\ 1'11 1111 W/III:I' IIIII.I 11m In unc" Juurn.tl ('flntcnl flnlnll' (rull-ll' \I), Sllh~( 111011 .. 11111\ H'J!I\kr \\JlII 1IIJ.:1'lIlu rur rrCl du: lrulllC ddIH'I ) \Ji! Ihe In lCrncl, '\/IIl- \u l"lnh,'r\ ('un uhlmn rull -h ,, 1 urlld"\.1\ IlIn-,ll' r -' 11' \\ \ , I n 'l'h \c;trch ;/IJ.:t'IlII1'\ Jllul'n.ll u,II.IIIU\I' (m .Irl\ \,111111 ,u'.ll Juurnnh) 1" 1,1'\ \\411 u , 1j.!J~ , Zlnd lIulhor (Jl)urnul :lrlldl uh,lnJCh :l\utluhl:,'oñhne rr('(' IhllJlI¡.:h 111/.:('11111) .

the diversity of life

Now in paper!

M

Tadpoles . ¡ 111 mOl tl l , \ UI \C\l J( \:->! \1(\ \1

Tadpoles The Biology of Anuran Larvae Edited by Roy W. l'IcDiarmid and Ronald A.ltig

"A brillianf compilarian of information on

che world's most commOll aquatic amphib­ians."-Richard Wassúsug, Copad

"The prcscm volume, a Hue magTlllrtl opus,

is a compr~hcnsive account of larval anu­mn biology and wjll no doubt hecome a

reference tome comparable with Dudlman and -¡-rueb's dassic on amphibian biology.

Tadpo/~s is a substantial contribmion [O

rhe ¡iterature and a wdcomc addition to my bookshdf. ... IAI rieh mine of infor­

m:Hion ro which many of liS w ill be refer­

ring fOf years to come."-Trevor Beche, Trends 1n Ec%gy and Evolution 458 pagas 11 8 halftones, 181 1ine drawings, 23 tables Paper $40.00

The Ecology of Invasions by Animals and Plants Charles S. Elton ",,¡¡J, 11 fort1wrd by Dlmitl SimbniojJ

"A hook has ro be rcally special to be

republished 40 years afrer ir fi rst appcarcd. Elton's book ¡s. The ropie is even more

important roday than ir was back rhen,

bccause invasivc spccies may now be the

mas! important cause of specics cxrinction. This book is a classic thar I would likc all of my studcnts lO rcad. ~ -Stuart Pinun, University ofTcnncsscc, Knoxvillc Paper $13.00

New from Chicago

In Search of the Golden Frog l'Iarty Crump

"[Tl ltis is nor jusr a book abollr frogsi fascinaring storics abotlr birds, insects, plants, lizards, and snakcs a[so abound. Complicatcd biologicll .~tor i cs are told concisely bm rhoroughly. 3nd her humor :H dcscribing natural history rraits makes them very rcadablc. . . Thc story of Marty Crump'~ life in the ficld is fitscinaring, and

mIel with respect foc both the readcr and rhe narufa! world . 'rhis is an cxrrcmely !ikable and rcadable story of a dcdicated scicmist in search of her go!dcn frog." -So/loroll H~rp(fo¡ogico 320 pages 16 color plates, 60 halftones, 19 maps Cloth $27.00

".

A Naturalist's Guide to the Tropics Marco Lambertini

"This well~i1!lIs[r:Hcd book takes a broad view of a remarbble range of tropical anribll tcs: di mate, 50ils, flora, F.lUlla, man­groves, forcsts and so on. Many exa mplcs are highlighred in detail. ... $0 while a guidc ro individual species is impossible, an cnlighecning cxp!anarion of much to interese any tropical travellcr is dearly pre­sented .... Emhu$iasm for rhe subjecr is obvious: rhe sheer bri!iiancc and variery of tropical life astonishing.~-Birds 348 pages 11 cotor ptates, 51 color photos, 21 maps, 76 line drawings Paper $25.00

FOUNDER AND EDITOR Craig Hassapakis Amphibian and Reptile Conservation Modesto, California

ASSOCIATE EDITOR Jack W. Sites, Jr. Departmenl 01 ZooIogy Brigham Young UnlverSlty

Allison C . Alberts Center fOf Reproduclion 01 Endangered Species Zoologlcal Soclety 01 San Diego

Jonathan D. Ballou Nabooal ZooIoglcal Park Smllhsonian Institution

Aaron M. Bauer Department 01 8iology Villanova Universlty

Andrew R. Blaustein Department 01 ZOOogy Oregon Stale Unlversity

Harold G. Cogger Australlan Museum Sydney. AUSTRALIA

C. Kenneth Dodd, Jr. Biologlcal Resources Division U S Geologlcal Survey

Lee A. Fitzgerald Department 01 Wildllle and Fisheries Sc!ences TexasA&M Unlversity

Julian C. Lee Department 01 BioIogy Umverslty 01 Mlami

ADVISORY BOARD

Joseph T. Collins Natural History Museum Universityof Kansas

Carl Gans Adjunct ProfesSO( 01 ZooIogy University 01 TeKas al Austin

Roy W. McDiarmid Herpetology. 8iological Resources Division U.S. Geological Survey

Russell A. Mittermeier Presidenl, Conservalion International Washington, O.e

EDITORIAL REVIEW BOARD

Harvey B. Lillywhite Departmentol Zoology University 01 Florida

Peter V. Lindeman Departmen t 01 Biology and Health Services Edinboro University 01 Pennsylvania

Joseph C . Mitchell Department 01 Biology University 01 Rlchmond

Henry R. Mushinsky Departmentof Biology UniverSlty 01 Florida

SECTION EDITORS

George B. Rabb Presldent. Chicago ZooIoglcal Socie\)' Vice chalr, Commun,eabons. Speaes Survlval Commlssion, The Wor1d ConseNatlor.lJmon

Hobart M, Smith Departmeot of EnVlronmental. PopuIatlon and Organlsmlc Blology UniverSI\)' of Colorado

Michael Soulé Research Prolessor. University 01 California at Santa Cruz Presiden!. The Wildlands Project

Jaime E. Péfaur Ecologla Animal , Facultad de CienCias Universidad de Los Andes, Mérida, VENEZUELA

Christopher J. Raxworthy Department 01 Herpetology Amenean Museum 01 Natural History

Andrew T. Storfer Wildllfe EcoIogy and Conservabon Unlverslty 01 Florida

Robert J. Wiese AssIstant Olrector 01 Animal Programs FortWorthZoo

AMPH IBIAN ECOLOGYAND CONSERVATION Jamie K. Reaser

STATISTICSAND EXPERIMENTAL DESIGN Anthony J, Krzysik

U S Departmenl of State. Bureau of Oceans and Intemational Enwonmental. and Scienbfic AIfaIrs

School 01 AAs and $clences Embry-Riddle Aeronautical Unlverslty

OfIice of EcoIogy and Terrestrial Conservabon. Washington. O C .

COPY EDITOR Mark l. Goodwin

COLUMNIST Craig M. Hoover Senior Program Officer, TRAFFIC North Amenea, Worfd Wildlife Fund Washnlgton. O.C

ARGENTINA Maria E. Bridarolli In$llMo ArgentIno de InvestigaCIones de las Zonas Andas. Centro Regional de InvestJgaoones Cientificas yTecnoI6glCas, Mendoza

GALAPAGOSARCHIPELAGO (ECUADOR) Marco Altamirano Museum olSouthwestern 8iology, The University 01 New Mexlco and Charles Darwin Research Station Galapagos. ECUADOR

INDIAAND MALAYSIA Indraneil Das Instltule of BIodiversity and Environmental Conservation

CONTRIBUTORS

GRAPHIC DESIGN Kenneth W. Sholar

INDEXING AND ABSTRACTING CONSULTANTIE-PUBLlSHING

Michael J . Tavares

WEBSITE DEVELOPMENT AND CONSULTANT

Jay G. Henry

COUNTRY L1AISONS

ITALY AND MADAGASCAR Franco Andreone

COMPUTER SUPPORT Mark Warren

ZOOLlAISON Chris Banks Curat()(of Herpetofauna, lnvertebrates & Educanon Animal! and Co-ordlnator South East ASlan Conservabon Programs MelboumeZoo. AUSTRALIA

TA IWAN David McLeod

Sezione di Zoologia. Museo Regionale di Scienze Naturali . TonnO, lTALY

Nabonal Expenmental Hlgh School al $clenee Based Industnal Park, HUlflchu

INDONESIA Darmawan Liswanto Executive Director, Yayasan Titian Jakarta

PERU Antonio W. Salas Museo de Historia Natural Universidad Rieardo Palma

U.S. VIRGIN ISLANDS Father Alejandro J . Sanchez Munoz Pastoral Saint Anne's Pansh Saint Thomas

VENEZUELA Jaime E. Péfaur Ecologla Animal , Facultad de Cienoas UnlVet"$ldad de Los Andes, Ménda

UnlVef$rti Malaysia Sarawak, Sarawak, MALAYSIA SOUTH AFRICA Marius Burger National Coordinator. SouthAfrican Frag Atlas Project. University 01 Cape Town

Cop), i¡¡ht e 2000 A"'p~jhj~ .. " .. .1 R.p,d. C" ...... atio ... AII ,ights ,ese"ed, ARC a utho,i l es pl>otoc,QP)inj\ ro, inte,nal o' ¡l<',son.1 use p,ovided ¡he .pp'Op,i 'l ' f.e is p. id di"'II) 10 ¡he COPJ"/l~' (/~"TU"C' Cu,~,. 1",'. 222 Rose"ood 1), .. Oan""$. 101 ,\ 01923·4599. USA T~I, (978) 7S0·8400¡ f .. : (971) 7~O·4no¡ e .... il : i"foacOf'}Tj¡:h'com; ".bsil~: .... ' .. '.c"P>riX~,.c"'"

Amphibian and Reptile Conservat ion 2(2):40.

Editorial WORDS FROM TI-lE EDITOR­With the complction of thi s issue, we have success full )' publi shed our second volume. Our mi ss ion is funher set, and I think it easy to visuali ze our intended mission, based on the observation of each new issue. Amphibian and Reptile Con­servation (ARC) now numbers over 700+ subscribcrs and has printed and distributed 3,000 copies each ofboth issues 2 and 3, thus making ARC the largest distributed, Engl ish language, herpetological scient ific journal in the world . 1 view my role with the journal as "carrying the torch" as editor and publi sher, ",hile many interested in ass isting the journal have come "on board" as ad visors or editors, specia l section editors , contributors, in-country liaisons, and in other "'ays. 1 am gratefu l for all of th is assistance, and ",ould li ke to invite others with a special interest or talenl to contact me for more informat ion and di scuss ion. 1 am always open to feedback, new input, and ideas regarding the journal andl or the eonserva lion of amphibians and reptiles worldwide, as "'ell as all life. Again , I invite alllo participate

Helping right a wrong . Our first day filming a documentary al Helch Hetchy reservoir (background) and holding one of our extras. a bullsnake Pituophis melanO/eucus, which wandered across our trail. Reference : I1'wlI'.hetcl1lw/chy. o/"g

proved to be fartoo few as \Ve quick ly ran out of stoek in aboul February of 1997, as ",e continued 10 ha ve a strong interest in the journal. After a move to the University of AriLona in Tueson, Arizona from my alma mater of BYU in Provo, Utah, in the late faH of 1997 as Ihe first snow began to fal l in Pravo, the journal and myse lf \Vcrc exposed to a new and exciting landscape. Work on the journal was agonizing ly s lo"', bul 1 continued lo pre5evere and make some progress. To help move the journal for\Vard, 1 again packed up and moved lO my hometown of Modesto , Ca li l"orn ia in the lale summer of 1999, 10 be closer lo my fami ly and their influencc. Al each step ofthe \Vay, throughollt the years, I continued in good failh to sleadil y \Vork and make needed associat ions to further the \vork on the journal , though not all of these assoc iations \Vere fruitfu1. 1 have left out many of the gory detail s for lack of space here, and /or left to some future conference gathering (informally), l can attest to the fae l thal il hasn't been an easy path to fol low \Vith many lIncertai nli cs a nd sc tbaeks. O nc

in what 1 feel is olle of the mOSI important conccrns for a ll herpclologisls, nOI just ones doing conservation related work. Now I "'ould like 10 briefly oulline our progress thus far.

The idea for a jOllrnal devoted strictl y to Ihe conservat ion of amphibians and rept il es evolved after 1 sadly left a brief sti nt at dental school in 1991 , much too early to ever praclice. Thollgh it ",as a ralher down time for me then (and nOl the last hurdle 1 would have to overcome), 1 found that it ",ould be severa l more years before 1 \Vou ld determine whal my lifc's "'ork couJd be. Afler spending about three ycars doing a lot ofbook",ork in the libraries of primarily Utah afier leaving dental schoot , 1 met a publisher, zoologi5t, and a formcr classmate of mine [in my first graduate level class as a studenl at Brigham Young University (BY U»), Jay Vilhena. It \Vas he that most direcled my interest in herpetology inlo someth ing more concrete and credit should be kindly given. On a long road trip 10 New Mexico, to deliver a computer system, 1 told Jay 1 needed hclp wil h focu s in g my ideas of writing, research, conservation , and herpelology. Afie r a long and contemplatcd diseuss ion, as \Ve1l as his background as a long-time friend and assoe iale, Jay an nounced, Ihal 1 should publi sh a newslctter dcvotcd to m)' inlcresls. This was 1994. After much thought, more lime, and finally, some slrong persuasions ofa spiritual naturc, I agrced lo undertakc the task. Event ua ll y, 1 progressed the idea I"rom a ne",sleuer, to a scielllific journal , lo last ly, what l no", refer to as a magazine stylcd, scientificjournal. In October of 1996 \Ve published our firs l modest journal. Fiftecn hundrcd copies of Ihe first issue

ce rt ainty is my unwavering commilment to cont inue forward regardless of ",hat mllst be overcome along the way. Further, as conservation biotogist Michael Soulé so accuralely stated by lelling me in a telephone conversion, it " takes a lot of gllmption" to start ajourna l as I have done, and 1 \Vould add, cven more 10 cont1 nue in Ihe face o f advcrsity and uncertainty. Howcver, wi lh all slrllggles and tribulations comes moments 01' greal joy and satis faclion for those who cont inue on. With lhe con linllcd pub liealion of Ihi s journal I in vite all to share in our sllccess. It is my rock so l id, determined, and solcmn de votion lO continllc on with the idea that there should be a journal accessibte by as many people as poss ible, devoted 10 Ihe conservation and prescrvation of amphibians and repti les, and their habitats worldwide. Words ofmy favori le musica l arti st, whom I had the pleasure to meet just before 1 left Tucson, comes to mind as he eloquen lly sang in the album, Late/or rheSky, and whal 1 truly feel about my miss ion with the journal: ·'Keep a Jire hum '/1 i/1 y01l1' eye ... Don 't let rhe uncertainty tllrnyolI al'Ound .. 00 ahead (md throw sorne seeds o/ yOtlr own (lIId somewhere hetween the time you arrive and the time you go. may Iie a reason yOIl were a/ive." (Jackson Browne 1974, se lect lines from the song, Fo,. a Dance,.). It is wilh lhe publicat ion of lhis iss lIe Ih<lt ",e gain momenlutll in contributing to the conservation 01' arnphibians ami repti les worldwide.

Craig I fassapakis FOllnde/: EditO!; and PubJisher

Authors JAIME E. I>ÉFAUR is Professo r of Ecology at the Fac ulty of Sc iencc, Un iversi ly o f Los Andes in Mérida , Ve nezue la. He graduated as a Veterinarian from the Univers ity ofChi le and reec ivcd a Masters and Ph.D. ( in 1979) degrees frolll the Univc rs ily ofKansas. Dr. Péfaur has published over 90 papers dea ling wilh taxonom y and eco logy o f South American vertebrates, as \-Vell as edited several books. He was lhe Exccul ivc Secrelary o flhe II Lat in American Congress of l-Ierpeto logy in 1990 and o flhe III Latin Amcri can Congress of Eco logy in 1995. He \Vas a member oflhe Int crnal ional Union for Conservalion ofNalurc (IUe )'5 De­c1ining Amphibian Populations Task Force Board o f Directors.

JUAN A. RIVERO is Dislingu ished Professor al lhe Un iversily of Puerto Rico, Mayagücz. I-Ie recci vcd hi s Ph.D. from Ilarvard University in 1953. H is dissertalion \Vas on lhe Salientia of Venezuela, which has become lhe c1ass ic text book for any herpeto logy studenl of Lalin America. I le has publi shed ove r 200 pape rs, mainly dca ling wilh frogs ofVenezuela, Colombia, and Ecuador. He has also aulhorcd a book on lhe HerpelOfauna of Puerto Rico.

40

CONTENTS Volume 2 Number 2

42

40

SPECIAL FEATURE ISSUE

VENEZUE Distribution, species-richness, endemism, and conservation of Venezuelan amphibians and reptiles Jaime E. Péfaur and Juan A. Rivera

DEPARTMENTS EDITORIAL Words from the editor Craig Hassapakis

Authors

Background screen photograph (see in color on page 69): A view of the ¡nside of San Eusebio cloud forest, in the Venezuela n Andes. Tree ferns and bromel ids are characteristics. Venezuela , Mérida. Piloto: Jaime E. Péfall/:

Cover The polkadot tree Irog (Hyla punetata) is a common inhabitant 01 the largest South American ecosystem , the Amazon , and it is also lound within the borders 01 Venezuela. It dwells in both primary and secondary humid lorests and is common in marshy areas along river borders , lakes , and Ilooded grasslands 01 lowland South America lrom Venezuela and Trinidad south to the Paraguayan Chaco, east 01 the Andes mountains . This treelrog beco mes almost entirely red at night. The ir diet consists 01 small inver­tebrates . Photo kindly provided by Goil Shumway.

SCOpt: Amphibian and Replilt Conul'\'afion (ISSN: I083-446X) [ARC] and ¡he accompanying online edition ( ISSN: 1525·9153) is a popular!)' accusible. peer-reviev.ed scientific joumal of inlemational scopc. "hich is dc\'oted lO the worldwide presco'ation and managcmcnt of amphibian and rcptilian di\'ersity. Online rdit ion: The full-te:d online edition i5 a\'ailable 10 substribers FREE-OF-CHARGE as POF (Ponable [)oo;:umcn! Fonnat) files throug.h ¡rtgtllltl at: www.ingenla.com.Theontineeditionma).\·ar}.slighll}. from the print edition due to our reducing file 5izes fOl' efficienl downloading O\'cr the Internet. Sorne background streen5 (photographs) are removed whkh are dccmed nOI essential lo Ihe conlent oflhe atticle{s). Thcre is also sorne I05S of darit) 10 photographs in redudng article file sizes to a minimum. Ir clarity of phot05 i5 11. problem. the print edition or lhe journal should be consulted. Auditnce: ARC is intended for a wide readenhip from nonprofe!5iOllal to professional herprlologists. the general public. II.nd sc ientisls. Frequeney : ARC publishes two issues per )'e31 ($emi-annuall)"). Focus: ARC concentrates on publishing timdy infonnation in Ihe fonn or fealure atticles, original papen and data. re\ iews. repons. shon communicatiOllS. columns. commentaries. boot.: re"icws. editorials , and news and notes. Oinribulioo: ARC is distributed worldwide b) 5ubscription as well as quaJit} newsstands. boot.:storC"S. and selec! vendOD. Iklh el)' is guaranteed. ARC is also available as par-for-"iew rull -text artic les online wilh illgenltl. Includcd and 8\'ailabte ontine al ingenIa are full ab5tracls and complete bibliographic entries. Much ofthis Jame material plus more. such as FREE samplc art iclc(s). are available al the ARe website (www.herpetofalma.C"om). Publishtr: Craig Hassapakis. Amphlbiart and Repti/e Comen·arion. 2525 lowa Ayenue. Modesto. California 9~3~8-9467. USA. Fax: (509) 695-8747; email: publi$her@herpetofauna.eom; wcbslle : www.herpetofauna.com. PO!l ma, te r : Picase send change of addrns 10 ARC eighl \\'eet.:s prior to mo\'e. Copyright : AH cont~nlS copyright e 2000 Amphibi(1I1 (Jnd Reprile ConJen'(Jtion. All riglllS reseo·ed. No pan of Ihis publication ma)' be reproduced, slored in a retrieval s)"stem or lransmined in any form or by any means. electronic. mechanical. photocopying. recording, or olherwise wilhoul prior ~nni"ion. PbOloeopyiog: ARC aUlhorizes pholocop)'ing rOl' internal or personal use pro\ided the appropriate fee is paid direcll}' to Ihe Copyright Clearance Center. Ine .• 222 Rosewood Orive. Oanven. MauachusellS 01923-4599. USA. Tel: (978) 750-8400; fu: (978) 7504470; \\eb5ite: "'_' cop)Tight.com; email: Info@;cop)Tighl.com. CommuDiea tioos: h is rrcornmended Ihal all cOTTespondence be conducted "ia cmail: ARC@herpelOfauna.com .... hene-\"l.rp05sible.Allmhcr t)"pes of correspondener should be direeled lo ARC using Ihe conlacl information abol'e under publishcr. Sobseriptioos: AH SUNcriben receÍ\'e fOUT consecutively disuibutcd issues with a minimum ofl\\"o copies per ycar. Calegorln: Sub5crlption nlle categories are indi\'idual United Stales (US)$20 and institutional $40 (ineludes poslage). Al! subscriptions oulside Ihe US include $7 for shipping and handling 10 regular subscriptiOll rale (non-US nI!e: individual $27 and inslÍlutional $47). Sin¡:lt Issue Pun:blUe: Sub5criplions to lhe journal are encouraged to reduce shippin, COSIS and paper work bUI JQme ma)' \\"ish 11,1 purchase single issues as they bt-cumr avoilable. The eost ror singlc copies is US$S.95 plus t\\'u dollars shipping wilhin the United Slate5. Oulside the US plea5e mquire about shipping costs. Pa)"-for-\ ie\\" melhod of securing articles oolio1.' is available through ingtnla at ww ..... ingenta.r:om. Arti1.'lc price charge is $8 except ror larger tilles, which increase in priee. Further pricing infonnalion available al ingenta. Blck luuu : Individual copies are available from Ihe publisher rol' SI 2 each (Note: Volume 1. Number 1 is out-uf·print). Facsimiles (photocopies) are avai1able for $10 eath once i!sue! are out-of-print. Copies of individual artides can be purchased for S8 each excepl for artidcs co\'ering an cntire issue and these sell rOl' Ihe regular single-issue price. Duna llool : ConlributiOlls for the journal are gladl)" accepted. These specifie resources are used to furthcr Ihe conselVation of amphibians and reptiles "ilh the- goals of the journal. Cl le¡:orlu inelude: Fricnds of ARC $40. Supporling $60. Contributing $100. Patron 5250. Sustaining 5500. and Foundu (ineludes lirelime subseriptiun to ARC) $ 1.000. AII other conlributors. excluding Founden. reeei\·e four eonseeulivcl}' distribulcd issues. PI)'ment: AII journal orden must be paid in US cUrTenc)" dra\\ll on a US bant.: (credit cardo check. or international money order). C r t dil Cl rd pa)'m'nl : Cards accepled are Visa. MilJterctm:!. DiscO\·er. and American Expnll. Wtbsilt : ....... ""herpetofauna.cam. Produclion: Manufaclured and printed in Ihe Unitcd States of Ameriea using rccycJed paper (containing 100/. posl consumed fibcr) and soy bascd int.:s.

@ Thls pape!" meets !he requlrements of ANSIINISO Z39.48-1992 (Recom~rmed 1997) [American National Standard fof Permanence of Paper ror P~blicalions and Documenta in libraries and ArchIVEIS]

In_ud by 8tO$I$ Prevlttw$ (includes BioIogieaII AbliIraClt) [w"." b.brit"'lll, eBSCO Onlloe (""""-111 ~bIco,,,,,,,.',",,I,, ... ): ingem¡I ¡ .......... Ilt$.MI .• -om); Maruzen Kna«ledgewOfker (,,_"''''''' .... '''' 1¡>¡"".,~lot>Itw·hlml): MRr'IIa e~onic €. Ont.ne SorvioM ( .. ." ....... """u<M)". OnIIne ~ Llbrary c.n.. FntSeardl ( .. ." ... ,,,."I,·oog:)hluaodr): ROMICom Infoonation auesl ( ........ ri¡, ...... ): SwetsoeI N;rv;g.,tOr ( ........ .ft . .-tJ,>.&rho..u. ...... ): IIId ZooIogK:aI Record ( ........ ""'" .. ""JI ,

•

Pseudoboa neuweiddii, Local ion: Venezuela, Guatopo.

Al! plwfugmpll\' UI1J ¡1I1/\'(ral¡()/lS prol'ido:d hy ¡he 0/111/01', J(jime E P(;filw; l/lIle,x slUled olherll'ise

Cop~right 10 1000 A.mpl';l>ia~ and R'pril, C<m" .. ','a'iQn. AII ri ghts re,e rved. II RC author ile, pho!ocop)'¡~g fo. ¡nle mol or p.nonal u'" providod lhe "ppropriate f •• il p. id dir.011)' lO ,he Cop)right Cl,",an,e Ce nter. lnc.. 222 Ro,"" ood Dr.. DanHrs. MA 0 1923_4599. USA. Te l (978) 750-8400: fax : 1975 ) 750·4470; cmoil : ¡njo'¡¡ C()Pyn'lIht.c()m: ".b. i,. : "'''''', COPY''lIht ''OIn

Amphibian and Reptile Conservation 2(2):42·70 .

Distribution, species-richness, endemism, and conservation of Venezuela n amphibians and reptiles

JAIME E. PÉFAUR,,2 AND JUAN A. RIVERO'

IEcologia Animal, Facultad de Ciencias, Universidad de Los Andes, Mérida. VENEZUELA JDepartamemo de Biología, Universidad de Puerlo Rico, Mayagüez. PUERTO RICO

Abstract.~ This report assesses the macrodistribu t ion of amphibian and reptile taxa, and compares the species-richness of the various biogeographical zones in \\'hich Venezuela is he rein divided. The macrodist ribution of 252 amphibia ns and 299 reptiles species was established for the eight defined biogeographical regions. On the basis of the presence or absence of species, these regions have been categorized into three units: a) those with fewer than 100 herpetological species (Maraca ibo Lake, Lara-Falcón, and the Isla nds), b) those with more than 100 but less than 200 spccies (Andes, eoastal Range, Llanos, and Amazonas), and c) with more than 300 species (Guayana). An index of species-rich ness shows that sorne regions of Venezuela a re among the most distinguished herpetological places worldwide; the Andes regio n is the highest with all index of 0.34. E ndemics are given particular attention in the light of the current knowledge of the geographical distribution of every taxon. There are 181 amphibian and 119 reptile species endemic to Venezuela. In terms of conserva­tion , tu rtles a nd c rocodiles are the most threatened species, but so rne frogs, particularly those of the high lands, are also endangered. T he existence of protected arcas, together with manageme nt and scientific projects designed to protect Ve nezuela 's amphibians and reptiles, and the increasing international concern for them, strengthen the hope of preserv­ing its valuabJe he r petofauna populations, and other animal resources , for future generations.

ReSumen.~La macrodistribución de 252 especies de anfibios y 299 especies de rept iles fue establecida para las ocho

regiones biogeográficas en las que Venezuela se divide corrientemente. Sobre la base de la presencia o ausencia de espec ies las regiones se han subdividido en tres unidades: a) las que tienen menos de 100 especies (Lago de Maracaibo, Lara-Falcón y las Islas), b) las que tienen más de 100 pero menos de 200 especies (Andes, Cordillera de la Costa, Llanos, y A mazonas), y e) con más de 300 especies (Guayan a). El índice de riqueza de especies demuestra que algunas de las regiones de Venezuela están entre las regiones herpetológicas m ás notorias en el ámbito mundial; particularmente Los Andes sobresa len con un índice de 0.34. Se presta una atención particular a las especies endémicas a la luz del actua l conocimiento de la distribución geográfica de cada taxón. Hay 181 especies de anfibios y 11 9 especies de reptiles cons ideradas como endémicas en Venezuela. En términos de conservación, las tortugas y los cocodrilos son los grupos más amenazados, pero alg unos anfib ios, particularmente los de alturas, también están en peligro debido a la declinación numérica de sus poblaciones. La existencia de áreas protegidas, unido a proyectos científicos y de manejo diseñados para proteger las espec ies amenazadas, y el aumento de la preocupación internacional, fortalecen la esperanza de que estos recursos faunísticos puedan preservarse para futuras generaciones.

Key words. Checklist, distribution, species·richness, endemism, conservation, Venezuela, amphibians, reptiles, herpetofauna

Introduction The great variety of ecosystems in South America has induced the development ofa greatl y di versifíed herpetofauna which is about a fourth of the total number of amphibian and reptile spec ies in the world.

Venezuela is one of the 13 countries comprising South America. It Hes in its northemmost portion and receives the c\imatic in fluences oflhe Caribbean Sea and Atlantic Ocean. In the north and northwest it is encased by the Andean Cordillera which in the Mérida Andes may attain a height of 5000 m. The extensive savannas in the central pon ion and Ihe Amazonian forest o f the south , con lri bute 10 a complex climate with an ab undance ofrainfa ll throughout most ofthe coun try, a lthough there are dri er areas in the reg ion of Fa lcón, lhe Goaj ira Penin·

2Correspo ndence . Fax: (58) (74) 401286: email: pefaur@ ciet/s.ula. ve

43

sula, and along the coast, as we ll. The terreslrial boundaries of Venezuela are continuous with Colombia on the west and south· west, Guyana on the east, and Brazil on lhe south. The existence ofmany large rivers and mountain ranges help make the country an cnvironmental mosa ic with a d iverse rauna of"amphibiansand reptiles (Fig. 1).

This rcport assesscs the macrodistribulion of am phib ian and repti le laxa, and compares the spec ies-richness and lhe fauni stical s im ilarity of the var ious biogeographical zones in whic h the country is div ided. Endemics are given panicular al· tention in the li ght o fth e current knowledge ofthe geograph ical distribution of every taxon. Remarks on the conservat ion of these faunas are d iscussed.

EtTorts have been made by several researchers to unravel the macrodistribut ional pattem s ofthe di verse Venezuelan ver· tebrate groups. The di stribut ional pattem of fí shes were de­scribed by Mago-Leccia ( 1970); of amphi bians by Ri vero (196 1,

VENEZUELA Venezuela is on\! afthe 13 countrics orSouth America. with an arca ofof91~.047 km~ (slightly more than l\\iice me size orCalifomia), It lies in the northl.!m pan of lhat cOnlinentjusl nonh orlhe Equator (ge-ographic coordinates: 8°00 N, 66°00 W). is dívid.:d into 24 statcs (polítical divisions). and charnch:nzcd by a di\'erse assemblage of landscapes. where 3 rnyriad 01' rivers and walcr bodics exist. Thc c1imal'" is tropical. hor. Md hllmid in lhe 10\\land5 bul more moderat\:! in lhe high"mds: some dI')" envimnments are common along lhe Co.1St. Venezuela is one oflhe main producers of oil in lhe \\'orld. hUI al50 produces natural gas.l'oal. iron nrl.!, gold. bauxite. diamonds, other mincrals, hydropo\\'er. ánd agriculturnl goods. such 3!) sugar c<.me, callee. rice, corno "'001.1 3111.1 rhum, among others. The fauna and flora are divcrse and it ranks among the tap ten countries in rcgards to biodiversity \\orldwide being classilil!d as a "mcgadiversity" countr)'. There cxist abollt ¡S.OOO species ofplants, more than 1,200 spl!cics offreshwatcr lishes,about 250 spccies of amphibians, 300 spccies of reptiles. more than 1,500 species ofbirds, and dose lo 350 spc:cics oC mammals. With about 20.000.000 inhabitants and ninety pc:rccllt oCthe population li\'ing Ilorth oflhe Orinoco River, the country sho~vs a series of criticalenvironmental problems, such as s()il em:-;ion and deforeslutioll in lhe Andean and wesl central n:gions when: aglicullural activity is ¡ntense. Olhcr environmemal issues ofconccm are mining opcrations in prolcclcd arl!as. sewage pollution 01' Lago dc Valencia. oil and urban polllltion of Lago de: Maracaibo, dcforcstatioll. urban and industrial pollution, especially along lhe Caribbean cuasl. Land tenure, hunting. and tires are also problems. Rights of ownership are not clarified in la\\'. and continucd occupalion.l1c\\ colonizalion and cont1icl wilhin protected areas b. commoll. The pctroleum scétor dominates the econolll~ thus. is of greal el1\'ironmental conccm as \Vell as a potcmiallhrcat 10 (he environmem. Many oflhl!se problems Slem from lhe lo\\' prioril)' given lo conservatioll by ¡he govcmmL'nt as weH as loc" ofequipmcnt and ¡rained Slan' ror dTectivc prolcction ofnalural resources. There are howcver.largc arl!as of\\ ildcrncss intaet and mor(' than a third 01' the counlry's surface 8re protcctcd lands consisting l)f Nalional Parks and Sam.:luaries.

Figure 1. Relief map of Venezuela. The density of the stippled areas shows land elevations. Only the larger rivers are shown.

1963a. b. e. 1964 •. b. e). DlIe llman ( 1988), Frost (1985), and La Marca (1992): orrep ti les by Roze (1966), Medem (1981. 1983). Pr itcha rd and Trebbau (1984). and Lancini ( 1986): and ofrnmnmals by Eiscnberg and Redford (1979) and Bodini and Pérez* l-l ern imdcz (1985). SOllle comprehensive studics 0 11 lhe herpctofauna frol11 several arcas of Venezuela have becll provided by Staton and Di xoll ( 1977). Duellllli.lll ( 1979), Il oogmoed (1979). Hoogmoed and Gorzu la (1979). Rivero* l3Ianco and Dixon ( 1979). Péfaur and Díaz de Pascual ( 1982).

44

Durant and Díaz (1996), and Yústi¿ (1996). Man) otiler conlribut ions about lhe di striblllion 01' o rders (i.e .. Brame and Wakc 1963), familics ( i.c., Di xon ,lIld Hendricks 1979). genera (Le. , Dixon 1980; Di Bernardo 1992: Pé rallr 1993; Scñaris el al. 1994), and/or species (i.e., Gallardo 1965. 1969; Dixon and Michaud.1992) have been conslIlled. Additional re ferenccs can be checked in Van¿olini (1978), Ducllm8n and Trueb (1986), La Marca (1992). Péraur ( 1992). and Ducllman (1995). ílmong others.

VENEZUELAN AMPHIBIANS AND REPTILES

VIII \ .0.

.. ~ / \ 111

\ ,/

\_ - ---; / / -

IV ------ ---- -- -- --/ /

,/,/ 11 / /

/' /'

/

/ -'1 I I I \

\

V -- -----------/--/"

VI

"-"-.

"-. "-

VII " " "-.

Figure 2. Sketch map of Venezuela, showing the main biogeographical regions· 1. Maracaibo Lake, 11. Andes , 111. Falcón-Lara , IV. Coa sIal Range, V. Llanos, VI. Guayana, VII. Amazonas, and VIII. Islands.

Materials and methods In order lO assess lhe distributional rccords of each species, a map of Venezuela wilh its here ín defined biogeographical re­giotls \Vas lI sed. A biogcographic sketch of Venezuela is pre­sented in Fig. 2. lt contains Ihe eight biogeographical zones cOtllmonly acceptcd for lhe country. This physiographic en­semble takcs into account mainly lhe re lief, climate, and vege ta­lion (Marrero 1964; E\\Ie! and Madriz 1968; Huber and Alarcón 1988). A1though the boundaries of each region are not well de· fincd , especially in some ecologica l nodllles, they serve Ollr pur· poses wel!. Different researchers have used Ihis framework for their studies (Ri vero 1963, 1964; Eisenberg and Redford 1979; Bisbal 1988; Péfallr and Ri yero 1989).

The exlent of each reg ion is shown in Table l . The land bordering Maraca ibo Lake corresponds to Region 1, which is covered by seasonall y dry tropical forest in the north al1d by Iropica llowland rain rorest in the south . Wetlands also cover an extensive secl ion ofthi s region representing more thaI1 3,500,000 ha. The sOllthern extenl of the Maracaibo Lake reg ion is bor· dered by Region 11 , corresponding lO thc elevated 1l1011nlains of lhe Cordillera de Mérida, and by lhe Cordill era de Perijá, on Ihe northwest. BOlh ranges compríse the Andes region. For Ihi s

45

sludy, alllands aboye 500 m are considered within Ihe Alldean llnit alld ílS biola as Alldeall; lallds below 500 m are considcred as piedmont. The Andes are environmenlal ly complex (Ewel and Madriz 1968; Díaz el al . 1997), alld may ínclude premolllane, dry and humíd foresls, rnontane \Vel and cloud foresls, xero· phytic va ll eys, hol and dry landscapes, and lhe impressive Páramos, highland tundras, occurring aboye 3000 111 . This region covers about 4,200,000 ha of lhe country's surface.

The Falcón·Lara reg ion 0 1' Region 111 also borders Ihe northeaslern part of the Maracaibo Lake reg ion. This land is affccted by the easterly drying winds of lhe Caribbean Sea and ex hibit mos tl y a xerophyt ic landscape, ca tal og lled as prcmontanc dry shrub or dry fo res!. A large portion of Ihe north sea líne of Venezue la is bordered by ¡he Coastal Range (Region IV), made up o f forested l11ountains, w ith eleyations IIp to 2765 111 , and covered by premontane tropical rain and monlane clolld loresls. The Andes and lhe Coasta l Range, s lopc down ínlo Ihe lowlands orthe Llanos, Regíon V, which ex tends 10 Ihe Orinoco Ri ver in Ihe sOll th , 10 Ihe border wi lh Colombia on Ihe west, and eXlending lO Ihe Orinoco Rive r delta in the ea sI. The approx imately 27,000,000 ha orlhe Llanos are cov· ered by savannas or exle ll sive prairies inlenningled with dry

Plate 3 ----._--....

• -Plate 5 Plate 6

Plate captions: 2. Ate/opus carbonerensis. Once very abundant, this bufonid frog is now extinct from the Andean cloud forests . Venezuela, Mérida. 3. Bufo granulosus. An inhabitant of all Venezuelan biogeographical zones. Venezuela, Aragua. 4. Bufo marinus. One of the largest toads from Venezuela and the one with the largest distribution; it remains abundant country wide. Venezuela , Táchira. 5. Bufo typhonius. Venezue la, Guatopo. Pholo courfesy 01 Lallrie 1. Viu. 6. Dendrobates leucome/as. This attractive frog is found in the southern part of the country, south of the Orinoco River. Venezuela , Bolívar.

46

VENEZUELAN AMPHIBIANS ANO REPTILES

Table 1. Exlent of land surface and number of tife zones present in the biogeographical regions of Venezuela. (Sourcc: 13isbal 1988; Ewcl and Madriz 1968.)

Estimated land surface

Biogeographical region

I Maracaibo Lake

11 Andes

111 Falcón·Lara

IV Coasta 1 Range

V Llanos

VI Guayana

VII Anw zonas

VIII Islands

Total

km ' 35,000

42,000

30,000

68,000

270,000

350.000

105 ,000

1,500

901 ,500

%

3.88

4.66

3.33

7.54

29.95

38.82

I 1.65

0.17

100.00

Number of Life Zones

5

16

6

10

3

10

2

6

22

Table 2. Surface of altitudinal belts in Venezuela . (Source: Ewcl and Madriz 1968.)

Area

Altitudin.1 belt km ' %

Tropical lowlands 640,283 7 1.30

Prcmontanc 227,390 25.00

Lo", monlane 27,9~7 3.05

Montane 4,570 0.52

Subalpine and alpi ne 1,270 0. 13

Total 9 01 ,500 100.00

forests ami ri vcrin e ga ll ery rorests. The largcs t biogeogra phi ca l region of Ve nezuela is Ih e

Guayana, Regioll VI, which includes about 35,000,000 ha, mas! of whi ch a re part of Ihe Venezue lan Guaya na shi e ld . The landseape is com plex, includi ng lowland , pre monlane and Illon lane wet and rain forests; savannas; a nd wet oases, ca ll ed moricha/es. r ich in palms and herpetofauna. The land· scape is defincd by pro found va ll eys bordered by fepuys , lable mountain remai ns of gco log ical la nds of the Precam· brian·Pa leozo ic era. T he a rea be longing lO the O ri noco Ri ver 's de lta is also incllldcd in th is reg ion. Some 10,500,000 ha of tropical forests covering lhe lowlands of lhe lI pper Orinoco River basi n compri sc Region VI I, Amazonas, which is conti nuolls with South America's largc green con:, the Amazonia. An important part o fthi s arca is eovcrcd by wet trop ica l fore s!.

A smal l portion of lhe counlry is made up of islands. AII of thclll are inclllded in Region VIII, which for the purpose of Ihe distribulion of turt les also includes the Venezue lan Carib· bean Sea. This region is ineluded lo call att ention to Ihe par· lieu lar di st ribulion oflh e island herpelofauna , especia lly Ihal ofl sla Margari ta, as we ll as mar ilime herpelOfauna.

Each ofthese biogeographica l regions is eeo logica ll y di­verse and complex. In their ecological map, Ewel and Madriz ( 1968) deseribed 22 life zones for Venezue la (Tab le 1). The mosl hOll1ogeneolls biogeographica l reg ion, Amazonas, has on ly I\VO li fe zones, and lhe most complcx o f all, Ihe Andes. includes 161ife zones in its relat ive ly small le rr ilOry. Dive r­s ity 01" eeosyslems is proportiona l lo the Sleepness 01" ve r· sant s, Ihe sides o f Ihe mounlains, in the Irop ies , bu l most of Venezue la is lowland. Abollt three·fourths of the eountry is eompri sed o f lands be low 500 111 ; lands aboye 1000 m do not

47

cxeecd 10% oC Ihe terrilory, yel Ihese are the mosl eeologi­ca ll y di verse o C a ll (Table 2) , a situat ion w ilh slrong con se­quences on Ihe biologieal featurcs oflhe count ry.

Sevcral melhods have been used lO obla in Ihe da la pre­senled in th is paper: museum record s, bibliographic informa­lion , and pe rsonal observations. Reports made on the hold­ings ofrelevanl muse ul1l s in Ve nezucla, Pucrto Rico, Colom· bia , Brazil, Franee, and Ihe Uni ted States have been taken il1l o accounl , bul in mosl instances Ihey have been im proved w ith no tes take n by Ihe aulhors du ring their Irips and mu­se llm vis its. In o rder 10 updale previolls reporls on the Ven­ezuelan amphi bians and repti les (Péfaur 1992: Péfaur and Rivero 1989), !le", lists had lO be produccd. To the best o f om knowledge Ihe e hecklisl database (Appcndix 1) conlains a l! known spccies tlp lO Deccmber 31, 1996.

The di stribllti onal aspecls of Ihe lisl are givcn by the presence o r abscnee of dala fo r every species in a biogco­graphieal reg ion. Thc eight reg ions were Ihus dcli11liled by Ihe known inforlllation rcgarding Ihe spec ies' geographical di stri­bUl ion. The number ofspee ies present in a region is an ind i­calion ofi ls spee ies diversily and oflhe geologica l, climato­logica l, ami bio logica l evollllion oflhe faunal elemenls con­cerned. However, Ihe index of spceies· richncss. ralhcr than the simple number 0 1" spec ies, is a better expression of evolu­lionary Irends. A Species-Riehness Index (SR I) \Vas caleu­lated by cons idering Ihe number of cxtant spcc ies divided by Ihe a rea mult ip li ed by (x) 100. To oblain Ihe faullistieal si mi · lar ilY bel\\leen reg ions, a four-fold eonlingeney table \\ as cre· aled for eve ry pai r 01" compared rcgions and the Dice coeffi ­cient , as showll by Hayek (1994), \Vas calelllalcd.

Ende mism. an ecological tc rm, was determined by find· ing whelher Ihe re \Vas a uniqlle relal ionship be twcen a spe·

JAIME E. PÉFAUR AND JUAN A. RIVERO

Table 3. Taxonomic compasitlon of the Venezuelan herpetofauna.

Class/Order Common name Families Genera Species

Amphibia

Anu ra

Caudata

GymJlophiona

Subtota l

Reptilia

Tesludines

Crocodylia

Am ph isbacnia

Lacertilia

Serpcntcs

Sublolal

Total

Frogs and toads

Salamanders

Caec il ians

Tun les and lOrlOises

Crocodi les and all igators

Amphisbacnians

Liza rds

Snakes

10

3

14

7

2

1

8

8

26

40

42 238

2

7 12

50 252

14 23

3 6

2 6

37 11 3

56 151

112 299

162 551

Table 4. Allocation 01 amphibian and reptile species in the biogeograph ical regions of Venezuela.

Biogeographical region

Taxa 11 111 IV V VI VII VIII Total

Frogs 18 72 17 63 36 127 53 5 238

Salamanders O I O I O O O O 2

Caeci lians 2 2 2 O 7 2 O 12

Turtl es 6 1 3 6 9 13 I 1 6 23

Crocodilcs O 1 2 2 3 5 6

Alllphisbaenians 1 2 2 2 2 5 4 O 6

Lizards 20 26 25 39 19 59 36 16 113

Snakes 44 38 47 75 43 96 74 22 151

Total 92 142 96 190 111 310 185 50 551

Table 5. Species-Richness Index (SRI) value calcu lations for herpetofaunal species in the biogeographica l reg ions of Venezuela. SR I = (Species number/area) x 100. The area to calculate the Index is provided in Table 1.

Biogeographic Amphibian species region n SRI

1 20 0.05

11 75 0.18

111 18 0.06

IV 66 0. 10

V 36 0.01

VI 134 0.04

VII 55 0.05

Total 252 0.03

cies and a gcographical region. As used in thi s study, when­ever a spec ies dwelled and appare nlly o ri g inatcd in a s ing le reg ion, il \Vas considered a biological endemic. However, ira spec ies \Vas considered endemic beca use it occupied a s ing le reg íon in Venezue la bul al so extended into another counlry. il was conside red as apolifical endemic \V ith respect lo Ihe firsl country. In Ihe case ofVenezuc la, ¡here are mainly biological endemics. bul Ihere are <l Iso scveral political endemics, Ihal

48

Reptile species Total Total n SRI n SRI

72 0.20 92 0.26

67 0. 16 142 0.34

78 0.26 96 0.32

124 0. 18 190 0.28

75 0.03 111 0.04

176 0.05 3 10 0.09

130 0.1 2 185 0.18

299 0.03 551 0.06

is, spec ies t!xte nd ing beyond Ihe boundari es 10 so rn e ne igh­boring counlr ies, such as Colombia, Braz il , and/or Guyana in the ma inland , or Tr in idad-Túbagú. and lhe Dutch Islands (Bona ire , Aruba, C uracao), in Ihe Caribbean Sea.

Tú determin e the conse rvalion s tatus of Ihe herpe­lofauna, only a few quantitati ve assessmenls are avai lable. Thus, a genera l impress ion rather than an accuralc ccnsus supports Ihe ca laloging for the specjes cons.idcrcd.

Plate 8

Plate 10

Plate 12

Plate 13 Plate 14

Plate captions: 7. Mannophryne colfaris. The ventral side of the females of th is species present a black collar and a yellow guiar region. Venezuela, Merida. 8, Nephefobates alboguttatus. A very abundant species in the past, today it has disappeared from their geographical range. Venezuela, Merida. 9. Nephefobates haydeeae, This frog is found only in the western state of Táchira, where its poputations are declining. Venezuela, Táchira . 10. Nephefobates meridensis. This was the largest dendrobatid frog of western Venezuela. This species is probably extinct. Venezuela, Mérida. 11 , Nephefobates serranus. A very restricted trog , inhabiting only a part of the Sierra Nevada mountains. Venezuela, Merida. 12. Flectonotus pygmaeus. A marsupial frog that inhabits the northem mountains. Venezuela , Merida. 13. Hyfa crepitans. This species dwells in most environments in the country. It remains very common. Probably conforms a taxonomical group of related species. Venezuela , Táchira. 14. Hyla fanciformis. This large and slender brown frog has a wide distribution in the country. Venezuela, Táchira.

JAIME E. PÉFAUR AND JUAN A. RIVERO

Table 6, Matrix of amphibian similarity among Venezuela n biogeographica l regions. Note: This matrix shows in bold the amount of species 01 every regían. Above the diagonal line are the similarity values between a pai r of regions , whi le under it are the numbers of shared species. (Source 01' data: Appendix 1.)

REGION

11 111 IV V VI VII

20 0.15 0.32 0.26 0.43 0.1 7 0.27 R 11 7 75 0.15 0.18 0.18 0.11 0.15 E 111 6 7 18 0 .33 0.33 0.14 0.22 G

IV 1 11 13 14 66 0.45 0.25 0.28

O V 12 1I 9 24 36 0.36 0.46 N VI 13 12 1I 25 31 134 0.40

VII 10 10 9 17 21 38 55

Table 7. Matrix of reptile similarity among Venezuelan biogeographical regions. Note: See comments under Table 6. (Source 01' data: Appendix 1.)

REGION

11 111 IV V VI VII

72 0.47 0.64 0.56 0.52 0.30 0.32

R 11 33 67 0.51 0.44 0.37 0.42 0.24 E 111 48 37 78 0.66 0.58 0.3 7 0.39 G

IV 1 55 42 66 124 0.56 0.47 0.46

O V 38 26 44 56 75 0.56 0.62 N VI 37 28 47 71 65 176 0.72

VII 32 24 41 59 59 110 130

Table 8. Number 01 endemic species and percentage of endemism in the herpetofauna 01 the biogeographic regions of Venezuela. (Sourcc Data: Appendix l.)

Amphibians

Regions Species by region Endemics

1 20 4 11 75 57 111 18 O

IV 66 32 V 36 I VI 134 75 VII 55 12 Country 's total 252 181

Resu lts

Taxonomic composition Thc herpcto logical component o f the Venezuelan fauna con­sists of 55 1 species of which 252 are amph ibians and 299 are reptiles. A taxonomic li st of spec ies is provided by a country checklist (Appendix 1) and sUITImari zed in Table 3. The am­phibians of Venezuela are included in !hree orders, of which !he larges! is Anura. The mos! specious I"amil)' is Hy lidae, contai ning 77 species, rollowed by Leptodac lylidae with 72 species. Olher ralher large ramilies are Buronidae , Centrolenidae, and Dendrobatidae. The res! orlhe frog rami lies have a ver)' low numher 01' spec ies. The ramilies 01' salamanders and caeci lians are also of minor numher.

The reptiles are comprised within flve orders orwhich Ihe largest are Serpentes and Lacertilia . Turtles and tor-

50

Reptiles

% Species by region Endemics %

2.2 72 12 10.1

31.5 67 20 16.8

0.0 78 5 4.2

17.7 124 19 16.0

0.6 75 2 1. 7

41.4 176 48 40.3

6.6 130 13 10.9

60.3 299 119 39 .7

toises have 23 species contained within seven familie s . C rocodi les and alligators have s ix spec ies belonging to !wo families. The arnphishaenian s are represented by onl)' !\vo genera and s ix species. The lizards have 11 3 species com­pr isi ng eight families, \Vith Gonalodes as its most specious genus, with 13 species. Snakes make up the most diversi­fied group with 151 species belong ing to e ight familie s. li s larges! ramily, the Colubridae, contains 104 species. A H other fam ilies 01" Ih is class have less than 40 spec ies each . It s mos! speciolls genus is Atrae/us with 16 species; other quite large genera are Liophis and Micrurus \\'ilh 10 and 12 spec ies, respec tivel )'.

Distribution The di slrihution 01' every species in lhe e igh! biogeographical regions, as considered in Appendix 1, is sUITImarized in Table 4.

VENEZUELAN AMPHIBIANS ANO REPTILES

By far, lhe largest assemblagc of amphibians alld reptil e species is fOlllld in Regían VI, Guayana, foll o\-ved by Ihe Coastal Range. Amazonas, and lhe Andes. The lowest number 01' spe­cíes in the country is found in the northern regions of Maracaibo Lake and Falcón-Lara. In the rnaril imc islands regían Lhefe are 50 species of amphib ians and reptiles.

Species-richness As surface area differences among Ihe biogeograph ical divisions do no! allo\\' a direct comparison 01' lhe biodiversity by Ihe number of herpctofaunal species alone, an index (S RI ) is re­qu ircd lo more accurately expreS$ the results. The SR! index valucs are showll in Table 5.

The country ¡Iself has a lo\\' valuc for Ihe index, 0.06. I-I owcvcr, the four smaltest regions achievc lhe highest SRI and lhe opposi le three largest regions have lhe lowesl indexes.

Faunistic similarity lhe herpelological simi larity betwecn the different biogeographi­cal regions ofVenezuela is presented in lable 6 and 7. Simi larity bascd on amphibian species is rather 10w(Table 6); the least pair bound of regions is the Andes-Guayana, which share 12 species and have a similarity va luc of 0.1 1; the strongest bound is lhe Llanos-Amazonas pair, which ha ve 2 1 specics, with a simi larity va lue of 0.46, füll owed by the Llanos-Coastal Range and Ll­anos-Maracaibo Lake pai rs.

Regional similarily based on Ihe rcpti le component oflhe fauna produces difTerent resul ls (Table 7). In general , the Si111i­larity n:gion based on reptile species has higher va lues than when amphibians are cons idered. Here, the leasl similar pair region is Ihe Andes-Amazonas, with a similarity value of O.24, while the strongest bounded pair is Gllayana-Amazonas, wilh 110 shared species and a similarity value ofO. 72.

Endemism There are 181 species ofamphibians and 119 ofrepl iles consid­ered lo be biologica/ andlor po/irica/ endemics (Table 8). Overal1 there are more endemic speeies alllong the amphibians Ihan among reptiles. However, four regions have less endemic amphibians Ihan endcmic rept iles. lhe Andes, the Coasta l Range, and Ihe Guayana are lhe places with more endemics in bOlh taxa.

Conservation Actual data on Ihe conserval ion status of lhe herpelofauna is relatively scarce in Venezuela. The most relevant infonnation on population status is provided by Ramo ( 1982), Praderio (1985), Si lva et al. ( 1985), Péfaur and Díaz de Pascual ( 1987), Péfaur et al. (1987), Silva and Valdéz ( 1989), La Marca and Rein tha ler (1991), Thorbjarnarson ( 1991), Péfaur and Pérez ( 1995), and Durant and Díaz (1996), alllong others. The Red Data Book of Venezuela (Rodriguez and Rojas-Suárez 1995) \Vas helpful in eSlablishi ng herpetofauna endangennent status.

The report elaborated by the senior author to the Decl ining Amphibian Populations Task Force (DAPlF) of lhe World Conservalion Union [IUCNJ (in Vial and Saylor 1993), on Ihe declining status of amphibians was al so used lo generale a lisl of endangered species (Appendix 2). Severa l species of amphib­ians, alllong which there are seven species of BlI tonidae (5 Are/opus, 2 Oreap/1Iynel/a), four Centrolenidae (2 Centro/ene, 2 HyalinobatrachiulI/), 15 Dendrobalidae (5 C%stelhus, 2 Man/1op/¡ryne, and 8 Nephe/obales), five Hylidae (1 Gasrrorheca,

51

4 /-Iy/a) , one AlIophrynidae ( 1 A/lophryne), sev en LeplOdactyl idae (2 Cerarophrys, 3 E/eurherodacrylus , 1 Leplodactylus, 1 Pselldopaludieola). and two Plethodontidae (2 Bolitog/ossa) are reponed as having a decl ine in lheir popu lation nUlllbers and th us species of special concern. Alllong reptiles there are severa! spccies on the vcrge of cxt inction, such as CroeodY/lIs inrermedius. C. aeurus. Caiman nigef: and Podoc flemis expansa, as well as all the sea turtles.

Discussion

Taxonomic composition lhe taxonomy of the Venezuclan herpctofauna changes con­linuously due to new research findings and systematic rear­rangemcnts. Sllbstantial changes in the number ofknown spe­cies for every region have taken place in Venezuela during recenl years. Descriplions of numerous new taxa ha ve occurred in the last decades mainly amollg amph ibians (Péfaur 1985; Rivero 1982a, b, 1985; Ayarzagüena 1992, a1llong others). Importan! changes ha ve recen tI y been inlrodueed to the tax­onomy ofthe country 's herpetofauna lhus. changing the sys­tematic scenery. Among frogs, lhe Centrolenid family has been divided inlo several genera (Cenrro/ene. Cochranella. and Hyalinobatrachium) by Ru íz-Carranza and Lynch (1 991 ). The Dendrobatidae was a1so generically rea rranged with the intro­duction of Epipedobates and Minyobates by Myers (1987), Aromabares by Myers el al. (1991). and MannophrYl1e and Nephe/abares by La Marca (1995). lhe Hylidae has also suf­fered some modification wilh lhe rearrangcment of so me Olo/ygon il1to the resurrected gel1us ScinQx (Duellman and Wiens 1992), and the elaboration of a new genus Tepuif1yla by Ayarzagüena el a l. ( 1992b). Among the caecil ians. a general laxonomic rearrangement offamilies and genera was proposed by Nussbaum and Wilkinsoll ( 1989) and Wilkinso l1 (1996).

i\mong the repti les, there have also been some taxonomie changes. Within the lizards, Iguan idae was divided i!lto several farnilies (Corytophanidae, Iguanidae, Pol ychrotidae, and Tropiduridae) by Frosl and Etheridge (1989), while Ihe genus Anolis was divided into five genera, ofwhich IWO are present in Venezuela, Dacry/oa and Norops (Savage and Guyer 1989). lhe Teiidae was also divided into two units. Gymnophlhalmidae (srna 11 teiids) and Tei idae [macroteiids] (Presch 1980). Within the snake group, sOlne eolubr id genera have been revised modi­fying the taxonorny for the Venezuelan members; for instan ce, Dixon (1989) rev icwed Liop/¡is and other associated genera. Among the Crotal idae there was al so a slrong change with the partitioning of Borhrops into several genera: Borhriechis, Both,.iopsis , Bothrops. and PorrhidiufIl, with considerable ctTeel to the VenezueJan fauna (Campbell and Lamar 1989). Of course, classi fication \Vi11 continue 10 change as it is a dynamic sc ience, and new systems and names wi11 be introduced changing subse­quent li sts of Venezuelan amphibian and repti le species. AII systematic allocations and changes should be taken as tempo­rary arrangements Ihat \Vi iI be modi fied by lhe collection ofnew data and insighls by researchers.

Distribution There is no single area in Venezuela where an amphibian andlor a rept ile species is not present. From the most Iuxllrious tropical wet forest of Amazonas to the vegetationally depauperate Páramos at the top oflhe Andean 1l1ounlains. where nllmerous

Plate 16

Plate 17 Plate 18

Plate 19

Plate 21

Plate captions : 15. Hyla luteocellata. Venezuela , Guatopo. PI/Olo courlesy 01 Jona/ee P Ca/{/¡rell. 16. Hyla microcephala. This sma" frog has one of the largest biogeographical distribution in northern South America . Venezuela, Trujillo . 17. Hyla vigilans . One of the smallest frogs of the country ; in ha bits the Lago de Maracaibo Zone. Venezuela , Zulia. 18. Phyllomedusa trinitatis. Venezuela Guatopo. PholO cour les)' 01 La urie 1. ViII. 19. Scinax rostratus. A medium sized frog with a distribution along the northern lowlands. Venezuela , Mérida. 20. Eleutherodactylus lancini;. An inhabitant of the páramos cold streams. Venezue la, Mérida. 21 . Eleutherodatylus lentiginosus . Asma!! frog from the Andes versants. Venezuela , Mérida. 22. Eleutherodactylus vanadise. A small frog occurring in the cloud forests of the Venezuelan Andes. Venezuela, Mérida.

Plate 24

Plate 26

Plate 29

Plate captions: 23. Leptodactylus wagnen'. A very elusive frog from most Andean environments, that can be detected by its peculiar call. Venezuela, Mérida. 24. Pfeurodema brachyops. One of the most popular frogs in the country by its color and dots in the rear part 01 the body. Venezuela, Portuguesa. 25. Pipa pipa. This aquatic 1rog lives in the Llanos 01 Venezuela and Colombia. The picture shows a museum specimen with eggs imbedded in its back. Venezuela , Barinas. 26. Rana palmipes. A typical 1rog from the lowlands 01 westem Venezuela but that can occasionally seen in the forested valleys of the Andes. Venezuela , Tachira . 27. Pseudis paradoxus. This medium sized frog is to be 10und in the marshes of the country's lowlands. Venezuela, Apure. 28. Bolitoglossa orestes. One of salamanders that dwells in the cloud forests of the Andes. Venezuela , Mérida. 29. Caecifia subnigricans. A worm-rike amphibian 01 northern Venezuela , that extends 1tS range into Colombia. Venezuela, Tachira. 30. Rhinoclemmys pllnctllfaria. This small turtle inhabits the Lake 01 Maracaibo region . Venezuela , Zulia .

JAIME E. PÉFAUR AND JUAN A. RIVERO

frogs and lizards specics dwcll (Hoogmoed 1979; Ri vera-Blanco and Dixon 1979; Duran! and Diaz 1996; Díaz el al. 1997), species abound; howcvcr, snakes have no! invaded lhe páramos (Péfaur and Diaz de Pascua l 1982; Diaz e l al. 1997). Different physiological and behavioral adjustl11cnlS are found in lhe fauna of cach ecosyslcm, to cope wilh lhe conlrasLing climatologica l facto rs impinging on every zonc. Severa! or lhe more notewor­Ihy adapt ive slrategies are Ihe development ofscveral unique reproduclivc modes: developing eggs and tadpolcs in Irce cavi­líes or bromeli ads. or eggs on dorstll11 o f remales, or tadpoles carricd on dorsum of males, such as OCC Uf in 31lUfallS (Ducllman 1985), or Ihe use ofco ll ect ive nest dcposits under rocks, as occurs in the li zards o fth e Páramos.

Newly d iscovered and new locality record s 01' spec ies have changed the known di stributional pallerns fo r many 3mphibians and reptiles. For instance, in the Andean rcgion, spec ies lOta ls have changed from 16 frog, 4 lizard, anel 10 snake spec ies (as reported by Ducllman 1979), to 56 frog and 15 liza rd spccics (as reported by Péfaur and Díaz de Pascual 1982). lO 72 frog , 26 li za rd. and 38 snake spccies repo rted in lhi s slUd y. The well·documcnted work by Lanci n i ( 1986) reports 133 snakc spec ies for the country lO 151 species reported here . Our knowlcdge oflhe di s lribut ion ofthe Ven· ezuelan amphi bians and reptiles is quite accepta ble al the present I ¡me. but an in crease could be expected as new data is be ing co ll ec led a lll he time by rcscarchcrs in the field. 8 0 th Ihe Andes and Ihe Guayana rcgions have been activel y ex· plored by groups of resca rchc rs from the Un ivers ity of Los Andes in Mérida and from Musco de Ciencias La Salle in Caracas. respectively (AyarzagUcna et a l. 1992a, b; Péfaur 1993; Scfla ri s e L al. 1994; Durant and Díaz 1996), wh ile ac · ti ve research on Ihe hcrpe lofauna of Ihe Coastal Range is underway by researchcrs from Central Uni ve rs ity, Museu1l1 oflhe Agrarian Zoology Institmion in Maracay (Manzanilla el a l. 1995, 1996).

The di s lribu lional range of certa in spec ies is of conccrn. There is a d rmna lic difference belween the di stribul ion o f SO I1lC laxa cX lcnding over Ihe whole CO Unlry (e.g .. Bllfo marinlls. flyla crepilans) compared to lhe punctual dis tribu­lion of those lepui.associated laxa (e.g., Oreop/u)'nella huberi, o. vasqlle=i), or lhe narrow d ist riblltio n of dendroba tids in the Mérida Andes. Many examples of lhese are knov·m in any biogeographical region.

The fasl change in the Venezuelan landscapes will ha ve a severe impact in the di slri bul ion of lhe herpclo fauna. In less than hal f a ccntu ry mOSI foresls in Region I have becn cut down and replaced by prairics. Amphibian species prevai l in the south. whi le rept iles dominate the no rth. In gene ral , most, but mai nl y the xerop hyti e faun a, is shared with Ihal o f xe ric Regio n 11 1, a relalionship cx tending to the rich and large va l­ley of Ihe Colombian Magda lena river. The va ll ey of Ihe Calatumbo river mighl have played a role in aC ling as a pass· way belween the faunas of Ihese regions.

The existence o fmany humid environmenls in Ihe Andes (Region 11 ) permils Ihe prcsence of a large numbcr o f frog species (Duellman 1979: Péfaur and Diaz de Pascual 1982). This region has fe\\' biogeograph ícal contacts w ith olher parts of the COUnlr). a1though il has se rved as a pass\Vay for dis· persi ng faunas (Péfau r and Pérez 1995; Ri vero 1979; Ri ve ro and Solano 1977). Somelhing simi lar occurs in the \Vel and e levated lands ofRegion lV, Ihe Coaslal Ran ge. The Andes do 11 0t have wil hin thern any importanl large geographieal bar-

54

ricr, bUI there are rieh mierocl ima tes thal aC I as ecological refuges. In lhe Coastal Range region, ho\Vever, there are at least two \Ve l1 differentiated k inds of lands: \Ve l e levated and xerophy tic lowlands close to Ihe Caribbcan Sea (Ri vero 1 964a: Manzan illa el al. 1995, 1996). Moreover. an import anl selec· I¡ve biogeographica l barr ier has aC lcd in thi s regio n, as is the case o f the Unare ri ver va lley, \Vhere many herpetofauna spe­eies di str ibutions are di scont inued pas l thi s point, such as Colosfel/¡us mande/orum, L. ¡nsu/arum. and Eleufherodacl)'lus lerraebo/ivaris, arnong others (R ivero 1964a: La Ma rca 1992).

In Ihe Ll anos (Regio n V) the re exis ts a fai d y large her­pcto log ieal fa una , which is comOlon wilh Ihe olhe r rcgio ns (S laton and Dixon 1977 ; Rivero·B la nco and Di xon 1979; Pé f~llJr ami Díaz de Pascua l 1987). Th e Llanos are c limali· ea lly and vcgctationa lly homogeneolls at Ihe rnacrogeograph­ica l level , altho llg h the y are inte rming led w ilh a web of ri­parian forest cOl1llllu nicating with Ihe o lhe r s ll rrou nding re· g ions. Bordered by Ihe Orinoco r ive r in the sou lh , thi s re· g ioll docs not separatc fauni s tica ll y from Regions VI and VII - the largesl Venezue lan ri ve r seems 11 0 t 10 be a se lec· live bi ogeograp hica l barrier fo r amphibians amI reptil es (Rivero 1961).

The soulh e rn regions o f Venezue la. Guayana and Amazonas (Regions VI and VII) are close r associated wit h Ihe Amazoni.an sector o fS olllh Ame ri ca than w ilh Ihe north· e rn areas or lhe country. The geolog ical changes, Ihe climatc, Ihe lOpography, and Ihe diverse vegelalio n makc these I\VO areas the r iches l in herpelOfau na spec ies. This is espccially true of Ihe Guayana where a mu1titude of Iwbilats faci lil alc the proeess 0 1' evo lulion. With the cxception of sa lam:mdcrs and croeod il ians, Ihe laxa are more nllmcrOllS in Ihc Guayana, than in any olher region. For Ihe rest ofl ll e groups, the I1UI11·

ber is almosl dOllbte in Ihe Guayana, as comparcd to any of the olhcr biogeographica l regions. The ccologica l cornplexi ly Oflhi5 reg io n has played several ro les in lhe evolutio n ofthe biola . On the one hand , it has eonstiWled a selcct ivc barrier je r Ihe: eXlcnsio n o f somc faunal elcments frollllhe Brazi liall Amazonian lowlands lO Ihe Llanos of Venezue la and v ice ve rsa; o n lhe other hand, it is the seat for l1lany e ndemic species ( I-I oogmocd 1979; this study). At the same t ime, il has shared spec ies with o ther regions of Venezuel a and Brazi l (Avila· l>ires 1995). Dne particular exceplion is Leplodaclyllls labyril1lhicus. This specics Ii ves north and soulh 01' Ihi s re· gion, Icaving a dist ributio na l hi at us in the Guayana shiel d (PUaur and Sierra 1995) .

Depcnding on the total number of species prcsent in cach rcg ion , thrce eatego ri es of reg io ns can be proposcd: a) those wilh lcss than 100 species, eomprised by the Maracaibo Lake, the La ra·Falcón, and Ihe Is land5 reg ions: b) Ihose \V ilh morc Ihan 100 bUI less than 200 species, which incJudes Ih~ Andes. Ihe Coasta l Range, Ll anos, and Amazonas; and c) w ith more Ihan 300 species, rep resented by Ihe Guayana regíon . Mo reover, Ihe lim its ofevery biogeog raphical region. are ve ry imprec ise. For instance, lhere is a sec lor where Rc­gions 11. 111. and V converge , and thus there is a n uncertain l)' about Ihe reg ion 10 which some species be lo ng. AnoLher bio· geographica l nodule is rhe secto r wherc Reg io ns 111 and IV mcrge. There is g reater need for a more delai led ana lys is 01' presence spec ies in lhese connic li ng seclOrs. On Ihe o lher hand , col lecl ing record s are ve ry scarce in the Cordill era de Pcrijá in western Ve nezue la and lhe Orinoco de lta region in easte rn Venezuela. It is poss ible that lhe Delta mighl be con-

VENEZUELAN AMPHIBIANS ANO REPTILES

sidered a scparalc biogeographica l region when more biologi­cal information is gathe red and analyzed.

Species-richness The f~ILJn i s ti c richllcss of an arca would be bcttcr cxpresscd by an Indcx of Spec ies-Rich ncss (SRI) tha! takes into account both Ihe area and Ihe numhcr ol'specics. In cssence, Ih is index is a spcc ies-density index- that ¡s, al lhe same number of spcc ics, those biogeograph ical arcas wilh small surfaces \V iII have higher va lues in SRI Ihan areas with larger surfaces. SR¡ nUlll hcrs allo\\' Ihe compar ison 0 1' an y arca Dr region of Ihe world with respec! to spccies-richness . In Ihe case ofVenezu­cla, il s large size and lhe ex istence ofcx tcnsive territories, ~uch as Ihe Llanos, w ilh a lo\\' nUlllber ofspecies, account fo r a low SRI va lue (0.06), s imilar lo Ihe one known for the Yucatán Peninsula in Mex ico (Lee 1980). This stalisti cal artifacl warns aboul Ihe use ofsuch index for counlries with a high diversity of landscapes. but lo use it ro r part icul ar regions on slllall cOunlri es. For instance, among published data, Oaxaca in Mexico has an SRI val ue of0.3 7 and is lIsually cOlls idered one of lhe richest herpelological reg ions in the worl d (Casas-AndrclI el al. 1996), whereas Costa Ri ca has been re ported as having Ihe largesl SRI vallle (0.71) in the world (Johnson 1989).

[f both amphibians and repti les are analyzed together, the highes l SRI va llle is found in the Andes, fo llowcd by the Fa lcón- Lara regíon. The SR I va llles o f0 .34 and 0.32, respec­li vely (Table 8), are di slinglli shed even when comparcd 10 Ihe SRI richesl regions ofthe wo rld. When the analys is is carri ed OU I separately fo r amphibiall spee ies, Ihe Andes is the onl y region Ihal s lands out as the 1110st di verse region. This re la­li ve ly .small area, with severa l e leva lional be lt s and an abun­dance 01' hUlllidity and \Vet lire zones , has a eondensed den­si ly 0 1' amphib ian spec ies. In turn, when the rcpli le fauna is analyzcd scparalely, Ihe largest va lue is found in Regíon 11 [ (Fa lcón- Lara), which is also a small arca covcred with xe ro­phylic vegetation- a fittin g place for reptiles.

Faunistic simUarity Similarity is a concept that brings logelhcr biogcographical e le­lllenls and evolutionary aspccts o rthe fauna. [n the comparison of the seven regions of Venezuela, all obtained val ues for am­phi bians are under 0.50, indicaling a low degree ofspecies shared between biogeographical regions. Thc most similar amphibian faunas are those of Regions IV-V and V -VII- the values o rtheir s imilarilY indexes are lhe highesl (0 .4 5 and 0.46, respeclive Iy). It is nOI a surprise lo find these relationships, fo r there is a geographieal continllity belwecn these reg ions, but \Vhal is

Table 9. Causes 01 amphibian population decline.

1. Environmental changes due to human activity a) Destruction and/or fragmcllIation o r hab itats b) Agriculture fro lltier ex pansion c) Inadcquate use ofp laguicides d) Waler pollution e) Ozone's eover destrucl ion or weakness 1) Acid rain g} Inlrodue lion of predator and/o r competilOr spec ics h) Expansion and/or introduclion of di scases

55

biogeographica lly rel evant is the high similarity between the Llanos and the Maraea ibo l ake region. whieh are separaled by ranges and dry lands, but that share SOIll C species \Vith disjul1ct di striblltions such as Leplodaclyflls bolivianlls, L wagneri, Physa/aeflllls pl/sll/foSIIS , and Pseudis paradoxa, 31ll0ng others. On the olher hand , elements of Region 11 differ considerably from the other arcas, an ind ication or the grealer amphi bian independence o r Ihe Andes with respecl to the other regions in VenezlIela.A simi lar pattcm is provided by the Guayana region, wh ich exhibits an independenee as a biolie souree.

Becallse reptile species usually have a rather ample di slri­bution, lhe nlll1lber of shared spec ies and lhc si milarity valllcs are higher in thi s grollp than in the amph ibians. The overa Jl s imi larity vallles are higher, espec ially between Regions VI and VII , and wilh the lowest leve l between Ihe Andes and Amazonas. Consideri ng the reptiles, the Andes does not stands out as a fallnislica lly independent region beeause of lhi s region's rela­lively strong rc lalionships lO Ihe Maracaibo Lake and Falcón­Lara regions.

Endemism The formation orne\\' animal species, as Ihe resul l of genetic and ecologica l processes, has been one of Ihe major evolut ionary featllres in l11al1 y arcas ofVenezuela. Whcrevcr the d istribut ion of a species is redllced in space, an endcmisl11 process is al work.

l-I erpelOlogical enJemism is difTerent in lhe several biogeo­graphical zones ofVenezuela. The Andes and the Guayana are the regions with the highesl pereentages of amphibian endem ics, most oflhem biofogical endemics, renccti ng an active spec iat iol1 proeess, something Ihat has been aeknowledged in olher animal grollps (Brown et al. 1974). On one sidc, at Ihe Andes there has been an acti ve process or contracti on and expansion offorests as consequence o rlhe g lac iation periods, and on Ihe OIher s ide, al Ihe Guayana , therc has been an appearancc o r difTerent vegeta­tional fonnations oriented by a long hi story of erosive changes. In the Coastal Range a lmos! half ofthe am phibian species are biological endemics, too. The faet thal mOSI amphibian endemics are found in elevated lands is evidence ravori ng the close rcla­tionship between abundance of life zones and diversity of am­phibians. The Andes has 16 life zones, and both the Coasta! Range ami the GlIayana regions have 10 each.

The siluatioll is d iffe relll with Ihe rept il es. Though there are 119 specics in Ihe country considered as biological or polili­cal endemics, only one region has a greatesl number of endemies, the Guayana, with 40% oflhe ir total species endemic. Reptiles are less restri cted 10 a geographica l place and have a wider eco­logical tolerancc, thus eX lending the ir distributions inlo di rrerenl

11. Environmental changes due to astronomic factors a) Globa l elimmie ehanges

1) in tcmperature pattems 2) in preei pitat ion and relati ve humidity pallerns

b) Ultrav io lel radiation ine reases c) Not-yc l-eva luated factors

1) Cosmic dusl impact 2) Micromeleors 3) X- rays 4) Gamma-rays

Plate 34

Plate 36

Plate 37 Plate 38

Plate capt io ns: 31. Geochelone carbonaría. A common and popular tu rtle Most peasants rise this species as food and peto Venezuela, Guárico. 32. Thecadactylus rapicaudus. A gekkonid lizard with an ample distribution in the country. Venezuela, Mérida . 33. Norops nitens. Venezuela, Guatopo. PIToro cOllrlesy 01 Lallrie .!. Via. 34. Polychrus marmoratus. A chameleon-type lizard, common in the wet forests. Venezuela, Aragua. 35. Anadia bitaeniafa. A member of a taxonomically very complex group of lizards from the Andean páramos and cloud forests. Venezuela, Mérida. 36. Gymnophthalmus speciosus. One ofthe smallest lizards; it lives in the litter of most types of forests. Venezuela, Mérida. 37 . Ameiva bifrontata. A medium sized lizard with an ample distribution in the lowlands. Venezuela , Táchira. 38. Cnemidophorus lemniscatus. A ground dwelling lizard that probably has the largest distribution in all ecological zones of the country. Venezuela, Zulia.

VENEZUELAN AMPHIBIANS ANO REPTILES

n.:gions and [ife zones \\ ¡thin a region. The Falcón-Lara and lhe Llanos llave Lhe sma llesl Tlumbers

of endcmics in lhe COlllllry. These arcas ¿Ife macroenvironrnclllally hOl1logenous, 110l favoring speciation as l1111ch as in heterogeneoLls clcv<ll cd lallds with unstable cnvironmc11ls. Both rcgiolls do no! have strong biogcograph ical barricrs; Ihis u"anslates ¡ma a more cXlcnsive sharing orherpetological spccics belwecn bordering re­gions. We hYPolhesiLe that any zoological laxa \Vi II ha ve lewer cndemic specics in lhe lands of Falcón-Lara ar lhe Llanos of Venezuela. n lC Maracaibo Lakc Region is ullusua1. as Ihe amphib­ian cndcmic numbers are quite [0\\1 c01llpared lo lhe rcpl ile en~

dClllic specics and can be explaincd by Ihe exlensive dry lands. Thc finding of only a 10\\ numbcr o f 3mphibian cndcmics in Amazonas is probably duc lo insllffic icnl research in ils Icrrilory.

I:.xcmpliricd by Ihe lilcrature. an imal grou ps cxhibi l more endcl11ics al clevatcd lands primarily duc to isolation. For in­stan te, whcn the mammal ian faunn, bats cxc ludcd, is sludied , the Andes region sho\\'s Ihe largcsI pcrccnlagc of endcmic spce ies of Icrrcstrial malllmals (Eiscnbcrg and Redford 1979). Supposedly, any laxon closel) assoeiated wit h el geographical tcrritory (as opposcd lo a loose <l ssoc iation , as in bals, birds, ny ing insccls, cte.) has experi enced a high cl egrce ofspcciation in Ihe Andes, mainly due lo Ihe g laciation processes du ring Ihe Ple istoccne (Schubert and Vivas 1993) and Ihe correspondi ng vegctal iona l responses 01' expansion and con lraclions (Ouellman 1982). A general rcview is nccdcd. but the preliminary work by Diazct al. (1997) seems tO be an indication that th is aClually oecurs.

Conservation Venel.ue la has a human poplllation wit ll a high degree 0 1' envi­ron mental coneem. but it is a lso a country wilh a ralher slrong push for development. This contrad ictory sÍluation has statisti ­cal express ion s in the high and rap id clestruction ofnalural cnvi­ronmcnlS, with arate 01'600,000 ha of de forestatiol1 yearly and Ihe ex islence, at the prcsent time. or 43 Naliollal Parks, Iwo Biosphere Reserves. and seven Wildli le Refuges (MARNR 1992). Such a greal Tlwnber 01' prolec led areas shou ld help lo conserve tile herpctorauna within Iheir limits.

Unlortunately, actual st ud y cases on the dedinc 01' animal populations are rew (Ojasti 1967; La Marca and Rcintll<ll cr 199 1). Fe\\ docul11en tcd cases are kllown ",here herpetological spec ies have been kno\\ n to be on the verge o rcxtinct ion or have vcrifiablc populatiol1 declines. Undoubtedly. Ihe large reptiles llave beell the most alTectcd by [111 cxtracti ve comlll crce (e .g .. co ll ccling for the skin trade,mcat consulllplion) devcloped prior to Ihe 1980s wll en several regulatiolls "'ere eSlab lishcd. Today strict regulations are in action to protccl crocod ilcs and lurtlcs.

Amphibialls art! also imperi led, as can be dcrivcd from Ihe provided li st ofendangcred spccies (Appendi x 2). As illll1any olher plaees ofthe \Vorlel ( l3J austein and Wakc 1990), a dccirna­tion ofalllphibians population is occurring in the h igh lands 01' Vcnezuela. The loss or reduclion 01" élm ph ibian species may ha ve some importanl ccological consequences, for Ihey are crucial to food clwins. Thll s. the ene rgy linkage. as weH Ihe predalor and/o r the prey POpul,lIions \Vou ld be a1terecl. Th is cco logical biodiversi ty irnpoverishment wou ld carry a loss 01" genetic material 01' potcntial use as \Ve ll as a wasle 01' food andl o r phannacological resources. The on ly pattern fo und among most orthe Venalle lan declining <1mphibian species is Ihat CI

high 1l1ll11ber are roun e! at elevation. Why lhi s occurs is nol totally understood blll lllay be due to water cOllt.amimuion or

57

highcr UV radialioll levels at elevat ion. thlls exhibiling vu lner­ab ility of a mph ib i.an s pecies to this reg ion particularly (Blaustcin et al. 1994). Many causes can be responsible lor Ihe amphibiCln Clnd olher verlebrale's decline, bUl most can be framed into two kinds 01" env ironmental changes: hurnéln activi lics and ¡:lslronomic faclors (Table 9: Péfaur 1993).

Many conserval ion problems an.: faeed ¡¡Ild intcnded to be so lved at the intcrnationa l leve l and at Ihe nationalleve l by diITcrel1t gove rnmellt offices. As a natio n. Venezuela has par­ticipatcd in severa l inlernational agrccments 10 prolecl thc cnvirolllll enl and/o l" the fauna. Among Ihe more imporlant o nes a re the Biodivcrsity Convention and The Ama/onic Co­opera ti o n Treaty. designed lo protect la rge areas: and the In tcralllerican Agrcemcnt for Pro leetion and Conservation of Sea Turtles, as we ll as the Fresh\\ aler Convention. \\ hich is conccrned with quality and quantil y orIlle resourcc. il s ba­s ins. tll e arca dr<lined by a river and ilS branches. ¡me! its fauna. Proj ec ls 10 p rotecl c rocodilcs (Gorzula 1985: Thorllbjarnarso ll 1991: Velaseo and Ayarzagücna 1995; Baquero de Pcdre t alld Quero de Ilella 1996; and many Icch­nical reporls listed under Seijas 1993) or lurlles (Licma et al. 1996) under Ihe respons ib ilit y ofthe Venczuclan Mini slry of Ihe Env irollmcn t (MARN R) shollld he lp cnsurc propcr pro­lecti an and prese rvation orthesc animals. Al severa l Venezu­elan univers ity laboratories. there are projects studying the biolagical and ecological aspects of amphibian and rcplile species providing grounds lO protcct these va lllable rCSOllrces. !\s a co rollary, il is usuall y acccptcd w isdom thal the better we know a fauna al1(l it s associatcd ecosystems. the better protection we can olfl.! r.

A list 01' im pcriled species (Ihis study). lhe existence 01' protected areas, logelher \\ ith managcl1lcl1t and seientific projects dcsigned lO protecl Venezuela's amphibians and reptiles. alld the increasillg internat iollal COllcenl ror thcl11. strcnglhcn the hope of preserving its va luable herpetofauna populations. and other resources, lor I"uture gcncrat ions.

Conclusions The existence orelevatcd ranges, extensive fo resls, and a myriad of ri vers and olher aq llatic habitats, makes Venelue la an ellvi rolllllcll­lallllosaic where a divcrsc fauna of252 amphibiallssJll-"'Cit'Salld 299 rcpliles speeies live. Each ofthecight current biogeographieal lones has a relatively nUlllerous herpctofauna the most diversc are the Guayana and lhe Andes, wilh 310 and 142 spec ies, respcctively.

Oue lo the large arca covered by certai n territori es. such as the Llanos and GlIayana, tlle obtai ned value for the cOlll1try 's Spccics-Richness Index is relatively 10\\. Ilo\\cver, lhe regions s iluated in northern Venezuela appear high in Ihe \\orld\\ide spccics-richness ranking.

Thc similarity among the biogcographical regions is ratller lo\\' wltcn comparing s ll ared am ph ibian spec ies, bul it is higher \.vhcn comparing reptiles. Se lective biogeographieal balTicrs \\ork mainly in the mounta inolls regions and are Icss cOcctivc in the cl:Ologica l continuolls landscape 01' the rest of lhe countr) . The higher <lmphibian s imilarity is found ml10ng the Llanos·AmClzonas regions, wh il e for Ihe reptiles is fO lllld among the Guayana­Amazonas regions.

Endemislll in amphibians dcvl! lops more in e lcvaled lands. such as in the Guayana, the Andes. and the Coaslal Range. There are less endemic reptiles than amphibians. due to thcir grcatcr abi lity tO di sperse. The arca with more endemic reptiles

Plate 44

Plate 45 Plate 46

Plate captions; 39. Tupinambis teguixin. The largest lizard of the country, is common in the Llanos and in the lake Maracaibo zone. Venezuela, Zulia. 40. Leptotyphlops affinis. This small ground dwelling snake, can be found in restricted parts of the Andean region. Venezuela, Mérida. 41 . Helminthophis flavolerminatus. This curious little snake distributes in many environments of the northwestern states. Venezuela , Mérida. 42. Leptodeira annulata. This species is probably one of the most common snakes and the one with the largest distribution in northern South America . Venezuela , Táchira. 43. Oxybelis fulgidus. This colored snake inhabits the southern lowlands. Venezuela , Bolívar. 44. Phylodryas viridissimus . An attract ive snake distributing in the southern sta tes of the country. Venezuela, Bolívar. 45. Micrurus mipartitus. Venezuela , Guatopo. P/¡OfO cOllrtesy o/ Ul/lrie 1. ViII . 46 . Bothrops venezuefensis. A large snake usually found in the forested envi ronments of the northern mountains. Venezuela , Táchira.

JAIME E. PÉFAUR AND JUAN A. RIVERO

IS Ihe Guayana reglon. In tenns of conservation, il is knowll thal Ihe more com­

mon environmenta l changes, destruction and rragmentation oC habitats and agricu lTura! expansion, \Vil! have severe cOllsequences for Ihe herpetofaulla in Venezuela. As many species are endemics and restricted lo 5m31! areas, lhe destruction of a few kilometers of Ihe habitals cou ld eliminate several spec ies. The rap id de­struct ion and/or contamination ofnatural env irollments are caus­ing a decline in herpetorauna po pulations and numbers so as lo pUl sorne species al extinclion ri sk . Mas! imperil ed amphibians are ¡hose living in Ihe highlands of northern Venezuela, whil e the mos! endangered rept il es are Ihe marine turlles, crocodi li ans, alligators, and \Un les. [1 is hoped lhal lhe existence of seve ral Nat ional Parks and other protected arcas, and the increas ing awarcness of the ci tizens, wou[d he lp lo preserve the Venezu­clan herpetofauna.

Acknowledgements.-We thank the people ofthe Animal

Eco logy grollp at the Uni vers idad de Los Andes and ol' the Department 01' Bio logy at the Uni versidad de Puerto Rico, Mayagüez. fOI" their permanenl support lO conti nll e with our herpeto log ical stud ies in Venezuela . In particular \Ve are in­debted to William E. Duellman, James R. Dixon, Pedro Durant, Naney M. Sierra, AlberlO Ve loso, Jesús Manzanilla , Gustavo Casas-Andreu, Césa r Moli na, and Ihe late Adao J. Cardoso for their help and continuous prov is ion 01' data and references. Thanks are ex tended to the curators of the vis ited museums, espec ially to Ihe Colección de Vertebrados, Universidad de Los Andes. Mérida. Venezuela; Estación Biológica Rancho Grande del Minister io del Ambiente y de los Recursos Natu ­rales, Maracay. Venezuela; Musco de 1-1 istoria Natural La Sa lle, Caracas. Venezuela; Museo de Biología, Facu ltad de C iencias, Universidad Central de Venezuela, Caracas, Venezue la; Musco de Zoología Agríco la, Unive rs idad Cen tral de Venezuela, Maracay, Venezuela; Museo de Biolog ía, Universidad de Puerto Ri co, Mayagüez, Puerto Rico; Instituto de Ciencias Natura les, Universidad Nacional, Bogotá, Colombia; Instituto de Biolog ia, Uni vers idade Estadual de Campinas, Sao Paul o, Brazil; Mu­seulll ofNatura l I-li story, The Univcrsity ofKansas, La\\lTenee, Kansas, USA; Fi eld Musculll o f Natural I-li story, Chicago, Illinois, USA; Departmenl of Wi [di ife and Fisheries Seiences, Texas A&M Un iversi ty, Co llege Station, Texas, USA ; Mu­seum of Vertebrate Zoology, The Uni vers ity 01' California, Berkeley, Cal ifornia , USA ; Museum ofComparative Zoology, I-Iarvard Universily, Cambridge, USA ; and Museum National d'Histoire Naturel le, Pari s, Franee. C0111ments and correct ions made by Jaek Si te s, William W. Lamar. Craig I-I assapaki s , and t\Vo anony mous revie\vers are deepl y appreciated. Errors re-111ail1 ours. Be lk is Rivas and Yhilda Paredes were helpful in the prepa ration of previous drafts . Mari se la Angelino prepared Ihe final version or lhe manuscript. This study has been granted by CDCHT-Universidad de Los Andes (C-805-96) anel CON IC IT (PI-O 11) .

References Avila-Pires, T. C . S. 1995. Lizards 01' Braz ilian Amazonia (Rep­

tilia: Squamata). Zoologisr.:he Verl/(/ndelillgell (Leiden) 299 : 1- 706.

Ayarzagüena . .l. 1992. Los Centro lénidos d e la Guayana Venezolana. Publicaciones Amigos de Dofiana (Sevilla) 1 AS p.

Ayarzagüena. J., Señaris. J. C .• and Gorzula , S. 1992. El grupo

59

OSleor.:eplwlus rodriglle=i de las tierras altas de la Guayana venczolana: descripción de cinco nuevas especies. Memorias de la Sociedad de Ciencias Na/urales (Caraca.~). Tomo L II 137 :\\3-\42.

Ayarzagüena. J .. Sci'laris, J. e, and Gorzula, S. \992. Un nuevo género para las especies del "Grupo OSleot.:ephal ll .~

rodri~lIe=i" (Anura: Hylidc¡e). Memorias de la Sociedad de Ciencias Na/m'ales (Caracas). Tomo L[I 138 :213-221.

Baquero de Pedret, B. and Quera de Per'la, M. [996. Manejo del programa de zoocriaderos de la especie Baba (Caiman eroeodilll s) en Venezuela. ZOO(."l"imJe/"Os 1(1 ):0[-06.

Bisbal, F. J. 1988. Impa cto hum~no sobre los habilat de Venezu­ela . Inferciencia 13(5) :226-232 .

B[austein, A. R. and Wake, D. B. \990. Declining amphibian populalions: a global phenomenon? Trends in Ec%JO' alltl EVO/Ulio/l 5(7) :203-204.

Blaustein. A. R. , Ho ffrnan . P. D .. Hokit. D. G., Kksccker. J . M .. Walls. S. c., and lI ays, J. 13. \994. UV repair and rcsistance to solar UV-B in amphibians cggs: a link 10 population declines? Proceedings o/Ihe /l/alional Academy o/ Sci­ences 91 :1791-1795.

13 odin i, R. and Pérez- Hernández, R. 1985. Proposición de regiones biogcográri cas para Venezuela en base a la distribución de [os Cebidos, p. 323-333 in A Primatolog.ia no Brasil 2. Allais do ] 0 Congreso Brasileira de Primaralagio. Campinas.

Brame. A. 1-1. and Wake, D. B. \963. The salamande rs of Soulh A meri ea . COnlribu/iofls in Science. Na/llral HislOry Mu­selllll o/ Los Angeles Cowlly 69: 1-72.

Brown, K . S .. 1r., Sheppard. P. M .. and Turner, R. G. 1974. Quaternary refuges in tropical America: evidence frorn race rormal io n in Helicollills butternies. Proceedings of ,he Royal Socie/y o/ LOlldon-(fJ) 18 7 :369-378.

Campbcll , J . A. and l.amar, W. W. 1989. Tlle Venomolls Rep/ile:s o{ La/in Amer¡ca. Corncl1 Univers ity Press, 1thaca, Ncw York. 425 p.

Casas-Andreu, 0.. Méndez-de la Cruz. F. R., and Camarillo. 1. L. \996 . Anfib ios y reptiles de Oaxaca: liSIa, distribución y con­servación. AC/a Zoológica Mexicana (nueva serie ) 69 :1-35.

Di Bernardo . M. 1992. Reva[idatian of the gcnus EchinaJllhera Cope. 1894, and it s conceptual amp[ification (Scrpentes, C o lubridac). Comunicaciones del MI/ seo de Ciéneias PUCRS, S'érie Zoologia (POrlO AfeRre) 5( 13 ):225-256.

Diaz, A .. Péfaur. J . E., and Durant, P. \99 7. Ecology of South American Paramos with emphas is on the fauna of the Ven­ezuelan Páramos. p. 263-310 in Wielgolaski. F. E. and Gaoda[l, D. (editors). Ecosysrems o/ Ihe World, Volurne 3. Polar and Alpine Tundra . E[scvier Science, Arnslcrdarn.

Dixon, 1. R. 1980. The nealrapica! co!ubrid snake genus Liophis: lhe generic concept. Milwaukee Public Muse lll/J. Cun/ri­bu/ions in Biology and Geo logy 31 :1-40.

Dixon, J. R. 1989. A kcy and chccklist 10 the neotropical snake genus Liophis with eountry list s and map s . Smi/lrsonial/ Herpewlogieal Informa/iol/ Service 79 : [-28.

Di xon, J. R. and Hendrie ks. F. S. 1979. The wortllsnakes (family Typhlopidae) of the Neotropic s , exclusi\e of Ihe Antill\!s . Zo%gi!}'(:he Verlwndelingen (Leiden) 173 :3-39.

Dixon , J. R. and Michaud, E. J. 1992. Shaw's black-backed snakc (Lioplris melallotrls) (Serpent\!s: Colubridac) of northern Sou th America . ./ourna/ o/ Herpelology 26(3) :250-259.

Duellman , W. E. 1979. The herpctofauna of the Andes: pat­ter ns of distribution. o r igino differentiation and prcsc nt communities, p. 371-459 in Duellman. W. E. (editor). Tire SOIlIIl American Herpelofaul1a: ilS origino evolu/ion (tl/d dispersa!. M/lselllll o/ Nalllral Hi.\·/oIY n/ ¡he UnÍl'ersi lY of KOfl.\·as Mon()graplls 7 .

Due llman, W. E. 1982. Compresió n climática cuaternaria en los Andes, p. 184-201 in E/ee/os sobre la e!}peci(l ción. AClas VI [1 Congreso Latinoamericano de Zoología, Mérida.

Ducllman, W. E. 1985. Rcproductive mode s in anuran amphib-

VENEZUEt AN AMPHIBIANS ANO REPTILES

¡nns: phylogcnclic signi ficance 01' ndn pti"c stratcgics. SOIllIl AfriclIlI./o/lrl/af ofScil!llee 8 1:17.l·178.

Duc1Jman. W. F. 1988. Pattcrns of spc:cies di\'crs il} in .m uran amphibians in American tropie s. Al/l/ah o/ ,he Missullr; BO/(lI/icfll (Jarden 75 :79- 104.

lJucllrmut. W. E. 1995. Temporal nuclUalions 3nd abundance 01' • muran umphibians in :1 scason:.1l AmuLoninn rainforcsl. .IQllmal 01 l-terpelUlog) 29( 1): 13-21.

Ducllman. VV. E. and Trucb. L. 1986. Biology 01 Amphibiam. McGraw-11i11 Book eo,. Nc\\' York.

Ducllman. \'./. E. and Wic ns. J. J. 1992. The s talUs of ¡he 11) lid frog gl'I1US O/o/ygulI and Ihe recognition 01' ScillllX Wng ler. 1830. (}c('(lsiollaJ Papers o/I/te MlIselll1l o/ N{lllIml fli!¡· 101J' (JI,lte Univer:;ir.\' o/ Kalls'I ,~ 15 1: 1-23.

Duranl. P. <lnd Dia z. A. 1996. Hcrpdorauna de cinco cuencas andino \cnezolana:-.. p. 351-375 in Péraur. J. E. (edilOr). HerpelO/ox;o Neolropical. Universidad dc 1.0" Andes. Consejo de Publicaciones. Mérida .

Eisenberg. J. r. ;lnd Redrord. K. 1979. A biogeographic analys is or Ihe mamrnalian fauna 01' Venezuela, p. 3 1-36 i/l Eisenberg. J. F. (cdilor). Vertebrate Ecolog)' ;n Ihe Norlll­an Neo/rop;cs. alion:ll Zoological Park. Sm it hsonian Institution Press. Washington. D.C. 271 p.

Ewel. J. J. ami M •• dril.. A. 1968. ZOllas de I'ida de Velle=lIe/o: lIIellloria explicaliva .l'ohre el lIIopa ecoló~ic(). MAC. Edi­lorial Sucre. Caracas. 265 p.

Fro:-.1. D. R. 1985. Ampllihiall.\ Species of Ihe "'orlll' o 1lI.\'O//olllic ami Ke/lgraphical refae"ce. Associ.ltion or Sy:.tell1mic Col­Icctions Hlld AlIen Pn .. s~, Ine .. kl\\rcnce. Kansas.

Frosl. D. R. lllle! Elhcridge. R. 1989. A phylogcnclic ana lysis and laXOnorn) or iguanian lil.ards (Rcptilia: Squarnata) . Mu:.ellm 01 Na/llral Hi.\lory o( Ih(' VI/i l'er!>'i t)' 01 Kall slI.\ kli.~cellolleol/s PI/Micolio"s 8 1: [-65.

Gallardo. J. M. 1965. A propósito dc los [.cptodaCI)lidae (Am­phibia. Anura). PopJi,1 IIvlIlso.l' dc Zoolo:<!ia (SÜ() Palllo) 17:77-87 .

Gallardo. 1. M. 1965. Thc spccics !Julo grllnulo.w .\' Spi...: (Salientia: Bufonidae ) und ils gcographic \ ariation. DI/I/c­lin (~( Ihc MlI.I' ell lll uf Cumpurll lil 'e Zool(}~y (I/un'ard) 134 :107-138.

G:.nardo. J. M. [969. La distribución de las sub..::spcc ies de Bufo grwlI/losl/l Spi:\: MI fidelidad a los s istemas hidrográricos Sudamericanos. Ciellcio e II1l'e,lliKoC;oll (/Juellos Aires) 25 :406-416.

<.iofí'ul::!. S. 1985. The managcmclll ni' crocodi1ians in Vcncí'uel::!. p. 91-101 ;11 Wl·bb. G.. Manolis. C .. and Whitdlead. P. (cd i­lors). IVi/dlije MW/aj.!l!lIIel// : aocodile,\ l/lid 1II1iROlIJr.\. Sur­re)' Ikally <lnd SOIlS P .. LId .. Chipping Norton. Au slralia.

I la)'ck. L-A .C. 1994. Analysis 01' 3mphibians biocl ivcrs ity data. p. 207-269 ;11 Ileycr. W. R .. Donncll y. M. A .. McD iarrnid , R. W .. Ila)ek. l.-A . C .. and Foslcr. M. S. (cditors). Meas///'­il/K al/d MOl/ilOrillj.! BioloKicol Dil'cr ... ity: sf{¡lIdard lI1e/h­ods for lIlIIp/¡ibillll.l. Smilhsonian Institulion Prcss. Wash­in gton. 1).(.

I loog mocd. M. S. 1979. The hcrpetorauna or Ih..: Ciuianan re~

g ion. p. 241-280 ill [)uellman, W. E. (editor). Tite SO/lflt American I/erpeloflllma. ¡,s origill. el'ollllion l/mi dispersal. Muselllll 01 lite Nalllml f-/isIOI:1' (JI lite Univerúly o[ KlIflsas MOl1of!.raplts 7.

Iloogmoed. M. S. ami GorL.lIla. S. F. 1979_ Chccklis t of Ihe sa\ arma inhabiling frogs or thc El Manteco region \\ ith notes on lheir ecolog) and the description or a !le\\' spccic<, or Ircc-rrog ( I-Iylidae. Anura). ZoologiscJrc /Hedcddillf!.c 54 : 18 3-2 16 .

!Iuber. O. and Alarcón. C. 1988. Mapa de Vegc/acirin de Ven­(';lIe10. Oscar Todlm:mn Editores. Caracas.

John son, J. D. 1989. A biogcogr:.phical analy~is or Ihc herpctofauna 01' Northwestern nuclear Ccnlral Ame rica . Mi/l\'tlllk.ee PIIMi" MII.lell/ll COl1lribllliol/s ;11 JJ;O/OKY (11/(/

GeoloKY 76 : 1-66.

60

La Marca. E. 1992. Ca ta logo taxonómico. biogcogrMico y bibliogrMico de las fa na s de Venezuela. CUllderllo .~

Geográficos (Merida) 9: 1- 197. [.a Marca. E. 1995. Biological and ~)slemati c s~nopsis 01' a

gcnus 01' rrogs from northcrn mounlain s 01' Soulh Amcril:u (An u ra: [) cndroba l idae: Manllop/¡ryn e). BII/lelill of lite Marylolld Herperoloj!ic(¡J Sude/y 3 1 (2) :40-77 .

La Marca. E. and Rcinlha ler. H. 1). 1991. Populatio n chang.cs in Alelop/ls specics 01' Ihe Cordillcra de Merida. Vcnc,mcla. I/erpefOlogical Rel'iell' 22 (4): 12 5·128.

Lancini. A. R. 1986. Serpiellles dc Vel1c;lIela. Erneslo Armitano editor. Ca racas.

I.ce. J. C. 1980. An ccogcographical analysis 01' the hcrpcloraun:. 01' the Yuca tan Peninsula. M/I.\'ell ll/ of Na/lIral l1i\'IO/:\' o/ /he Ulliversily oI KlIIl,I'(/,\' Miscellllll eOIlS Pllhliclllioll .\ 67 : 1-75.

I. ical:l. L .. Galvez. S .. Marín. E .. Rebolledo. N .. lJ"eche. E .. and Pérc7. F. 1996. Bases para el manejo de la tortuga arra u (Podl1cllellli.~ expllllslI) en el Orinnco medio. p. 379-417 i/l Pé t~ lI r. J. E. (editor). lIerpe/olof!Ja Neolropical. Uni"ersidad Los An des . Consejo de Publicaciones. Merida.

Mago- Leccia. F. 1970. LisIo de lo:. peces de Jlene=uela : im:luyemlu /111 esllIllio preliminur sobre la icriogeuKralill del/mis. Ministerio de Agricullura ~ Cri¡¡. Oticina Nacional de Pcsca . Caracas. 283 p.

Manza ni lla. J.. Ferná ndcz- Hadillo. A .. La Marca. E .. and Visbal G. IC 1995. Fau na del Parque Nacional I lcnri Pittier. Ven­ezucla: composición y di s tribución de los anfibios. ,1 CI(¡

Cienlij'iclI Jlelle=()/mw 46 :294- 302. Manzanilln. J.. FcrntindC7-Badillo. A .. and Vishal (l .• R. 1996.

F:lUna del Parquc Nacional ll cnri Pillier. Vene/ucla: composic ión y di st ribución de los repliles. AC/(l Ciemi{icCl Velle;ololla .t7:1 - 12.

MA RNR. 1992. AreM Nml/m/c:. Protegida.~ de l'elle;ue!(/ . . ~cr;e lIspeclos com:eplllule.\· y meloc!o!uKicos. D(iSPOA/ ACM I 01. Caracas.

Marn:ro. L. 1964. Velle;ueltl y SII.\' Recllnos. Cultural VenC7olalla. Sociedad Anónima. Caracas. 700 p.

Mcdelll. F. 1981. Los Cmcodylia lh- SlIr Alllérica. Volume 1. Los Crocody lia de Colombia. Colección de Ciencia ... Mini!'tterio de Educnción Naciona[. Fondo Colombiano de ln\cs lig:le iones Cientificas > I'ro~c<:los E!'tpl'cialcs "l'. J. de Caldas." Bogot:í.

Mcdclll. F. 1983. Lo.\ Cro(:od)'lia de SlIr América. Volulllc [1.

Venezue la , Trinidad. Tobago. Gu)una. Surinallle, GlIa~ana Francesa. Ecuador. Perl!. 1J01i " ia. Brasil. Paragua). Argen­lina. U rugua~ . Colección de Ciencias. Ministerio de EduCa¡; ión Na¡;ional. I;ondo Colombiano de Im es tigacioncs Cientificas ) Proyectos Especi'llc:. "F. J. de Caldas:' BOgot{L

Myers. C. \V. 1987, New genc r ic namcs ror <¡ome ncolropk:..1 po iso n rrogs (Dcndrobatidal'). Pa/J/!is AI'/¡/WJI de Zoolo,:,ia (Stio P(lIIlo) 36(25 ) :301-306.

l\'l ~crs. C. W .. l'aolill0. A .. and Dal). J. W. 1991. Di sco\er~ 01' a dcfcnsivcl) rnalodorolls and nocturnal rrog in ¡he famil) Dcn{lroba ti dac: ph ylogenctic significance 01' a n¡,:\\ gcnus and !>pl.:c ics rrorn Ihc Vcnezuclan Andes. VQ\'iWle.\·. Alllt'ri­call MlI.I'eUIII of Nalural Nis/OI:I' (Nell' )'1"'10 3002. 33 p.

Nussbaulll. R. A. flnd Wilkinson. M. 1989.011 thl.: classification am[ ph) logcny 01' Caecilians (Amphibia: G) rnnophiona). a critical revic\\'. l/erpelO lo¡.:,ic:lIl MUflofVaplt.l' 3 : 1-42 .

Ojasli. J . [967 . Considcraciones sohre 1:. ccologia ~ COl1senar.:iÓn dc la tortuga Podocllemi~ eXIU/ma (Chclonia. Pclo­medlls inac). AI(I.~ do Simpa:..id ~ohre (/ Biala Amll;ollicu

7 :201-206. Péfaur, .l . E. 1985. Ncw spccie~ 01' Venezuclan ('olO\'lellllll

( J)cndrobalidae). Joul'1lal nf I/erpl.'lulogy 19(3 ):321- 327 . P~faur. J. E. 1992. Lisl ¡lnd bibliograph) (1960-85) 01' the Vcn­

czuc!an hcrpctofauna . Smi//uonioll Herlu!lolo;:,it-ul In/o/'­lII(lfiOI/ Service 89 : 1-54.

Péfaur. J . E. 1993. Dcscriplion or a l1e\\ Colol/elhul (Dcnd roba t idac) \\ ith sorne natural hislor~ corn ments on

JAIME E. PÉFAUR ANO JUAN A . RIVERO

¡he gcnus in Venezuela. Aly'e.~ 11 (3) :88-96. Péfuur. J. E., and Diaz de Pascual, A. 1982. Aspectos

biogcográficos de las comunidades de anfibios y saurios de los Andes venezo lanos, p. 229-262 in Salinas. P. (ed itor). Zoología Neorropical. Actas VI I I C on greso Latino­americano de Zoología, Mérida.

Pélilu r, J. E. and DiaL de Pascual, A. 1987. Distribución ewlóg ica y variación temporal de los anfibios del Estado Harinas, Ven­ezue la. Revista de Ecología L(lr¡noamericana I (3-4) :9- 19.

Pda ur, J. E. and Ilérez. R. 1995. Zoogcografía y variación espacial y temporal de algunos vertebrados cp igcos de la zona xerófila de la cuenca media de l rí o Charna, Me rida, Venezuel a. Eco/ rópicos 8( 1): 15-35.

Péfaur. J. E. and Sierra, N. M . 1995. Status of Lep/odac/yius /abyrimhiclls (Ca lf Frog, RWIlI lemero) in Venezuela. Her­pelOlogical Reviell' 26(3) : 124-127.

p¿raur, J. E. and Rivero, J. A. 19 89 . B iogeograna de la herpetofauna venezolana. Re.~lÍmelles i COlIgre.\'o Lmino­americano de Ecología 7 p.

Pé r:1ur. J. E., Pére.l. R. , Sierra, N., an d Godoy, F. 1987. Density reappraisal of caec ilians in the Andes of Venezuela. Jour­nal 01 Hape/oloKY 2 1(4) :414-419.

Praderio. M. J . 1985. AspeclOs ecológicos de u/w pohlación de Co los tethus herminae (Dendrobafidae) en la qllebrada de Gllariquira. Esrado Miranda. Tesis Especial de Grado, Universidad Simón Bolívar, Caracas.

Presch, W. 1980. Evo lutionary h isto ry of the South America n Microteiid lizards (Teiidae: Gym nopht hal mi nae). Copeia 1980( 1 p6-56.

Pritchard , P. C. 11. and Trebbau, P. 1984. The TI/r/les uf Venezu­ela. Society for the Study 01" Amphibi an s and Reptiles, Cushing-Malloy, Ine., Ann tubor, Mi c hi gan.

Ramo , C. 1982. Biología del galárago (Podocl/clllis vog/i ) (Muller, 1935) en el Hato El Frío. Llano de Apure (Venezu­ela). Doílana ACf(¡ Ver/ebra/a (Se villa) 9(3) : 1- 157.

R ivero. J. A. 1961. Sa lie ntia 01" Venezuela. Bullefin of/he MII­seum 0f Compararil'e Zoology (IJarvard) 126 : 1-207.

Rivero, J. A. 19633. The dislribulion of Venezucl an frogs. r. T he Maracaibo basi n. Carihhean ./olll'llal oI Scú:llce 3 :7- [3.

Ri vera. J . A. 1963b. The di Slribution 01" Venezue!an frogs. 11 . The Venezuelan Andcs. Caribbean Joumal 01 Science 3 :87-102 .

Ri ve ra, J. A. 1963c. The di s tribulion of Venezuc1an frogs. 11 1. The Sierra de Pe rijá and the Falcón region. Caribbean Jourlwl of Sciellee 3: 197-199.

Ri vera, J. A. 1 964a. The distrib uti on of Ve nezuclan frogs. IV. Thc Coastal Ibngc. Caribbean Jotlrlutl of Science 4 :307-319.

Ri vero. J . A. 1964b. The distribution 01" Venezuelan frogs. V. The Venezuelan Guayana. Caribbeall JOllnwl 01 Se iellce 4 :411-420.

Ri ve ro , J. A. 1964c. The distribution of Venezuelan frogs. VI. The Llanos and Delta region. Carihhean JOl/mal oI Sei­e nee 4:491-495.

Rive ro. J. A. 1979. Sob re el origen de la fauna parame ra de anfibios venezola nos. p . 165-175 in Sa lgado-Laboriau, M. L. (editor). El medio ambiellle páramo. IDEA , Caracas .

Ri ve ro. J. A. 1982a. Los Elell/herodac /ylll s (Amphi bia , Leptodacly lidae) de los Andes Venezolanos. 1. Especies del páramo. Memorias de la Sociedad de Ciencias Na /li Nd es La Salle (Caracas) 42( 11 8) :9- 16.

Rivera, J. A. 1982b. Los Eleu/h e rodacryllls (Amp h ibia. Lep tod ac ty lidae) de los Andes Venezo lanos. 11 . Especies subparameras. Memorias de la Sociedad de Ciellcias Naru­rales La Salle (Car(Jcas) 42( 118) :57-132.

Ri vera, J. A. 1985. Nuevos eenlrolénidos de Colombia y Ve n­ezuela. Brenes ia 23 :335-373.

Rive ro. J. A. and Solano, H. 1977 . Or igen y evolUl.:iÓn de los Elel//herodactyllls (Am phibia : Leptodacty li dae) de los Andes venc701a nos. MC'1II0rill.~ de la Sociedad de Ciencias Naturales La Salle (Caracas) 37:265-282.

Rive ro-Blanco. C. and Dixon, J. R. 1979. Origin an d di s lribu-

61

lion of the hcrpetol"auna of the dry lowlands regions 01' northern South America. p. 2 8 1-298 ill Duellman, W. E. (editor) . Tlle Somlt American Herpetofauna: i/s origino evo/u/ion and dispenal. Musellm of NalUral Hislory of ,he V niversi/y of Kall.ws Monographs 7.

Rodríguez, J. P. and Roja s-Suáre z, F. 1995. Libro rojo de la fauna venezolana . PROV/TA . Fundaciól/ Polar, Caracas.

Rozc, J. 1966. Taxollomíll y =oogeogralía de los ofidios de Vel/ e=ueJa. Editorial Biblioteca, Un ivers idad Central Ven­ezuela, Caracas. 362 p.

Rui /.-Carranza, P. M. and Lynch , J . D. 1991. Ranas Cenlrolcnidae de Colo mbia 1. Propuesta de una nueva clas ificación genérica. Lozania (Bogo/á) 57 :30 p.

Savage, J. M. and Guycr. C . [989. Infrageneric class ificatio n and spceies composition 01" the anole genera, AIIOlis, C/el/Oll/s, Dael)'loa. Norops a nd Semillrl/s (S auria: Iguan idae) . Amphibia-Rep/ i1ia 10 : I 0 5- 116.

Schuberl, C. and Huber, O. 1989. La Gran Sabana, Panorámica de /lila región. Cuadel'l1Qs Lagoven Lagoven S. A., Caracas. 108 p.

Scbubcrl, C. and Vivas, L. (editors) . 1993. El Cualerntlrio de la Cordillera de Merida. Andes Velle=olallos. Un iversidad de Los Andes/Fundación Polar, Mérida .

Seijas . A. E. 1993. Li s tado bibliográfico comentado sobre los crocodylia de Venezuela. Biblioapullles. documenfo técnico de Biodoc (Cllallare) 1: I O p.

Scijas, A. E . 1996. La conse rvación y manejo de Crocodílidos en la región neotro piea[ , p. 419-4 27 il/ Pél"aur. J. E. (edi ­tor). Herpetología NeOlropical. Un ive rs idad de Los Andes. Consejo de Publi cac iones, Méri da.

Sei'lari s, J. c., Ayarzagüena, J., and Gorzula, S. 1994. Los sapos de la familia l3u fonidae (Amphibia : Anura) dc las tierras alIas de la Guayana ve nezolan a: de sc ripc ión de un nuevo género y tres especies. Public(Jcione~' de la Asociación de Amigos de Do/jana (Sevilla) 3 : 1-37.

Si lva, J. L., Valdez. J .• and Ojas ti. J. 19&5. Algunos aspectos de una comunidad de ofid ios del Nortc de Venczuela. Bio/ropica ' 7(2) , " 2- ' 25.

Slalnn, M. A. and Dixon, J. R. 1977. The he rpctofauna nI" the central Ll anos: noteworth y rcco rd s, a ten tative ehccklist and ecological notes. Journal 01 flerpe/ology I1 ( 1) : 17- 24 .

Thorbjarnarson, J. 1991. An ana lysis 01" the spectac led ca iman (Caiman crocodilus) harvest progra rn in Venezucla, p. 217-235 in Robinson, J. and Redford . K. (editors). Neofropicol Wildlife Use alld Conserva/ion. Uni vers ity 01" Chicago Press, Ch icago, Il1 inois.

Van zolini, P. E. 1978. On Sou th American Hemidacrylll.\· (Sauria, Gekkoni dae). Pap eis AVlllos de ZooloKia (Sao P(lIdo) 3 1(20) ,30 7-343.

Vela sc o. A. and Ayarzagüena. J. 1995. Situación aClU al de las poblac iones de baba (Caiman crocodil lls) so metidas a aprovec hamiento co me rcial en lo s Llanos venezo lanos. Pllblicac iolles de la A.HJciaciÓ n de Amigo.\· de DOIj(Jlla (Sevilla) 5. 71 p.

Via l. J. L and Saylor. L. 1993. The s tatus of amphibian popula­tions: a compila tion and anal ys is. A repor' 0l,he Dedining Ampltibiall Poplllo/Íons Ta.~k Force (DAPTF) (O /he SSC/ IVCN Working Document l . 98 p.

Wilkin son , M . 1996. Reso lution of thc taxonomic s talU s of Neclocaecilia Iwydee (Roze) and a revised key to th e gen­era 01" the Typhlonectidae (Amphibia: Gyrnnophiona). Jour­l/al of HerpelOluKJI 30(3) :41 3- 41 5.

YÚ stiz. E. 1996. Aspectos biogeog ráfi cos de la herpetofauna de la cuenca hidrogdfica del Río Turbio (Estado Lara-Ve n­ezuela), p. 317-349 in Pél"aur. J. E. (editor). lferpe/oloKía Neorropicol. Unive rsi da d de Lo s Andes . Consejo d e Publicaciones. Mérida.

Mallllurip/ received : 26-April-1997 A ccepfed: 15-Jafllfary-1998

'" ..,

APPENOIX 1: CHECKLlST OF VENEZUELAN AMPHIBIANS ANO REPTILES T

IAMPHIBIA I ANURA

l' I~E

lArdo" ... '

A x

. RI"ro lA. lA.

I d.a Mar" 1996

lA. lA. lA lA. lA. 11M" ' I/J.

. 1 ,"d Manen, 1856) ,Rlvero 1974

1903 , , VillC/IIROi Rivero 1980 '""",ro' La Marca 1983

"'"""O" La Marca~ I! 1 · 1882

, S,I, 1824

IIJ. I IJ. 18. 18.

, .. "roo .. , SeI", 1799 m{jf¡'ws Linnaeus 1758) IIl1.\in/S Werner 1903

IIJ. r"/1lroo',,,, G'nthc~

, 1758)

A

1990

, Señar's. ''',,"'''0 SeOarl, 1993 I",b,,'

. aud Com,l. 1994

10.

IH. 111. IH. 111. 111. 111. 1/1. 111 . 1/1. 111. 111. IH.

"

198í 1900

",,,,, Sefrar',. 'I,,,/elrii

'"''',,''' SeO,d,.

. a"d Gorzul, 1994 1895)

. a"d Cur""a 1')94

rPNlnAF

',~ ""d'''''''' (Rlvero 1968) O .. áley' 1882) ,o" .. la~)992)

! .. osum (Riv~~() 1_9~~ , (SeOarl' ,,,,) ,

1992) I . 1992)

, , , ' (GOl" 1963) d,,,,,,,r ' (R'vero 1985)

'1~92) i I ~I (Rivero 1985)

0,;" .. ,,1'.' (Rlvero 1968) IRlvero 1985)

""/I"I,,,u (RI"ro 1985)

~er01985)

, 1993)

I zanes 11 I 1m 11V IVI VI IVII I VIII

C. (:uJllrinl'llsis Péraur 1993 C. dunni (Rivero 1961) C. hwni/is Rivero 1978 C. leo/Jar¡/alis Rivero 1976 C. mwulelorum (Schmidl 1932) C. parkeme Meinhardt and Parmelee 1996 C. so/IUCIrS;S Rivero 1978 C. smlluarlilli Rivera. LaIlJJ;one. and Pri,gioni 1986 C. ~J1rI?\'ei (Rivero 1961) D(!/Idmba/es leucnmelas Steindachner 1864 D. rlifll/us Gorzula 1990 Eprpedobares piwlS (fschudi 1938) E. IrÍl'il/afl/s (Spix 1824) MallllO ,/rr\"lIe col/Mis (Boulenger 1912) M. cordilleriana La Marca 1994 M. /terminae (Boen2cr 1893) M. ¡arQ/u/illa Yústlz 1991) M. /wlJ/i lla (fesr 1956) M. oblillera/a (Rivero 1984) M. ril'eroi (Donoso-Barros 1965) M. /rini/mis (Garman 1887) M. \'us/izi (La Marca 1989) Millvolxues Sle)"t' rm(lrki (Rivero 1971) Nephelolxlles alhogl.llllllus (Boulell~er 1903) N. dlmm/; (Péfaur 1985) N. Iwwleeae (Rivero 1976) N. IIwWlrgai (Rivero 1978 N. meridensis (Dole and Duralll 1972) N. lIIo/inarii (La Marca 1985) N. oros/onw (Rivero 1976) N. serruflus (Péfaur 1985)

IIYLlDAE Aparasphenodorr verrewlanus (Mertens 1950) FEee/OIIO/us fr/zxeraldi Parker 1934 F. '" '//WCIIS (BoeuRer 1893) CllSlrutlleca hdemre Dunn 1944 G. lIicefori Caige 1933 G. ol'i(cra (Weinland [854) G. lralkeri Duellman 1980 G. wil /iamsoni Caige 1922 H Y/ll l,lbomem:illC,la Spi;.,: 1824 H. (ll~p,mcww Soix 1824 H. alemwri Rivero 1964 H. emmw/ica AyarzaRúena and Señaris 1993 H. barlersbvi Rivero 1961 H. Imlile:; I~ivero 1961 H. bow,~ U/tnaeus 1758) N. ea/camw Troschel 1848 H. ere ,i/(I/IS Wied 1824 H. ¡:eo 'ra ,hica Soi;.,: 1824 H. srlllw.'·(I Boulenger 1882 H. ¡"van ¡u'si AyarzaRuena and Seilílris 1993 H. ·"llIIi Rivero 1961 11. kwwimo Coin and Woodley 1969

*Shading repreSen!s presence ofparticular species wilhin lhe respective b iogeographica l zone.

a<

'"

IN. '~) I H. ',,,d,,", Rive<o I H. ',,"ai Rivero 1972 I H. . Rivero 1961 IN., , Roux 1927 111. , , lLau,,",i 1768) I H. I ', ; (Rivero 1961 IN. " Cope 1886 11-1, I ! ! Rivero 1971 I H. mi"",o 'eteos 1872 I H . r I :;ü!lther 1858 ! H. l>"i'¡"a Duellm,o 1989 'H. . 1905 N. 1185, N. 1"""'''''0 . 1799) N. m",im" Duellm,o ,nd ! , 1992 N. ,iI,,,,,, Ri",o 1972 N. "i, a,,,,, Sol,oo 19,

I H. 1\'(lrreni 1 . ¡mIZ IN. w'''',ioi P"keo

0 , 0,

lo.

' 1', P. r . 1'. p,

. , I

IlI/ck/eyi I ~,nd Seo;,d, 1996

I,,,,i,,,,;; ID"m" il "ni Bibron 1831

'",,,im" ~~18~62 ______ ~~~~~~~~~ "',,,,'O", Lau"oti 1768f , ,bicolor(Bodd,en J772:

, " ID'udin 18U mediuai (t 1962 ,""n" ICope 1868)

r , I reone 1868) ni/litaris t I

(Rivera 196 I I Goin 1966]

,'a",,, ai""

cO~~~----------~~-+-+-r-r~~ '1m'o 1986)

"el",'o"",,, ISOiX¡ ,""m"', Peteo, 6~31-----------------+--t--r~r-+--t--r--r--l ",b" IL,,,renti ]' , , d ,od Co"", I, 1979)

eoi"i Rivem 196' I f I I (Rivero 196])

,im' Duellm'" ,ni Hoo.moed 1984

",'ooi , i

.. SeO"i" ",d Cm",ul, 1992 . i i. 'nd ~0,,,,1~2

. SeO,d" ,,,d Co,,,,I, 1992 . Se""i,. ,od Co,,"I, 1992 ~ d Coezul, 1992

IRivero .!..!!2!!l.

ALLOPIIRYNIDA E Allo ,lm'lle rUlhl'l'IIi Caige 1926 I I

LEPTODACTYLlDAE Addopltrrlll' gll/turosa Hoogmoed and Lescure 1984 AdCllomera wrdrclU' (Muller 1923) A. h't/acdaCf)'/a (Cope 1868) Ct'rCIIQ Jh",s calcan/la Boulenger 1890 C. wrmllu (Linnaeus 1758) lJischi(/Ollo(' /vlus duide/lsis (Rivero 1968) ». CO/OIII!elloi ~~r4!&!:!ena 1983 Eleulherodacr \'lu .~ lUlO/is Wal ker and Test 1955 E. bicumulllS (peters 1863) 1':. borO/we/lsis Rivero and M-ªy"o.!1@ 1973 E, briw!i (Boulenger 1903) E. cwuiu/Ils Myersand Donnellv \996 E., d!/orosoma Rivero 1982 E. colos/idlOs La Marca and Smith 1982 E. l'onwl llS (Jiménez de la Espada 1871) E. ---.EJncsi (Rivero 1964) E. 'oIIllSlol1e; Barbour 19 14 E. Icmdnii Donoso-Barros 1968 E. lemiJú/wsus Rivero 1982 E. mamWrcllllS (Bou le~r 1900) E. IfIwusi (Boeuger 1893) E. melcmoproclIIs Rivero 1982 E. mondol ¡j Rivero 1982 E. nir:cfori Cochrán and Coin J 970 E. pClf(¡men,S Rivero 1982 E, '/Cllrostri(jIllS Rivero 1982 E. Jrolixodimls Lynch 1978 E. )n,inmus Myers and Donnelly 1996 E. )/l/ido; Rivero 1982 E. Juh·inallu Rivero 1968 E. rer icldaws Walker and Test 1955 E. riveroi !:1:'!lch and La Marca 1993 E. ro ;:e i Rivero 1961 E. s/e/wdiscus Walker and Test 1955 E. lerraebolil'(¡ri .~ Rlvero 1961

E. wbenulS/u Rivero 1982 E. HlrUmilJllÍrensis Rivero 1961

E. urichi (Boe~r 1894) E. mnadise La Marca 1984 E. l'il(.lfsi (MeHn1941) E. '(II' jensis ~rs and Donne!'!y'1996 1':. ::.eucrordlls Lynch and Hoogmoed 1977 Le J/m/¡¡crvl,u bolil·iwlUS Boulem.ter 1898 L dieltrus Heyer 1994 L USCII .~ Schneider 1799) L. ¡'IS/t!anm¡ Barbour 1906 L k/Uulseni H~~r 1972 L IlIbialis (Cope 1877 L 11Ibyrinrhicu~ (Spix 1824) L !!E!.odClc/vloidt!s (Andersson 1945) L li¡honaeles H~er 1996 L /ollgirostris Boulenger 1882

O> ..

, 1 1926 .• (Sp" 18241

, L"IZ 1930

",'mil '(~I , (Cope 1

r 1883 "mo' . Pvb"m 1983 ,,,,,,,,,, Noble 1923

94 . Heyer 19'

"'"'''' (Peoers 1862 1

"""'fa, , 1 Sol",.. and lIe""ole 1965 11'. ""',,ri 1896) 11'. ,(cope~ ., , , (Cope

,boli,"a"" Pa.ke. 1927 11'. '"'''''''' LYOlcl~ , 1'. ",,,iII,, IROII"en 19

MICROIIYLlOAE I Zweife

~9) , , "a,·, , , O"ali, E. , D",d'n 1802)

• 1 ~ robllSllI , 1900

PIPIOAE

""'" ""''''",Ii I"ck,ohn 1976 p. ''''''''a Müller 1924 p. ''''''''' ROI"ven and Galge 1923 1'. "'pa ,U,m"'" 1758)

,,,n,"-, O<:"OO"I"'E

1/)~eudis ¡x"mloxa ; 1758)

I CAUDATA

lfigTl lODONTIDAE , boob,,,,,,,, frap'do 1942

11l. a,,"". j Wake 1962

ICA"rOl 'AIf)AE C(¡('cili(¡. , , Ro" and Sol'no 1963

"'ac"" Shaw 1802 , , ,Oonn 1942 , , 1 Unn"n' 1758

ji ,

, . , mber (Ro" an~ I r (Mikan

IRII 'TIDAE , ,(0"'00 1942)

Iw,,", , """,il 1 . 1882)

Pmomotyplll,IS kUl¡pii (Berthold 1859) T)'Vlllonce/es compressicmu/cl (Dumeril and Bibron 1841) T. IUI/aIlS (Fisher 1879) T. \'Cllc;::lIelenst' Fuhnnann [914

REPTlLIA

TESTUDlNES

PELOMEDUSIDAE Po(locllemi_~ e""llrocc"hu/u (Spix 1824) P. e.lfJWIsa (SchweÍlzaer 1812)

". IewVQlla Dumeril 1852 P. 1111; .'is Troschel 1848 p. \'08/i Mullcr 1935 Pe/'oce ,halllS dwneri/ian!u (Schwei~er 1812)

CHELYIDAE Clte/II" rmhrjatus (Schneider 1783 Plrr)'lU))S (Ph,,'/IO IS) f{eof r()(lIlIIS (Schweigger 1812) P. (Mesode mmw;) 'ihbllS (Schweh.!l!er 19/2 P. (B(Umclremvs) /WSJlWS Schwei¡;&er 1912) P. zulim! Pritchard and Trebbau 1984

EMYDIDAE Pkuemvs ¡/lJIycepJra/cl (Schneider 1792) RlriIlOcll'IIlIll}'J dilldelllllUl (Menens 1954) R. Jlmcwlaria (Daudin 1801 Tradll'm\',~ ,~c ri)/(/ (Gray 1855)

TESTUDINIDAE Geocllc/u/re /Chclollo ides) carbOllllri(1 (Spix 1824) I I . 1 I G. delltiC'J/I(Ha (Linnaeus 1766) I I I KINOSTERNIDAE

J I K¡'wsterrrOIl scor ,ioides (Lillnaeus 1766) I I I I

DERMOCHEL YIDAE I I I DerfllocJIt~hs coriacea (Linnaeus 1766) I I I

CHELONIIDAE CJrelm¡ja m\'das Linnaeus 1758)

I I I I I I

Carelw ('(I¡-('I/(/ (Linnaeus 1758) Erellllocliel\l' imbriclJIa (Linnaeus 1766 Le"idudrdWi uliWlcea (Eschscholz 1829

CROCODYL1A

CROCüDYLIDAE CrOClXl\'/u:> ClcullU (Cuvier 1807) I I C. illlefmediu,\ Graves 1819 I

ALLlGATORIDAE COilllCllI c/'Owdillu Linnaeus 1758 Cctimwt lri 'u Linnaeus 1758 J>a/euSllclrus /)(11 ¡ebrosus (Cuvler 1807)

I I I P. lri 'O/la/US (Schneider 1801)

AMPlilSBAENIA

I I I AMPHISBAENIDAE

1\111 ,hislx.ella alha Unnaeus 1758 A llliginosa Linnaeu$ 1758 A. graci/ is Slrauch 1881) A. rote; Lancini 1963 A. SJU ,relli Boulemzcr 191 5 Mesolxll:'no IlUeblleri Menens 1925

LACERTILlA

SCINCIDAE MlIbuva bis/rima ($pix 1825) M. CIIn,alhoi Reboucas-$Dieker and Vanzolini 1990 M. croi;:a/i Horton 1973 M. mabOlI}'o (Lacé ,edc 1788

GE KK ON IDAE ColeOlllICI )'/US amllymicus (Anderson 1918 e sep/elllr;O/w{is Yanzolini 1980 GOlla/(xle.~ alboJ!ularis (Dumeril and Bibron 1836) G. u/lIwlaris Boulenger 1887 G. a nrillensis (Van Lirlth de Jeude 1887) G. bodi"j; I~iyero- Blanco 1964 G. ceci/ill€ Donoso-Barros [966 G. conónll(l/l/S (O'Shaughnessy 1881) G. lI/couellsis Shreve 1947 G. Iwmera/ís (Guichenol 1855) G. ocella/lls (Gray 1831) G. ,etersi Donoso~Barros 1967

N. N.

N. N. N.

N. N.

P. P.

1'. P. P.

, 1834) <Id"" (WIIH,m, 1974)

, • (O",n,,11

,lbb'''I'' (Cope 186') """" (Waglec 1830) .,0<'" 1875

i r· Barros. r ';:.,=,.-= ",,, :;-1 "V¡"iill° "Ol,"ill' ,"' 199""6-t--+-+-t--+-+-t---t---t

I ·1 ¡ Mvers. WilHams. OI nd i i 1993 Ilkdori Dunn 1944 "",,11 Ba/m,. Willi'ms. and Vilocla 1996

Po/ychru., I 1 I ¡ 1758)

I IIR lI1AE

""'''''""' ; 1758) I T. 1/0,,,'1 Ro," 1958 1"1. ¡,,",pi<l,,, (Splx 1825)

/O",,,,,i,, j My'" 1991 "11,,, (Linnae", 1758) ,,,,,10m (Linna,", 1758)

·1903

~1'I"k' i seh~m¡.dl.193Z ~mg ~ r I i (Peters 1862) G. seiJ,!/ jei Donoso-Barros 1966

G. /(mim' Roze 1963 G. "ill(¡HU (Lichlcnslcin 1856 Hcmitlm_'f\'lus m{/bOlú(l (Moreau de Jonnes 1818) H. )(¡I(lje/,/lltlS Kluge 1969 Le úclobleplUlrjs mon/ecanocnsis Markezich and Taphorn 1994 Ph 'Ilodac/vlu s c/l.'(Olli Rivero-Blanco and Lancini 1967 P. ,·elHmlis O·Shau~hnessv 1875 PseutlogOltalOdes guianensis Parker 1935 P. /wlUJCllus (Roux 1927) S/)lwcnxülCly/us mo/ei Boe((&~r 1894 TheclU/o,·' 'IrIS mpictllu/us (Hou((uYIl 1782)

COR YTOPHANIDAE fJ(uiJiscus 1J(!.~ilisnj.\ (Lillllaeus 1758) I

IGUANIDAE 1 'UCII'" i.el/mw (Linnacus 1758) I I

POL YCHROTIDAE DaCI ·joofrenaUl (Cope 1899) /J. ·{ICl"C (Boulelll!cr 1903) D. "iKm )llIZela/(¡ WilIiarns 1974) /J. )/IIIe/a/(/ (Daudin 1802 /J. rOl/ud (Bonnaterrc 1789 D. squClmuJaw (PClcrs 1863) /J. ,iR'ilw (PClcrs 1863) D. /rClIIsversa/is (Dumeril 185 J) Noro)s wmeClens (Williams 1974) N. Cl/lrOIllS Daudin 1802

I

I

¡ 1903) h"b""i La Marca and 1990 , , (C"y 1846

11 I I Dunn 1 """i Nledcn 191'

, ",i<ulala 1881 Donnel1y.

',"/ni (Bun and Bun 193 J) " and Mye" 1992

""''''gil Van Lidlh de J .. "le 1904 ¡¡",hi" bieoloc (Cope 1896)

lB. , 789) 18. " , 1 and Dlxon 1977 I 8. 1"'''0,,,, i l . , 1856 lB., (Daudln 180Z)

" 0"""''' W,gloc 183'

lE. lE.

IN. IN. IN.

., (Pele" 1862) ,,,,I,,i MvelS and lonnellv 1996

Umcini 1968 1 lIS c,,·pru,~ 1 1, Cole. and

\IJCciOSI/J 1 II 1861 , , Cea'" 1958

I 1 I (MuHer 192~

., (Linna"" 1758) m""i.,i Ro," 1958 n,di> 1 . 1900 '0,,1 (Bun and Bun 1931

i (Laneiu i 1968)

1992

'" '"

P. cl'" lu¡/(Jlill e{l'u.~ García-Pércz and Yústiz 1995 P. luclUosus (Pcters 1862) PI 'e/lO IO.mu ku leri Roux [927 P. brevi(wllwli.\' (Boulenger 191 2) Río/ama leucos¡icU4S (Boulen~er [900) Tre/iosrinClU bi(asó(J/us (Dumeril 1851) T. oriximinensi,t A\fila-Pir~ 1995

TEIIDA E Amcivn ml,eim (Linnaeus 1758) A. bi rOl/lUla CODC 1862 C/lemiaoplrorlu cryprus Cole alld Dessauer 1993 C. lemnisClIIIU (Li nnaeus 1758) C. .l!Rlmil'lI/tu.\ McCrVSI(ll and Dixon 1987 C. nigricolor Pclers 1873 C,ocodi'llrlrs lacertil1llS Daudin 1802) Kentro 'YX alllmU/::,o/lica CODe 1876 K. /)Qrckimw Peters 1869 K colcara/a Spix 1825 K. pe/l'iCel)S COlle 1868 K. .""im(l (Daudin 1802) TllpillClmbis legui:C;1I (Linnaeus 1758)

SERPENTES

BOIDAE 80a cO/!SIric/Or Linnaeus 1758 CorallllS canimu (Lillnaeus 1758) C. I!OrllllWIUS (Linnaeus 1758) Epicfllles cencllfia (Linnacus 1758) Eunt'c/es murirms Llnnaeus 1758

ANIUIDAE Alliljus sc\'ulle Roze 1958

LEPTOTYPHLOPIDAE LeplO/)'pll/ops aJfillis (Boulcn,gcr 1884) L albi ro/u' (Wa~ler 1824) L allwZQllicu,f Ore'as-Miranda 1969 L dimidia/lis (Jan 186!) L >olU/vlli (Dumeril and Bibron 1854) L IIU/emle ,i.\' (Petcrs 1857) L sep/ems/rilllus (Schncidcr 1801) L lenellus Klauber 1939

ANOMALEPIDAE Helmillllw,lIisJ1aI'OlemlillcllIU' (Peters 1857) LiOl\'phlo 's alJ,iros/ris (Peters 1881)

TYPHLOPIDAE Tvph/ops brulIRersmjwws Vanwlini 1972 1·. lelllleri Roux 1926 T minuisqlUJrlHu Dixon and Hendricks 1979 T. re/icl¡/lIIus (Linnaeus 1758)

COLUBRIDAE Almclus /uulius (Boie 1827) A. t/uidellsis Roze 1961 A. elaps (C unther 1BSID

. 1 1 1

1 1 1

1 1 1

1 1

~ 1 1 f ~ 1

LA· lA. lA. lA. lA. lA. lA. lA. lA.

"' lA. lA. lA .

1911 1903

I 1845)

''''p'd"., Roze 1961 Imlcinji Roze 196i

I U,.,C,,,, 1969

""''0' Roze 1961

:~~--------------+--r~--+--r~~~1-~ 1905

" ,,,,,", I 1894 , '58)

exol"", ; 1, 58) 1",,", (Unn"u' 1158 "'0."'<010 Roze 1952

I W,lke' 1943 .~Cllml lus (WaRler 1824)

Cld'u ""'a (D,ud'" 1803) (Schle.eI 1837)

D. "ud",Ji, (Pete" 18641

D'p"" '"".""" lSentzen 1796) D. <ope' (Gü" ther 1872) D. D.

D. D. ,

1905) pm'o" ',,u Sehle.el 1837

(A lemán 1953) ,',,,'ego," (Dumer'l, Blbron. and Dmner'l 1854)

, CO"'" (Bo'e 182, , , . , (pete" 186,

, ' (Linnaeu, 1758) , 1863

, Roze 1959

H,"",,,., ""''''0'''.' Linr'aeus 1758) H, haRlluwlli ~oux 1910 H. lrog" Lanc·n' 1964 H. Sehle.el 1837

H. """",0' Shrev, 1934 i ' . "",,,,, · ,n II

'd"""., (Hemnarm 1804) I Hwlrop, (Wagler 1824)

,ce"ch"" (~8) le""fen" ¡Cope-¡S9,j I 1 (COlle ! 860)

" '(Unn"u, F85) bak,,' (Ruthven 1936)

, 1859) , , (G"nlher 1859)

Li"""" b,,,""'" Cope 1860 CO~II", (U~~~~~ ____________ ~ __ +-1--+~ __ +-1--+~ "",oru, [Unnaeu, 1758)

· (~~i~5) ''''"a, (Unnaeu, 758)

" ,0mm~M"" 1991 "phi .. " ' ~Im,,,u, wll/l"",,1 (!lo" 1958)

, , " "~'bm,,, ",d Ou .. "" 1854 , ,,,,,,,,,,,11 (S,u,,' M, I 1 ¡ Raddi [820 M, lI"dd"",1 IS"",zeo 1796) M, pi,,; 10um" ,Slbcon, aod Oum,," 1854) Nillio O/rata (Hallowell 18· 15)

OnDeI;, "''''''' Waglm 24 O, h,"'d,,, iDa",lIo 180

" ",,,,la , ,,758) O, 1 "/1 ¡ DumeriL Bibron, and Dumeril 1854

, " Sh<eve 1947 h '/ "",d,,,,, Oonnelly end Mye" 1991

1', olfmll , 1823) I p, ::1 , Lloo,,", 1758)

, , , i Tm>chel lo ' , 1848) , , ,",ooal<, 1801

11', ncuwiedi )"",,,i,, Blbcon, aod O ... """ 1854) 11' , ' 1'11,,,,' 11> ( Llon"u, 1758)

I P" ",'W,' , :Cümh" l58) 11', , larbom aod ''''"al 19241 Ir ,IW"ler 1824 , , DO>'"III Aod",oo 1926 I R, , (ScoIJolI 1785) Sil"", ,,,1,,,10", ¡ 1758)

I S"ilvI<" ,mil",,,,, (Llnoaeos 1758) " , , , IB,nl/old 1846)

, " IPet", 1863) TlI/l/ilIa 1 Uoo,,", 17581

, Dume"" Slbrm" a,," Dumeril 1854) , , , s c!Jim(1/1ta Roze 1958 1'1', ' , ,Gormlaao~ 1995

,I"id" M,e" aod Ooonelly , '1, Gormla aod 1995

Ir 1,,,lIid,,,, (Llnoaeus 1758) stri.~¡lh 1787) 1m'l j Do"o,lIv 1996

" , n' (Dauc!in 1803) " ''"""",,, Roze 1803

IV",II,,,,,, , , ~ X"""lIm

,~e\'!>rllS 'Linnaeus 1758: , ,IDaudln 1803)

I El APIDAF , , ,mil",,, ISchleoci 18371

I A1I"""" "",,,,,11, (Dume~ Slbroo 1854) 1M. 1, I . Cope ,O) 1M. e/uIII."lli (Jan 1858) 1M,

";"""", ;~~;;,~ I AL 1M, , • 1 u (Linnaeu 1758) 1M, 1, Roze 198~ 1M , i , (D",n"rl , BlbrO;L "," O",neril 1854 ,

M. remv(w; Roze J 987 I -L M. E!!xi W~ler 1824 -L -L lit surinamensi,) (Cuvicr 1817) 1 1 VIPERIDAE Bolltriechis sclll~Ii~Bef(hold 1846) Borhriopsi.\ bilillca/(¡ (Wied 1821) R lI!ei!usa Stcrnfeld 1920 iI faelliaw (Wagler 1824) Ho/ llm ,.~ as 'er (Gasman 1883) B, (llrox (Linnacus J 758) R bru:ili Hoge 1953 iI cll!0:..d(w Salldner-Montilla 1976 B iS(lbdac Sandner-Montilla 1979 R \'enewi!lellsis Sandner-Montilla 1952 CrowJu.~J.?i. WlOnm l Sandner-Montilla 1980 e duri.ulIs Linnaeus 1758 LtlcllC.~js IIlU/(I (Linnaeus 1766) PorthidiUII! /wlSbe,- ·ji (Sdl~el 1841)

Caiman crocodilus. This large crocodile is common in masl waters in the lowlands of Venezuela and other surrounding countries. Venezuela , Apure.

-L -L J

CLASS AMPHIBIA

'" co

REPTILlA

APPENOIX 2: CHECKLlST OF ENOANGEREO VENEZUELAN AMPHIBIANS ANO REPTILES ORDER FAM ILY SPECIES

Anura Al10 hrynidae

AIIQ JI 'Ile rwhl'('/I¡

Bufonidae Arelo us cllrlxmerell.'ii.l

A. cruci er

A. mucuhu'iellsis

A. ox~'rh)'nduu

A. sorianoj · Oreo lIn'/leIla IlUberi

o. maeelme/Ji

Cenlrolcnidae Ce'llrole/!f~ altiwd¡,,,,lis· C. l/m/ilrus · Hval;IIQfJ(/fr(lc/l;U/II durwui * H. /t'I/rolil!Clltwll*

Delldrobatidae ColO.tletlws ca lllr¡II/:! II$;S'"

C. /wmili s

C. leo wrda/is

C. ma/Uldorum

C. sal/ue/lSis MOn/1O hryne col/aris

M. corllillerWlu* N, heloooles albo U1Wlu ~

N. (/u r(lIIl i

N. /wydec(H!

N. mavor tÚ

N. lIIeridel l.~il·

N. molillll rij *

N. oros/oma

N. serrll/IIIS

Hylidae

G(lslrorheCClllice or;-Hyla 'u/mi

H. /,m:ill;ll

H. meridellsis*

H. la -dac/y/a Le todact lidae

Cerato hr\'s calcarulll

C. co"ulla Le "odacl\'!r1S lab ',iluhiel/s· Elew/ler()(lactrllls ~ille)i

E. lauci/úi E. Hlmmem$

Pseudo alutlico/" J!ul/I,,· Caudata

Plelhodont idae Boliro ·loHa borlmrata

B. ores/es

Tesludines Pelomedusidae

DermochelYidae

Chelonidae

Crocod lia Crocodylidae

Alli atoridae

·AdditiOI1$ 10 ¡he previolls repon lO I)APT"'SSCtUCN

Podoorclllis ex lWISCI

p, lewwmll ·

Dennoche/\,$ corillcell

Cllrellll curerra ClrelO/tia 1Il)'das frellllochd)'.~ imbricllW

CrocoddllS llCIIIUS

C. illlemretliu.\

Caillllllr ni er'·

Plate 51 Plate 52

PI ate captions: 47. The Venezuelan Andes. The versanls of Ihe mountains are covered wilh lush foresls , mainly of cloud ty~; al Ihe top, the páramo develops. The Bolívar Peak, covered by ice, is the highest point in the country. Venezuela, Ménda. 48. Coasta1 Range in north-central Venezuela. The northern tradewinds dry u~ the sea-lacing environments producing a semixeric or xeric ecosyslem. Venezuela, Aragua. 49. The Llanos of Venezuela is one of the largest ana more homogeneous biomes of the country. These f1atlands develop between the Andes and Ihe Coastal Range lo the north, and the Apure-Orinoco rivers to the soulh. Venezuera, Bannas. 50. In the Venezuelan Guayana appears the tepuis , isola1es tabletop mountains, Ihal dominated Ihe landscape. The lower lands are covered by prairies, as the Gran Sabana, or by forests, as the largest part of the states of Bolivar and Amazonas. Venezuela, Bolívar. 51 . A view of the inside of San Eusebio claud foresttin Ihe Venezuelan Andes. Tree ferns and bromelids are characteristics. Venezuela, Mérída. 52. Where dense clouds caming from !he aribbean Sea hit the mauntains, a dense claud tores! appears at the upper parts af the eaastal Range. One af Ihe best known is Rancho Grande, on the road fram Maracay lo Ocumare. Venezuela, Aragua.

Become a member of Asiatic Herpetological Research

Volume 8' 1999

Edi,,,,

hmiZh,..,

(]",,,~J" 1"";'0'" of 8","'~). A"adern," ~",,<a. o.."~J<l, S,eh".n. China

"<llar A •• umn

l,c~'¡' & ('1", . C<~ksc. P(~,I."d. Ore1",n. l,;SA

J.II ........ \1 ..... ) Ikp"n","", "flh ,loH. W • .Jimgw" Ln¡"",,,). SI. Lv.;'_ Mi,,,,.,,i_USA

Ilcr.d<~_ Cal if...-m" USA

Kr.i. ,'~ ....

.:di,,,,ialll< .. ,d

.~",nlJI

n~" .. 111 ' '' '''';'1.1''''''., No,. Y",l. . L~_,

N.,.~" ". ~ ... nj<, ~ 7-<><>1'1'",1,.""",,.\,_ Ptt<"N'~_ Ru" ••

H'hu'U"n A",,,,,~,",". I I m"""l ..... uhu_ ·'"""'.0"".

1'",,_ """",,1 (<>I"r.:. 1l'"~/"'~'. 7""",,~,n,", I'H"'·t Ll Y;"", S,...,,,, en""", V ... ",. ,!u. ..... J. (l"n,

1 no"",," Qf"<,-:.! •. J ... , h~",. ~".-,b. U,,,

H~"'" \\. ""'1'h) W"pJ 00""., M""um, T,,,, .. ..,,o.,,,,,,,. C .......

"i~ .... " ,"'.' 1-"""",,<,1 In,,,m,,,." r"",,,,,,,_ W."" U;,t ....... hiO""

OI.,".~ ... J"","

I·j;."" eh..... J."", r. J' ....... ,",,,,"", ,,1 ~br;"" U.,""'" N"1Q1lal T.,~ .. {)o."'" Lo,,~,,"), L""_~,,,) "'L' •• ,r, .. ",,, 11<,"'<), l'.'''"'n,,, ' '" K«I .. ". T .. ~",C!o",.

" y. 1"",,>1.) _óIoSl>afi¡ 7"'~'>rK'"II~""",,. s" """'''"''t¡:. R",,. l'n''''''''1 "1 'iflohMl lJoh< .... , Km .... ". 1,...,

Ibj;"" ru~.d. I u .. "'", " .... \e"",u,¡s.",,,,~OO , H , ,,,ho-,.m.k,, ~ )<."_J",,,," ,,,,,.,,,,, l.""",,, ,. II",.,~), ~""¡""H. Son~', O<"f'"

S" ... ,,,o,m, e .rl ló'R, '<h ..... h \\er ... ' Un",,, ,, ,), ,,f \h d.~.", ""O ~.toc.-, '''''' .. pn, 1 <;~ 1l,1>«~ Loo"""),, J,,",,",,",,, 1" .. 1

11._ .. t-,l"""" K.n_'~n. ZI>aB hdd .'I" ",uln, '--11~'.¡:<>, 111 ,...,." l'S~ Su,"' ... R .. I~'.) T,,,,-,,,,,, (,uI .. ~" Su,hc.I. Jo ... ". CII,o,

Mi", ... 1I"¡>tIOW¡:"'~/ fI,,,,,,,(1t ¡, I"'hl,""",d h) ,i>< A"al", 1l<<p<",lo~"-al R<",,",eh Soc""r (AIlRSI."d thc Ch",c,", SO· C'''' } 1", ,he S'ud} of Amph,b¡.n< ."" M'p"lt, ,('SS,\!!.)., ,~< Mu~"'" 01 V~"cbt .. ~ ".<><,k'l!). I lo've .... "y uf C.hlom .. , 'Ihe ,oJ u"" {'Il""""'¡;<: .u'h" .... f'",n .11 ""uoU .. , lO >u!:>m;! '<I,d", <oncemmg bul ,"" limll..!'o A,i.n h<,I"'~)~'I!Y, AII COI"'""",· ,1<""" ,~¡"id< .,1 Chm,.nd "'''1 '.'''' r", ,ub",~¡[>I¡on ,1It .. 1<.I bt: "<,nI',, AH!!., ~f",,-u,o ,,¡ V.","" .. o b'-'",H, UO'''"'''!)' "f e, l¡ r".-,,, •• n .tI.<It),('.hf""n, •. USA 947]!:I. <><by .ma¡I 'o.'Í<Jh~".(jl"r¡;n~.h<r.~I".~du AII "'If""p<!ntkn« ""hi"C)Oi­na ,hoold be "'"' '" Enni 7.hao , Ed¡,o.-, CII<"gd. In",,",c '" Iholog), ",O, 11", 416. Ci><nSdll, S",hu,n f'm","co, Ch,,,,, Au_ !hol'> ,hot,1d co" , ull G""ülóne> F" Mm'kHTIf" I'rt'f"'""i",, a~,/ Sub",i""", .. 111< <.01 "flho< ,''U, "uhs<'rip'ion onU ,,,,'mb,,,,,hj~ n. ~25 1"" y~" (545 fo. l, bTon.,) , Po".so """"leof!h< liSA and Ch,"a_ pleo", aJ,J S~ 1"" '"'''' fo.-<un",-. ma,1 '" SIO 1'<' ¡,_,uc 1", .i, m.,), .\1 ""',' .h",h '" "")"<~ oro.", pa).b!c- '" US ,",re",) 'o ¡\HRS, If )"", do "'" h",'. a<c<"

,,, US ('''''"O"~, pie.", no"t)' U< .... <1 we ",11 "",l.o "'her aml .. ~<m<"'"

",""o< 11"I"wl,,~i<~1 N~ ... ",..~ Vol"m< 8",,'c~<d, Vol"",,, 7 publi,hc<l in 1997, Vol, """bli,h<d in 1?9~. Vol 5 ""b l"heJ in I"'J>, Vul 4 p"bl ,,,,.d In 1?9!. Vo! ~ [>\'hI"l>c" 'o 1'J90 .nJ C~ilf"~ 1I",../OlDgkg / fI~"GT<h Vol, 2, "h",h ~ .. , p.b 1,<J>.:d" lhe MU"-'um of V<neb.-a", 7..oo1o~)'_ 1988 19I19 o>'~ "",m.1 f". ,11< Ch,".", &""-'y f,~ ,he S'uJy ,,/ ... ,oph,h",,-., anJ !!.cptilr., Voh",,,, 2 "",,,,,,-.:I<;.! t'hiM$~ 1I~""''''¡''g;'-GI Il''~T<. 1')~7. pu(>h<~d fo,.i>< Ci><ngdu I",,,,,,,~ ~I lI,olo~)' hy ,he CI"ng~lIl~ H"",'h s.c,~n"¡¡c .,MI T<chnulog;col l.i'CTa'o,e l're". Chc",~~,n~_ Si<.-h.",. Ch,n" M'~ Iff'P"'''¡''pCG

~';" ...... 'c,,,,J p"hl",,,,,,,, ,,' ¡"oo. 1988

These articles will appear in Volume 9 - Due out in SPRING 2001

The amphibians and reptiles 01' Panay Island. Pbilippines.

The discovery of MauYelnys ¡versoni-Iike turtles at a breeding facility in Hainan Province, P.R.e.: The counlerfeil golden coin.

Lizards oftbe Northem Mongolian deserts.

A new species Q(JJqtrachfIPl!rus lToot nortb_tem China.

A new species ofEremlas from !he Highlands otKennanshah Province, Iran.

Asiatic Herpetological Research Museum ofVertebrate Zoology, University ofCalifornia , Berkeley, California 94720

Fax: (510) 643-7706; email : asiaherp@uclink.berkeley.edu

The B

This widely acclaimed series has

"been praised

throughout the world.

st of Its Kind

_oIR.,.--oI"'-oIf'lorio1o

IW THE SNAKES

HandbJoks of Reptiles and Amphibians of Florida by Ray E. Ashton Jr. and Patricia Sawyer Ashton

I'art 1 ...1nal:1M This series is comprised of 560 pazes

Part2

part 3

ordC~':;:bcr: WW· 033X ......... ... $ 18.95 and more than 300 color photogra phs and inc1udes

~, 1ürtles &; Crocodilians Order NUI"bcr: \VW· 03S, ............. $ 18.95

The Amphibians Order Number: WW· 0372 ............. $18 .95

deseriptions of specific species, as well as ranze charts and infonnatiol1 011 their habitats. Detailed drawinzs emphasis characteristics that aid in

identification. A must for al\lateur and professional herpetologists alike.

Dcvclopcd in cooperal iol1 wilh Ashton Biodiversity Research & Preservatian Institute, Ine.

www.goecitics.col11 / ash 1011 biodiversi ty

Windward Publishing A Division 01 Finney Company

, --. - . ,, " . . . -. .

. .