting should , R. L., andKirmse Peterson beavoided , D. ร. 1979 . Ecology and behavior of the mana - ciesandconstitutes whenever , p. 1%9. Notes Hartman possible . onVamforthe presence marker a good of pyrum spectrum , ) ( Publica the . Special Florida in manatus tee Trichechus false vampire , bat in . Panama t ree groups Captive . h ollow in given a Vampyrum have Can . J . Zool . 47:14015. . ofMammalogists Society tionno. 5. American . signals ofsocial v ariety a wide demonstrated Animals , ร.; Stiles Vehrencamp (Trichechus , F. G.; andBradbury manatee , ร. L. 1977 . TheWfestIndian Husar , R. ร.; Linsky , J. 1977 Casebeer , B. ร.; andNelson . , c. 1%3.The Observations , forforaging , forexample , fromtheirgroup on the Fish Wildl . Serv.t Wildl . Res . Rept . separated manatus ) . foraging behavior phyllostomid and , avian bats p rey Ectophylla alba and Vampyrum of theNeotropical totheroostbyaninterlocking ontheirreturn aregreeted 7:1-22. bat, Vampyrum specspectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . J . Mamm . 58:carnivorous . A rapidhigh inwolves .Review; ofmouths manatee muchlikethatfound , J. E. 1979 . Thesemisocial Reynolds 46978. , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group (VampiroFalso , times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight FalseVampireBat) singly . toforage appear ห , thesebatsarecarnivorous . Mostreabove Asnoted agree , feces , orroost litteranalyses . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andotherbatsarethemajor ofupto200 birds withadultweights animal It isanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong-term batsthatsharea occasion to be rare ofphyllostominine ina series extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae Allhavelargecomplete , andtrogons , parakeets , motmots , doves . formssuchas Micro- suchasanis . Smaller noseleaves elongate orroostin , intermediate Mostofthesepreyareeitherfairlysmelly plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge in weight . Thissamestudysuggested suchaslizards withsmall vertebrates sects Vampyrum cansome , feedeverynight , thata foraging suchasbirds substantial vertebrates Vampyrum favors within the anhourafterleaving , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , andthatthebatsdonotnecessarily withlargecanine teethanda rowofcar- roost inglyendowed the thatalthough ; theskullbears .Italsoshowed fromtheroost distances a pronounced nassiallike molars shearing in the pupin thestudygroupwasleftbehind . Were suckling jawmuscles thepowerful cresttoanchor sagittal “guardedbyoneadultorsubadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some member . fora smalldogin headform , inTrinidad . Earlyexplorerstheadult pairwasobserved ofacaptive nameis a misnomer male Thescientific . It behavior batssimply assumed thatthe andthismaybea common in thisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrum in theflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheirlargewings fortheenergy toaccount fanned theirvictims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices turesweresuction (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) . Itisclearthatthesebatshave adistinct possibility pires( Desmodus , Diaemus , andDiphylla donotfantheir seems with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe othermammalian Vampyrum spectrum roosts ingroups . carnivores suchascanids oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most darkupper reaches trees , withtheexception butisnever a function .Thisispresumably ofSaccopteryx tropics ciesavoid thesetrees dense dispersion . Thissparse . The ofitsstatusasa largecarnivore bilineata , whichmayroostnearthetreeopening skin of hasbeenexacerbated toconsist appear in recentyearsbymuseum thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors oneadultpairandtheirrecent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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MAMMALS

, andonenearly adult , but pup male , onehalf andperforms eats Snooks . Baby -grown . 1973 Anonymous -sized . . Tampa Tribune , 8 March continues thebats Intheroost problem . hadnotyetbred form female that a and contact manatees , o. ๐11935 . Notesconcerning BaiTett , and social tobehighly . Theyappear cluster . . J . Mamm . 16:216 dugongs -20 forlongperiods stable mayremain group composition . . When Costaricas saugethiere , A. von . 1869. Die Frantzius , thewhole asoft group emits the roost in disturbed ) 35:247 Archiv. Naturgesch . (Berlin -325. ofthespe ischaracteristic noise sound . This chittering , D. ป11979 . Ecology and behavior of the mana - cies Hartman for the presence marker a good and constitutes of Publica tee (Trichechus manatus ) in Florida . Special t ree groups Captive . h ollow in given a Vampyrum have . ofMammalogists Society tionno. 5. American . Animals signals ofsocial a widevariety (Trichechus demonstrated manatee , ป1L. 1977 . TheWfestIndian Husar , , for example for group their foraging , from separated Fish Wildl . Serv.t Wildl . Res . Rept . manatus ) . byaninterlocking totheroost ontheir return aregreeted 7:1-22. . A rapid inwolves high .Review; ofmouths likethatfound manatee much , J. E. 1979 . Thesemisocial Reynolds , pp wings . 93-%. June oftheclosed isa byvibration accompanied piping thewhole to agitate behavior thatseems contagious . Batsleaving a roosttofeedoftenemita loud group Vampyrum spectrum ( Vampiro Falso, times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear หว , these . Mostrebatsarecarnivorous above Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld that Vampyrum spectrum isthelargest , rodents , andother arethemajor ofupto200 birds withadult bats weights animal Itisanimposing constituents maybetaken . Somefruitandinsects isthe of thediet 80cm. The species on approaching ganda wingspan , butthisappears . Onelong a occasion batsthatshare toberare ofphyllostominine -term inaseries extreme . study prey back toa roost biasforlarger ofpreybrought ontheremains anda dietary in physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae complete large Allhave , andtrogons , parakeets , motmots , doves . forms suchasMicro- suchasanis . Smaller noseleaves elongate orroost fairly smelly , intermediate in Mostofthesepreyareeither fruit plus orinsects nycteris feedoninsects taken match thebats ofthespecies atnight in groups . Some forms suchasTrachops andChrotopterus mixlarger thatnotallbats , andthelarge inweight suggested study . Thissame aslizards such withsmall vertebrates sects Vampyrum cansome , feedevery , thata foraging night suchasbirds substantial vertebrates Vampyrum favors leaving after within the anhour , andother . ThejawsofVampyrum arestrik timesfillitsstomach bats rodents atgreat forage , and donotnecessarily roost thatthebats withlarge canine teeth anda rowofcar endowed ingly the although that ;theskull .Italsoshowed from theroost distances apronounced nassiallike molars bears shearing inthe wasleftbehind group pupinthestudy . Weresuckling jawmuscles thepowerful crest toanchor sagittal orsubadult “guarded byoneadult , itwasalways easily betakenroost a noseleaf, thisbatcould itnottohave by ofprovisioning evidence ofthegroup . Some form member . doginhead forasmall , inTrinidad wasobserved . Early pair explorers ofacaptive name is a misnomer theadult male Thescientific . It behavior batssimply assumed thatthe andthismaybea common inthisspecies talesof vampire hearing remains ofprey forthepresence certainly be“the** would must account species largest andmostfierce -looking insuf of Vampyrum intheflooroftheroost : these thatindividuals . Onereported vampire arequantitatively , and ficient ofallthebatsin needs withtheir fortheenergy toaccount large wings fanned their victims tosleep some enough butaresubstantial towarrant struc thegroup leafandassociated another thenose that suggested to back arebrought blood notallprey eaten . Since . Both explanation usedtoremove devices weresuction tures sitter (and , provisioning , since vam thebaby ofyoungsters thetrue speculation clearly were suggestions - ) - theroost have ) these bats that ( Desmodus , Diaemus , and . Itisclear adistinct Diphylla donotfantheirseems possibility pires with many parallels family livesthatmayshow complex victims andlapblood instead ofsuckit. ofuptofiveinthe other Vampyrum spectrum roosts ingroups . carnivores suchascanids mammalian oftheAmerican overa widerange ofhollow batspe Vampyrum occurs other . Most upper dark reaches trees , withtheexception butisnever afunction .Thisispresumably ofSaccopteryx tropics ciesavoid these dense trees dispersion . Thissparse carnivore asa large mayroostnearthetreeopening . The ofitsstatus bilineata , which skin years bymuseum of hasbeenexacerbated toconsist appear inrecent captured been thathave fewgroups who biologists , , field careless and . Onesuchgroup young in collectors pair oneadult andtheir recent petfanciers are Rica , onesucklingwillnotleave andCosta . Trinidad done thebats of thetwoadults consisted Guanacaste

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inwhich twosites Vampyrum roosts arenowkeptsecretGoodwin , G. G., and , A M 1% Greenhall 1. Areview forfearthatsome small museum willfeeltheneedto ofthebats of Trinidad and T obago . . Am. Mus.Nat . Bull complete itsskin collection attheexpense oftheremnantHist 122:191301 . . Fieldworkers populations areencouraged , A. M. 1% Greenhall toobserve 8. Notes and onthebehavior ofthe enjoy these large bats , butunnecessary falsevampire . J . Mamm . 49 bat disturbance : and 337 40 . collecting should , R. L., andKirmse Peterson beavoided whenever , p. 1% possible . 9. Notes onVampyrum spectrum , thefalsevampire , bat in . Panama Can . J . Zool . 47:14015. ,ป Vehrencamp 18Stiles , F. G.; andBradbury , R. ป18Linsky , J. 1977 Casebeer , B. ป18and . , . 1% N elson 3 . The Observations on the f oraging behavior phyllostomid and , Ectophylla alba andVampyrum a vian bats prey of theNeotropical carnivorous , Vampyrum specbat spectrum , inCosta Rica . J . Mamm . 44:186 -89. trum . SPECIES ACCOUNTS

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MAMMALS

, andonenearly adult , but pup male , onehalf eatsandperforms . BabySnooks -grown . 1973 Anonymous -sized , 8 March . Tribune . Tampa continues Intheroostthebatsform problem . female thathadnotyetbred a and contact manatees , o. . Notesconcerning 1935 BaiTett , and social to be highly . Theyappear cluster :216 . 16 . J. Mamm dugongs -20. forlongperiods stable mayremain group composition . . When Costaricas . Diesaugethiere , A. von. 1869 Frantzius , thewhole asoft group emits the roost in disturbed ) 35:247 . (Berlin . Naturgesch Archiv -325.ofthemana- chittering ofthespe ischaracteristic sound . Thisnoise , D. 1979 . Ecology andbehavior Hartman forthe presence marker a good and constitutes cies of ) inFlorida Publica . Special manatus tee(Trichechus t ree groups Captive . hollow in given a Vampyrum have . ofMammalogists Society tionno. 5. American . Animals signals ofsocial a widevariety (Trichechusdemonstrated manatee , L. 1977 . TheWfestIndian Husar , , for example for g roup their foraging , from separated ). . Res. Rept. .t Wildl . Serv FishWildl manatus byaninterlocking totheroost ontheir return aregreeted 7:1-22. ; ofmouths . A rapid inwolves high .Review likethatfound manatee much , J. E. 1979 . Thesemisocial Reynolds , pp. 93-%. wings June oftheclosed isa byvibration accompanied piping thewhole behavior thatseemsto agitate contagious . Batsleaving a roosttofeedoftenemita loud group ( Vampiro Falso, times itseveral while astheydepart screech andrepeat in , since ofthissound isunclear thebats .Thefunction flight False Vampire Bat) singly . toforage appear , these . Mostrebatsarecarnivorous above Asnoted agree , feces , orroost analyses litter . ports ofstomach batintheNewWorld isthelargest that Vampyrum spectrum , rodents , andother ofupto200 birds withadult batsarethemajor weights animal Itisanimposing constituents maybetakenon isthe of thediet. Somefruitandinsects 80cm. Thespecies approaching ganda wingspan , butthisappears . Onelong a occasion batsthatshare to be rare ofphyllostominine -term inaseries extreme . studyontheremains prey backtoa roostin biasforlarger ofpreybrought anda dietary physiognomy similar , and Guanacaste , canidlike muzzles biasfornonpasserine birds showed a heavy pinnae complete Allhavelarge , andtrogons , parakeets , motmots , doves - suchasanis . forms suchasMicro . Smaller noseleaves elongate orroostin fairly smelly , intermediate Mostofthesepreyareeither plusfruit orinsects nycteris feedoninsects taken match thebats atnight in groups . Someofthespecies mixlarger andChrotopterus forms suchasTrachops thatnotallbats , andthelarge inweight . Thissamestudysuggested aslizards such withsmall vertebrates sects Vampyrum , feedeverynight , thata foraging cansome suchasbirds favors substantial vertebrates Vampyrum arestrik timesfillitsstomach within the anhourafterleaving , andother . ThejawsofVampyrum bats rodents atgreat forage , andthatthebatsdonotnecessarily roost withlarge canine teethanda rowofcar inglyendowed the thatalthough ;theskull .Italsoshowed from theroost distances apronounced nassiallike molars bears shearing inthe wasleftbehind pupinthestudygroup . Were suckling jawmuscles thepowerful cresttoanchor sagittal orsubadult “guarded byoneadult , it wasalways easily betaken roost it nottohavea noseleaf, thisbatcould by ofprovisioning evidence ofthegroup . Some form member . doginhead forasmall , inTrinidad . Early pairwasobserved explorers ofacaptive name is a misnomer theadult male Thescientific . It behavior batssimply assumed thatthe andthismaybea common inthisspecies talesof vampire hearing ofpreyremains forthepresence certainly mustbe“the** would account species largest andmostfierce -looking insuf of Vampyrumintheflooroftheroost : thesearequantitatively thatindividuals . Onereported vampire , and ficient ofallthebatsin needs withtheir fortheenergy toaccount large wings fanned their victims tosleep some enough butaresubstantial to warrant struc thegroup another thatthenoseleafandassociated suggested backto arebrought blood notallpreyeaten . Since . Both explanation usedtoremove devices weresuction tures (andthebaby , since ofyoungsters thetruevam speculation wereclearly suggestions -sitter) - theroost, provisioning ) batshave thatthese (Desmodus . Itisclear donotfantheir seems adistinct , and , Diaemus Diphylla possibility pires with many parallels family livesthatmayshow complex victims andlapbloodinstead ofsuckit. ofuptofiveinthe other roosts Vampyrum ingroups . spectrum carnivores suchascanids mammalian oftheAmerican occurs overa widerange ofhollow batspe Vampyrum other . Most darkupper reaches trees , withtheexception butisnever afunction .Thisispresumably tropics ofSaccopteryx ciesavoid these dense trees dispersion , which . Thissparse carnivore asa large mayroostnearthetreeopening . The ofitsstatus bilineata skin yearsby museum of hasbeenexacerbated toconsist appear inrecent thathavebeencaptured fewgroups who biologists , , field careless and . Onesuchgroup young in collectors pairandtheir oneadult recent petfanciers Ricaare , onesucklingwillnotleave andCosta . Trinidad thebatsdone of thetwoadults consisted Guanacaste

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SPECIES ACCOUNTS

twositesinwhich Vampyrum roosts arenowkeptsecretGoodwin , G. G., andGreenhall , A M 1%1. A review forfearthatsomesmallmuseum willfeeltheneedto ofthebatsofTrinidad and T obago . Bull. Am. Mus.Nat. complete itsskincollection attheexpense oftheremnant Hist 122:191301 . . Fieldworkers populations areencouraged , A. M. 1%8. Notes toobserve and Greenhall onthebehavior ofthe enjoytheselarge bats , butunnecessary falsevampire bat. J. Mamm . 49 disturbance : and 337 40 . collecting should , R. L., andKirmse Peterson beavoided whenever , p. 1%9. Notes possible . onVampyrumspectrum , thefalse vampire , bat in . Panama Can.J. Zool. 47:14015. , Vehrencamp , F. G.; andBradbury Stiles , R. Linsky , J. 1977 Casebeer , B. andNelson . , . 1%3.The Observations on the foraging behavior phyllostomid and , Ectophylla a vian bats p rey alba and Vampyrum of theNeotropical carnivorous bat, Vampyrum spectrum , inCosta spec Rica . J. Mamm . 44 :186 -89. trum.

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ฺ์ ๑ ฺ ฺ ๑ไ ์ ไ ์ , 0 ด Tampa Tribune ด ๑ ๑ ด ์ ๖ ์ ฺด ด ฺ์ ด . ฐไ ๊ด ด J. Mamm ด ๑ด ฺ หด ด ด ๑ ฺ๑ ๑ ๖ ๑ ์ “ด « . ้ ิ ฐไ . Naturgesch Archiv ๑๑ ๑ ์ ด ofthemana ฺ andbehavior Ecology ฟด ด ๑ ๖ ์ ) inFlorida . manatus tee(Trichechus ไ๖ ๑ ด ด Vampyrum๖ ด ด « ด ๖ (Trichechus ๑ ฺ็ ด ด๑ ฺ ฺ ๑ ฺ ์ ). . Res. Rept. .t Wildl . Serv FishWildl manatus ๑ ๑ ์ ด “ ฐ ไ 5 ด ห ์ ์ ด Review ๑ ± ฺ 8 ด ด ด๑ ช ฺด ! ด ๑ ์ ๖ ไ ๑ ๑ ๑ ๖ ด ๖ Vampyrum spectrum ( Vampiro Falso , ๑ ์ ฺ ๑ ์ ๑ ด ๑ ) False Vampire Bat ด ด ไ ์ฺ ๑ ๑ ๑ ๑ ฺ ฺ ด ์ ๑ Vampyrum spectrum ๑ ๑๑ " ฺ ฺ์ ๑ ์ ๑ด ด๑ ๖ ์ ฺ ด๑ ไ ๑ ๖๑ ๖ ๑ ด ๖ ์ ๑ ๖ ฺ ๑ฺ ์ ๖ ๑ ฺ์ ด ฺ ฺ ฺ Micro ด ฺ ๑ ๑ nycteris ๑ ๑ ด Trachops ์ Chrotopterus ฺ์ ๑ ๑ ์ ด Vampyrum ฺ ฺ ๖ Vampyrum ๑ ฺ์ ด ๑ Vampyrum๑ ฺ์ ๑ ๖ ์ ์ ๑ ไ ๑ ๑ด ๖ ๑ ±๑ ๑ ๑ ด ๑ ฺ ์ ๑ ฺ ์ ๖ ๑ ด ด ๖ ฺ ด ๖ ๑ ๑ ๖ ๑ ด ์ ๑ ์ ๑ ๖ 9 9 ๑ ์ ๑ ์ ไ Vampyrum ๑ ๑ ฐ ๑๑ ด๑๑ ๑ ๑ ๑ ฺ์ ๑ ์ ]๑ ์๑ ๑ ์ ด ๑ ด ้ ์๑ ไ ิ ฺ ๑ ๑ ไ ๑ฺ ) ๑ (Desmodus ด ๖ , ์ , Diaemus Diphylla ์ ์ ์ ด ๑ Vampyrum ด spectrum ๖ ๑ ๑ ด ๑ Vampyrum ๖ Saccopteryx ด ์ ๑ฺ ๑ ๑ , ๖ ๑ ด๑ ด๑ bilineata ๑ ์ ฺ์ ฺ ์ ด๑ ๑ ์ ฺ๑ ์ ๖ ๑ ด ์๑ ๑๑

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. Musด Nat. Hist ฌ ฐ5 5 ด ด ฺ หด สด 5 ๑ 0 ! ด ์ ๑ ๑ ๑ ๑ฺ ๑ ๑ ด J. Mamm . ฐ“ ์๑ ฺ ด ไ วด็ ฺ ์ ฺด ด ์ 5! ด ฝ ด Vampyrumspectrum , ๑ ๑ ๑ ฺ ด Can.J. Zool. ฐ" ด ฺ ฺ ด ±์ อ จด ฺ ฺ วด 8 ด ด ฺ ๐ด ์ ฺ ] ด 5 ! ํ ด ๑ ๑ ์ ฺ Ectophylla alba ์ Vampyrum ๑ ฺ spectrum Vampyrum , ๖ด spec J. Mamm . ฐไ ๊ด trum. J. Mamm . ฐฝด

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ฺ์ ๑ ฺ ฺ ๑ไ ์ ไ ์ , 0 ด Tampa Tribune ด ๑ ๑ ด ์ ๖ ์ ฺด ด ฺ์ ด . ฐไ ๊ด ด J. Mamm ด ๑ด ฺ หด ด ด ๑ ฺ๑ ๑ ๖ ๑ ์ “ด « . ้ ิ ฐไ . Naturgesch Archiv ๑๑ ๑ ์ ด ofthemana ฺ andbehavior Ecology ฟด ด ๑ ๖ ์ ) inFlorida . manatus tee(Trichechus ไ๖ ๑ ด ด Vampyrum๖ ด ด « ด ๖ (Trichechus ๑ ฺ็ ด ด๑ ฺ ฺ ๑ ฺ ์ ). . . Res. Rept .t Wildl . Serv FishWildl manatus ๑ ๑ ์ ด “ ฐ ไ 5 ด ห ์ ์ ด Review ๑ ± ฺ 8 ด ด ด๑ ช ฺด ! ด ๑ ์ ๖ ไ ๑ ๑ ๑ ๖ ด ๖ ( Vampiro Falso , ๑ ์ ์ ฺ ๑ ๑ ด๑ False Vampire Bat )

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SPECIES

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ฺ จ ฺ์ ด จด ฺ หด สด 5! 5ด ห ๑ ์ ด . Am Bull . ด Mus Nat. ๑ Hist ฌ ฐ5 5 ด ด ๑ ฺ หด สด 5 ๑ 0 ! ด ์ ๑ ๑ ๑ ๑ฺ ๑ ๑ ด J. Mamm . ฐ“ ์๑ ฺ ด ไ วด็ ฺ ์ ฺด ด ์ 5! ด ฝ ด Vam pyrum spectrum , ๑ ๑ ๑ ฺ ด Can . J. Zool. ฐ" ด ฺ ฺ ด ±์ อ จด ฺ ฺ วด 8 ด ด ฺ ๐ด ์ ฺ ] ด 5 ! ํ ด ๑ ๑ ์ ฺ Ectophylla alba ์ Vampyrum ๑ ฺ spectrum Vampyrum , ๖ด spec J. Mamm . ฐไ ๊ด trum . J. Mamm . ฐฝด

Vampyrum

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๑ ๑ ด Trachops์ Chrotopterus ฺ์ ๑ ๑ ์ ด Vampyrum ฺ ฺ ๖ Vampyrum ๑ ฺ์ ด ๑ Vampyrum๑ ฺ์ ๑ ๖ ์ ์ ๑ ไ ๑ ๑ด ๖ ๑ ±๑ ๑ ๑ ด ๑ ฺ ์ ๑ ฺ ์ ๖ ๑ ด ด ๖ ฺ ด ๖ ๑ ๑ ๖ ๑ ด ์ ๑ ์ ๑ ๖ 9 9 ๑ ์ ๑ ์ ไ Vampyrum ๑ ๑ ฐ ๑๑ ด๑๑ ๑ ๑ ๑ ฺ์ ๑ ์ ]๑ ์๑ ๑ ์ ด ๑ ด ้ ์๑ ไ ิ ฺ ๑ ๑ ไ ๑ฺ ) ๑ (Desmodus ด ๖ , ์ , Diaemus Diphylla ์ ์ ์ ด ๑ Vampyrum ด spectrum ๖ ๑ ๑ ด ๑ Vampyrum ๖ Saccopteryx ด ์ ๑ฺ ๑ ๑ , ๖ ๑ ด๑ ด๑ bilineata ๑ ์ ฺ์ ฺ ์ ด๑ ๑ ์ ฺ ๖ ๑ ์ ด ๑ ๑ ์๑ ๑

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Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee (Trichechus manatus ) . บ .ร . Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrum spectrum ( Vampiro Falso, False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

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pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior phyllostomid bats , Ectophylla alba and Vampyrum and avian prey of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J . Mamm . 44:186-89. spectrum . J . Mamm . 58:469-78.

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**

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501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. ๐11935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ป11979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. . Animals Husar, ป1L. 1977. The Wfest Indian manatee (Trichechusdemonstrated a wide variety of social signals , , for example for group their foraging, from separated ). ร (พ( FishWildl . Res. Rept. .t Wildl . Serv manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud Vampyrum spectrum ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains vampire. One reported that individuals of Vampyrumin the floor of the roost: these are quantitatively insuf fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

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two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrumspectrum , the false vampire bat , in . Panama Can.J. Zool. 47:140 15. ,ป 18Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . ป18Linsky, B . ป18and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , Ectophylla alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum. J. Mamm . 58:469-78.

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**

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-

-

^ฬ

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but problem continues. Tampa Tribune , 8 March. BaiTett , o. พ. 1935. Notes concerning manatees and dugongs. J . Mamm . 16:216-20. Frantzius, A . von. 1869. Die saugethiere Costaricas. Archiv . Naturgesch . ( Berlin ) 35:247-325. Hartman , D. ร. 1979. Ecology and behavior of the mana tee (Trichechus manatus ) in Florida . Special Publication no. 5. American Society of Mammalogists. Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus manatus ) . ร ท- ท Fish Wildl . Serv .t Wildl . Res . Rept . 7:1-22. Reynolds , J. E. 1979. The semisocial manatee. Review; June, pp. 93-%. Vampyrum spectrum ( Vampiro Falso, False Vampire Bat) J . Bradbury

Vampyrum spectrum is the largest bat in the New World. It is an imposing animal with adult weights of up to 200 g and a wingspan approaching 80 cm. The species is the extreme in a series of phyllostominine bats that share a similar physiognomy and a dietary bias for larger prey. All have large complete pinnae , canidlike muzzles , and elongate nose leaves. Smaller forms such as Micronycteris feed on insects or insects plus fruit , intermediate forms such as Trachops and Chrotopterus mix larger in sects with small vertebrates such as lizards, and the large Vampyrum favors substantial vertebrates such as birds , rodents , and other bats. The jaws of Vampyrum are strik ingly endowed with large canine teeth and a row of carnassiallike shearing molars; the skull bears a pronounced sagittal crest to anchor the powerful jaw muscles. Were it not to have a nose leaf , this bat could easily be taken for a small dog in head form . The scientific name is a misnomer. Early explorers hearing tales of vampire bats simply assumed that the** largest and most fierce- looking species must be “ the vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and another suggested that the nose leaf and associated struc tures were suction devices used to remove blood . Both suggestions were clearly speculation , since the true vampires ( Desmodus , Diaemus , and Diphylla ) do not fan their victims and lap blood instead of suck it. Vampyrum spectrum roosts in groups of up to five in the dark upper reaches of hollow trees. Most other bat spe cies avoid these trees , with the exception of Saccopteryx bilineata , which may roost near the tree opening . The few groups that have been captured appear to consist of one adult pair and their recent young . One such group in Guanacaste consisted of the two adults , one suckling

-

-

-

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of Vampyrum in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals separated from their group , for example , for foraging, are greeted on their return to the roost by an interlocking of mouths much like that found in wolves . A rapid high piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats appear to forage singly. As noted above , these bats are carnivorous. Most reports of stomach , feces , or roost-litter analyses agree that birds , rodents, and other bats are the major constituents of the diet. Some fruit and insects may be taken on occasion , but this appears to be rare. One long - term study on the remains of prey brought back to a roost in Guanacaste showed a heavy bias for nonpasserine birds such as anis, doves , parakeets , motmots , and trogons. Most of these prey are either fairly smelly or roost in groups at night . Some of the species taken match the bats in weight . This same study suggested that not all bats feed every night , that a foraging Vampyrum can sometimes fill its stomach within an hour after leaving the roost , and that the bats do not necessarily forage at great distances from the roost. It also showed that although the suckling pup in the study group was left behind in the roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by the adult male of a captive pair was observed in Trinidad, and this may be a common behavior in this species. It would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufficient to account for the energy needs of all the bats in the group but are substantial enough to warrant some explanation . Since not all prey eaten are brought back to the roost , provisioning of youngsters ( and the baby-sitter ) seems a distinct possibility. It is clear that these bats have complex family lives that may show many parallels with other mammalian carnivores such as canids. Vampyrum occurs over a wide range of the American tropics but is never dense. This is presumably a function of its status as a large carnivore. This sparse dispersion has been exacerbated in recent years by museum skin collectors , pet fanciers, and careless field biologists who will not leave the bats done. Trinidad and Costa Rica are

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull . Am . Mus. Nat . Hist 122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J . Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in Panama. Can . J . Zool . 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp 1977. Observations on the foraging behavior phyllostomid bats , Ectophylla alba and Vampyrum and avian prey of the Neotropical carnivorous bat , Vampyrum spec spectrum , in Costa Rica. J . Mamm . 44:186-89. trum . J . Mamm . 58:469-78.

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**

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). . . Res . Rept .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) appear to forage singly. As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am . Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool . 47:140 15. , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on foraging behavior and avian prey phyllostomid bats , Ectophylla albaand Vampyrumof the Neotropicalthecarnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but March. ๖ ฺ8๑ problem continues. BaiTett , o. 1935. Notes concerning manatees and ด 16:216-20. dugongs. 8ด Frantzius, A . von. 1869. Die saugethiere Costaricas. ด ( Berlin ) 35:247-325. ด ๑ ไ ์ Hartman , D. 1979. ิ ้ Publica ด Special ๖ ๑ tion no. 5. American Society of Mammalogists. Husar, L. 1977. The Wfest Indian manatee ้

pup , one half -grown male, and one nearly adult-sized female that had not yet bred. In the roost the bats form a contact cluster. They appear to be highly social , and group composition may remain stable for long periods. When disturbed in the roost , the whole group emits a soft chittering sound . This noise is characteristic of the species and constitutes a good marker for the presence of in a given hollow tree. Captive groups have demonstrated a wide variety of social signals. Animals ิ ด ด ด separated from their group , for example , for foraging , ด ด ด บ .ร . are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost-litter analyses agree that It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds ไ such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in - groups at night . Some of the species taken match the bats and forms such as sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can somefavors substantial vertebrates such as birds , feed every night , that a foraging are strik - times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insufvampire. One reported that individuals of fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc- the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ิdo not fan their seems a distinct possibility. It is clear that these bats have ฺand ฺ pires ้ complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spetropics but is never dense. This is presumably a function cies avoid these trees , with the exception of ฺwhich ๖may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

two sites in which

SPECIES ACCOUNTS

roosts are now kept secret Goodwin , G. G . , and Greenhall , A. 1% 1. A review for fear that some small museum will feel the need bats of Trinidad and Tobago. M. ด ด . ด complete its skin collection at the expense of the remnantto of the 122 : 191 301 . populations. Fieldworkers are encouraged to observe and Greenhall , A . M. 1%8. Notes on the behavior enjoy these large bats, but unnecessary disturbance false vampire bat. 8ด ด49:337-40. of the and collecting should be avoided whenever possible. Peterson , R . L. , and Kirmse, p. 1%9. Notes on ไ ฺthe false 8ด ด47:140 15. vampire bat , in Panama. ด , Stiles, F. G.; and Bradbury, J . 1977. Casebeer, R . Linsky, B . and Nelson , c. 1%3. The Vehrencamp Observations on the foraging behavior and avian prey phyllostomid bats , and of the Neotropical carnivorous ฺin Costa Rica. 8 ไ ด ด44:186-89. ด 8ด ด58:469 78. bat ,

-

-

-

**

501

MAMMALS

Anonymous. 1973. Baby Snooks eats and performs, but pup , one half -grown male, and one nearly adult-sized , 8 March . Tribune problem continues. Tampa female that had not yet bred. In the roost the bats form a BaiTett , o. พ. 1935. Notes concerning manatees and contact cluster. They appear to be highly social , and . 16:216-20. dugongs. J. Mamm group composition may remain stable for long periods. Frantzius, A . von. 1869. Die saugethiere Costaricas. When disturbed in the roost , the whole group emits a soft . ( Berlin ) 35:247-325. . Naturgesch Archiv of the spe is - chittering sound . This noise characteristic ofthemana andbehavior Hartman , D. ร. 1979. Ecology for the presence marker a good and constitutes cies of ) inFlorida . Special Publicamanatus tee(Trichechus tree groups Captive . hollow in given a Vampyrum have tion no. 5. American Society of Mammalogists. demonstrated a wide variety of social signals. Animals Husar, ร. L. 1977. The Wfest Indian manatee (Trichechus from their group , for example , for foraging, separated ). . . Res. Rept .t Wildl . Serv FishWildl manatus are greeted on their return to the roost by an interlocking 7:1-22. ; of mouths much like that found in wolves . A rapid high Reynolds , J. E. 1979. The semisocial manatee. Review June, pp. 93-%. piping accompanied by vibration of the closed wings is a contagious behavior that seems to agitate the whole group. Bats leaving a roost to feed often emit a loud ( Vampiro Falso, screech as they depart and repeat it several times while in flight. The function of this sound is unclear, since the bats False Vampire Bat) to forage singly. appear J . Bradbury As noted above , these bats are carnivorous. Most reis the largest bat in the New World. ports of stomach , feces , or roost litter analyses agree that Vampyrum spectrum It is an imposing animal with adult weights of up to 200 birds , rodents, and other bats are the major constituents g and a wingspan approaching 80 cm. The species is the of the diet. Some fruit and insects may be taken on extreme in a series of phyllostominine bats that share a occasion , but this appears to be rare. One long - term similar physiognomy and a dietary bias for larger prey. study on the remains of prey brought back to a roost in All have large complete pinnae , canidlike muzzles , and Guanacaste showed a heavy bias for nonpasserine birds - such as anis, doves, parakeets , motmots, and trogons. elongate nose leaves. Smaller forms such as Micro nycteris feed on insects or insects plus fruit , intermediate Most of these prey are either fairly smelly or roost in mix larger in groups at night . Some of the species taken match the bats and Chrotopterus forms such as Trachops sects with small vertebrates such as lizards, and the large in weight . This same study suggested that not all bats can some favors substantial vertebrates such as birds , feed every night , that a foraging Vampyrum Vampyrum are strik times fill its stomach within an hour after leaving the rodents , and other bats. The jaws of Vampyrum ingly endowed with large canine teeth and a row of car- roost , and that the bats do not necessarily forage at great nassiallike shearing molars; the skull bears a pronounced distances from the roost. It also showed that although the sagittal crest to anchor the powerful jaw muscles. Were suckling pup in the study group was left behind in the it not to have a nose leaf , this bat could easily be taken roost , it was always “ guarded by one adult or subadult member of the group . Some evidence of provisioning by for a small dog in head form . The scientific name is a misnomer. Early explorers the adult male of a captive pair was observed in Trinidad, hearing tales of vampire bats simply assumed that the** and this may be a common behavior in this species. It largest and most fierce- looking species must be “ the would certainly account for the presence of prey remains in the floor of the roost: these are quantitatively insuf vampire. One reported that individuals of Vampyrum fanned their victims to sleep with their large wings, and ficient to account for the energy needs of all the bats in another suggested that the nose leaf and associated struc the group but are substantial enough to warrant some tures were suction devices used to remove blood . Both explanation . Since not all prey eaten are brought back to suggestions were clearly speculation , since the true vam- the roost , provisioning of youngsters ( and the baby-sitter ) ) do not fan their seems a distinct possibility. It is clear that these bats have , and Diphylla , Diaemus pires (Desmodus complex family lives that may show many parallels with victims and lap blood instead of suck it. roosts in groups of up to five in the other mammalian carnivores such as canids. Vampyrum spectrum Vampyrum occurs over a wide range of the American dark upper reaches of hollow trees. Most other bat spe tropics but is never dense. This is presumably a function cies avoid these trees , with the exception of Saccopteryx , which may roost near the tree opening . The of its status as a large carnivore. This sparse dispersion bilineata few groups that have been captured appear to consist of has been exacerbated in recent years by museum skin one adult pair and their recent young . One such group in collectors , pet fanciers, and careless field biologists who Guanacaste consisted of the two adults , one suckling will not leave the bats done. Trinidad and Costa Rica are

-

two sites in which Vampyrum roosts are now kept secret for fear that some small museum will feel the need complete its skin collection at the expense of the remnantto populations. Fieldworkers are encouraged to observe and enjoy these large bats, but unnecessary disturbance collecting should be avoided whenever possible. and

SPECIES ACCOUNTS

Goodwin , G. G. , and Greenhall , A. M. 1% 1. A review of the bats of Trinidad and Tobago. Bull. Am. Mus . Nat. Hist122: 191 -301 . Greenhall , A . M. 1%8. Notes on the behavior of the false vampire bat. J. Mamm . 49:337-40. Peterson , R . L. , and Kirmse , p. 1%9. Notes on Vam pyrum spectrum , the false vampire bat , in . Panama Can . J. Zool. 47:140 15. , ร.; Stiles , F. G .; and Bradbury, J . Casebeer, R . ร.; Linsky, B . ร.; and Nelson , c. 1%3. The Vehrencamp . Observations on the foraging behavior and avian1977 phyllostomid bats , Ectophylla prey alba and Vampyrum of the Neotropical carnivorous bat , Vampyrum spectrum , in Costa Rica. J. Mamm spec . 44:186-89. trum . J. Mamm . 58:469-78.

-

-

-

-

**

-

-

-

501

MAMMALS

ff -, •f

f

Tampa Tribune , F f

f

hZ /f f • »f f f 'f \ Archiv . Naturgesch . V [ h Z • Ecology and behavior of the mana !f ร, f ) in Florida . ( manatus tee Trichechus Z f Vampyrum f f \ (Trichechus • ร, «f f manatus ) . ร (พ( Fish Wildl . Serv .t Wildl . Res . Rept . f ' h Z 7 f Reviewg • y f ff 8 •f S f Z ( Vampiro

False

Vampire

f

P

f nycteris

Trachops

Vampyrum

Chrotopterus

f

•

•

•

g

f

•

f

•

Vampyrum spectrum

f

•

f

•

• •

f

•

fZ Z

Z Saccopteryx

•

• Ectophylla

alba

f J . Mamm .

•f 7 S J f

Vampyrum

hZ /f

Z

f

•

•

Z

Vampyrum

•

f Q Q

f

•

f

Z

h f

•

Vampyrum

f

f

spectrum ,

• 9f ร,‘

Z

f

•

f

f

• f ร,‘

f

f ff f Bull Am .

• »< 7S 7»

. Musf Nat . h7 v f 7 Z • »f

F

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , FZ

.

293

.

Zygophyllaceae 183

JZ

•150, •514 , 589

.

•»•5

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper PZ \> Yellow spotted byttnena borer, 7Z Yellow warbler, \Z Yiquirro, PZ Young , A . M . , ix , 724-26 Yuca , 66 , 0Z

.

f709-10

•

Yams , 70

• F

Xantusiidae . 359, 372

275

.

Zopilota , vZ• FZ Zopilote cabecirrojo, PZ Zorra mochila , FZ Zorro pelon FZ

•

•

•621 627 •147 , 208 416

•

.

•194 210, 241. 245 299, 323, 628, 633-34 •215, 7 •215 'Z f215 •215 Xylocopidae , 335, 633. See

.

4

Z

INDEX Vriesea ( com. ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheriaindica, 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewicziococcinea,

.

.

-

—

-

.

-,

.

Weinmannia, 144 45 , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla, 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia, 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia, 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastistimais, Xamhoxylumsetulosum, 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex, 275 Xestoblattahamata, 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus, 462 Zalophus californianus

. 428

fairchildiana, 349-50 skinneri, 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum, 147 mayanum, melanostichum, 144 panamense, 140 , (seealsoXan setulosum, thoxylumsetulosum) Zapote, 704 Zarhynchuswagleri. 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays, 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata, 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana, 150 , 589 Xiphorhynchus, 514 , Xylocopa,

.

Zonotrichiacapensis, 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones, turbata, , 208416 Xylosma, 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata, 215 gualenensis, 215777 -79 muscaria. 215 viridis, 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , f • 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

าพ ๖

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

/

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

• v • , J. f 0 J Zeledon Z , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , ว ้ 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

าพ

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

[

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

้ ๓ ้ , J. ว ฐ „ Zeledon ํ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

".

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , ห ฟ 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

--also

.

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22 9

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฟ ” ฟ , J. ห ว ณ Zeledon ส , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

.)

( com

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136

.

๖

-

H., 119 Wendland , c.,4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

,3, J. c. 4,502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

๖

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136

.

-

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

816

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

าพ ี

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

น. .

INDEX Vriesea orosiense

.)

( com

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 - , H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker - ป , 7056 stick Walking , 188 Walnut

Walter . K. ร,*282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 อ

.

1

-

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร, L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

น

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com. ) orosiense 593 Vulture 404 , 456, 502

.

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711-12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 -39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 -16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

Weinmannia, 144 45 Welfia georgii , 74 136, F

.

.

138-40, 146,

Wendland H . , 119 ฟ 4 , 6, 119 Werkle , ห West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498-500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 -83 White- banded sphinxlet . 680-81 White faced capuchin 458 -60 White faced monkey 202, 320, 329, 439, 458-60 , 764 White fronted nunbird , 587-90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481-83 . See also Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 211-12 Wild ginger 221-22

---

9

.

.. .

.

-

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 Waltheria indica , 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345-46 Wild papaya , 264 -65 , 633 Wild pineapple , 197 -200 Wild pitahaya , 252-53 Wild plantain 249-51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 Wilsonia pusilla , 526 Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 Witheringia , 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 -18 Wyeomyia, 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastis timais , Xamhoxylum setulosum, 195 Zamhoxylum Xantusiidae 359, 372 Xenarescus 709 Xenomyrmex, 275 Xestoblatta hamata , 693- 96

630

.

.

Ximenia americana , 150, Xiphorhynchus, 514 , 589 , Xylocopa,

.

Yams , 70 Year of Costa Rican Natural History, A ,

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750-5 / Yellow spotted byttnena borer, 681-82 Yellow warbler, 575-76 Yiquirro, 610-11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 -17

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 Zamia , 349-50 Zanate 601 3

.

-

Zancudo , 727 -29

751

Zamhoxylum, 147 , 586, mayanum, melanostichum , 144 panamense , 140 , ( see also Xan setulosum, thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 -69

147

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

ฟ ” ฟ ว ณ Zeledon , J. ห ส Zeledon , R . , 8

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8 Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 688-91

.

-

. See also

293

.

Zonotrichia capensis , 618 Zopilota , 392 93 , 398 -99 Zopilote cabecirrojo, 560-62 Zorra mochila , 468 -69 Zorro pelon 468 -69

.

-10

Xicote , 777 -79 Ximenez , F- , 1

621 627 208 416

Xylophones, turbata , Xylosma, 147 ,

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 gualenensis , 215 777 -79 muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

-

5

INDEX Vriesea orosiense

.)

( com

. 593

.

Vulture 404 , 456, 502-5, 507, 517, 552, 560

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

coccinea , 136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711 - 12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 - 39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 - 16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio

.

.

-

—

-

.

พรี

-,

, 144 45 Weinmannia Welfia georgii , 74 136,

.

.

138-40, 146,

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498 - 500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 - 83 White- banded sphinxlet . 680 - 81 White faced capuchin 458 - 60 White faced monkey 202, 320, 329, 439, 458 - 60 , 764 White fronted nunbird , 587 - 90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481 - 83 . See

--also

.

.. .

.

Deer Whitmore , J . L., 281 -82, 331-33 Wild bitter gourdlet . 211 - 12 Wild ginger 221 - 22

816

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 Waltheria indica , 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345 - 46 Wild papaya , 264 - 65 , 633 Wild pineapple , 197 - 200 Wild pitahaya , 252 - 53 Wild plantain 249 - 51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 pusilla , 526 Wilsonia Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 , 715 Witheringia Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 - 18 , 640 Wyeomyia

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum Zamhoxylum

.

630

Xantusiidae 359, 372 709 10 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 Xicote , 777 79

.

,

-

.

Yams , 70 Year of Costa

Rican

-

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 Zamia , 349 - 50 Zanate 601 3

.

. 428

- 50

-

Zancudo , 727 - 29 , 147 , 586, 751 Zamhoxylum mayanum , 147 , 144 melanostichum

, 140 panamense , ( see also Xan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 - 69

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia. 616 - 18

Zeledonia

coronata

, 616

- /8

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708

.

Zompopa , 688

- 91

-

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 , 215 777 - 79 gualenensis muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

, A,

History

.

.

- 96

293

Natural

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750 - 5 / Yellow spotted byttnena borer, 681 - 82 Yellow warbler, 575 - 76 Yiquirro, 610 - 11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 - 17

capensis , 618 Zopilota , 392 93 , 398 - 99 Zopilote cabecirrojo, 560 - 62 Zorra mochila , 468 -69 Zorro pelon 468 - 69

195. See also

Ximenia americana , 150, , 514 , 589 Xiphorhynchus , Xylocopa ,

.

, 621 627 147, 208 416

Zonotrichia

-

Ximenez , F- , 1

Xylophones turbata , Xylosma ,

-

5

INDEX Vriesea ( com. ) orosiense 593 Vulture 404 , 456, 502

.

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711-12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 -39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 -16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio coccinea,

.

.

-

—

-

.

-,

Weinmannia, 144 45 Welfia georgii , 74 136, F

.

.

138-40, 146,

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498-500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 -83 White- banded sphinxlet . 680-81 White faced capuchin 458 -60 White faced monkey 202, 320, 329, 439, 458-60 , 764 White fronted nunbird , 587-90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481-83 . See also Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 211-12 Wild ginger 221-22

---

816

.

.. .

.

-

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 Waltheria indica, 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345-46 Wild papaya , 264 -65 , 633 Wild pineapple , 197 -200 Wild pitahaya , 252-53 Wild plantain 249-51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 Wilsonia pusilla , 526 Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 Witheringia, 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 -18 Wyeomyia, 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastis timais, Xamhoxylumsetulosum, 195 Zamhoxylum Xantusiidae 359, 372 Xenarescus 709 Xenomyrmex, 275 Xestoblatta hamata, 693- 96

630

.

.

Ximenia americana, 150, Xiphorhynchus, 514 , 589 , Xylocopa,

.

Yams , 70 Year of Costa Rican Natural History, A ,

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750-5 / Yellow spotted byttnena borer, 681-82 Yellow warbler, 575-76 Yiquirro, 610-11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 -17

-

.

Zacryptocerus, 462 Zalophus californianus

. 428

fairchildiana, 349-50 skinneri, 349 -50 Zamia , 349-50 Zanate 601 3

.

-

Zancudo , 727 -29

751

Zamhoxylum, 147 , 586, mayanum, melanostichum, 144 panamense, 140 , ( see also Xan setulosum, thoxylum setulosum ) Zapote , 704 Zarhynchuswagleri . 614 - 16 Zarigiieya 468 -69

147

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8

.

Zeledonia 616 -18 Zeledonia coronata, 616 - / 8 Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 688-91

.

-

. See also

293

.

Zonotrichia capensis , 618 Zopilota , 392 93 , 398 -99 Zopilote cabecirrojo, 560-62 Zorra mochila , 468 -69 Zorro pelon 468 -69

.

-10

Xicote , 777 -79 Ximenez , F- , 1

621 627 208 416

Xylophones, turbata, Xylosma, 147 ,

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata, 215, 111 gualenensis, 215 777 -79 muscaria. 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

-

5

INDEX Vriesea ( com . ) orosiense 593 Vulture 404 , 456, 502

.

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711-12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 -39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 -16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

Weinmannia , 144 45 Welfia georgii , 74 136,

.

ี

.

138-40, 146,

Wendland H . , 119 ฺ ด 4 , 6, 119 Werkle , ] West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498-500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 -83 White- banded sphinxlet . 680-81 White faced capuchin 458 -60 White faced monkey 202, 320, 329, 439, 458-60 , 764 White fronted nunbird , 587-90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481-83 . See also Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 211-12 Wild ginger 221-22

---

๊

.

.. .

.

-

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 Waltheria indica , 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345-46 Wild papaya , 264 -65 , 633 Wild pineapple , 197 -200 Wild pitahaya , 252-53 Wild plantain 249-51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 Wilsonia pusilla , 526 Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 Witheringia , 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 -18 Wyeomyia , 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195 Zamhoxylum Xantusiidae 359, 372 Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

Ximenia americana , 150, Xiphorhynchus , 514 , 589 , Xylocopa ,

.

Yams , 70 Year of Costa Rican Natural History, A ,

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750-5 / Yellow spotted byttnena borer, 681-82 Yellow warbler, 575-76 Yiquirro, 610-11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 -17

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 Zamia , 349-50 Zanate 601 3

.

-

Zancudo , 727 -29

751

Zamhoxylum , 147 , 586, mayanum , melanostichum , 144 panamense , 140 , ( see also Xan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 -69

147

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

ฺ ฌ ฺ ู ํ ด Zeledon , J. ] ไ Zeledon , R . , 8

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8 Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 688-91

.

-

. See also

293

.

Zonotrichia capensis , 618 Zopilota , 392 93 , 398 -99 Zopilote cabecirrojo, 560-62 Zorra mochila , 468 -69 Zorro pelon 468 -69

.

-10

Xicote , 777 -79 Ximenez , F- , 1

621 627 208 416

Xylophones , turbata , Xylosma , 147 ,

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 gualenensis , 215 777 -79 muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

-

5

INDEX Vriesea ( com . ) orosiense 593 Vulture 404 , 456, 502

.

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711-12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 -39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 -16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

Weinmannia , 144 45 Welfia georgii , 74 136, F

.

.

138-40, 146,

Wendland H . , 119 Werkle , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498-500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 -83 White- banded sphinxlet . 680-81 White faced capuchin 458 -60 White faced monkey 202, 320, 329, 439, 458-60 , 764 White fronted nunbird , 587-90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481-83 . See also Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 211-12 Wild ginger 221-22

---

.

.. .

.

-

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 Waltheria indica , 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345-46 Wild papaya , 264 -65 , 633 Wild pineapple , 197 -200 Wild pitahaya , 252-53 Wild plantain 249-51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 Wilsonia pusilla , 526 Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 Witheringia , 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 -18 Wyeomyia , 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195 Zamhoxylum Xantusiidae 359, 372 Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

Ximenia americana , 150, Xiphorhynchus , 514 , 589 , Xylocopa ,

.

Yams , 70 Year of Costa Rican Natural History, A ,

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750-5 / Yellow spotted byttnena borer, 681-82 Yellow warbler, 575-76 Yiquirro, 610-11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 -17

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 Zamia , 349-50 Zanate 601 3

.

-

Zancudo , 727 -29

751

Zamhoxylum , 147 , 586, mayanum , melanostichum , 144 panamense , 140 , ( see also Xan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 -69

147

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

Zeledon , J. Zeledon , R . , 8

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8 Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 688-91

.

-

. See also

293

.

Zonotrichia capensis , 618 Zopilota , 392 93 , 398 -99 Zopilote cabecirrojo, 560-62 Zorra mochila , 468 -69 Zorro pelon 468 -69

.

-10

Xicote , 777 -79 Ximenez , F- , 1

621 627 208 416

Xylophones , turbata , Xylosma , 147 ,

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 gualenensis , 215 777 -79 muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

-

5

INDEX Vriesea orosiense

น. .

.)

( com

593 Vulture 404 , 456, 502-5, 507, 517, 552, 560

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner - , H . , 187, 239-40, 311-18 Wake , M . H., 400^*01 Walker -

.

.

ious insects Warscewicz , J . , 118

.

-

coccinea , 136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711 - 12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 - 39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 - 16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio

.

.

-

—

-

.

-,

, 144 45 Weinmannia Welfia georgii , 74 136, อ

.

1

.

138-40, 146,

Wendland H . , 119 • 4 , 6, 119 Werkle , f West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498 - 500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 - 83 White- banded sphinxlet . 680 - 81 White faced capuchin 458 - 60 White faced monkey 202, 320, 329, 439, 458 - 60 , 764 White fronted nunbird , 587 - 90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481 - 83 . See

--also

.

.. .

.

Deer Whitmore , J . L., 281 -82, 331-33 Wild bitter gourdlet . 211 - 12 Wild ginger 221 - 22 /

.

.

ป

Walking stick , 705-6 Walnut , 188 Walter K . ร,*282-92 Waltheria indica , 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345 - 46 Wild papaya , 264 - 65 , 633 Wild pineapple , 197 - 200 Wild pitahaya , 252 - 53 Wild plantain 249 - 51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 pusilla , 526 Wilsonia Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 , 715 Witheringia Wolf 500, 716 Wolfe , K ., 659 Wood , ร, L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 - 18 , 640 Wyeomyia

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

น

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum Zamhoxylum

.

630

Xantusiidae 359, 372 709 10 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 Xicote , 777 79

.

,

-

.

Yams , 70 Year of Costa

Rican

-

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 Zamia , 349 - 50 Zanate 601 3

.

. 428

- 50

-

Zancudo , 727 - 29 , 147 , 586, 751 Zamhoxylum mayanum , 147 , 144 melanostichum

, 140 panamense , ( see also Xan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 - 69

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

• v • 0 J Zeledon , J. f Z Zeledon , R . , 8

Zeledonia. 616 - 18

Zeledonia

coronata

, 616

- /8

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708

.

Zompopa , 688

- 91

-

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 , 215 777 - 79 gualenensis muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

, A,

History

.

.

- 96

293

Natural

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750 - 5 / Yellow spotted byttnena borer, 681 - 82 Yellow warbler, 575 - 76 Yiquirro, 610 - 11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 - 17

capensis , 618 Zopilota , 392 93 , 398 - 99 Zopilote cabecirrojo, 560 - 62 Zorra mochila , 468 -69 Zorro pelon 468 - 69

195. See also

Ximenia americana , 150, , 514 , 589 Xiphorhynchus , Xylocopa ,

.

, 621 627 147, 208 416

Zonotrichia

-

Ximenez , F- , 1

Xylophones turbata , Xylosma ,

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush ,] ฺ ด 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

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. 593

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9

๊

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฺ ฌ ฺ , J. ] ู ํ ด Zeledon ไ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria indica , 778 , 283 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 Witheringia , 715 Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly Weinmannia , 144 - 45 WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 า Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush ,] ฺ ด 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet Xenomyrmex , 275 -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 293 Ximenia americana , 150 peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

.

. 593

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9

๊

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican Natural

History , A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus Zalophus californianus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฺ ฌ ฺ , J. ] ู ํ ด Zeledon ไ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

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.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

๕๐

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria indica , 778 , 283 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 Witheringia , 715 Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly Weinmannia , 144 - 45 WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , ห ฟ 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet Xenomyrmex , 275 -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 293 Ximenia americana , 150 peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

.

. 593

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

พร

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican Natural

History , A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus Zalophus californianus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฟ ” ฟ , J. ห ว ณ Zeledon ส , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX [ f

V

f593

.

, 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 • 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 •136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 7Z Waspmantispid ; Ma , 505 Water Duck birds -6. ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect , /Z hyacinth Water , 427 opossum Water ; , 35 46. Dryseason Weather Seasonality , 52122 effect onbirds , \Z Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

.

.

-

—

-

.

Z , 136, 138-40, 146,

.

• 0 •74

.

;

า

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper . ' , FZP WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , vZ White fatima -banded White . PZ sphinxlet -banded White faced capuchinFZ , 329 , 439 , facedmonkey 202,320 White FZ•764 , 'Z•606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 7Zf , 263 White tailed deer248

.

---

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet . 7Z Wildginger7Z

816

, \Z Wildnutmeg , 0Z•633 Wildpapaya , 'ZP Wildpineapple , vZ Wildpitahaya Wildplantain4Z , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 •526 Wind38 40 42-44, 632 , 283 . Windpollination Polli , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula •715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , / Worm snake , 558 , 5057, 524 Wren -60 , />/Z Wrenthrush •640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

-

80

•

f709-10

, 'Z Xicote , F-, 1 Ximenez

•

•»•5

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow \> PZ -linedgrasshopper , 7Z borer byttnena Yellow spotted , \Z Yellow warbler , PZ Yiquirro , A. M., ix, 724 Young -26 , 66, 0Z Yuca

-

.

•462 •

f428

4Z

• 4Z , 4Z Zamia Zanate 7 J

.

Z

, 'Z Zancudo

, 751 147,586

•

•147

•144 140

• •

127,131(seealso Z •704 f 0Z [

. .Z

ZarigiieyaFZ

Zarza 'Z •

, 607 antshrike Zebra -striped , J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia . /Z

• /Z > F

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, FZ Zompopa

• F

.

183 Zygophyllaceae

JZ

293

•150 , , 589 •514

.

, 70 Yams

.

Xantusiidae . 359, 372

275

.

, vZ• FZ Zopilota , PZ cabecirrojo Zopilote , FZ Zorramochila Zorro pelon FZ

630 •195 . Seealso

•

•

•621 627 , 208416 •147

•

.

, 241 •194210 . 245299, 323, , 633 628 -34 , 7 •215 •215 ' Z f215 •215 , 335 , 633 . See Xylocopidae

.

4

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

าพ

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush ,] ฺ ด 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

๊

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฺ ฌ ฺ , J. ] ู ํ ด Zeledon ไ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

.

. -

. .

.

1

-

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , ว ้ 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22 [

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

้ ๓ ้ , J. ว ฐ „ Zeledon ํ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com. ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, ), 76 191 Wasp(aculeate , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall effectonclimate ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effectonbirds Web -footedfrog,415-16 Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia, 144 45 , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , ว ้ 4, 6, 119 Werkle West-Eberhard M. J., 758 -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded Whitefaced 458 -60 capuchin , 329 , 439 , 202,320 Whitefacedmonkey 458-60 , 764 ,587-90 , 606 Whitefronted nunbird peccary Whitelipped 471 , 388 Whitenecked heron , 547 White antbird -plumed , 481 -83 . See , 263 Whitetailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

[

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 ,M F . 191-93 Willson , D. E., 261-62, 426 Wilson ^ 7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , by wind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Woodboring beetle -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Woodduck , 546 Woodhaunter , 657 . See also Satyridae Woodnymph , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Woodpigeon Woodstork388 . See tree species accounts Woodtraits , 609 , 567 Woodwarbler Woodwren . 509 , 427 Woolyopossum WorldWarII, 6 67 , xi States WorldWildlife FundUnited , 406 Wormsnake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia, 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastis timais , Xamhoxylum setulosum, 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex, 275 Xestoblatta hamata , 693- 96

630

.

.

.

-10

, 777 -79 Xicote , F-, 1 Ximenez

293

, Ximenia americana , 150 , 589 Xiphorhynchus, 514 , Xylocopa,

.

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

621 627

Xylophones, turbata , , 208416 Xylosma, 147

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellowheaded vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellowspotted , 575-76 Yellowwarbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 Yuca, 66, 114 -17

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum, 147 mayanum, melanostichum , 144 panamense , 140 , (seealsoXan setulosum, thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

้ ๓ ้ , J. ว ฐ „ Zeledon ํ , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorra mochila Zorro pelon468 -69

-

.

.

183 Zygophyllaceae

-

5

ะ. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 ร * H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker ร พ , 7056 stick Walking , 188 Walnut

Walter . K. อ*เ 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland ,] ฺ ด 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22 ๊

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , อ*L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

ะ

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฺ ฌ ฺ , J. ] ู ํ ด Zeledon ไ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , ว ้ 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

[

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

้ ๓ ้ , J. ว ฐ „ Zeledon ํ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria indica , 778 , 283 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 Witheringia , 715 Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly Weinmannia , 144 - 45 WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet Xenomyrmex , 275 -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 293 Ximenia americana , 150 peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

.

. 593

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

พร

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican Natural

History , A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus Zalophus californianus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

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-

..

. -

-

-

.

.

-

.

-

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—

-

.

าพ

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria indica , 778 , 283 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 Witheringia , 715 Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly Weinmannia , 144 - 45 WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 า Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet Xenomyrmex , 275 -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 293 Ximenia americana , 150 peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

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9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican Natural

History , A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus Zalophus californianus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

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9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

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9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

Wendland H . , 119 Werkle , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 5 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

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9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

.

. -

. .

.

1

-

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c . , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

าพ

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c . , 3 - 4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

ะ. .

INDEX

( com

Vriesea orosiense

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 ร * H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker ร พ , 7056 stick Walking , 188 Walnut

Walter . K. อ*เ 282-92

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

indica ,

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , อ*L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

ะ

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus Xenomyrmex , 275 Xestoblatta hamata , 693 - 96

630

.

.

.

.

Rican Natural

.

-

.

, 462 Zacryptocerus Zalophus californianus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa capensis

, 618

.

.

183 Zygophyllaceae

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

History , A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

293

, Ximenia americana , 150 , 589 , 514 Xiphorhynchus , Xylocopa ,

, 621 627 147, 208416

, 70 Yams Year of Costa

Zonotrichia

-10

, 777 - 79 Xicote , F-, 1 Ximenez

Xylophones turbata , Xylosma ,

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX Vriesea orosiense

.)

( com

. 593

.

Vulture 404 , 456, 502-5, 507, 517, 552, 560

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

coccinea , 136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711 - 12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 - 39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 - 16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio

.

.

-

—

-

.

พร

-,

Weinmannia , 144 45 Welfia georgii , 74 136, F

.

.

138-40, 146,

Wendland H . , 119 Werkle , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498 - 500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 - 83 White- banded sphinxlet . 680 - 81 White faced capuchin 458 - 60 White faced monkey 202, 320, 329, 439, 458 - 60 , 764 White fronted nunbird , 587 - 90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481 - 83 . See

--also

.

.. .

.

Deer Whitmore , J . L., 281 -82, 331-33 Wild bitter gourdlet . 211 - 12 Wild ginger 221 - 22

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 Waltheria indica , 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345 - 46 Wild papaya , 264 - 65 , 633 Wild pineapple , 197 - 200 Wild pitahaya , 252 - 53 Wild plantain 249 - 51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 pusilla , 526 Wilsonia Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 Witheringia , 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 - 18 Wyeomyia , 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum Zamhoxylum

.

630

Xantusiidae 359, 372 709 10 Xenarescus Xenomyrmex , 275 Xestoblatta hamata , 693 Xicote , 777 79

.

,

-

.

Yams , 70 Year of Costa

Rican Natural

-

.

, 462 Zacryptocerus Zalophus californianus , 349 fairchildiana skinneri , 349 - 50 Zamia , 349 - 50 Zanate 601 3

.

293

. 428

- 50

-

Zancudo , 727 - 29 , 147 , 586, 751 Zamhoxylum mayanum , 147 , 144 melanostichum

panamense , 140 , ( see also Xan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 - 69

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

Zeledon , J. Zeledon , R . , 8

Zeledonia. 616 - 18

Zeledonia

coronata

, 616

- /8

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708

.

Zompopa , 688

- 91

-

.

.

Zygophyllaceae 183

- 96

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 , 215 777 - 79 gualenensis muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

History , A ,

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750 - 5 / Yellow spotted byttnena borer, 681 - 82 Yellow warbler, 575 - 76 Yiquirro, 610 - 11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 - 17

capensis , 618 Zopilota , 392 93 , 398 - 99 Zopilote cabecirrojo, 560 - 62 Zorra mochila , 468 -69 Zorro pelon 468 - 69

195. See also

Ximenia americana , 150, , 514 , 589 Xiphorhynchus , Xylocopa ,

.

, 621 627 147, 208 416

Zonotrichia

-

Ximenez , F- , 1

Xylophones turbata , Xylosma ,

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

.)

( com

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136

.

ห

-

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria indica , 778 , 283 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 Witheringia , 715 Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly Weinmannia , 144 - 45 WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush ,] ฺ ด 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet Xenomyrmex , 275 -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 293 Ximenia americana , 150 peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

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. 593

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9

๊

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican Natural

History , A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus Zalophus californianus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฺ ฌ ฺ , J. ] ู ํ ด Zeledon ไ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

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. 593

.

.

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9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , c . , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c . , 3 - 4 , 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

ป-

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria indica , 778 , 283 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 Witheringia , 715 Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly Weinmannia , 144 - 45 WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 า Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet Xenomyrmex , 275 -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 293 Ximenia americana , 150 peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

.

. 593

.

.

.

-

*

.

.

.

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.

.

*

.. .

-

..

. -

-

-

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.

-

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-

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—

-

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.

. -

. .

.

;

-

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican Natural

History , A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus Zalophus californianus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

าพ

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com. ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheriaindica, 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewicziococcinea,

.

.

-

—

-

.

-,

๖

.

Weinmannia, 144 45 , 13840, 146 , Welfia georgii , 74 136

.

-

H., 119 Wendland , c . , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

816

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla, 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia, 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia, 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastistimais, Xamhoxylumsetulosum, 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex, 275 Xestoblattahamata, 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus, 462 Zalophus californianus

. 428

fairchildiana, 349-50 skinneri, 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum, 147 mayanum, melanostichum, 144 panamense, 140 , (seealsoXan setulosum, thoxylumsetulosum) Zapote, 704 Zarhynchuswagleri. 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays, 88 92

, 607 antshrike Zebra -striped

, J. c . , 3 - 4 , 502 Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata, 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana, 150 , 589 Xiphorhynchus, 514 , Xylocopa,

.

Zonotrichiacapensis, 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones, turbata, , 208416 Xylosma, 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata, 215 gualenensis, 215777 -79 muscaria. 215 viridis, 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

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816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

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9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria indica , 778 , 283 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 Witheringia , 715 Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly Weinmannia , 144 - 45 WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 า Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush ,] ฺ ด 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet Xenomyrmex , 275 -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 293 Ximenia americana , 150 peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

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๊

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican Natural

History , A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus Zalophus californianus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฺ ฌ ฺ , J. ] ู ํ ด Zeledon ไ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya plantain Wild 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E, 546 47, 607 Willis ^01 -8 Walker ,MF . 191-93 Willson , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia K. 282 Walter Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Wind pollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Wasp mantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds 70 89 . See tree species accounts Wood traits 77, 97, 2089, 212,222,246, Weevil , 609 , 567 Wood warbler , 627 , 271 , 324 , 326 , 348 , 629 255 Wood wren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World FundUnited , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 West Indian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , faced 202,320 monkey White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wild ginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

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9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow grassquit -faced , 209 Yellow vireo -green , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

277 - 78 Zarza

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorra mochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX

ี. .

Vriesea ( com. ) orosiense 593 Vulture 404 , 456, 502

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner ๐1H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

-6 ๗ 282-92 Walter. K . ป1

.

.

Waltheria indica,

ious insects Warscewicz , J . , 118

-

136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711-12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 -39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 -16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio coccinea,

.

.

-

—

-

.

-,

Weinmannia, 144 45 Welfia georgii , 74 136, F

.

.

138-40, 146,

Wendland H . , 119 ฺ ด 4 , 6, 119 Werkle , ] West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498-500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 -83 White- banded sphinxlet . 680-81 White faced capuchin 458 -60 White faced monkey 202, 320, 329, 439, 458-60 , 764 White fronted nunbird , 587-90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481-83 . See also Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 211-12 Wild ginger 221-22

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.

.. .

.

-

.

.

Walker ๐ ๕ Walking stick , 705 Walnut , 188

778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345-46 Wild papaya , 264 -65 , 633 Wild pineapple , 197 -200 Wild pitahaya , 252-53 Wild plantain 249-51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 Wilsonia pusilla , 526 Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 Witheringia, 715 Wolf 500, 716 Wolfe , K ., 659 Wood , ป1L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 -18 Wyeomyia, 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

ี

-

Xamhomonas, 80 Xanthopastis timais, Xamhoxylumsetulosum, 195 Zamhoxylum Xantusiidae 359, 372 Xenarescus 709 Xenomyrmex, 275 Xestoblatta hamata, 693- 96

630

.

.

Ximenia americana, 150, Xiphorhynchus, 514 , 589 , Xylocopa,

.

Yams , 70 Year of Costa Rican Natural History, A ,

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750-5 / Yellow spotted byttnena borer, 681-82 Yellow warbler, 575-76 Yiquirro, 610-11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 -17

-

.

Zacryptocerus, 462 Zalophus californianus

. 428

fairchildiana, 349-50 skinneri, 349 -50 Zamia , 349-50 Zanate 601 3

.

-

Zancudo , 727 -29

751

Zamhoxylum, 147 , 586, mayanum, melanostichum, 144 panamense, 140 , ( see also Xan setulosum, thoxylum setulosum ) Zapote , 704 Zarhynchuswagleri . 614 - 16 Zarigiieya 468 -69

147

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

ฺ ฌ ฺ ู ํ ด Zeledon , J. ] ไ Zeledon , R . , 8

.

Zeledonia 616 -18 Zeledonia coronata, 616 - / 8 Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 688-91

.

-

. See also

293

.

Zonotrichia capensis , 618 Zopilota , 392 93 , 398 -99 Zopilote cabecirrojo, 560-62 Zorra mochila , 468 -69 Zorro pelon 468 -69

.

-10

Xicote , 777 -79 Ximenez , F- , 1

621 627 208 416

Xylophones, turbata, Xylosma, 147 ,

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata, 215, 111 gualenensis, 215 777 -79 muscaria. 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

-

5

ท. ว

INDEX

’

: ว

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 ร / H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker ร พ , 7056 stick Walking , 188 Walnut

Walter . K. บ/' 282-92 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . Nox coloration Warning

-

iousinsects , J., 118 Warscewicz

, 138, 140 ้136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , ™ํ Waspmantispid ; Ma , 505 Water Duck birds -6. ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect , [ํ hyacinth Water , 427 opossum Water ; , 35 46. Dryseason Weather Seasonality , 52122 effect onbirds , แํ Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

.

.

-

—

-

.

.

ํ , 136, 138-40, 146,

้ ฐ ้74

F

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper . ๙ , \ํุ WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , ๓ํ White fatima -banded White . ุํ sphinxlet -banded White faced capuchin\ํ , 329 , 439 , facedmonkey 202,320 White \ํ้764 , ๙ํ้606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , ™ํว , 263 White tailed deer248

.

---

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet . ™ํ Wildginger™ํ

816

.

. -

.

.

.

้ 778

.

, แํ Wildnutmeg , ฐํ้633 Wildpapaya , ๙ํุ Wildpineapple , ๓ํ Wildpitahaya Wildplantain๖ํ , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 ้526 Wind38 40 42-44, 632 , 283 . Windpollination Polli , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula ้715 , 716 Wolf500 , K., 659 Wolfe , บ/ L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , [ Worm snake , 558 , 5057, 524 Wren -60 , [”[ํ Wrenthrush ้640

*

.. .

-

-

.

-

.

.

. -

. .

ท

-

80

้

ว709-10

, ๙ํ Xicote , F-, 1 Ximenez

้

้3้5

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow แ” ุํ -linedgrasshopper , ™ํ borer byttnena Yellow spotted , แํ Yellow warbler , ุํ Yiquirro , A. M., ix, 724 Young -26 , 66, ฐํ Yuca

-

.

้462 ้

ว428

๖ํ

้ ๖ํ , ๖ํ Zamia Zanate ™ „

.

ํ

, ๙ํ Zancudo

, 751 147,586

้

้147

้144 140

้ ้

127,131(seealso ํ ้704 ว ฐํ :

. .ํ

Zarigiieya\ํ

Zarza ๙ํ ้

, 607 antshrike Zebra -striped , J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia . [ํ

้ [ํ ” \

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, \ํ Zompopa

้ \

.

183 Zygophyllaceae

„ํ

293

้150 , , 589 ้514

.

, 70 Yams

.

Xantusiidae . 359, 372

275

.

, ๓ํ้ \ํ Zopilota , ุํ cabecirrojo Zopilote , \ํ Zorramochila Zorro pelon \ํ

630 ้195 . Seealso

้

้

้621 627 , 208416 ้147

้

.

, 241 ้194210 . 245299, 323, , 633 628 -34 , ™ ้215 ้215 ๙ ํ ว215 ้215 , 335 , 633 . See Xylocopidae

.

๖

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

๕๐ 1

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

c. .

INDEX

Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 . - H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker . " , 7056 stick Walking , 188 Walnut

-R282-92 Walter . K. —

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria indica ,

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe ,— -L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

c

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 า Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c . , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

.

าพ ;

. -

. .

-

.

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c . , 3 - 4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush ,] ฺ ด 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

". ห

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

๊

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฺ ฌ ฺ , J. ] ู ํ ด Zeledon ไ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , ห ฟ 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22 9

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฟ ” ฟ , J. ห ว ณ Zeledon ส , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea(com.) 593 orosiense , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 ,778 Waltheria indica , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . SeealsoNox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 , 136 coccinea Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp

.636, 712, 737. , 615 297. 386 . 739 -40, 758-60, 760-62, 766

, 711-12 Waspmantispid ; Ma, 505 Water birds -6. SeealsoDuck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236-39 hyacinth Water , 427 opossum Water ; , 35 46. SeealsoDryseason Weather

—

Seasonality , 52122 effect onbirds , 415-16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

-

.

ป-

-,

.

,144 45 Weinmannia , 13840, 146 , ,74 136 Welfia georgii F

-

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682-83 White fatima -banded White .680-81 sphinxlet -banded White faced 45860 capuchin , 329 , 439 , facedmonkey 202,320 White 45860,764 ,587-90, 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481, 263 White tailed deer248 83.See alsoDeer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

---

.

.. .

.

-

816

, 34546 Wildnutmeg , 264-65,633 Wildpapaya , 197-200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 24951 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 , 526 pusilla Wilsonia Wind38 40 42-44, 632 , 283 . SeealsoPolli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula ,715 Witheringia , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . SeealsoSatyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . Seetreespecies Wood traits accounts , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 /6-18 Wrenthrush , 640 Wyeomyia

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

,80 Xamhomonas , timais Xanthopastis , 195See Xamhoxylumsetulosum also Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus ,275 Xenomyrmex ,69396 Xestoblatta hamata

630

.

, 777-79 Xicote , F-, 1 Ximenez

.

, 70 Yams ,A ,5 YearofCostaRicanNatural History

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750-5 / -linedgrasshopper , 681-82 borer byttnena Yellow spotted , 575Yellow warbler 76 , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114-17 Yuca

-

.

,462 Zacryptocerus 428 californianus Zalophus

.

,349fairchildiana 50 ,34950 skinneri , 349-50 Zamia Zanate 601 3

.

-

, 727-29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum, , 144 melanostichum , 140 panamense , (seealsoXan , setulosum ) thoxylum setulosum ,704 Zapote Zarhynchus wagleri .614-16 69 468Zarigiieya

147

127131

. .-

Zarza277-78

Zeamays,88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

.

Zeledonia 61618 ,616-/8 Zeledonia coronata

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, 150 , americana Ximenia , 589 , 514 Xiphorhynchus , , Xylocopa

.

,618 capensis Zonotrichia , 392 93,398-99 Zopilota , 56062 cabecirrojo Zopilote , 468-69 Zorramochila Zorro pelon468-69

-10

.

621 627

, Xylophones , turbata , 208416 , 147 Xylosma

.

, , 323 194210241. 245299 , 633 628 -34 , 111 , 215 fimbriata ,21577779 gualenensis muscaria .215 , 215 viridis , 335 , 633 . SeealsoXylocopa Xylocopidae

.

9

-

INDEX Vriesea ( com. ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheriaindica, 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewicziococcinea,

.

.

-

—

-

.

-,

.

Weinmannia, 144 45 , 13840, 146 , Welfia georgii , 74 136

.

ี

-

H., 119 Wendland , c.,4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

---

.

.. .

.

-

816

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla, 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia, 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia, 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastistimais, Xamhoxylumsetulosum, 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex, 275 Xestoblattahamata, 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus, 462 Zalophus californianus

. 428

fairchildiana, 349-50 skinneri, 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum, 147 mayanum, melanostichum, 144 panamense, 140 , (seealsoXan setulosum, thoxylumsetulosum) Zapote, 704 Zarhynchuswagleri. 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays, 88 92

, 607 antshrike Zebra -striped

,3, J. c. 4,502 Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata, 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana, 150 , 589 Xiphorhynchus, 514 , Xylocopa,

.

Zonotrichiacapensis, 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones, turbata, , 208416 Xylosma, 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata, 215 gualenensis, 215777 -79 muscaria. 215 viridis, 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

น. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 - , H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker - ป , 7056 stick Walking , 188 Walnut

Walter . K. ร,*282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร, L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush , 640 Wyeomyia

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

น

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

.

Rican

Natural

.

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa capensis

, 618

.

.

183 Zygophyllaceae

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

, A,

History

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

293

, Ximenia americana , 150 , 589 , 514 Xiphorhynchus , Xylocopa ,

, 621 627 147, 208416

, 70 Yams Year of Costa

Zonotrichia

-10

, 777 - 79 Xicote , F-, 1 Ximenez

Xylophones turbata , Xylosma ,

-

5

INDEX Vriesea ( com. ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheriaindica, 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewicziococcinea,

.

.

-

—

-

.

-,

.

Weinmannia, 144 45 , 13840, 146 , Welfia georgii , 74 136 อ

.

1

-

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla, 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia, 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia, 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastistimais, Xamhoxylumsetulosum, 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex, 275 Xestoblattahamata, 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus, 462 Zalophus californianus

. 428

fairchildiana, 349-50 skinneri, 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum, 147 mayanum, melanostichum, 144 panamense, 140 , (seealsoXan setulosum, thoxylumsetulosum) Zapote, 704 Zarhynchuswagleri. 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays, 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata, 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana, 150 , 589 Xiphorhynchus, 514 , Xylocopa,

.

Zonotrichiacapensis, 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones, turbata, , 208416 Xylosma, 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata, 215 gualenensis, 215777 -79 muscaria. 215 viridis, 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c.,4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

าพ

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

,3, J. c. 4,502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya plantain Wild 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E, 546 47, 607 Willis ^01 -8 Walker ,MF . 191-93 Willson , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia K. 282 Walter Wind38 40 42-44, 632 -92 Waltheria indica , 778 , 283 . See also Polli Wind pollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 Witheringia , 715 Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Wasp mantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds 70 89 . See tree species accounts Wood traits 77, 97, 2089, 212,222,246, Weevil , 609 , 567 Wood warbler , 627 , 271 , 324 , 326 , 348 , 629 255 Wood wren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly Weinmannia , 144 - 45 WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World FundUnited , 406 า Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 West Indian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet Xenomyrmex , 275 -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , faced 202,320 monkey White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 293 Ximenia americana , 150 peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wild ginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

.

. 593

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

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.

-

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-

.

—

-

.

.

. -

. .

.

;

-

.

---

-

.. .

-

.

.

-

.

.

.

9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican Natural

History , A ,

, 604 Yellow grassquit -faced , 209 Yellow vireo -green , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus Zalophus californianus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

277 - 78 Zarza

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorra mochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , ห ฟ 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

.

. -

. .

.

1

-

.

---

-

.. .

-

.

.

-

.

.

.

9

9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฟ ” ฟ , J. ห ว ณ Zeledon ส , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136 5

.

^

-

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

816

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

น. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 - , H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker - ป , 7056 stick Walking , 188 Walnut

Walter . K. ร,*282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

".

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

--also

.

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร, L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush , 640 Wyeomyia

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

น

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

.

Rican

Natural

.

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa capensis

, 618

.

.

183 Zygophyllaceae

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

, A,

History

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

293

, Ximenia americana , 150 , 589 , 514 Xiphorhynchus , Xylocopa ,

, 621 627 147, 208416

, 70 Yams Year of Costa

Zonotrichia

-10

, 777 - 79 Xicote , F-, 1 Ximenez

Xylophones turbata , Xylosma ,

-

5

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

ป- ^ -,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136 5

-

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

816

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

c. .

INDEX Vriesea orosiense

.)

( com

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 . - H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker . " , 7056 stick Walking , 188 Walnut

-R282-92 Walter . K. — indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136

.

ี

-

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

--also

.

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe ,— -L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush , 640 Wyeomyia

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

c

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

.

Rican

Natural

.

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa capensis

, 618

.

.

183 Zygophyllaceae

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

, A,

History

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

293

, Ximenia americana , 150 , 589 , 514 Xiphorhynchus , Xylocopa ,

, 621 627 147, 208416

, 70 Yams Year of Costa

Zonotrichia

-10

, 777 - 79 Xicote , F-, 1 Ximenez

Xylophones turbata , Xylosma ,

-

5

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136 5

.

^

-

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

816

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

с. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. р 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

ป-

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , f • 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

--also

.

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22 /

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , р L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush , 640 Wyeomyia

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

с

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

.

Rican

Natural

.

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

• v • , J. f 0 J Zeledon Z , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa capensis

, 618

.

.

183 Zygophyllaceae

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

, A,

History

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

293

, Ximenia americana , 150 , 589 , 514 Xiphorhynchus , Xylocopa ,

, 621 627 147, 208416

, 70 Yams Year of Costa

Zonotrichia

-10

, 777 - 79 Xicote , F-, 1 Ximenez

Xylophones turbata , Xylosma ,

-

5

INDEX Vriesea orosiense

น. .

.)

( com

593 Vulture 404 , 456, 502-5, 507, 517, 552, 560

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner - , H . , 187, 239-40, 311-18 Wake , M . H., 400^*01 Walker -

.

.

ious insects Warscewicz , J . , 118

.

-

coccinea , 136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711 - 12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 - 39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 - 16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio

.

.

-

—

-

.

าพ

-,

, 144 45 Weinmannia Welfia georgii , 74 136, F

.

.

138-40, 146,

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498 - 500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 - 83 White- banded sphinxlet . 680 - 81 White faced capuchin 458 - 60 White faced monkey 202, 320, 329, 439, 458 - 60 , 764 White fronted nunbird , 587 - 90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481 - 83 . See

--also

.

.. .

.

Deer Whitmore , J . L., 281 -82, 331-33 Wild bitter gourdlet . 211 - 12 Wild ginger 221 - 22

816

.

.

ป

Walking stick , 705-6 Walnut , 188 Walter K . ร,*282-92 Waltheria indica , 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345 - 46 Wild papaya , 264 - 65 , 633 Wild pineapple , 197 - 200 Wild pitahaya , 252 - 53 Wild plantain 249 - 51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 pusilla , 526 Wilsonia Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 , 715 Witheringia Wolf 500, 716 Wolfe , K ., 659 Wood , ร, L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 - 18 , 640 Wyeomyia

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

น

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum Zamhoxylum

.

630

Xantusiidae 359, 372 709 10 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 Xicote , 777 79

.

,

-

.

Yams , 70 Year of Costa

Rican

-

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 Zamia , 349 - 50 Zanate 601 3

.

. 428

- 50

-

Zancudo , 727 - 29 , 147 , 586, 751 Zamhoxylum mayanum , 147 , 144 melanostichum

, 140 panamense , ( see also Xan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 - 69

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia. 616 - 18

Zeledonia

coronata

, 616

- /8

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708

.

Zompopa , 688

- 91

-

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 , 215 777 - 79 gualenensis muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

, A,

History

.

.

- 96

293

Natural

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750 - 5 / Yellow spotted byttnena borer, 681 - 82 Yellow warbler, 575 - 76 Yiquirro, 610 - 11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 - 17

capensis , 618 Zopilota , 392 93 , 398 - 99 Zopilote cabecirrojo, 560 - 62 Zorra mochila , 468 -69 Zorro pelon 468 - 69

195. See also

Ximenia americana , 150, , 514 , 589 Xiphorhynchus , Xylocopa ,

.

, 621 627 147, 208 416

Zonotrichia

-

Ximenez , F- , 1

Xylophones turbata , Xylosma ,

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 456 , 502 , 552 , 517 404 , Vulture , 197 - 200 Wildpineapple -5, 507 560 , 252 - 53 Wildpitahaya plantain Wild 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 , 239 WagnerH., 187 , G. B., 224 Williamson -40,311 -18 -2525659 , M. H., 400 Wake , E, 546 47, 607 Willis ^01 -8 Walker ,M F . 191 Willson -93 , 705 stick Walking , D. E., 261 Wilson -6 ^7, 47778 -62, 426 , 188 Walnut pusilla , 526 Wilsonia K. 282 Walter Wind384042-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Wind pollination , 525 , ,517 , 507 , 523 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76,613 -14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston ious insects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138 , 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283 (aculeate , ),76191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K.,659 Wolfe 739 , L., 76869 Wood -40, 758 -60, 760 -62, 766 ,711 - 12 Wasp mantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 ,631 ,678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rine birds , 768 681 -69 -82, 719 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 3546 . See also Dryseason Weather , 614 , , 50455758 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon ,415 - 16 Web frog 388 Wood stork -footed Weeds 7089 . See tree species accounts Wood traits , 222 , 246 , 77, 97,2089, 212 Weevil , 609 , 567 Wood warbler , 627 , 271 ,324 , 326 , 348 ,629 255 Wood wren . 509 , 763 , 701 , 719 , 698 631 , 427 opossum Wooly , 144 - 45 Weinmannia WarII,6 67 World , 138 , , 136 Welfia georgii , 74 , xi States Wildlife World FundUnited -40,146 , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , ห ฟ 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas ,498 - 500 West Indian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . See Xamhoxylum also setulosum , 195 -67 , 428 Whales Zamhoxylum ,66, 70 Wheat , 372 Xantusiidae . 359 ,682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , ,320 faced 202 monkey White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 ,481 - 83 . See , 263 White tailed deer 248 ,111 fimbriata , 215 also Deer , 215 777 - 79 gualenensis , J. L., 281 , 33133 Whitmore -82 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wild ginger , 335 , 633 221 - 22 .Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

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9

9

Xylophones turbata , Xylosma ,

, 621 627 , 208416 147

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow grassquit -faced , 209 Yellow vireo -green ,561 Yellow headed vulture Yellow 750 - 5 / grasshopper -lined ,681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler ,610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66 , 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 ,349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

,727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (see also Xan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

277 - 78 Zarza

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฟ ” ฟ , J. ห ว ณ Zeledon ส , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

,688 - 91 Zompopa

Zonotrichia

capensis

, 618

,392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote ,468 -69 Zorra mochila Zorro 468 - 69 pelon

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle , 640 Wyeomyia West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 293 Ximenia americana , 150 peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

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. 593

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9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

Natural

, A,

History

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX

—. .

Vriesea ( com . ) orosiense 593 Vulture 404 , 456, 502

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner พ. H . , 187, 239-40, 311-18 Wake , M . H., 400^*01 Walker พ า Walking stick , Walnut , 188

.

.

.

Waltheria indica ,

ious insects Warscewicz , J . , 118

-

136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711-12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 -39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 -16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

Weinmannia , 144 45 Welfia georgii , 74 136, า

.

;

.

138-40, 146,

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498-500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 -83 White- banded sphinxlet . 680-81 White faced capuchin 458 -60 White faced monkey 202, 320, 329, 439, 458-60 , 764 White fronted nunbird , 587-90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481-83 . See also Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 211-12 Wild ginger 221-22

---

816

.

.. .

.

-

.

.

705-6

Walter K . ร., 282-92 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345-46 Wild papaya , 264 -65 , 633 Wild pineapple , 197 -200 Wild pitahaya , 252-53 Wild plantain 249-51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 Wilsonia pusilla , 526 Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 Witheringia , 715 Wolf 500, 716 Wolfe , K ., 659 Wood , ร. L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 -18 Wyeomyia , 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

—

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195 Zamhoxylum Xantusiidae 359, 372 Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

Ximenia americana , 150, Xiphorhynchus , 514 , 589 , Xylocopa ,

.

Yams , 70 Year of Costa Rican Natural History, A ,

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750-5 / Yellow spotted byttnena borer, 681-82 Yellow warbler, 575-76 Yiquirro, 610-11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 -17

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 Zamia , 349-50 Zanate 601 3

.

-

Zancudo , 727 -29

751

Zamhoxylum , 147 , 586, mayanum , melanostichum , 144 panamense , 140 , ( see also Xan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 -69

147

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8 Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 688-91

.

-

. See also

293

.

Zonotrichia capensis , 618 Zopilota , 392 93 , 398 -99 Zopilote cabecirrojo, 560-62 Zorra mochila , 468 -69 Zorro pelon 468 -69

.

-10

Xicote , 777 -79 Ximenez , F- , 1

621 627 208 416

Xylophones , turbata , Xylosma , 147 ,

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 gualenensis , 215 777 -79 muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

-

5

—. .

INDEX Vriesea orosiense

.)

( com

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136

.

ี

-

H., 119 Wendland , c . , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

--also

.

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c . , 3 - 4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

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*

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9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , ห ฟ 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22 9

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฟ ” ฟ , J. ห ว ณ Zeledon ส , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX

—. .

Vriesea ( com . ) orosiense 593 Vulture 404 , 456, 502

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner พ. H . , 187, 239-40, 311-18 Wake , M . H., 400^*01 Walker พ า Walking stick , Walnut , 188

.

.

.

Waltheria indica ,

ious insects Warscewicz , J . , 118

-

136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711-12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 -39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 -16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

Weinmannia , 144 45 Welfia georgii , 74 136, F

.

.

138-40, 146,

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498-500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 -83 White- banded sphinxlet . 680-81 White faced capuchin 458 -60 White faced monkey 202, 320, 329, 439, 458-60 , 764 White fronted nunbird , 587-90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481-83 . See also Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 211-12 Wild ginger 221-22

---

816

.

.. .

.

-

.

.

705-6

Walter K . ร., 282-92 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345-46 Wild papaya , 264 -65 , 633 Wild pineapple , 197 -200 Wild pitahaya , 252-53 Wild plantain 249-51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 Wilsonia pusilla , 526 Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 Witheringia , 715 Wolf 500, 716 Wolfe , K ., 659 Wood , ร. L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 -18 Wyeomyia , 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

—

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195 Zamhoxylum Xantusiidae 359, 372 Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

Ximenia americana , 150, Xiphorhynchus , 514 , 589 , Xylocopa ,

.

Yams , 70 Year of Costa Rican Natural History, A ,

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750-5 / Yellow spotted byttnena borer, 681-82 Yellow warbler, 575-76 Yiquirro, 610-11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 -17

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 Zamia , 349-50 Zanate 601 3

.

-

Zancudo , 727 -29

751

Zamhoxylum , 147 , 586, mayanum , melanostichum , 144 panamense , 140 , ( see also Xan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 -69

147

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8 Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 688-91

.

-

. See also

293

.

Zonotrichia capensis , 618 Zopilota , 392 93 , 398 -99 Zopilote cabecirrojo, 560-62 Zorra mochila , 468 -69 Zorro pelon 468 -69

.

-10

Xicote , 777 -79 Ximenez , F- , 1

621 627 208 416

Xylophones , turbata , Xylosma , 147 ,

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 gualenensis , 215 777 -79 muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , ห ฟ 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฟ ” ฟ , J. ห ว ณ Zeledon ส , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush ,] ฺ ด 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

๕๐

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

๊

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฺ ฌ ฺ , J. ] ู ํ ด Zeledon ไ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

น. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 - , H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker - ป , 7056 stick Walking , 188 Walnut

Walter . K. ร,*282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร, L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush , 640 Wyeomyia

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

น

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

.

Rican

Natural

.

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa capensis

, 618

.

.

183 Zygophyllaceae

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

, A,

History

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

293

, Ximenia americana , 150 , 589 , 514 Xiphorhynchus , Xylocopa ,

, 621 627 147, 208416

, 70 Yams Year of Costa

Zonotrichia

-10

, 777 - 79 Xicote , F-, 1 Ximenez

Xylophones turbata , Xylosma ,

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

๖

Wendland H . , 119 ้ 4 , 6, 119 Werkle , ว West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

[

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

้ ๓ ้ ฐ „ Zeledon , J. ว ํ Zeledon , R . , 8

Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

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*

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าพ ๖

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---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya plantain Wild 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E, 546 47, 607 Willis ^01 -8 Walker ,MF . 191-93 Willson , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia K. 282 Walter Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Wind pollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Wasp mantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds 70 89 . See tree species accounts Wood traits 77, 97, 2089, 212,222,246, Weevil , 609 , 567 Wood warbler , 627 , 271 , 324 , 326 , 348 , 629 255 Wood wren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World FundUnited , 406 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , ห ฟ 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 West Indian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , faced 202,320 monkey White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wild ginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

. 593

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.

9

9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow grassquit -faced , 209 Yellow vireo -green , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

277 - 78 Zarza

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฟ ” ฟ , J. ห ว ณ Zeledon ส , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorra mochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

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. 593

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-

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9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya plantain Wild 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E, 546 47, 607 Willis ^01 -8 Walker ,MF . 191-93 Willson , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia K. 282 Walter Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Wind pollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Wasp mantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds 70 89 . See tree species accounts Wood traits 77, 97, 2089, 212,222,246, Weevil , 609 , 567 Wood warbler , 627 , 271 , 324 , 326 , 348 , 629 255 Wood wren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World FundUnited , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c . , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 West Indian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , faced 202,320 monkey White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wild ginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

.

. -

. .

.

1

-

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow grassquit -faced , 209 Yellow vireo -green , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

277 - 78 Zarza

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c . , 3 - 4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorra mochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea orosiense

—. .

.)

( com

593 Vulture 404 , 456, 502-5, 507, 517, 552, 560

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner พ. H . , 187, 239-40, 311-18 Wake , M . H., 400^*01 Walker พ า Walking stick , Walnut , 188

.

.

ious insects Warscewicz , J . , 118

.

-

coccinea , 136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711 - 12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 - 39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 - 16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio

.

.

-

—

-

.

-,

, 144 45 Weinmannia Welfia georgii , 74 136, F

.

.

138-40, 146,

Wendland H . , 119 ้ 4 , 6, 119 Werkle , ว West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498 - 500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 - 83 White- banded sphinxlet . 680 - 81 White faced capuchin 458 - 60 White faced monkey 202, 320, 329, 439, 458 - 60 , 764 White fronted nunbird , 587 - 90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481 - 83 . See

--also

.

.. .

.

Deer Whitmore , J . L., 281 -82, 331-33 Wild bitter gourdlet . 211 - 12 Wild ginger 221 - 22 [

.

.

705-6

Walter K . ร., 282-92 indica , 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Waltheria

Wild nutmeg , 345 - 46 Wild papaya , 264 - 65 , 633 Wild pineapple , 197 - 200 Wild pitahaya , 252 - 53 Wild plantain 249 - 51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 pusilla , 526 Wilsonia Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 , 715 Witheringia Wolf 500, 716 Wolfe , K ., 659 Wood , ร. L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 - 18 , 640 Wyeomyia

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum Zamhoxylum

.

630

Xantusiidae 359, 372 709 10 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 Xicote , 777 79

.

,

-

.

Yams , 70 Year of Costa

Rican

-

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 Zamia , 349 - 50 Zanate 601 3

.

. 428

- 50

-

Zancudo , 727 - 29 , 147 , 586, 751 Zamhoxylum mayanum , 147 , 144 melanostichum

, 140 panamense , ( see also Xan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 - 69

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

้ ๓ ้ ฐ „ Zeledon , J. ว ํ Zeledon , R . , 8

Zeledonia. 616 - 18

Zeledonia

coronata

, 616

- /8

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708

.

Zompopa , 688

- 91

-

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 , 215 777 - 79 gualenensis muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

, A,

History

.

.

- 96

293

Natural

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750 - 5 / Yellow spotted byttnena borer, 681 - 82 Yellow warbler, 575 - 76 Yiquirro, 610 - 11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 - 17

capensis , 618 Zopilota , 392 93 , 398 - 99 Zopilote cabecirrojo, 560 - 62 Zorra mochila , 468 -69 Zorro pelon 468 - 69

195. See also

Ximenia americana , 150, , 514 , 589 Xiphorhynchus , Xylocopa ,

.

, 621 627 147, 208 416

Zonotrichia

-

Ximenez , F- , 1

Xylophones turbata , Xylosma ,

-

5

INDEX

—. .

Vriesea ( com. ) orosiense 593 Vulture 404 , 456, 502

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner พ. H . , 187, 239-40, 311-18 Wake , M . H., 400^*01 Walker พ า Walking stick , Walnut , 188

.

.

.

Waltheria indica,

ious insects Warscewicz , J . , 118

-

136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711-12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 -39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 -16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio coccinea,

.

.

-

—

-

.

-,

Weinmannia, 144 45 Welfia georgii , 74 136, F

.

.

138-40, 146,

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498-500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 -83 White- banded sphinxlet . 680-81 White faced capuchin 458 -60 White faced monkey 202, 320, 329, 439, 458-60 , 764 White fronted nunbird , 587-90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481-83 . See also Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 211-12 Wild ginger 221-22

---

816

.

.. .

.

-

.

.

705-6

Walter K . ร., 282-92 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345-46 Wild papaya , 264 -65 , 633 Wild pineapple , 197 -200 Wild pitahaya , 252-53 Wild plantain 249-51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 Wilsonia pusilla , 526 Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 Witheringia, 715 Wolf 500, 716 Wolfe , K ., 659 Wood , ร. L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 -18 Wyeomyia, 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

—

-

Xamhomonas, 80 Xanthopastis timais, Xamhoxylumsetulosum, 195 Zamhoxylum Xantusiidae 359, 372 Xenarescus 709 Xenomyrmex, 275 Xestoblatta hamata, 693- 96

630

.

.

Ximenia americana, 150, Xiphorhynchus, 514 , 589 , Xylocopa,

.

Yams , 70 Year of Costa Rican Natural History, A ,

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750-5 / Yellow spotted byttnena borer, 681-82 Yellow warbler, 575-76 Yiquirro, 610-11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 -17

-

.

Zacryptocerus, 462 Zalophus californianus

. 428

fairchildiana, 349-50 skinneri, 349 -50 Zamia , 349-50 Zanate 601 3

.

-

Zancudo , 727 -29

751

Zamhoxylum, 147 , 586, mayanum, melanostichum, 144 panamense, 140 , ( see also Xan setulosum, thoxylum setulosum ) Zapote , 704 Zarhynchuswagleri . 614 - 16 Zarigiieya 468 -69

147

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8

.

Zeledonia 616 -18 Zeledonia coronata, 616 - / 8 Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 688-91

.

-

. See also

293

.

Zonotrichia capensis , 618 Zopilota , 392 93 , 398 -99 Zopilote cabecirrojo, 560-62 Zorra mochila , 468 -69 Zorro pelon 468 -69

.

-10

Xicote , 777 -79 Ximenez , F- , 1

621 627 208 416

Xylophones, turbata, Xylosma, 147 ,

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata, 215, 111 gualenensis, 215 777 -79 muscaria. 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

". .

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

F

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX Vriesea orosiense

.)

( com

. 593

.

Vulture 404 , 456, 502-5, 507, 517, 552, 560

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

coccinea , 136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711 - 12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 - 39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 - 16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio

.

.

-

—

-

.

-,

, 144 45 Weinmannia Welfia georgii , 74 136, F

.

.

138-40, 146,

Wendland H . , 119 Werkle , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498 - 500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 - 83 White- banded sphinxlet . 680 - 81 White faced capuchin 458 - 60 White faced monkey 202, 320, 329, 439, 458 - 60 , 764 White fronted nunbird , 587 - 90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481 - 83 . See

--also

.

.. .

.

Deer Whitmore , J . L., 281 -82, 331-33 Wild bitter gourdlet . 211 - 12 Wild ginger 221 - 22

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 Waltheria indica , 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345 - 46 Wild papaya , 264 - 65 , 633 Wild pineapple , 197 - 200 Wild pitahaya , 252 - 53 Wild plantain 249 - 51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 pusilla , 526 Wilsonia Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 , 715 Witheringia Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 - 18 Wyeomyia , 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum Zamhoxylum

.

630

Xantusiidae 359, 372 709 10 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 Xicote , 777 79

.

,

-

.

Yams , 70 Year of Costa

Rican

-

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 Zamia , 349 - 50 Zanate 601 3

.

. 428

- 50

-

Zancudo , 727 - 29 , 147 , 586, 751 Zamhoxylum mayanum , 147 , 144 melanostichum

panamense , 140 , ( see also Xan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 - 69

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

Zeledon , J. Zeledon , R . , 8

Zeledonia. 616 - 18

Zeledonia

coronata

, 616

- /8

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708

.

Zompopa , 688

- 91

-

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 , 215 777 - 79 gualenensis muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

, A,

History

.

.

- 96

293

Natural

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750 - 5 / Yellow spotted byttnena borer, 681 - 82 Yellow warbler, 575 - 76 Yiquirro, 610 - 11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 - 17

capensis , 618 Zopilota , 392 93 , 398 - 99 Zopilote cabecirrojo, 560 - 62 Zorra mochila , 468 -69 Zorro pelon 468 - 69

195. See also

Ximenia americana , 150, , 514 , 589 Xiphorhynchus , Xylocopa ,

.

, 621 627 147, 208 416

Zonotrichia

-

Ximenez , F- , 1

Xylophones turbata , Xylosma ,

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c . , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c . , 3 - 4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

Vriesea orosiense

—ห. ;

ณ ฟอ ฟ ฟ ฟ” ฟอ ว ฟ 9ส ” ู

ู หห ห ส ฟ ฟ ห ”

ู 9 ส ฟ — ” หหร., ส ฟ

ห

— ฟอ ฟอ ฟฟ ู ู ณฟ ณ ูฟส ส ฟ ห — See also indica ,

ส

ฟ— ฟ ห

ฟ— ฟ; 9 ฟฟ ฟณ ฟ ๊ ” ๊ ห ฟห ฟ อ อ 9ห ูฟ 9” ห ฟส —ฟ ส ;ฟ 9 ส

Warscewiczio

,

coccinea

ฟ 1

/

ฟ 711 - 12

ส

ฟส See also ห ู 9 ฟส ณ

ห249

พ า

ห See

หห

หหห ฟ ฟส ฟ หห ฟส ู ; 9 ฟฟ

ห ฟห ฟ

หส

ฟ ร. ฟ — ฟส ฟ หห ณ

ฟฟ — ๊ ส ส ฟ9 ฟ ว ฟ 9 ฟอ ห See also ฟ ว อฟ ฟ ว ๊ฟ ; ส ฟ— —

ห

ห หส ห ห

ส ฟฟ ฟ9 ฟ ฟฟ — ” ” ห ส ฟอ ฟ๊ ฟว ฟ9 ฟ — ฟห ู

See tree species

ฟ ฟอ

ห

ฟ๊ ฟฟ ณ ฟ— ๊

ฟ หห ฟส — ส หหห 557 ห ฟ ฟ ฟ c., ว 9

ฟ 498 - 500

ฟส ฟห ห ู ฟ—

ฟฟ

ส ส

ส ส 458 ส ส ส ส ส also

680 - 81 ห

ฟ 682

83

ฟ 9 ฟ 406

ส ฟ6 - 18

accounts

6/

,

60

90 ,

ส

,

,

gualenensis

- 22

viridis ,

,

ฟู ฟ

79

ห ณ also

Xylocopa

9

- 82

—

- 50

- 29

อ

๊

อ ๊

หห -

277 78 Zea mays , 88 92

ห

616 - 18 ห

, 616 - / 8 ฟส ฟู 9 ฟ— ฟ๊

coronata

capensis

Xan

ฟอ

ฟc., 3-4 , 502 ฟห ฟ —

, 618

ฟ 392 - 93 , 398 - 99 ฟ 560 - 62

ฟ 468

,

481

.

ฟฟ , อ 9 Zamhoxylum mayanum , , ว melanostichum , ; panamense ฟ/ setulosum , ) thoxylum setulosum ว Zapote , Zarhynchus wagleri . 614 - 16 468 - 69

Zonotrichia

,

Xiphorhynchus

ห ฟ 727

”

ห—

60 ,

587

681

ฟ 688 - 91

; ห ู

.

750 - 5 / ห ฟ

ฟ 575 - 76

, 349 fairchildiana skinneri , 349 - 50 ฟ 349 - 50 601 3

;

ห

ว

ฟ

ฟ 610 - 11 ฟหห ฟฟส ว ฟฟ 114 - 17

Zeledonia

, Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , Zamhoxylum ฟ ” Xenarescus , ู Xenomyrmex Xestoblatta hamata , 693 - 96

, A,

History

Natural

ฟ ฟ๊

ส

ฟ

ฟส ฟู ฟว — อ Wyeomyia

Rican

ส ส ส ส

อ

—

ห

ฟ Year of Costa

ส

458 ห ฟฟ ” ; ฟฟ ฟ ห 777 - 79 ว ฟส ฟ ๊ ฟ9 ฟณ ; Ximenia americana ๊ ห ฟ ว ฟ— ฟห ว ; ู ฟฟ ณ Xylocopa ๊ ห ห ฟอ ฟ —ส ณ ฟฟ ณ - 83 . See ห— ฟ ู 111 fimbriata 777 ู ห ฟ๊ ฟ ฟ๊ หห ส ู muscaria . ส 211 - 12 ห ู

221

816

ฟ

45 -ฟ ฟส ฟ 9 —ฟ 9 F

ห

ฟ ๊ ฟ๊ ฟส —

also

๊ อ ฟ— 9 อ

,

, Zacryptocerus californianus Zalophus

ณ 9 ู ฟว ฟ อ , ู ฟ 9 ;

Witheringia

ฟ 415 - 16

, 144 Weinmannia Welfia georgii ,

ส ส

สฟ ส ฟฟ

ฟ๊

ส

หส

ฟห ฟส ห ว 9 ฟฟ ส — 9ฟ อ ส ฟ ห ส ๊ ฟ๊ ฟ 9อ ฟอ ฟห ห ส 9 ๅ pusilla , Wilsonia ฟ ” ส ฟณ ห See also

ฟอ

ฟ

51

อ

ฟ

ฟ 236 - 39

Xylophones turbata , Xylosma ,

ฟ ณ 264 - 65 , ฟ 197 - 200 ฟ 252 - 53

ฟส ฟหห —

ฟอ ฟส ฟ๊ พ. ห ส ฟ; ฟหห ๅ

Waltheria

ฟ 345 - 46

.)

( com

468 - 69 ห หณ 69

-

ู

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

๕๐

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

.

. -

. .

.

1

-

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

INDEX Vriesea ( com. ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^ *01 Walker

, 705 stick Walking -6 , 188 Walnut K. 282 Walter -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Wasp mantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds 70 89 77, 97, 2089, 212,222,246, Weevil , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia, 144 45 , 13840, 146 , Welfia georgii , 74 136 อ

.

1

-

H., 119 Wendland , c.,4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 West Indian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , faced 202,320 monkey White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wild ginger 221-22

.

---

.. .

.

-

816

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya plantain Wild 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E, 546 Willis -47, 607-8 ,MF . 191-93 Willson , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Wind pollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Wood warbler Wood wren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World FundUnited , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia, 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastis timais , Xamhoxylum setulosum, 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex, 275 Xestoblatta hamata , 693- 96

630

.

.

.

-10

, 777 -79 Xicote , F-, 1 Ximenez

293

, Ximenia americana , 150 , 589 Xiphorhynchus, 514 , Xylocopa,

.

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

621 627

Xylophones, turbata , , 208416 Xylosma, 147

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow grassquit -faced , 209 Yellow vireo -green , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum, 147 mayanum, melanostichum , 144 panamense , 140 , (seealsoXan setulosum, thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

277 -78 Zarza

Zea mays , 88 92

, 607 antshrike Zebra -striped

,3, J. c. 4,502 Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorra mochila Zorro pelon468 -69

-

.

.

183 Zygophyllaceae

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , c., 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , ว ้ 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

[

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

้ ๓ ้ , J. ว ฐ „ Zeledon ํ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

าพ

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , ว ้ 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

[

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

้ ๓ ้ , J. ว ฐ „ Zeledon ํ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. ว

INDEX

’

: ว

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . Nox coloration Warning

-

iousinsects , J., 118 Warscewicz

, 138, 140 ้136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , ™ํ Waspmantispid ; Ma , 505 Water Duck birds -6. ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect , [ํ hyacinth Water , 427 opossum Water ; , 35 46. Dryseason Weather Seasonality , 52122 effect onbirds , แํ Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

.

.

-

—

-

.

.

ํ , 136, 138-40, 146,

้ ฐ ้74

F

.

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper . ๙ , \ํุ WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , ๓ํ White fatima -banded White . ุํ sphinxlet -banded White faced capuchin\ํ , 329 , 439 , facedmonkey 202,320 White \ํ้764 , ๙ํ้606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , ™ํว , 263 White tailed deer248

.

---

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet . ™ํ Wildginger™ํ

816

.

. -

.

.

.

้ 778

.

, แํ Wildnutmeg , ฐํ้633 Wildpapaya , ๙ํุ Wildpineapple , ๓ํ Wildpitahaya Wildplantain๖ํ , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 ้526 Wind38 40 42-44, 632 , 283 . Windpollination Polli , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula ้715 , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , [ Worm snake , 558 , 5057, 524 Wren -60 , [”[ํ Wrenthrush ้640

*

.. .

-

-

.

-

.

.

. -

. .

—

-

80

้

ว709-10

, ๙ํ Xicote , F-, 1 Ximenez

้

้3้5

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow แ” ุํ -linedgrasshopper , ™ํ borer byttnena Yellow spotted , แํ Yellow warbler , ุํ Yiquirro , A. M., ix, 724 Young -26 , 66, ฐํ Yuca

-

.

้462 ้

ว428

๖ํ

้ ๖ํ , ๖ํ Zamia Zanate ™ „

.

ํ

, ๙ํ Zancudo

, 751 147,586

้

้147

้144 140

้ ้

127,131(seealso ํ ้704 ว ฐํ :

. .ํ

Zarigiieya\ํ

Zarza ๙ํ ้

, 607 antshrike Zebra -striped , J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia . [ํ

้ [ํ ” \

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, \ํ Zompopa

้ \

.

183 Zygophyllaceae

„ํ

293

้150 , , 589 ้514

.

, 70 Yams

.

Xantusiidae . 359, 372

275

.

, ๓ํ้ \ํ Zopilota , ุํ cabecirrojo Zopilote , \ํ Zorramochila Zorro pelon \ํ

630 ้195 . Seealso

้

้

้621 627 , 208416 ้147

้

.

, 241 ้194210 . 245299, 323, , 633 628 -34 , ™ ้215 ้215 ๙ ํ ว215 ้215 , 335 , 633 . See Xylocopidae

.

๖

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

าพ

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136 5

.

^

-

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

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9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

F

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

. 593

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.

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—

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พร

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---

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.

-

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.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

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-

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9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

าพ ๖

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX ิ ๖้ ด ๑593 Vulture 404 , 456, 502

ด

.

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner H . , 187, 239-40, 311-18 Wake , M . H., 400^*01

.

.

ious insects Warscewicz , J . , 118

.

-

.

-

—

-

.

ไ , 136, 138-40, 146,

.

ฺ ู ฺ74

.

;

า

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. “ West Indian manatee , 0ไ" Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , ฌไ White- banded sphinxlet . "ไ White faced capuchin 0ไ White faced monkey 202, 320, 329, 439, 0ไฺ764 White fronted nunbird , “ไฺ606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 5ได ๑ Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 5ไ Wild ginger 5ไ

---

816

.

.. .

.

-

..

.

ฺ136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 5ไ Water birds , 505-6. ๑ Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , ๊ไ Water opossum , 427 Weather, 35 46. ๑ Dry season; Seasonality effect on birds , 521 22 Web-footed frog , «ไ Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

.

.

Walker

Walking stick , 705-6 Walnut , 188 Walter K . 282-92 ฺ 778 ๖ Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. ๑ Nox

Wild nutmeg , «ไ Wild papaya , ูไฺ633 Wild pineapple , “ไ" Wild pitahaya , ฌไ Wild plantain ฝไ Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 ฺ 526 ๖ Wind 38 40 42-44 , 632 Wind pollination , 283. ๑ Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 ฺ ๖ 715 Wolf 500, 716 Wolfe , K ., 659 Wood , L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. ๑ Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. ๑ Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , ๊ Wren , 505 7, 524 , 558-60 Wrenthrush , ๊ฒ๊ไ ฺ640

. -

.

*

.

-

.

-

-

.

.

. -

. .

-

80

ฺ

630

ฺ

195. See also

ฺ

Xicote , ไ Ximenez , F- , 1 ๖

.

ฺ462

293

ด428

ฺ๖ฝ ไ ฺ ฝไ ฝ ไ 5ํ

Zamia , Zanate ไ Zancudo , “ไ

.

147, 586, 751

ฺ

ฺ147

144

ฺ

๑ฺ140

127, 131 ( see also

ฺ

ิ

704

ฺ

ด ไ . ไ Zarza. “ไ ๖ ู

Zarigiieya

0

ไ

ฺ

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8 Zeledonia.

๊ไ ฺ ๊ไ ฒ 0

๖

Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 0ไ

.

ฺ 0

.

Zygophyllaceae 183

ํไ

ฺ150, ๖ ฺ514 , 589

.

-

.

ด709-10 275

ฺหฺ5

๖

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper "ไ «ฒ Yellow spotted byttnena borer, 5ไ Yellow warbler, «ไ Yiquirro, "ไ Young , A . M . , ix , 724-26 Yuca , 66 , ูไ

๖

Xantusiidae . 359, 372 ฺ

Yams , 70

.

208 416

Zopilota , ฌไฺ 0ไ Zopilote cabecirrojo, "ไ Zorra mochila , 0ไ Zorro pelon 0ไ

ฺ

๖ “

621

ฺ

ฺ 627 ๖ ๖ฺ147 ,

.

ฺ194 210, 241. 245 299, 323, 628, 633-34 ฺ ๖215, 5 ฺ215 “ไ ด215 ฺ215 Xylocopidae , 335, 633. See

.

ฝ

ไ

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136 5

.

^

-

H., 119 Wendland , c., 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

816

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4, 502 Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c . , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

.

. -

. .

.

1

-

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c . , 3 - 4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

.

. -

. .

.

1

-

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com. ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^ *01 Walker

, 705 stick Walking -6 , 188 Walnut K. 282 Walter -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Wasp mantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds 70 89 77, 97, 2089, 212,222,246, Weevil , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia, 144 45 , 13840, 146 , Welfia georgii , 74 136 อ

.

1

-

H., 119 Wendland , c. , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 West Indian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , faced 202,320 monkey White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wild ginger 221-22

.

---

.. .

.

-

816

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya plantain Wild 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E, 546 Willis -47, 607-8 ,MF . 191-93 Willson , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Wind pollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Wood warbler Wood wren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World FundUnited , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia, 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastis timais , Xamhoxylum setulosum, 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex, 275 Xestoblatta hamata , 693- 96

630

.

.

.

-10

, 777 -79 Xicote , F-, 1 Ximenez

293

, Ximenia americana , 150 , 589 Xiphorhynchus, 514 , Xylocopa,

.

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

621 627

Xylophones, turbata , , 208416 Xylosma, 147

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow grassquit -faced , 209 Yellow vireo -green , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum, 147 mayanum, melanostichum , 144 panamense , 140 , (seealsoXan setulosum, thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

277 -78 Zarza

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorra mochila Zorro pelon468 -69

-

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

.

พร1

. -

. .

-

.

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

. -

. .

.

าพ ห

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com. ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, ), 76 191 Wasp(aculeate , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall effectonclimate ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effectonbirds Web -footedfrog,415-16 Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

าพ

-,

.

Weinmannia, 144 45 , 13840, 146 , Welfia georgii , 74 136

-

.

H., 119 Wendland , 4, 6, 119 Werkle West-Eberhard M. J., 758 -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded Whitefaced 458 -60 capuchin , 329 , 439 , 202,320 Whitefacedmonkey 458-60 , 764 ,587-90 , 606 Whitefronted nunbird peccary Whitelipped 471 , 388 Whitenecked heron , 547 White antbird -plumed , 481 -83 . See , 263 Whitetailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

---

.

.. .

.

-

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 ,M F . 191-93 Willson , D. E., 261-62, 426 Wilson ^ 7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , by wind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Woodboring beetle -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Woodduck , 546 Woodhaunter , 657 . See also Satyridae Woodnymph , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Woodpigeon Woodstork388 . See tree species accounts Woodtraits , 609 , 567 Woodwarbler Woodwren . 509 , 427 Woolyopossum WorldWarII, 6 67 , xi States WorldWildlife FundUnited , 406 Wormsnake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia, 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastis timais , Xamhoxylum setulosum, 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex, 275 Xestoblatta hamata , 693- 96

630

.

.

.

-10

, 777 -79 Xicote , F-, 1 Ximenez

293

, Ximenia americana , 150 , 589 Xiphorhynchus, 514 , Xylocopa,

.

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

621 627

Xylophones, turbata , , 208416 Xylosma, 147

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellowheaded vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellowspotted , 575-76 Yellowwarbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 Yuca, 66, 114 -17

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum, 147 mayanum, melanostichum , 144 panamense , 140 , (seealsoXan setulosum, thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorra mochila Zorro pelon468 -69

-

.

.

183 Zygophyllaceae

-

5

ADDRESSES OF CONTRIBUTORS

ร. M . Smith

Department of Biology Mount Holyoke College South Hadley, Massachusetts 01075 N. Smythe Smithsonian Tropical Research Institute APO Miami 34002 L. K . Sowls " Arizona Cooperative Wildlife Research Unit University of Arizona Tucson , Arizona 85721 M . A . Staton Feather Crest Farms Route 1 , Box 2250 Douglass, Texas 75943 G . Stevens Department of Biology University of Pennsylvania Philadelphia , Pennsylvania 19104 F. G . Stiles Escuela de Biolog fa Universidad de Costa Rica Ciudad Universitaria Costa Rica D. E. Stone Department of Botany Duke University Durham . North Carolina 27706 J . Stout Department of Zoology Michigan State University East Lansing , Michigan 48824 J . G . Strauch , Jr. University Museum University of Colorado Boulder, Colorado 80309 D. R . Strong Department of Biological Sciences Florida State University Tallahassee , Florida 32306 c. A . Todzia Department of Botany University of Texas Austin , Texas 78712 D. c. Timer Ethologie und Wildforschung University Zurich

Birchstrasse 95 8050 Zurich Switzerland J. F. Utley Department of Biological Sciences University of New Orleans New Orleans , Louisiana 70122 J . H . Vandermeer Division of Biological Sciences University of Michigan Ann Arbor, Michigan 48109 786

ADDRESSES OF CONTRIBUTORS

R . พ. Van Devender Department of Biology Appalachian State University Boone , North Carolina 28608 M . Vargas V. Escuela de Microbiolog fa Universidad de Costa Rica Ciudad Universitaria Costa Rica A. Vasquez Morera Unidad de Suelos Ministerio de Agricultura y Ganaderia San Jos6 Costa Rica ร. L. Vehrencamp Department of Biology, C-016 University of California La Jolla , California 92093 ร. B . Vinson Department of Entomology Texas A . & M . University College Station , Texas 77843 H . K . Vbris Field Museum of Natural History' Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605 H . B. Wagner Smithsonian Tropical Research Institute APO Miami 34002 พ. H . Wagner Department of Botany University of Michigan Ann Arbor, Michigan 48109 M. H . Whke Department of Zoology University of California Berkeley, California 94720 K. ร. Walter New York Botanical Garden Bronx , New York 10458 M . J . West - Eberhard Escuela de Biolog fa Universidad de Costa Rica Ciudad Universitaria Costa Rica R . M . Wetzel Florida State Museum University of Florida Gainesville , Florida 32611 J . L. WTiitmore บ.ร. Department of Agriculture International Forestry p o Box 2417 Washington , D .c. 20013 R. H . Wiley Department of Biology University of North Carolina Chapel Hill , North Carolina 27514 .

G . B . Williamson Department of Botany Louisiana State University Baton Rouge , Louisiana 70803 Willis UNESP Zoologia 13500 Rio Claro, s . p Brazil

—

M . F. Willson Department of Ecology, Ethology, and Evolution University of Illinois at Urbana-Champaign Champaign , Illinois 61820 . D E. Wilson Museum Section บ.ร. Fish and Wildlife Service National Museum of Natural History, NHB 382 Washington , D.c. 20560 D. M . Windsor Smithsonian Tropical Research Institute APO Miami 34002

M . Winston Department of Biological Sciences Simon Fraser University Burnaby, British Columbia V5A 1 S6 Canada

ร. L Wood

Life Sciences Museum Brigham Young University Provo, Utah 84602 T. K . Wood Department of Entomology and Applied Ecology University of Delaware Newark , Delaware 19711 A . M . Young Milwaukee Public Museum 800 Wfest Wells Street Milwaukee , Wisconsin 53233 G. R. Zug National Museum of Natural History Smithsonian Institution Washington , D . c . 20560

.

787

ADDRESSES OF CONTRIBUTORS

ADDRESSES OF CONTRIBUTORS

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ADDRESSES OF CONTRIBUTORS

ADDRESSES OF CONTRIBUTORS

M. Smith of Biology Department College Holyoke Mount 01075 , Massachusetts Hadley South N. Smythe Institute Research Tropical Smithsonian APO Miami34002 L. K. Sowls" Unit Research Wildlife Cooperative Arizona ofArizona University 85721 , Arizona Tucson M. A. Staton Farms Crest Feather Route 1, Box2250 , Texas75943 Douglass G. Stevens ofBiology Department ofPennsylvania University 19104 , Pennsylvania Philadelphia F. G. Stiles fa deBiolog Escuela Rica deCosta Universidad Universitaria Ciudad Rica Costa D. E. Stone of Botany Department University Duke 27706 Carolina . North Durham J. Stout ofZoology Department University State Michigan 48824 , Michigan EastLansing , Jr . J. G. Strauch Museum University ofColorado University , Colorado 80309 Boulder D. R. Strong Sciences of Biological Department University State Florida , Florida 32306 Tallahassee c. A. Todzia ofBotany Department ofTexas University , Texas78712 Austin D. c. Timer undWildforschung Ethologie University Zurich

95 Birchstrasse Zurich 8050 Switzerland J. F. Utley Sciences of Biological Department of NewOrleans University 70122 , Louisiana NewOrleans J. H. Vandermeer Sciences of Biological Division of Michigan University , Michigan 48109 AnnArbor ๊

R. VanDevender of Biology Department University State Appalachian 28608 , North Carolina Boone V. M. Vargas fa deMicrobiolog Escuela Rica deCosta Universidad Universitaria Ciudad Rica Costa Morera A. Vasquez deSuelos Unidad y Ganaderia deAgricultura Ministerio 6 SanJos Rica Costa L. Vehrencamp , C-016 ofBiology Department ofCalifornia University 92093 , California LaJolla B. Vinson ofEntomology Department A. &M. University Texas , Texas77843 Station College H. K. Vbris ' History of Natural FieldMuseum Drive Shore Road atLake Roosevelt , Illinois 60605 Chicago H. B. Wagner Institute Research Tropical Smithsonian Miami34002 APO H. Wagner of Botany Department of Michigan University 48109 , Michigan AnnArbor M. H. Whke ofZoology Department ofCalifornia University 94720 , California Berkeley K. Walter Garden NewYorkBotanical , NewYork10458 Bronx M. J. West -Eberhard fa deBiolog Escuela Rica deCosta Universidad Universitaria Ciudad Rica Costa R. M. Wetzel Museum State Florida ofFlorida University 32611 , Florida Gainesville J. L. WTiitmore of Agriculture Department Forestry International Box2417 , D.c. 20013 Washington R. H. Wiley ofBiology Department Carolina of North University 27514 Carolina , North Hill Chapel

G. B. Williamson ofBotany Department University State Louisiana , Louisiana 70803 Rouge Baton . ๐พ๐Willis UNESPZoologia , s.p RioClaro 13500 Brazil

M. F. Willson , Ethology , andEvolution ofEcology Department atUrbana ofIllinois University -Champaign , Illinois 61820 Champaign . Wilson E . D Section Museum Service FishandWildlife , NHB 382 History Museum of Natural National , D.c. 20560 Washington D. M. Windsor Institute Research Tropical Smithsonian APO Miami34002

M. Winston Department Sciences of Biological Simon Fraser University V5A 1S6 , British Columbia Burnaby Canada

L Wood

Museum LifeSciences University Young Brigham , Utah84602 Provo T. K. Wood andApplied Ecology ofEntomology Department ofDelaware University , Delaware 19711 Newark A. M. Young Museum Public Milwaukee Street 800WfestWells , Wisconsin 53233 Milwaukee G. R. Zug of Natural History Museum National Smithsonian Institution , ฟด] " ด Washington

ด ด

“

ADDRESSES OF CONTRIBUTORS

ร. M . Smith

Department of Biology Mount Holyoke College South Hadley, Massachusetts 01075 N. Smythe Smithsonian Tropical Research Institute APO Miami 34002 L. K . Sowls " Arizona Cooperative Wildlife Research Unit University of Arizona Tucson , Arizona 85721 M . A . Staton Feather Crest Farms Route 1 , Box 2250 Douglass, Texas 75943 G . Stevens Department of Biology University of Pennsylvania Philadelphia , Pennsylvania 19104 F. G . Stiles Escuela de Biolog fa Universidad de Costa Rica Ciudad Universitaria Costa Rica D. E. Stone Department of Botany Duke University Durham . North Carolina 27706 J . Stout Department of Zoology Michigan State University East Lansing , Michigan 48824 J . G . Strauch , Jr. University Museum University of Colorado Boulder, Colorado 80309 D. R . Strong Department of Biological Sciences Florida State University Tallahassee , Florida 32306 c. A . Todzia Department of Botany University of Texas Austin , Texas 78712 D. c. Timer Ethologie und Wildforschung University Zurich

Birchstrasse 95 8050 Zurich Switzerland J. F. Utley Department of Biological Sciences University of New Orleans New Orleans , Louisiana 70122 J . H . Vandermeer Division of Biological Sciences University of Michigan Ann Arbor, Michigan 48109 786

ADDRESSES OF CONTRIBUTORS

R . พ. Van Devender Department of Biology Appalachian State University Boone , North Carolina 28608 M . Vargas V. Escuela de Microbiolog fa Universidad de Costa Rica Ciudad Universitaria Costa Rica A. Vasquez Morera Unidad de Suelos Ministerio de Agricultura y Ganaderia San Jos6 Costa Rica ร. L. Vehrencamp Department of Biology, C-016 University of California La Jolla , California 92093 ร. B . Vinson Department of Entomology Texas A . & M . University College Station , Texas 77843 H . K . Vbris Field Museum of Natural History' Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605 H . B. Wagner Smithsonian Tropical Research Institute APO Miami 34002 พ. H . Wagner Department of Botany University of Michigan Ann Arbor, Michigan 48109 M. H . Whke Department of Zoology University of California Berkeley, California 94720 K. ร. Walter New York Botanical Garden Bronx , New York 10458 M . J . West - Eberhard Escuela de Biolog fa Universidad de Costa Rica Ciudad Universitaria Costa Rica R . M . Wetzel Florida State Museum University of Florida Gainesville , Florida 32611 J . L. WTiitmore บ.ร. Department of Agriculture International Forestry p o Box 2417 Washington , D .c. 20013 R. H . Wiley Department of Biology University of North Carolina Chapel Hill , North Carolina 27514 .

G . B . Williamson Department of Botany Louisiana State University Baton Rouge , Louisiana 70803 E. o. Willis UNESP Zoologia 13500 Rio Claro, s . p Brazil

—

M . F. Willson Department of Ecology, Ethology, and Evolution University of Illinois at Urbana-Champaign Champaign , Illinois 61820 . D E. Wilson Museum Section บ.ร. Fish and Wildlife Service National Museum of Natural History, NHB 382 Washington , D.c. 20560 D. M . Windsor Smithsonian Tropical Research Institute APO Miami 34002

M . Winston Department of Biological Sciences Simon Fraser University Burnaby, British Columbia V5A 1 S6 Canada

ร. L Wood

Life Sciences Museum Brigham Young University Provo, Utah 84602 T. K . Wood Department of Entomology and Applied Ecology University of Delaware Newark , Delaware 19711 A . M . Young Milwaukee Public Museum 800 Wfest Wells Street Milwaukee , Wisconsin 53233 G. R. Zug National Museum of Natural History Smithsonian Institution Washington , D . c . 20560

.

787

INDEX Vriesea ( com. ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^ *01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

๖

.

Weinmannia, 144 45 , 13840, 146 , Welfia georgii , 74 136

.

-

H., 119 Wendland ,] ฺ ด 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , faced 202,320 monkey White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

๊

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E, 546 Willis -47, 607-8 ,M F . 191-93 Willson , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , by wind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Woodboring beetle -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae Woodnymph , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Woodpigeon Wood stork388 . See tree species accounts Woodtraits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World FundUnited , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia, 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastis timais , Xamhoxylum setulosum, 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex, 275 Xestoblatta hamata , 693- 96

630

.

.

.

-10

, 777 -79 Xicote ,F Ximenez -, 1

293

, Ximenia americana , 150 , 589 Xiphorhynchus, 514 , Xylocopa,

.

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

621 627

Xylophones, turbata , , 208416 Xylosma, 147

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow vireo -green , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum, 147 mayanum, melanostichum , 144 panamense , 140 , (seealsoXan setulosum, thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฺ ฌ ฺ , J. ] ู ํ ด Zeledon ไ , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorra mochila Zorro pelon468 -69

-

.

.

183 Zygophyllaceae

-

5

-

Vriesea ( com. ) orosiense

345 46 264 -65 , 197 -200 252 -53 249 51

.

Xylophones, turbata , Xylosma,

-

Year of Costa Rican Natural History, A ,

-

750 5 / 681 -82 Wilsonia pusilla ,

-

575-76

610 11 Waltheria indica ,

See also

114 -17 See also

Warscewiczio coccinea ,

Zacryptocerus , Zalophus californianus

fairchildiana , 349-50 skinneri , 349 -50 349 -50 601 3

Witheringia ,

-

-

711 12 See also

Zamhoxylum, mayanum, melanostichum , panamense , setulosum, thoxylum setulosum ) Zapote , Zarhynchus wagleri . 614 - 16 468 -69

-

See also See also

-

-

415 16

277 78 Zea mays , 88-92

See tree species accounts

ฺ ฌ ฺ ู ] ํ ด ไ

-

๖

Weinmannia, 144 45 Welfia georgii ,

688 -91

Xamhomonas, Xanthopastis timais , Xamhoxylum setulosum, Zamhoxylum

-

682 83 680 -81 458 -60 587 -90 ,

-

211-12

-

.

-

Ximenia americana , Xiphorhynchus, Xylocopa,

481 83 . See

also

Zonotrichia capensis , 618 392 93 , 398 - 99 560 -62 468 -69 468 -69

Xenarescus Xenomyrmex, Xestoblatta hamata , 693- 96 777 79

458-60 ,

๊

-

6 / 6 18

Wyeomyia,

498-500

-

616 -18 Zeledonia coronata , 616 - / 8

406

557

221 22

-

727 29

236 39

ฺ ด ]

.

fimbriata , gualenensis , muscaria viridis ,

.

111 777 -79 9

also Xylocopa

-

Xan

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

าพ ๖ -,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136

-

.

H., 119 Wendland , ห ฟ 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

.

---

.. .

.

-

9

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฟ ” ฟ , J. ห ว ณ Zeledon ส , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

-

Vriesea ( com. ) orosiense

345 46 264 -65 , 197 -200 252 -53 249 51

.

Xylophones, turbata , Xylosma,

-

Year of Costa Rican Natural History, A ,

-

750 5 / 681 -82 Wilsonia pusilla ,

-

575-76

610 11 Waltheria indica ,

See also

114 -17 See also

Warscewiczio coccinea ,

Zacryptocerus , Zalophus californianus

fairchildiana , 349-50 skinneri , 349 -50 349 -50 601 3

Witheringia ,

-

-

711 12 See also

Zamhoxylum, mayanum, melanostichum , panamense , setulosum, thoxylum setulosum ) Zapote , Zarhynchus wagleri . 614 - 16 468 -69

-

See also See also

-

-

415 16

277 78 Zea mays , 88-92

See tree species accounts

• v • 0 J f Z

-

Weinmannia, 144 45 Welfia georgii ,

ี

616 -18 Zeledonia coronata , 616 - / 8

406

-

6 / 6 18

Wyeomyia,

688 -91

557

Xamhomonas, Xanthopastis timais , Xamhoxylum setulosum, Zamhoxylum

498-500

-

682 83 680 -81 458 -60 587 -90 ,

-

-

221 22

211-12

-

.

-

Ximenia americana , Xiphorhynchus, Xylocopa,

481 83 . See

also

Zonotrichia capensis , 618 392 93 , 398 - 99 560 -62 468 -69 468 -69

Xenarescus Xenomyrmex, Xestoblatta hamata , 693- 96 777 79

458-60 ,

/

-

727 29

236 39

• f

.

fimbriata , gualenensis , muscaria viridis ,

.

111 777 -79 9

also Xylocopa

-

Xan

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 456 , 502 , 552 , 517 404 , Vulture , 197 - 200 Wildpineapple -5, 507 560 , 252 - 53 Wildpitahaya plantain Wild 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 , 239 WagnerH., 187 , G. B., 224 Williamson -40,311 -18 -2525659 , M. H., 400 Wake , E, 546 47, 607 Willis ^01 -8 Walker ,M F . 191 Willson -93 , 705 stick Walking , D. E., 261 Wilson -6 ^7, 47778 -62, 426 , 188 Walnut pusilla , 526 Wilsonia K. 282 Walter Wind384042-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Wind pollination , 525 , ,517 , 507 , 523 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76,613 -14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston ious insects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138 , 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283 (aculeate , ),76191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K.,659 Wolfe 739 , L., 76869 Wood -40, 758 -60, 760 -62, 766 ,711 - 12 Wasp mantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 ,631 ,678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rine birds , 768 681 -69 -82, 719 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 3546 . See also Dryseason Weather , 614 , , 50455758 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon ,415 - 16 Web frog 388 Wood stork -footed Weeds 7089 . See tree species accounts Wood traits , 222 , 246 , 77, 97,2089, 212 Weevil , 609 , 567 Wood warbler , 627 , 271 ,324 , 326 , 348 ,629 255 Wood wren . 509 , 763 , 701 , 719 , 698 631 , 427 opossum Wooly , 144 - 45 Weinmannia WarII,6 67 World , 138 , , 136 Welfia georgii , 74 , xi States Wildlife World FundUnited -40,146 , 406 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush ,] ฺ ด 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas ,498 - 500 West Indian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . See Xamhoxylum also setulosum , 195 -67 , 428 Whales Zamhoxylum ,66, 70 Wheat , 372 Xantusiidae . 359 ,682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , ,320 faced 202 monkey White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 ,481 - 83 . See , 263 White tailed deer 248 ,111 fimbriata , 215 also Deer , 215 777 - 79 gualenensis , J. L., 281 , 33133 Whitmore -82 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wild ginger , 335 , 633 221 - 22 .Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

.

. 593

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

.

.

-

-

—

.

.

.

. -

. .

-

-

.

ป- ๖

-

.

.

---

-

.. .

-

.

.

-

.

.

.

9

๊

Xylophones turbata , Xylosma ,

, 621 627 , 208416 147

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow grassquit -faced , 209 Yellow vireo -green ,561 Yellow headed vulture Yellow 750 - 5 / grasshopper -lined ,681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler ,610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66 , 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 ,349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

,727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (see also Xan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

277 - 78 Zarza

Zea mays , 88 92

, 607 antshrike Zebra -striped

ฺ ฌ ฺ , J.] ู ํ ด Zeledon ไ , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

,688 - 91 Zompopa

Zonotrichia

capensis

, 618

,392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote ,468 -69 Zorra mochila Zorro 468 - 69 pelon

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com. ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^ *01 Walker

, 705 stick Walking -6 , 188 Walnut K. 282 Walter -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Wasp mantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds 70 89 77, 97, 2089, 212,222,246, Weevil , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

-,

.

Weinmannia, 144 45 , 13840, 146 , Welfia georgii , 74 136 อ

.

1

-

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 West Indian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , faced 202,320 monkey White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wild ginger 221-22

.

---

.. .

.

-

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya plantain Wild 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E, 546 Willis -47, 607-8 ,MF . 191-93 Willson , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Wind pollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Wood warbler Wood wren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World FundUnited , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia, 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas, 80 Xanthopastis timais , Xamhoxylum setulosum, 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex, 275 Xestoblatta hamata , 693- 96

630

.

.

.

-10

, 777 -79 Xicote , F-, 1 Ximenez

293

, Ximenia americana , 150 , 589 Xiphorhynchus, 514 , Xylocopa,

.

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

621 627

Xylophones, turbata , , 208416 Xylosma, 147

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow grassquit -faced , 209 Yellow vireo -green , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum, 147 mayanum, melanostichum , 144 panamense , 140 , (seealsoXan setulosum, thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

277 -78 Zarza

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorra mochila Zorro pelon468 -69

-

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

.)

( com

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 า

.

;

-

H., 119 Wendland , c . , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

816

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c . , 3 - 4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 อ Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

.)

( com

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

.

. -

. .

.

1

-

.

---

-

.. .

-

.

.

-

.

.

.

9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , c. , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

าพ ี

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

816

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. c., 3-4 , 502 Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

าพ

-,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

---

.

.. .

.

-

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

INDEX , 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 502 404, 456 Vulture , 197 - 200 -5, 507, 517,552, Wildpineapple 560 , 252 - 53 Wildpitahaya Wildplantain 249 - 51 H. 57576 Wagner , R. H., 558 Wiley -60 M., 2, 119 Wagner WileA 7 Wagner H., 187,239-40, 311-18 , G. B., 224 Williamson -25 25659 , M. H., 400 Wake , E , 546 47, 607 Willis ^01 -8 Walker , M F. 191 Willson -93 , 705 stick Walking , D. E., 261-62, 426 Wilson -6 ^7, 47778 , 188 Walnut pusilla , 526 Wilsonia WalterK. 282 Wind38 40 42-44, 632 -92 Waltheria , 283 indica , 778 . See also Polli Windpollination , 525 , , 517, 523 , 507 Warblers 505 , bywind nation 575 , D M., 705 Windsor -76, 613-14 -6, 76062 , 358 . See also Nox coloration Warning , M- , 94-95 Winston iousinsects , 295 , 183 , 236 Winteraceae , J., 118 Warscewicz , 684 , 627 Wissadula , 138, 140 coccinea , 136 , 715 Witheringia Warscewiczio , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 716 Wolf500 , 712 , 737 , 615636 297386 , K., 659 Wolfe 739 , L., 76869 Wood -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid , T. K., 773 Wood -75 ; Ma , 505 Water birds , 641 , 631 , 678 Wood beetle boring -6. See also Duck -79 ; Shorebirds rinebirds 681-82, 719, 768-69 , 733 Waterbug , 546589 Woodcreeper -34 , 45 Waterfall onclimate effect , 554 Wood duck ,236 - 39 hyacinth Water , 546 Woodhaunter , 427 opossum Water , 657 . See also Satyridae nymph Wood ; , 35 46. See also Dryseason Weather , , 50455758, 614 329 Woodpecker Seasonality 691 -92, 770 , 52122 effect onbirds , 608 Wood pigeon , 415 - 16 Web frog Wood stork388 -footed Weeds70 89 . See tree species accounts Wood traits Weevil77, 97, 2089, 212,222,246, , 609 , 567 Woodwarbler , 627 , 271 , 324 , 326 , 348 , 629 255 Woodwren . 509 , 763 631, 698, 701, 719 , 427 opossum Wooly , 144 - 45 Weinmannia WarII, 6 67 World , 136, 138-40, 146, Welfia georgii , 74 , xi States Wildlife World Fund United , 406 F Worm snake , 558 , 5057, 524 Wren -60 H., 119 Wendland , 6 / 6 - 18 Wrenthrush , 4, 6, 119 Werkle Wyeomyia , 640 West M. J., 758 -Eberhard -60 Western sandpiper .557 , 80 Xamhomonas , 498 - 500 WestIndian manatee timais , 630 Xanthopastis , R. M., 465 Wetzel . Seealso Xamhoxylum setulosum , 195 -67 , 428 Whales Zamhoxylum , 66, 70 Wheat Xantusiidae . 359, 372 , 682 - 83 White fatima Xenarescus . 70910 -banded White .680 - 81 sphinxlet , 275 Xenomyrmex -banded White faced 458 - 60 capuchin Xestoblatta hamata , 693 - 96 , 329 , 439 , facedmonkey 202,320 White , 777 - 79 Xicote 458 - 60 , 764 , F, 1 Ximenez ,587 - 90 , 606 White fronted nunbird , 150 , 293 Ximenia americana peccary lipped White 471 , 589 , 514 Xiphorhynchus , 388 White necked heron , 241245299 , , 323 Xylocopa , 194210 , 547 White antbird , 633 628 -plumed -34 , 481 - 83 . See , 263 White tailed deer248 , 111 fimbriata , 215 also Deer , 215777 - 79 gualenensis , J. L., 281 Whitmore -82, 33133 muscaria . 215 Wildbitter gourdlet .211 - 12 viridis , 215 Wildginger , 335 , 633 221 - 22 . Seealso Xylocopa Xylocopidae Vriesea orosiense

( com

.)

. 593

.

.

.

.

-

*

.

.

.

.

.

.

*

.. .

-

..

. -

-

-

.

.

-

.

-

.

—

-

.

าพ

.

-

.

---

. -

. .

.

-

.. .

-

.

.

-

.

.

.

9

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

—. .

INDEX Vriesea orosiense

( com

.)

593 , 502 404, 456 Vulture -5, 507, 517,552, 560

.

Wagner . HM..,575 -76 Wagner 2, 119 พ. H., 187,239-40, 311-18 Wagner , M. H., 400 Wake ^*01 Walker พ า , 7056 stick Walking , 188 Walnut

Walter . K. ร., 282-92 indica ,

778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

Waltheria

.

-

iousinsects , J., 118 Warscewicz

, 138, 140 coccinea , 136 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711 - 12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 - 39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 - 16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

Warscewiczio

..

.

.

-

—

-

.

-,

.

, 144 45 Weinmannia , 13840, 146 , Welfia georgii , 74 136 F

-

.

H., 119 Wendland , f • 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498 - 500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 - 83 White fatima -banded White .680 - 81 sphinxlet -banded White faced 458 - 60 capuchin , 329 , 439 , facedmonkey 202,320 White 458 - 60 , 764 ,587 - 90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 - 83 . See , 263 White tailed deer248

.

--also

.. .

.

Deer , J. L., 281 Whitmore -82, 331 -33 Wildbitter gourdlet .211 - 12 Wildginger 221 - 22 /

, 345 - 46 Wildnutmeg , 264 - 65 , 633 Wildpapaya , 197 - 200 Wildpineapple , 252 - 53 Wildpitahaya Wildplantain 249 - 51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 pusilla , 526 Wilsonia Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula , 715 Witheringia , 716 Wolf500 , K., 659 Wolfe , ร. L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 - 18 Wrenthrush Wyeomyia , 640

.

.

.

.

. *

.. .

-

-

.

-

.

.

. -

. .

—

-

, 80 Xamhomonas timais , Xanthopastis Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae 709 Xenarescus , 275 Xenomyrmex Xestoblatta hamata , 693 - 96

630

.

.

.

-10

, 777 - 79 Xicote , F-, 1 Ximenez

, 150 , 293 514,589 , , 323 194210, 241245299 , 633 628 -34 , 111 fimbriata , 215 , 215777 - 79 gualenensis muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa 9 Xylocopidae

Ximenia americana , Xiphorhynchus Xylocopa ,

.

.

.

.

Xylophones turbata , Xylosma ,

, 621 627 147, 208416

.

, 70 Yams Year of Costa

Rican

, A,

History

Natural

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 - 5 / -linedgrasshopper , 681 - 82 borer byttnena Yellow spotted , 575 - 76 Yellow warbler , 610 - 11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 - 17 Yuca

-

.

, 462 Zacryptocerus californianus Zalophus , 349 fairchildiana skinneri , 349 - 50 , 349 - 50 Zamia Zanate 601 3

.

. 428

- 50

-

, 727 - 29 Zancudo

751

, 586 , , 147 Zamhoxylum mayanum , , 144 melanostichum , 140 panamense , (seealsoXan setulosum , ) thoxylum setulosum Zapote , 704 Zarhynchus wagleri . 614 - 16 468 - 69 Zarigiieya

147

127131

. .-

Zarza277 - 78

Zea mays , 88 92

, 607 antshrike Zebra -striped

• v • , J. f 0 J Zeledon Z , R., 8 Zeledon

Zeledonia .616 - 18

Zeledonia

coronata

, 616

- /8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales

.

, 688 - 91 Zompopa

Zonotrichia

capensis

, 618

, 392 - 93 , 398 - 99 Zopilota , 560 - 62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 - 69

.

.

183 Zygophyllaceae

-

5

INDEX Vriesea ( com . ) orosiense 593 , 5025, 507, 517,552, 404, 456 Vulture

.

.

560

-

.

Wagner . HM..,575 -76 Wagner 2, 119 Wagner H., 187,239-40, 311-18 , M. H., 400 Wake ^*01 Walker

, 705 stick Walking -6 , 188 Walnut WalterK. 282 -92 Waltheria indica , 778 , 525 , , 517, 523 , 507 Warblers 505 575 -76, 613-14 , 358 . See also Nox coloration Warning

.

.

-

iousinsects , J., 118 Warscewicz

136,138, 140 , 261 , 202 , 283, (aculeate ), 76 191 Wasp , 712 , 737 , 615636 297386 739 -40, 758-60, 760-62, 766 , 711-12 Waspmantispid ; Ma , 505 Water birds -6. See also Duck ; Shorebirds rinebirds , 733 Waterbug -34 , 45 Waterfall onclimate effect ,236 -39 hyacinth Water , 427 opossum Water ; , 35 46. See also Dryseason Weather Seasonality , 52122 effect onbirds , 415 -16 Web frog -footed Weeds70 89 Weevil77, 97, 2089, 212,222,246, , 627 , 271 , 324 , 326 , 348 , 629 255 , 763 631, 698, 701, 719

..

Warscewiczio coccinea ,

.

.

-

—

-

.

าพ

-,

.

Weinmannia , 144 45 , 13840, 146 , Welfia georgii , 74 136

-

.

H., 119 Wendland , 4, 6, 119 Werkle West M. J., 758 -Eberhard -60 Western sandpiper .557 , 498-500 WestIndian manatee , R. M., 465 Wetzel -67 , 428 Whales , 66, 70 Wheat , 682 -83 White fatima -banded White .680-81 sphinxlet -banded White faced 458 -60 capuchin , 329 , 439 , facedmonkey 202,320 White 458-60 , 764 ,587-90 , 606 White fronted nunbird peccary lipped White 471 , 388 White necked heron , 547 White antbird -plumed , 481 -83 . See , 263 White tailed deer248 also Deer , J. L., 281 Whitmore -82, 33133 Wildbitter gourdlet .211-12 Wildginger 221-22

---

.

.. .

.

-

, 345 -46 Wildnutmeg , 264 -65 , 633 Wildpapaya , 197 -200 Wildpineapple , 252-53 Wildpitahaya Wildplantain 249-51 , R. H., 558 Wiley -60 WileA 7 , G. B., 224 Williamson -25 25659 , E , 546 Willis -47, 607-8 , M F. 191 Willson -93 , D. E., 261-62, 426 Wilson ^7, 47778 Wilsonia pusilla , 526 Wind38 40 42-44, 632 , 283 . See also Polli Windpollination , bywind nation , D M., 705 Windsor -6, 76062 , M- , 94-95 Winston , 295 , 183 , 236 Winteraceae , 684 , 627 Wissadula Witheringia , 715 , 716 Wolf500 , K., 659 Wolfe , L., 76869 Wood , T. K., 773 Wood -75 , 641 , 631 , 678 Wood beetle boring -79 681-82, 719, 768-69 , 546589 Woodcreeper , 554 Wood duck , 546 Woodhaunter , 657 . See also Satyridae nymph Wood , , 50455758, 614 329 Woodpecker 691 -92, 770 , 608 Wood pigeon Wood stork388 . See tree species accounts Wood traits , 609 , 567 Woodwarbler Woodwren . 509 , 427 opossum Wooly WarII, 6 67 World , xi States Wildlife World Fund United , 406 Worm snake , 558 , 5057, 524 Wren -60 , 6 / 6 -18 Wrenthrush Wyeomyia , 640

.

.

.

. -

.

*

.. .

-

-

.

-

.

.

. -

. .

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195Seealso Zamhoxylum , 372 359 Xantusiidae Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

.

, 70 Yams Year of Costa Rican Natural History, A ,

, 604 Yellow -facedgrassquit , 209 Yellow -greenvireo , 561 Yellow headed vulture Yellow 750 -5 / -linedgrasshopper , 681 -82 borer byttnena Yellow spotted , 575-76 Yellow warbler , 610-11 Yiquirro , A. M., ix, 724 Young -26 , 66, 114 -17 Yuca

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 , 349-50 Zamia Zanate 601 3

.

-

, 727 -29 Zancudo

751

, 586 , Zamhoxylum , 147 mayanum , melanostichum , 144 panamense , 140 , (seealsoXan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 468 -69 Zarigiieya

147

127131

. .-

Zarza277-78

Zea mays , 88 92

, 607 antshrike Zebra -striped

, J. Zeledon , R., 8 Zeledon

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8

, 616 , 505 Zeledoniidae -18 , 708 , 621 Zingiberaceae 708 Zingiberales , 688-91 Zompopa

.

-

.

.

.

183 Zygophyllaceae

293

, Ximenia americana , 150 , 589 Xiphorhynchus , 514 , Xylocopa ,

.

Zonotrichia capensis , 618 , 392 93 , 398 -99 Zopilota , 560-62 cabecirrojo Zopilote , 468 -69 Zorramochila Zorro pelon468 -69

-10

, 777 -79 Xicote , F-, 1 Ximenez

621 627

Xylophones , turbata , , 208416 Xylosma , 147

.

, , 323 194210241. 245299 , 633 628 -34 , 111 fimbriata , 215 gualenensis , 215777 -79 muscaria . 215 viridis , 215 , 335 , 633 . Seealso Xylocopa Xylocopidae

.

9

-

5

INDEX

—. .

Vriesea ( com . ) orosiense 593 Vulture 404 , 456, 502

560

-5, 507, 517, 552,

.

Wagner. H. 575-76 Wagner M ., 2, 119 Wagner พ. H . , 187, 239-40, 311-18 Wake , M . H., 400^*01 Walker พ า Walking stick , Walnut , 188

.

.

.

Waltheria indica ,

ious insects Warscewicz , J . , 118

-

136, 138 , 140 Wasp ( aculeate), 76 191 , 202, 261 , 283, 297 386, 615 636, 712, 737 739-40, 758-60, 760-62, 766 Wasp mantispid , 711-12 Water birds , 505-6. See also Duck; Ma rine birds; Shorebirds Waterbug , 733-34 Waterfall effect on climate , 45 Water hyacinth , 236 -39 Water opossum , 427 Weather, 35 46. See also Dry season; Seasonality effect on birds , 521 22 Web-footed frog , 415 -16 Weeds 70 89 Weevil 77 , 97, 208 9, 212, 222, 246 , 255, 271, 324 , 326 , 348, 627, 629 631 , 698 , 701, 719, 763

..

Warscewiczio coccinea ,

.

.

-

—

-

.

พร^ -,

Weinmannia , 144 45 Welfia georgii , 74 136, 5

.

.

138-40, 146,

Wendland H . , 119 Werkle , c. , 4 , 6, 119 West - Eberhard M . J . , 758-60 Western sandpiper. 557 West Indian manatee , 498-500 Wetzel , R . M . , 465-67 Whales , 428 Wheat , 66 , 70 White- banded fatima , 682 -83 White- banded sphinxlet . 680-81 White faced capuchin 458 -60 White faced monkey 202, 320, 329, 439, 458-60 , 764 White fronted nunbird , 587-90 , 606 White lipped peccary 471 White necked heron , 388 White- plumed antbird , 547 White tailed deer 248, 263, 481-83 . See also Deer Whitmore , J . L., 281 -82, 331 33 Wild bitter gourdlet . 211-12 Wild ginger 221-22

---

816

.

.. .

.

-

.

.

705-6

Walter K . ร., 282-92 778 Warblers 505, 507, 517, 523, 525, 575-76, 613-14 Warning coloration , 358. See also Nox

Wild nutmeg , 345-46 Wild papaya , 264 -65 , 633 Wild pineapple , 197 -200 Wild pitahaya , 252-53 Wild plantain 249-51 Wiley, R . H . , 558-60 Wile A 7 Williamson , G. B . , 224-25 256 59 Willis , E , 546-47, 607-8 Willson , M F. 191-93 Wilson , D. E. , 261-62, 426^ 7, 477 78 Wilsonia pusilla , 526 Wind 38 40 42-44 , 632 Wind pollination , 283. See also Polli nation , by wind Windsor, D M . , 705-6, 760 62 Winston , M - , 94 -95 Winteraceae , 183, 236, 295 Wissadula , 627, 684 Witheringia , 715 Wolf 500, 716 Wolfe , K ., 659 Wood , ร. L., 768 69 Wood , T. K . , 773-75 Wood boring beetle, 631 , 641, 678-79 681 -82, 719 , 768-69 Woodcreeper, 546 589 Wood duck , 554 Woodhaunter, 546 Wood nymph , 657. See also Satyridae Woodpecker 329, 504 557 58, 614, 691-92, 770 Wood pigeon , 608 Wood stork 388 Wood traits. See tree species accounts Wood warbler, 567, 609 Wood wren . 509 Wooly opossum , 427 World War II , 6 67 World Wildlife Fund United States , xi Worm snake , 406 Wren , 505 7, 524 , 558-60 Wrenthrush , 6 / 6 -18 Wyeomyia , 640

..

. -

.

*

.

-

.

-

-

.

.

. -

. .

—

-

Xamhomonas , 80 Xanthopastis timais , Xamhoxylum setulosum , 195 Zamhoxylum Xantusiidae 359, 372 Xenarescus 709 Xenomyrmex , 275 Xestoblatta hamata , 693- 96

630

.

.

Ximenia americana , 150, Xiphorhynchus , 514 , 589 , Xylocopa ,

.

Yams , 70 Year of Costa Rican Natural History, A ,

Yellow-faced grassquit , 604 Yellow-green vireo , 209 Yellow headed vulture, 561 Yellow- lined grasshopper 750-5 / Yellow spotted byttnena borer, 681-82 Yellow warbler, 575-76 Yiquirro, 610-11 Young , A . M . , ix , 724-26 Yuca , 66 , 114 -17

-

.

Zacryptocerus , 462 Zalophus californianus

. 428

fairchildiana , 349-50 skinneri , 349 -50 Zamia , 349-50 Zanate 601 3

.

-

Zancudo , 727 -29

751

Zamhoxylum , 147 , 586, mayanum , melanostichum , 144 panamense , 140 , ( see also Xan setulosum , thoxylum setulosum ) Zapote , 704 Zarhynchus wagleri . 614 - 16 Zarigiieya 468 -69

147

127 131

Zarza

. .- -

277 78 Zea mays , 88 92

Zebra-striped antshrike , 607

Zeledon , J. c., 3-4, 502 Zeledon , R . , 8

.

Zeledonia 616 -18 Zeledonia coronata , 616 - / 8 Zeledoniidae , 505, 616-18 Zingiberaceae , 621 , 708 Zingiberales 708 Zompopa , 688-91

.

-

. See also

293

.

Zonotrichia capensis , 618 Zopilota , 392 93 , 398 -99 Zopilote cabecirrojo, 560-62 Zorra mochila , 468 -69 Zorro pelon 468 -69

.

-10

Xicote , 777 -79 Ximenez , F- , 1

621 627 208 416

Xylophones , turbata , Xylosma , 147 ,

.

Zygophyllaceae 183

.

194 210 241. 245 299, 323, 628, 633-34 fimbriata , 215, 111 gualenensis , 215 777 -79 muscaria . 215 viridis , 215 Xylocopidae , 335, 633. See also Xylocopa

.

9

-

5