•
Pseudoboa neuweiddii, Localion: 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
Distribution, species-richness, endemism, and conservation of Venezuela n amphibians and reptiles
JAIME E. PÉFAUR,,2AND 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 macrodistribution of amphibian and reptile taxa, and compares the species-richness of the various biogeographical zones in \\'hich Venezuela is herein divided. The macrodistribution 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 (Maracaibo 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-richness 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 conserva tion, tu rtles a nd c rocodiles are the most threatened species, but sorne frogs, particularly those of the highlands, 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 preserv ing its valuabJe her petofauna populations, and other animal resources, for future generations.
ReSumen.~La macrodistribución de 252 especies de anfibios y 299 especies de reptiles fue establecida para las ocho regiones biogeográficas en las que Venezuela se divide corrientemente. Sobre la base de la presencia o ausencia de especies 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 sobresalen con un índice de 0.34. Se presta una atención particular a las especies endémicas a la luz del actual conocimiento de la distribución geográfica de cada taxón. Hay 181 especies de anfibios y 11 9 especies de reptiles consideradas como endémicas en Venezuela. En términos de conservación, las tortugas y los cocodrilos son los grupos más amenazados, pero algunos anfibios, 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 especies 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 sula, and along the coast, as well. The terreslrial boundaries of The great variety of ecosystems in South America has induced Venezuela are continuous with Colombia on the west and south· the development ofa greatly di versifíed herpetofauna which is west, Guyana on the east, and Brazil on lhe south. The existence about a fo urth of th e total number of amphibian and reptile ofmany large rivers and mountain ranges help make the country species in the world. an cnvironmental mosaic with a diverse rauna of"amphibiansand Venezuela is one of th e 13 countries comprising South reptiles (Fig. 1). America. It Hes in its northemmost portion and receives the This rcport assesscs the macrodistribulion of am phibian c\imatic in fluences oflhe Caribbean Sea and Atlantic Ocean. In and repti le laxa, and compares the species-richness and lhe the north and northwest it is encased by the Andean Cordillera fauni stical sim ilarity of the various biogeographical zones in which in the Mérida Andes may attain a height of 5000 m. The which the country is divided. Endemics are given panicular al· extensive savannas in the central pon ion and Ihe Amazonian tenti on in the li ght ofthe current knowledge ofthe geographical forest of the south, conlri bute 10 a complex climate with an distribution of every taxon. Remarks on the conservation of abundance ofrainfall throughout most ofthe coun try, although these faunas are discussed. there are drier areas in the region of Falcón, lhe Goaj ira Penin· EtTorts have been made by several researchers to unravel the macrodistributional pattem s ofthe di verse Venezuelan ver· 2Correspondence. Fax: (58) (74) 401286: email: pefaur@ tebrate groups. The di stributional pattem of fí shes were de ciet/s.ula. ve scribed by Mago-Leccia ( 1970); of amphi bians by Ri vero (196 1,
43 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). DlI ellman ( 1988), Frost (1985), and Durant and Díaz (1996), and Yústi¿ (1996). Man) otiler La Marca (1992): orrepti les by Roze (1966), Medem (1981. conlribut ions about lhe distriblllion 01' ord ers (i.e .. Brame 1983). Pritchard and Trebbau (1984). and Lancini ( 1986): and Wakc 1963), familics (i.c., Di xon ,lIld Hendricks 1979). and ofrnmnmals by Eiscnberg and Redford (1979) and Bodini genera (Le. , Dixon 1980; Di Bernardo 1992: Pérallr 1993; and Pérez* l-l ern imdcz (1985). SOllle comprehensive studics Scñaris el al. 1994), and/or species (i.e., Gallardo 1965. 1969; 0 11 lhe herpctofauna frol11 several arcas of Venezuela have Dixon and Michaud.1992) have been conslIlled. Additional becll provided by Staton and Di xoll ( 1977). Duellllli.lll ( 1979), referenccs can be checked in Van¿olini (1978), Ducllm8n and Il oogmoed (1979). Hoogmoed and Gorzula (1979). Rivero* Trueb (1986), La Marca (1992). Péraur ( 1992). and Ducllman l3Ianco and Dixon ( 1979). Péfaur and Díaz de Pascual ( 1982). (1995). ílmong others.
44 VENEZUELAN AMPHIBIANS AND REPTILES
VIII .0. \ .. \ 111 / ~ \ ,/ \_ - ---; IV / / ------/ / ,/,/ 11 / / V / ------/' /------/' /" --- -'1 / I VI I I \ \ "- "-. "-. "- 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 sludy, alllands aboye 500 m are considered within Ihe Alldean In order lO assess lhe distributional rccords of each species, a llnit alld ílS biola as Alldeall; lall ds below 500 m are considcred map of Venezuela wilh its hereín defined biogeographical re as piedmont. The Andes are environmenlal ly complex (Ewel giotls \Vas lI sed. A biogcographi c sketch of Venezuela is pre and Madriz 1968; Díaz el al . 1997), alld may ínclude premolllane, sented in Fig. 2. lt contain s Ihe eight biogeographical zones dry and humíd foresls, rnontane \Vel an d cloud foresls, xero· cOtllmonly acceptcd for lhe country. This ph ysiographic en phytic vall eys, hol and dry landscapes, and lhe impressive semble takcs into account mainly lhe relief, climate, and vegeta Páramos, hi ghland tundras, occurring aboye 3000 111 . This region lion (Marrero 1964; E\\Ie! and Madriz 1968; Huber and Alarcón covers about 4,200,000 ha oflh e country's surface. 1988). A1though the boundaries of each region are not well de· The Falcón·Lara region 0 1' Region 111 also borders Ihe fincd , especially in some ecological nodllles, they serve Ollr pur· northeaslern part of the Maracaibo Lake region. This land is poses wel!. Different researchers have used Ihi s framework for affccted by th e easterly drying winds oflhe Caribbean Sea and their studies (Ri vero 1963, 1964; Eisenberg and Redford 1979; exhibit mostl y a xerophyt ic landscape, catalogll ed as Bi sbal 1988; Péfallr and Ri yero 1989). prcmontanc dry shrub or dry fo res!. A large portion of Ih e The exlent of each region is shown in Table l. The land north sealíne of Ven ezuela is bordered by ¡he Coastal Range bordering Maracaibo Lake corresponds to Region 1, which is (Region IV ), made up o f forested l11ountains, w ith eleyations covered by seasonall y dry tropical forest in the north al1d by IIp to 2765 111 , and covered by premontane tropical rain and Iropicallowland rain rorest in the south . Wetlands also cover an monlane clolld loresls. The Andes and lhe Coastal Range, slopc extensive seclion ofthi s region representing more thaI1 3,500,000 down ínlo Ihe lowlan ds orthe Llanos, Regíon V, which extends ha. The sOllthern extenl of the Maracaibo Lake region is bor· 10 Ihe Orinoco Ri ver in Ihe sOll th , 10 Ihe border wi lh Colombia dered by Region 11 , corresponding lO thc elevated 1l1011n lains of on Ihe west, and eXlending lO Ihe Orinoco River delta in the lhe Cordillera de Mérida, and by lhe Cordillera de Perijá, on Ihe ea sI. The approxim ately 27,000,000 ha orlhe Llanos are cov· northwest. BOlh ran ges compríse the Andes region. For Ihi s ered by savannas or exlell sive prairies inlenningled with dry
45 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. Venezuela, 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 km ' % Number of Life Zones I Maracaibo Lake 35,000 3.88 5 11 Andes 42,000 4.66 16 111 Falcón·Lara 30,000 3.33 6 IV Coasta 1 Range 68,000 7.54 10 V Llanos 270,000 29.95 3 VI Guayana 350.000 38.82 10 VII Anw zo nas 105 ,000 I 1.65 2 VIII Islands 1,500 0.17 6 Total 901 ,500 100.00 22
Table 2. Surface of altitudinal belts in Venezuela . (Source: Ewcl and Mad riz 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 Mo ntane 4,570 0.52 Subalpi ne and alpi ne 1,270 0. 13 Total 9 01 ,500 100.00 forests ami ri vcrin e gall ery rorests. cxeecd 10% oC Ih e terrilory, yel Ihese are th e mosl eeologi The largcst biogeogra phi cal region of Ve nezuela is Ih e call y di verse o Ca ll (Table 2), a situation w ilh slrong con se Guayana, Reg ioll VI, which includes about 35,000,000 ha, quences on Ihe biologieal featurcs oflhe country. mas! of whi ch are part of Ih e Venezuelan Guayana shi eld . Sevcral melhods have been used lO oblain Ih e dala pre The landseape is compl ex, includi ng lowland, premonlane senl ed in th is paper: museum record s, bibliographic informa and Ill onlane wet and rain forests; savannas; and wet oases, lion, and personal observati ons. Reports made on the hold call ed moricha/es. rich in palms and herpetofauna. The la nd· ings ofrelevanl museul1l s in Ve nezucla, Pucrto Rico, Colom· scape is defincd by profound vall eys bordered by fepuys, bia, Brazil, Franee, and Ihe United States have been taken lable mountain remains of gcolog ical lands of the Precam· il1l o accounl, bul in mosl instances Ihey have been im proved brian·Paleozoic era. T he area belonging lO the O ri noco w ith notes taken by Ihe aulhors during their Irips and mu Ri ver 's delta is also incllldcd in th is region. Some 10,500,000 sellm visits. In order 10 updale previolls reporls on the Ven ha of tropical forests covering lhe lowlands of lhe lI pper ezuelan amphibians and repti les (Péfaur 1992: Péfaur and Orinoco River basi n compri sc Region VI I, Amazonas, which Rivero 1989), !le", lists had lO be produccd. To the best o f is continuolls with South America's largc green con:, th e om knowledge Ihe ehecklisl database (Appcndix 1) conlains Amazonia. An important part o fthis arca is eovcrcd by wet a l! known spccies tlp lO Deccmber 31, 1996. tropical fores!. The di stribllti onal aspecls of Ihe lisl are givcn by the A smal l portion oflh e counlry is made up of islands. AII presence or abscnee of dala fo r every species in a biogco ofthclll are inclllded in Region VIII, which for the purpose of graphieal region. Thc eight regions were Ihus dcli11liled by Ihe Ihe distribulion ofturt les also in cludes the Venezuelan Carib· known inforlllation rcgarding Ihe species' geographical di stri bean Sea. This region is ineluded lo call att ention to Ih e par· bUl io n. The number ofspeeies present in a region is an ind i lieular di st ribulion oflhe island herpelofauna, especially Ihal calion ofils speeies diversily and oflhe geological, climato ofl sla Margarita, as well as marilime herpelOfauna. logical, ami biological evollllion oflhe faunal elemenls con Each ofthese biogeographical regions is eeologicall y di cerned. However, Ih e index of spceies·richncss. ralhcr than verse and complex. In th eir ecological map, Ewel and Madriz the simple number 0 1" species, is a better expression of evolu ( 1968) deseribed 22 life zones for Venezuela (Table 1). The lionary Irends. A Species-Riehness Index (SR I) \Vas caleu mosl hOll1ogeneolls biogeographi cal region, Amazonas, has lated by consid ering Ihe number of cxtant spccies divided by on ly I\VO li fe zones, and lh e most complcx o f all, Ihe Andes. Ihe area multip li ed by (x) 100. To oblain Ihe faullistieal si mi · includes 161ife zones in its relative ly small lerr ilOry. Diver larilY bel\\leen regions, a four-fold eonlingeney table \\ as cre· s ity 01" eeosyslems is proportional lo the Sleepness 01" ver· aled for every pai r 01" compared rcgions and th e Di ce coeffi sants, Ihe sides o f Ihe mounlains, in the Iropies, bul most of cient , as showll by Hayek (1994), \Vas calelllalcd. Venezue la is lowland. Abollt three·fourths of the eountry is Endemi sm. an ecological tcrm , was determined by find· eompri sed o f lands be low 500 111 ; la nds aboye 1000 m do not ing whelher Ihere \Vas a uniqlle relalionship betwcen a spe·
47 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 Frogs and toads 10 42 238 Ca ud ata Salamanders 2 GymJl ophiona Caec il ians 3 7 12 Subtota l 14 50 252 Reptilia Tesludines Tunles and lOrlOises 7 14 23 Crocodylia Crocodi les and all igators 2 3 6 Am ph isbacnia Amphisbacnians 1 2 6 Lace rtilia Li za rd s 8 37 11 3 Serpcntcs Snakes 8 56 151 Sublolal 26 112 299 Total 40 162 551
Table 4. Allocation 01 amphibian and reptile species in the biogeogra ph ical regions of Ve nezuela.
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 Alllphis baenians 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 herpetofau na l species in the biogeographical reg io ns of Venezuela. SR I = (Species number/area) x 100. The area to calcu late the Index is pro vided in Table 1.
Biogeographic Amphibian species Reptile species Total Total region n SRI n SRI n SRI 1 20 0.05 72 0.20 92 0.26 11 75 0.18 67 0. 16 142 0.34 111 18 0.06 78 0.26 96 0.32 IV 66 0. 10 124 0. 18 190 0.28 V 36 0.01 75 0.03 111 0.04 VI 134 0.04 176 0.05 3 10 0.09 VII 55 0.05 130 0.1 2 185 0.18 Total 252 0.03 299 0.03 551 0.06
cies and a gcographical region. As used in thi s study, when is, species t!xte nd ing beyond Ihe boundaries 10 sorn e neigh ever a species dwelled and apparenlly o ri ginatcd in a sing le boring counlries, such as Colombia, Brazil , and/or Guyana in region, il \Vas considered a biological endemic. However, ira the mainland, or Trin idad-Túbagú. and lhe Dutch Islands spec ies \Vas considered endemic beca use it occupied a single (Bonaire, Aruba, C uracao), in Ihe Caribbean Sea. regíon in Venezuela bul al so extended into another counlry. il Tú determine the conservalion status of Ihe herpe was considered as apolifical endemic \V ith respect lo Ih e firsl lofauna, only a few quantitative assessmenls are available. country. In Ihe case ofVenezucla, ¡here are mainly biological Thus, a general impression rather than an accuralc ccnsus endemics. bul Ih ere are 48 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 biogeographical regions. Note: This matrix shows in bold the amount of species 01 every regían. Above the diagonal line are the sim ilarity 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 11 14 0.45 0.25 0.28 1 13 66 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 0.4 7 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 en demism in the herpetofa una 01 the biogeographic regions of Venezuela. (Sourcc Data: Appendix l.) Amphibians Reptiles Regions Species by region Endemics % Species by region Endemics % 1 20 4 2.2 72 12 10.1 11 75 57 31.5 67 20 16.8 111 18 O 0.0 78 5 4.2 IV 66 32 17.7 124 19 16.0 V 36 I 0.6 75 2 1. 7 VI 134 75 41.4 176 48 40.3 VII 55 12 6.6 130 13 10.9 Country's total 252 181 60.3 299 119 39.7 Resu lts toises have 23 species contained within seven families . C rocodi les and alligators have six species belonging to !wo Taxonomic composition families. The arnphishaenian s are represented by onl)' !\vo Thc herpctological component of the Venezuelan fauna con genera and six species. The lizards have 11 3 species com sists of 55 1 species of which 252 are amphibians and 299 are prisi ng eight families, \Vith Gonalodes as its most specious reptiles. A taxonomic li st of species is provided by a country genus, with 13 species. Snakes make up the most diversi checklist (Appendix 1) and sUITImari zed in Table 3. The am fied group with 151 species belong ing to eight families. lis phibians of Venezuela are included in !hree orders, of which larges! ramily, the Colubridae, contains 104 species. A H !he larges! is Anura. The mos! specious I"amil)' is Hy lidae, other fam ilies 01" Ihis class have less than 40 species each. contai ning 77 species, rollowed by Leptodaclylidae with 72 It s mos! speciolls genus is Atrae/us with 16 species; other species. Olher ralher large ramilies are Buronidae, quite large genera are Liophis and Micrurus \\'ilh 10 and 12 Centrolenidae, and Dendrobatidae. The res! orlhe frog rami lies species, respectivel)'. have a ver)' low numher 01' species. The ramilies 01' salamanders and caecilians are also of minor numher. Distribution The reptiles are comprised within flve orders orwhich The di slrihution 01' every species in lhe eigh! biogeographical Ihe largest are Serpentes and Lacertilia. Turtles and tor- regions, as considered in Appendix 1, is sUITImarized in Table 4. 50 VENEZUELAN AMPHIBIANS ANO REPTILES By far, lhe largest assemb lagc of amphibians alld reptile 4 /-Iy/a) , one AlIophrynidae ( 1 A/lophryne), seven species is fOlllld in Regían VI, Guayana, foll o\-ved by Ihe Coastal LeplOdactyl idae (2 Cerarophrys, 3 E/eurherodacrylus, 1 Range. Amazonas, and lh e Andes. The lowest number 01' spe Leplodactylus, 1 Pselldopaludieola). and two Plethodontidae (2 cíes in the country is found in the northern regions ofMaracaibo Bolitog/ossa) are reponed as having a decl ine in lheir popu lation Lake and Falcón-Lara. In the rnar il imc islands regían Lhefe are 50 nUlllbers and th us species of special concern. Alllong reptiles species of amphibians and reptiles. there are severa! spccies on the vcrge of cxt inction, such as CroeodY/lIs inrermedius. C. aeurus. Caiman nigef: and Species-richness Podocflemis expansa, as well as all the sea turtles. As surface area differences among Ihe biogeograph ical divisions do no! allo\\' a direct comparison 01' lhe biodiversity by Ihe Discussion number of herpctofaunal species alone, an index (S RI ) is re qu ircd lo more accuratel y expreS$ the results. Th e SR! index Taxonomic composition val ucs are showll in Table 5. lhe taxonomy of th e Venezuclan herpctofauna changes con The country ¡Ise lf has a lo\\' valuc for Ih e index, 0.06. linuously due to new research findings and systematic rear I-I owcvcr, the four smaltest region s ac hievc lhe hi ghest SRI and rangemcn ts. Sllbstantial changes in the number ofknown spe lhe opposile three largest region s have lh e lowesl indexes. cies for every region have taken place in Venezuela during recenl years. Descriplions of numerous new taxa ha ve occurred Faunistic similarity in the last decades mainly amollg am ph ibians (Péfaur 1985; lhe herpelological simi larity betwecn the different biogeographi Rivero 1982a, b, 1985; Ayarzagüena 1992 , a1ll ong others). cal regions ofVenezuela is presented in lable 6 and 7. Simi larity Importan! changes ha ve recen tI y been inlrodueed to the tax bascd on amphibian spec ies is rather 10w(Table 6); th e least pair onomy ofthe country's herpetofauna lhus. changing the sys bound of regions is the An des-Guayana, which share 12 species tematic scenery. Among frogs, lhe Centrolenid family has been and have a similarity va luc of 0.1 1; the strongest bound is lhe divided inlo several genera (Cenrro/ene. Cochranella. and Ll anos-Amazonas pair, which ha ve 2 1 specics, with a simi larity Hyalinobatrachium) by Ru íz-Carranza and Lynch (1 991 ). The va lue of 0.46, füll owed by the Llanos-Coastal Range and Ll Dendrobatidae was a1so generically rea rranged with the intro anos-Maracaibo Lake pai rs. duction of Epipedobates and Minyobates by Myers (1987), Regional similarily based on Ihe rcpti le component oflhe Aromabares by Myers el al. (1991). and MannophrYl1e and fauna produces difTerent resulls (Table 7). In general , the Si111i Nephe/abares by La Marca (1995). lhe Hylidae has also suf larity n:gion based on reptile species has higher va lues than fered some modification wilh lhe rearrangcment of so me when amphibians are considered. Here, the leasl similar pair Olo/ygon il1to the resurrected gel1us ScinQx (Duellman and Wiens region is Ihe An des-Amazonas, with a similarity value ofO.24, 1992), and the elaboration of a new genus Tepuif1yla by whil e the strongest bounded pair is Gllayana-Amazonas, wilh Ayarzagüena el al. ( 1992b). Amo ng the caecilians. a general 110 shared species and a si milarity val ue ofO. 72. laxonomic rearrangement offamilies and genera was proposed by Nussbaum and Wilkinsoll ( 1989) and Wilkinso l1 (1996). Endemism i\mong the repti les, there have also been some taxonomie There are 181 spec ies ofamphibians and 119 ofrepl iles consid changes. Within th e lizards, Iguan idae was divided i!lto several ered lo be biologica/ andlor po/irica/ endemics (Table 8). Overal1 farnilies (Corytophanidae, Iguanidae, Pol ychrotidae, and there are more endemic speeies alllong the amphibians Ihan among Tropiduridae) by Frosl and Etheridge (1989), while Ihe genus reptiles. However, four regions have less endemic amph ibi an s Anolis was di vided into five genera, ofwhich IWO are present in Ih an endcmic rept il es. lhe Andes, the Coasta l Ran ge, and Ihe Venezuela, Dacry/oa and Norops (Savage and Guyer 1989). lhe Guayana are lhe places with more endemics in bOlh taxa. Teiidae was also divided into two units. Gymnophlhalmidae (srna 11 teiids) and Tei id ae [macroteiids] (Presch 1980). Within Conservation the snake group, sOln e eolubrid genera have been revised modi Actual data on Ihe conserval ion status of lh e herpelofauna is fying the taxonorn y for the Venezuelan members; for instan ce, relatively scarce in Venezue la. Th e most relevant infonnation on Dixon (1989) rev icwed Liop/¡is and other associated genera. population status is provi ded by Ramo ( 1982), Praderio (1985), Among the Crotal idae there was al so a slrong change with the Si lva et al. ( 1985), Péfaur and Díaz de Pascual ( 1987), Péfaur et partitioning of Borhrops into several genera: Borhriechis, al. (1987), Silva and Val déz ( 1989), La Marca and Rein thaler Both,.iopsis, Bothrops. and PorrhidiufIl, with considerable ctTeel (1991), Thorbjarnarson ( 1991), Péfaur and Pérez ( 1995), and to the VenezueJan fauna (Campbell and Lamar 1989). Of course, Durant and Díaz (1996), alllong others. The Red Data Book of classi fication \Vi11 continue 10 change as it is a dynamic sc ience, Venezuela (Rodriguez and Rojas-Suárez 1995) \Vas helpful in and new systems and names wi11 be introduced changing subse eSlablishi ng herpetofauna endan gennent status. quent li sts of Venezuelan amphibian and repti le species. AII Th e report elaborated by the senior author to the Decl inin g systematic allocations and changes should be taken as tem po Amphi bian Populations Task Force (DAPlF) of lhe World rary arrangements Ihat \Vi iI be modi fied by lhe collection ofnew Conservali on Union [IUCNJ (in Vial and Saylor 1993), on Ihe data and in sighls by researchers. declining status of amphibians was al so used lo generale a lisl of endangered species (Appendix 2). Severa l species of amp hib Distribution ians, alllong which there are seven species of BlI tonidae (5 There is no sin gle area in Venezuela where an amphibian andlor Are/opus, 2 Oreap/1Iynel/a), four Centroleni dae (2 Centro/ene, 2 a reptile species is not present. From the most Iuxllrious tropical HyalinobatrachiulI/), 15 Dendrobalidae (5 C%stelhus, 2 wet forest of Amazonas to the vegetationally depauperate Man/1op/¡ryne, and 8 Nephe/obales), five Hylidae (1 Gasrrorheca, Páramos at th e top oflhe Andean 1l1ounlains. where nllmerous 51 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. Venezuela, 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, Mérid a. 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 ricr, bUI there are rieh mieroclimates thal aC I as ecological and Dixon 1979; Duran! and Diaz 1996; Díaz el al. 1997), refuges. In lhe Coastal Range region, ho\Vever, there are at species abound; howcvcr, snakes have no! in vaded lhe páramos least two \Ve l1 differentiated kinds of lands: \Ve l e levated and (Péfaur and Diaz de Pascual 1982; Diaz e l al. 1997). Different xerophytic lowlands close to Ihe Caribbcan Sea (Ri vero 1964a: physiological and behavioral adjustl11cn lS are found in lhe fauna Manzanilla el al. 1995, 1996). Moreover. an import anl selec· of cach ecosyslcm, to cope wilh lhe conlrasLing climatologica l I¡ve biogeographical barrier has aC lcd in this regio n, as is the factors impingi ng on every zonc. Severa! orlhe more notewor case o f the Unare ri ver vall ey, \V here many herpetofauna spe Ihy adaptive slrategies are Ihe development ofscveral unique eies di stributions are di scontinued pasl this point, such as reproduclivc modes: developing eggs and tadpolcs in Irce cavi Colosfel/¡us mande/orum, L. ¡nsu/arum. and Eleufherodacl)'lus líes or bromeli ads. or eggs on dorstll11 o f remales, or tadpoles lerraebo/ivaris, arnong others (R ivero 1964a: La Marca 1992). carricd on dorsum of males, such as OCC Uf in 31lUfallS (Ducllman In Ih e Ll anos (Regio n V) the re exists a fai d y large her 1985), or Ihe use ofcoll ective nest dcposits under rocks, as pctologieal fa una, which is comOlon wilh Ihe olhe r rcgions occurs in th e li zards ofthe Páramos. (S laton and Dixon 1977; Rivero·B la nco and Di xon 1979; Newly discovered and new locality record s 01' species P é f ~llJr ami Díaz de Pascua l 1987). The Ll anos are c limali· have changed the known distributional pallerns fo r many eally and vcgctationally homogeneolls at Ihe rnacrogeograph 3mphibians and reptiles. For instance, in the Andean rcgion, ical level, altholl gh they are inte rming led w ilh a web of ri species lOta ls have changed fr om 16 frog, 4 lizard, anel 10 parian forest cOl1llllu ni cating with Ih e olhe r sll rrounding re· snake species (as reported by Ducllman 1979), to 56 frog and gions. Bordered by Ihe Orinoco river in the soulh, this re· 15 lizard spccics (as reported by Péfaur and Díaz de Pascual g ioll docs not separatc faunisticall y from Regions VI and 1982). lO 72 frog , 26 li zard. and 38 snake spccies repo rted in VII- the largesl Venezuelan ri ver seems 11 0t 10 be a se lec· lhis slUd y. The well·documcnted work by Lanci ni ( 1986) live bi ogeographical barrier fo r amphibians amI reptiles reports 133 snakc species for the country lO 151 species (Rivero 1961). reported here. Our knowlcdge oflhe di slribution ofthe Ven· The soulhe rn regions o f Venezue la. Guayana and ezuelan amphi bians and reptiles is quite acceptabl e al the Amazonas (Regions VI and VII) are closer associated with present I¡ me. but an in crease could be expected as new data is Ihe Amazoni.an sector ofSolllh Ame ri ca than w il h Ihe north· bein g coll ecled a lllhe time by rcscarchcrs in the field. 8 0th e rn areas orlhe country. The geological changes, Ihe climatc, Ihe Andes and Ihe Guayana rcgions have been actively ex· Ih e lOpography, and Ihe diverse vegelalio n makc these I\VO plored by groups of rescarchcrs from th e Un iversity of Los areas the ric hesl in herpelOfauna species. This is espccially Andes in Mérida and from Musco de Ciencias La Salle in true of Ih e Guayana where a mu1titude of Iwbilats faci lilalc Caracas. respectively (AyarzagUcna et a l. 1992a, b; Péfaur th e proeess 0 1' evolulion. With the cxception of salam:mdcrs 1993; Scflari s e Lal. 1994; Durant and Díaz 1996), wh ile ac· and croeodil ians, Ihe laxa are more nllmcrOll S in Ih c Guayana, ti ve research on Ih e hcrpelofauna of Ih e Coastal Range is than in any olher region. For Ih e rest oflll e groups, the I1UI11· underway by researchcrs fr om Central Uni vers ity, Museu1l1 ber is almosl dOllbte in Ihe Guayana, as comparcd to any of oflhe Agrarian Zoology Institmion in Maracay (Manzanilla the olhcr biogeographical regions. The ccological cornplexily el al. 1995, 1996). Oflhi5 regio n has played several roles in lhe evoluti o n ofthe The di slribuli onal range of certain species is of conccrn. biola . On the one hand, it has eonstiW led a selcctivc barrier There is a d rmnali c difference belween th e distribulion of je r Ihe: eXlcnsio n o f somc faunal elcments frollllhe Brazi liall SO I1lC laxa cX lcnding over Ih e whole CO Unlry (e.g .. Bllfo Amazoni an lowlands lO Ihe Llanos of Venezuela and vice marinlls. flyla crepilans) compared to lhe punctual distribu versa; o n lhe other hand, it is th e seat for l1lan y e ndemic lion ofthose lepui.associated laxa (e.g., Oreop/u)'nella huberi, species ( I-I oogmocd 1979; this study). At the same time, il o. vasqlle=i), or lhe narrow d istriblltio n of dendrobatids in has shared species with oth er regions of Venezuela and Brazil the Mérida Andes. Many examples of lhese are knov·m in any (Avila·l>ires 1995). Dne particular exceplion is Leplodaclyllls biogeographi cal region. labyril1lhicus. This specics Ii ves north and soulh 01' Ihi s re· The fasl change in the Venezuelan landscapes will ha ve a gion, Icaving a distributio na l hi atus in the Guayana shield severe impact in th e di slribulion of lhe herpclofauna. In less (PUaur and Sierra 1995). than hal f a ccntury mOSI foresls in Region I have becn cut Depcnding on the total number of species prcsent in down and replaced by prairics. Amphibian species prevail in cach rcgion, thrce eategori es of regio ns can be proposcd: a) the south. whi le reptiles dominate th e no rth. In general, most, those wilh lcss than 100 species, eomprised by the Maracaibo but mai nl y the xerophyti e fauna, is shared with Ih al o f xeric Lake, the La ra·Falcón, and Ihe Island5 regions: b) Ihose \V ilh Regio n 11 1, a relalionship cx tending to the rich and large val morc Ihan 100 bUI le ss than 200 species, which incJudes Ih~ ley of Ihe Colombian Magdalena river. The vall ey of Ihe Andes. Ih e Coasta l Range, Ll anos, and Amazonas; and c) Calatumbo river mighl have played a role in aC ling as a pass· w ith more Ihan 300 species, represent ed by Ihe Guayana way belween the faunas of Ihese regions. regíon. Mo reover, Ihe lim its ofevery biogeographical region. The existence o fmany humid environmenls in Ihe Andes are very imprecise. For instance, lhere is a seclor where Rc (Region 11 ) permils Ihe prcsence of a large numbcr of frog gions 11. 111. and V converge, and thus there is a n uncertainl)' species (Duellman 1979: Péfaur and Diaz de Pascual 1982). about Ihe region 10 which some species belo ng. AnoLher bio· This region has fe\\' biogeographícal contacts w ith olher parts geographical nodule is rhe sector wherc Regio ns 111 and IV of the COUn lr). a1though il has served as a pass\Vay for dis· mcrge. There is greater need for a more delai led analysis 01' persi ng faunas (Péfaur and Pérez 1995; Ri vero 1979; Ri vero presence species in lhese connicling seclOrs. On Ihe olher and Solano 1977). Somelhing simi lar occurs in the \Vel and hand, collecling record s are very scarce in the Cordillera de e levated lands ofRegion lV, Ihe Coaslal Range. The Andes do Pcrijá in western Ve nezue la and lhe Orinoco delta region in 11 0t have wilhin thern any importanl large geographieal bar- easte rn Venezuela. It is possible that lhe Delta mighl be con- 54 VENEZUELAN AMPHIBIANS ANO REPTILES si dered a scparalc biogeographical region when more biologi biogeographically rel evant is the high similarity between the cal inform atio n is gathered and analyzed. Llanos and the Maraeaibo l ake region. whi eh are separaled by ranges and dry lands, but that share SOIll C species \Vith disjul1ct Species-richness di striblltions such as Leplodaclyflls bolivianlls, L wagneri, The f ~ILJn i s ti c richllcss of an arca would be bcttcr cxpresscd by Physa/aefllll s pl/sll/foSIIS , and Pseudis paradoxa, 31ll 0ng oth ers. an Indcx of Species-Richncss (SRI) tha! takes into account On the olher hand, elements of Region 11 differ considerably both Ih e area and Ihe numhcr ol'specics. In cssence, Ih is index from the other arcas, an indication or the grealer amphibian is a spcc ies-density ind ex- th at ¡s, al lhe sam e number of independence o r Ihe Andes with respecl to the other regions in spccics, those biogeographical arcas wilh small surfaces \V iII VenezlIela.A simi lar pattcm is provided by the Guayana region, have hi gher values in SRI Ih an areas with larger surfaces. SR¡ wh ich exhibits an independenee as a biolie souree. nUlll hcrs allo\\' Ihe comparison 0 1' an y arca Dr region of Ihe Becallse reptile species usually have a rather ample di slri world with respec! to spccies-richness. In Ihe case ofVenezu bution, lhe nlll1lber of shared species and lhc si milarity valllcs cla, il s large size and lhe ex istence ofcxtcnsive territories, ~ u c h are higher in this grollp than in the amph ibians. The overaJl as Ih e Ll anos, w ilh a lo\\' nUlllber ofspecies, account fo r a low simi larity vallles are hi gher, especiall y between Regions VI and SRI value (0.06), similar lo Ihe one known for th e Yucatán VII , and wilh the lowest level between Ihe Andes and Amazonas. Peninsula in Mexico (Lee 1980). This stalistical artifacl warns Consideri ng th e reptiles, the Andes does not stands out as a aboul Ihe use ofsuch index for counlries with a hi gh diversity fallnislica lly independent region beeause of lhi s region's rela of landscapes. but lo use it ror part icul ar regions on slllall lively strong rclalionships lO Ihe Maracaibo Lake and Falcón• cOunlri es. For in stance, among published data, Oaxaca in Mexico Lara regions. has an SRI val ue of0.37 and is lIsually cOlls idered one of lhe richest herpelological regions in the worl d (Casas-AndrclI el Endemism al. 1996), whereas Costa Ri ca has been re ported as having Ihe The formation orne\\' animal species, as Ih e resul l of genetic and largesl SRI vallle (0.71) in the world (Johnson 1989). ecological processes, has been one of Ihe major evolut ionary [f both amphibians and repti les are analyzed together, featllres in l11al1 y arcas ofVenezuela. Whcrevcr the distribut ion the highesl SRI vallle is found in the Andes, foll owcd by the of a species is redllced in space, an endcmisl11 process is al work. Falc ón-Lara regíon. The SR I vallles o f0.34 and 0.32, respec l-I erpelOlogical enJemism is difTerent in lhe several biogeo li vely (Table 8), are dislinglli shed even when comparcd 10 Ihe graphi cal zones ofVenezuela. The Andes and the Guayana are SRI richesl regions ofthe world. When the analysis is carried the regions with the highesl pereentages of amphibian endem ics, OU I separately fo r amphibiall speeies, Ihe Andes is th e onl y most oflhem biofogical endemics, renccting an active speciat iol1 region Ihal slands out as the 1110st di verse region. This rela proeess, something Ihat has been aeknowledged in olh er animal li ve ly .small area, with several elevalional be lt s and an abun grollps (Brown et al. 1974). On one sidc, at Ih e Andes there has dance 01' hUlllidity and \Vet lire zones, has a eondensed den been an active process or contracti on and expansion offorests as si ly 0 1' amphibian species. In turn, when the rcplile fauna is consequence o rlhe glaciation periods, and on Ih e OIher side, al analyzcd scparalely, Ih e largest value is found in Regíon 11[ Ih e Guayana , therc has been an appearancc o r difTerent vegeta (Falcón-Lara), whi ch is also a small arca covcred with xero tional fonnations oriented by a long hi story of erosive changes. phylic vegetati on- a fitting pl ace for reptiles. In the Coastal Range almos! half ofthe amphibian species are biological endemics, too. The faet thal mOSI amphibian endemi cs Faunistic simUarity are found in elevated lands is evidence ravoring the close rcla Si milarity is a concept th at brings logelh cr biogcographical ele tionship between abundance of life zones and diversity of am lllenls an d evolutionary aspccts o rthe fauna. [n the comparison phibians. The Andes has 16 life zones, and both the Coasta! of the seven regions of Venezuela, all obtained val ues for am Range ami the GlIayana regions have 10 each. phi bians are under 0.50, indicaling a low degree ofspecies shared The siluatioll is diffe relll with Ihe reptil es. Though there between biogeographical regions. Thc most similar amphibian are 119 specics in Ihe country considered as biological or polili faunas are those ofRegions IV- V and V -VII- the values ortheir cal endemics, only one region has a greatesl number of endemies, similarilY indexes are lhe hi ghesl (0.4 5 and 0.46, respeclive Iy). the Guayana, with 40% oflheir total species endemi c. Reptiles It is nOI a surprise lo find th ese relationships, for th ere is a are less restri cted 10 a geographical place and have a wider eco geographi eal continllity belwecn these regions, but \Vhal is logical tol erancc, thus eX lending their distributions inlo di rrerenl Table 9. Causes 01 amphibian population decline. 1. Environmental changes due to human activity 11. Environmental changes due to astronomic factors a) Destructi on and/or fra gmcllIation o r habitats a) Global elimm ie ehanges b) Agriculture fro lltier ex pansion 1) in tcmperature pattems c) Inadcquate use ofplagui cides 2) in preei pitat ion and relative humidity pallerns d) Waler pollution b) Ultrav iolel radiation inereases e) Ozone's eover destrucl ion or weakness c) Not-ycl-evaluated factors 1) Acid rain 1) Cosmic dusl impact g} Inlrodueli on of predator an d/or competilOr spec ics 2) Mi cromeleors h) Expansion and/or introduclion of di scases 3) X- rays 4) Gamma-rays 55 Plate 34 Plate 36 Pl ate 37 Plate 38 Plate captions: 31. Geochelone carbonaría. A common an d 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 cO llrlesy 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. highcr UV radialioll levels at elevation. thlls exhibiling vu lner The Falcón-Lara and lhe Llanos llave Lhe sma llesl Tlumbers ability of a mphibi.an s pecies to this region particularly of endcmics in lhe COlllllry. These arcas ¿I fe macroenvironrnclllally (Blaustcin et al. 1994). Many causes can be responsible lor Ihe hOl1logenous, 11 0l favoring spec iation as l1111ch as in heterogeneo Ll s amphibiCln Clnd olher verlebrale's decline, bUl most can be framed clcv 57 Plate 44 Pl ate 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 fou nd in restricted parts of the An dean 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. Th is colored snake inhabits the southern lowlands. Ve nezuela, Bolívar. 44. Phylodryas viridissimus. An attractive snake distributing in the southern sta tes of the country. Venezuela, Bolívar. 45. Micrurus mipartitus. Venezuela , Guatopo. P/¡OfO cO llrtesy o/ Ul/lrie 1. ViII . 46 . Bothrops venezuefensis. A large snake usually found in the forested environments of the northern mountains. Venezuela , Tá chira. JAIME E. PÉFAUR AND JUAN A. RIVERO IS Ihe Guayana reglon. OSleor.:eplwlus rodriglle=i de las tierras altas de la Guayana In tenns of conservation, il is knowll thal Ih e more com venczolana: descripción de cinco nuevas especies. Memorias mon environmental changes, destruction and rragmentation oC de la Sociedad de Ciencias Na/urales (Caraca .~). Tomo habitats and agricu lTura! expansion , \Vil! have severe cOllsequences L II 137:\\3-\42. Ayarzagüena. J .. Sci'laris, J. e, and Gorzula, S. \992. Un nuevo for Ihe herpetofaulla in Venezuela. As many species are endemics género para las especies del "Grupo OSl eot.:ephal ll .~ and restricted lo 5m31! areas, lhe destruction of a few kilometers rodri~lIe=i" (Anura: Hylidc¡e). Memorias de la Sociedad de of Ih e habitals could eliminate several species. The rapid de Ciencias Na/m'ales (Caracas). Tomo L[I 138:213-221. struction and/or contamination ofnatural envirollments are caus Baquero de Pedret, B. and Quera de Per'la, M. [996. Manejo del in g a decline in herpetorauna po pulations and numbers so as lo programa de zoocriaderos de la especie Baba (Caiman pUl sorne species al extinclion ri sk . Mas! imperiled amphibians eroeodilll s) en Venezuela. ZOO(."l"imJe/"Os 1(1 ):0[-06. Bisbal, F. J. 1988. Impacto hum~no sobre los habilat de Venezu are ¡hose living in Ihe highlands of northern Venezuela, while the ela. Inferciencia 13(5):226-232 . mos! endangered rept il es are Ihe marine turlles, crocodi li ans, B[austein, A. R. and Wake, D. B. \990. Declining amphibian alligators, and \Un les. [1 is hoped lhal lhe existence of several populalions: a global phenomenon? Trends in Ec%JO' alltl Nat ional Parks and other protected arcas, and the increasing EVO/Ulio/l 5(7) :203-204. awarcness of the ci tizens, wou[d he lp lo preserve the Venezu Blaustein. A. R. , Hoffrnan . P. D .. Hokit. D. G., Kksccker. J. M.. clan herpetofauna. Walls. S. c., and lIays, 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/ Sci Acknowledgements.-We thank the people ofthe Animal ences 91 :1791-1795. Ecology grollp at th e Uni versidad de Los Andes and ol' the 13 odin i, R. and Pérez- Hernández, R. 1985. Proposición de Department 0 1' Biology at the Uni versidad de Puerto Rico, regiones biogcográri cas para Venezuela en base a la Mayagüez. fOI" their permanenl support lO continll e with our distribución de [os Cebidos, p. 323-333 in A Primatolog.ia herpetological stud ies in Venezuela. In particular \Ve are in no Brasil 2. Allais do ] 0 Congreso Brasileira de debted to William E. Duellman, James R. Dixon, Pedro Durant, Primaralagio. Campinas. Naney M. Sierra, AlberlO Ve loso, Jesús Manzanilla, Gustavo Brame. A. 1-1. and Wake, D. B. \963. The salamanders of Soulh A meri ea. COnlribu/iofls in Science. Na/llral HislOry Mu Casas-Andreu, César Moli na, and Ihe late Adao J. Cardoso for selllll o/ Los Angeles Cowlly 69: 1-72. their help and continuous provision 01' data and references. Brown, K . S .. 1r., Sheppard. P. M .. and Turner, R. G. 1974. Thanks are extend ed to the curators of the visited museums, Quaternary refuges in tropical America: evidence frorn race especially to Ihe Colección de Vertebrados, Universidad de rormal ion in Helicollills butternies. Proceedings of ,he Los Andes. Mérida. Venezuela; Estación Biológica Rancho Royal Socie/y o/ LOlldon-(fJ ) 18 7 :369-378. Grand e del Ministerio del Ambiente y de los Recursos Natu Campbcll, J. A. and l.amar, W. W. 1989. Tll e Venomolls Rep/ile:s rales, Maracay. Venezuela; Musco de 1-1 istoria Natural La Salle, o{ La/in Amer¡ca. Corncl1 Univers ity Press, 1thaca, Ncw York. 425 p. Caracas. Venezuela; Museo de Biología, Facultad de C iencias, Casas-Andreu, 0.. Méndez-de la Cruz. F. R., and Camarillo. 1. L. Universidad Central de Venezuela, Caracas, Venezuela; Musco \996. Anfibios y reptiles de Oaxaca: liSIa, distribución y con de Zoología Agrícola, Universidad Central de Venezuela, servación. AC/a Zoológica Mexicana (nueva serie) 69:1-35. Maracay, Venezuela; Museo de Biología, Universidad de Puerto Di Bernardo. M. 1992. Reva[idatian of the gcnus EchinaJllhera Ri co, Mayagüez, Puerto Rico; Instituto de Ciencias Naturales, Cope. 1894, and it s conceptual amp[ification (Scrpentes, Universidad Nacional, Bogotá, Colombia; In stituto de Biologia, C o lubridac). Comunicaciones del MI/ seo de Ciéneias Uni versidade Estadual de Campinas, Sao Paul o, Brazil; Mu PUCRS, S'é rie Zoologia (POrlO AfeRre) 5( 13 ):225-256. seulll ofNatural I-li story, The Univcrsity ofKansas, La\\lTenee, Diaz, A .. Péfaur. J. E., and Durant, P. \997. Ecology of South American Paramos with emphasis on the fauna of the Ven Kansas, USA; Fi eld Musculll of Natural I-li story, Chi cago, ezuelan Páramos. p. 263-310 in Wielgolaski. F. E. and Illinois, USA; Departmenl of Wi [di ife and Fi sheries Seiences, Gaoda[l, D. (editors). Ecosysrems o/ Ih e World, Volurne 3. Texas A&M Un iversity, Coll ege Station, Texas, USA; Mu Polar and Alpine Tundra . E[scvier Science, Arnslcrdarn. seum of Vertebrate Zoology, The Uni versity 01' California, Dixon, 1. R. 1980. The nealrapica! co!ubrid snake genus Liophis: Berkeley, California, USA; Museum ofComparative Zoology, lhe generic concept. Milwaukee Public Muse lll/J. Cun/ri I-Iarvard Universily, Cambridge, USA; and Museum National bu/ions in Biology and Geology 31 :1-40. d'Histoire Naturel le, Pari s, Franee. C0111ments and corrections Dixon, J. R. 1989. A kcy and chccklist 10 the neotropical snake genus Liophis with eountry lists and maps . Smi/lrsonial/ made by Jaek Sites, William W. Lamar. Craig I-I assapakis, and Herpewlogieal Informa/iol/ Service 79 : [-28. t\Vo anonymous revie\vers are deeply appreciated. Errors re- Di xon, J. R. and Hendrieks. F. S. 1979. The wortllsnakes (family 111ail1 ours. Be lk is Rivas and Yhilda Paredes were helpful in the Typhlopidae) of the Neotropics, exclusi\e of Ihe Antill\!s. preparation of previous drafts. Marisela Angelino prepared Zo%gi!}'(:he Verlwndelingen (Leiden) 173:3-39. Ihe final version orlhe manuscript. This study has been granted Dixon, J. R. and Michaud, E. J. 1992. Shaw's black-backed snakc by CDCHT-Universidad de Los Andes (C-805-96) anel (Lioplris melallotrls) (Serpent\!s: Colubridac) of northern CONIC IT (PI-O11) . South America . ./ourna/ o/ Herpelology 26(3) :250-259. Duellman, W. E. 1979. The herpctofauna of the Andes: pat terns of distribution. o rigino differentiation and prcscnt References communities, p. 371-459 in Duellman. W. E. (editor). Tire Avila-Pires, T. C. S. 1995. Lizards 01' Braz ilian Amazonia (Rep SOIlIIl American Herpelofaul1a : ilS origino evolu/ion (tl/d tilia: Squamata). Zoologisr.:he Verl/(/ndelillgell (Leiden) dispersa!. M/lselllll o/ Nalllral Hi.\·/oIY n/ ¡he UnÍl'ersilY of 299 : 1- 706. KOfl.\·as Mon()graplls 7 . Ayarzagüena . .l. 1992. Los Centro lénidos d e la Guayana Due llman, W. E. 1982. Compresión climática cuaternaria en los Venezolana. Publicaciones Amigos de Dofiana (Sevilla) Andes, p. 184-201 in E/ee/os sobre la e!}peci(l ción. AClas 1 AS p. VI [1 Congreso Latinoamericano de Zoología, Mérida. Ayarzagüena. J., Señaris. J. C .• and Gorzula, S. 1992. El grupo Ducllman, W. E. 1985. Rcproductive modes in anuran amphib- 59 VENEZUEt AN AMPHIBIANS ANO REPTILES ¡nns: phylogcnclic signi ficance 01' ndn pti"c stratcgics. SOIllIl La Marca. E. 1992. Cata logo taxonómico. biogcogrMico y AfriclIlI./o/lrl/af ofScil!llee 8 1:17.l·178. bibliogrMico de las fa nas de Venezuela. CUllderllo .~ Duc1Jman. W. F. 1988. Pattcrns of spc:cies di\'crsil} in .m uran Geográficos (Merida) 9: 1- 197. amphibians in American tropies. Al/l/ah o/ ,he Missullr; [.a Marca. E. 1995. Biological and ~)s lemati c s~nopsis 01' a BO/(lI/icfll (Jarden 75 :79- 104. gcnus 01' rrogs from northcrn mounlains 01' Soulh Amcril:u lJucllrmut. W. E. 1995. Temporal nuclUalions 3nd abundance 01' (An u ra: [)cndrobalidae: Manllop/¡ryne). BII/lelill of lite •muran umphibians in :1 scason:.1l AmuLoninn rainforcsl. Marylolld Herperoloj!ic(¡J Sude/y 3 1( 2) :40-77 . .IQllmal 01 l-terpelUlog) 29( 1): 13-21. La Marca. E. and Rcinlhaler. H. 1). 1991. Populatio n chang.cs in Ducllman. VV. E. and Trucb. L. 1986. Biology 01 Amphibiam. Alelop/ls specics 01' Ihe Cordillcra de Merida. Vcnc,mcla. McGraw-11i11 Book eo,. Nc\\' York. I/erpefOlogical Rel'iell' 22 (4): 12 5·128. Ducllman. \'./. E. and Wicns. J. J. 1992. The stalUs of ¡he 11) lid Lancini. A. R. 1986. Serpiellles dc Vel1c;lIela. Erneslo Armitano frog gl'I1US O/o/ygulI and Ihe recognition 01' ScillllX Wng ler. editor. Caracas. 1830. (}c('(lsiollaJ Papers o/I/te MlIselll1l o/ N{lllIml fli!¡· I.ce. J. C. 1980. An ccogcographical analysis 01' the hcrpcloraun:. 101J' (JI,lte Univer:;ir.\' o/ Kalls'I ,~ 15 1: 1-23. 01' the Yuca tan Peninsula. M/I.\'ellll/ of Na/lIral l1i\'IO/:\' o/ /he Duranl. P. Mi/l\'tlllk.ee PIIMi" MII.lell/ll COl1lribllliol/s ;11 JJ;O/OKY (11/(/ Péfaur. J . E. 1993. Dcscriplion or a l1e\\ Colol/elhul GeoloKY 76 : 1-66. (Dcndrobatidac) \\ ith sorne natural hislor~ cornments on 60 JAIME E. PÉFAUR ANO JUAN A . RIVERO ¡he gcnus in Venezuela. Aly'e.~ 11 (3):88-96. lion of the hcrpetol"auna of the dry lowlands regions 01' Péfuur. J. E., and Diaz de Pascual, A. 1982. Aspectos northern South America. p. 28 1-298 ill Duellman, W. E. biogcográficos de las comunidades de anfibios y saurios de (editor). Tll e Somlt American Herpetofauna: i/s origino los Andes venezolanos, p. 229-262 in Salinas. P. (ed itor). evo/u/ion and dispenal. Musellm of NalU ral Hislory of ,he Zoología Neorropical. Actas VI II C ongreso Latino V niversi/y of Kall.ws Monographs 7. americano de Zoología, Mérida. Rodríguez, J. P. and Rojas-Suárez, F. 1995. Libro rojo de la fauna Pélilur, J. E. and DiaL de Pascual, A. 1987. Distribución ewlógica venezolana. PROV/TA. Fundaciól/ Polar, Caracas. y variación temporal de los anfibios del Estado Harinas, Ven Rozc, J. 1966. Taxollomíll y =oogeogralía de los ofidios de ezuela. Revista de Ecología L(lr¡noamericana I (3-4):9- 19. Vel/ e=ueJa. Editorial Biblioteca, Un iversidad Central Ven Pda ur, J. E. and Ilérez. R. 1995. Zoogcografía y variación ezuela, Caracas. 362 p. espacial y temporal de algunos vertebrados cpigcos de la Rui /.-Carranza, P. M. and Lynch, J . D. 1991. Ranas Cenlrolcnidae zona xerófila de la cuenca media del rí o Charna, Merida, de Colombia 1. Propuesta de una nueva clasificación genérica. Venezuel a. Eco/rópicos 8( 1): 15-35. Lozania (Bogo/á) 57:30 p. Péfaur. J. E. and Sierra, N. M . 1995. Status of Lep/odac/yius Savage, J. M. and Guycr. C. [989. Infrageneric classification /abyrimhiclls (Calf Frog, RWIlI lemero) in Venezuela. Her and spceies composition 01" the anole genera, AIIOlis, pelOlogical Reviell' 26(3): 124-127. C/el/Oll/s, Dael)'loa. Norops a nd Semillrl/s (Sauria: p¿raur, J. E. and Rivero, J. A. 1989 . B iogeograna de la Iguanidae). Amphibia-Rep/i1ia 10 : I 0 5- 116. herpetofauna venezolana. Re .~lÍmelles i COlIgre.\'o Lmino Schuberl, C. and Huber, O. 1989. La Gran Sabana, Panorámica de americano de Ecología 7 p. /lila región. Cuadel'l1Qs Lagoven Lagoven S. A., Caracas. 108 p. Pé r:1ur. J. E., Pére.l. R. , Sierra, N., and Godoy, F. 1987. Density Scbubcrl, C. and Vivas, L. (editors). 1993. El Cualerntlrio de la reappraisal of caecilians in the Andes of Venezuela. Jour Cordillera de Merida. Andes Velle=olallos. Un iversidad de nal 01 Hape/oloKY 2 1(4) :414-419. Los Andes/Fundación Polar, Mérida. Praderio. M. J. 1985. AspeclOs ecológicos de u/w pohlación de Seijas . A. E. 1993. Li stado bibliográfico comentado sobre los Colostethus herminae (Dendrobafidae) en la qllebrada de crocodylia de Venezuela. Biblioapullles. documenfo técnico Gllariquira. Esrado Miranda. Tesis Especial de Grado, de Biodoc (Cllallare) 1: I O p. Universidad Simón Bolívar, Caracas. Scijas, A. E. 1996. La conservación y manejo de Crocodílidos Presch, W. 1980. Evolutionary h istory of the South American en la región neotropiea[, p. 419-427 il/ Pél"aur. J. E. (edi Microteiid lizards (Teiidae: Gym nophthalmi nae). Copeia tor). Herpetología NeOlropical. Univers idad de Los Andes. 1980( 1 p6-56. Consejo de Publicaciones, Mérida. Pritchard, P. C. 11. and Trebbau, P. 1984. The TI/r/les uf Venezu Sei'laris, J. c., Ayarzagüena, J., and Gorzula, S. 1994. Los sapos ela. Society for the Study 01" Amphibian s and Reptiles, de la familia l3u fonidae (Amphibia: Anura) dc las tierras Cushing-Malloy, Ine., Ann tubor, Mi c hi gan. alIas de la Guayana venezolana: descripc ión de un nuevo Ramo, C. 1982. Biología del galárago (Podocl/clllis vog/i) género y tres especies. Public(J c ion e~' de la Asociación de (Muller, 1935) en el Hato El Frío. Llano de Apure (Venezu Amigos de Do/jana (Sevilla) 3 : 1- 37. ela). Doílana ACf(¡ Ver/ebra/a (Sevilla) 9(3): 1- 157. Si lva, J. L., Valdez. J .• and Ojasti. J. 19 &5. Algunos aspectos de Rivero. J. A. 1961. Sa lie ntia 01" Venezuela. Bullefin of/he MII una comunidad de ofidios del Nortc de Venczuela. Bio/ropica seum 0f Compararil'e Zoology (IJarvard) 126: 1-207. ' 7(2), " 2- ' 25. Rivero, J. A. 19633. The dislribulion of Venezucl an frogs. r. T he Slalnn, M. A. and Dixon, J. R. 1977. The herpctofauna nI" the Maracaibo basin. Carihhean ./olll'llal oI Scú:llce 3:7- [3. central Ll anos: noteworthy rccord s, a tentative ehccklist Ri vera. J. A. 1963b. The di Slribution 01" Ve nezue!an frogs. 11 . The and ecological notes. Journal 01 flerpe/ology I1 ( 1): 17- 24 . Venezuelan Andcs. Caribbean Joumal 01 Science 3 :87-102 . Thorbjarnarson, J. 1991. An analysis 01" the spectac led caiman Ri vera, J. A. 1963c. The distribulion of Venezuc1an frogs. 11 1. (Caiman crocodilus) harvest prograrn in Venezucla, p. 217- The Sierra de Pe rijá and the Falcón region. Caribbean 235 in Robinson, J. and Redford. K . (editors). Neofropicol Jourlwl of Sciellee 3: 197-199. Wildlife Use alld Conserva/ion. Uni versity 01" Chicago Press, Ri vera, J. A. 1964a. The distributi on of Ve nezuclan frogs. IV. Thc Chicago, Il1 inois. Coastal Ibngc. Caribbean Jotlrlutl of Science 4:307-319. Van zolini, P. E. 1978. On South American Hemidacrylll.\· (Sauria, Ri vero. J. A. 1964b. The distribution 01" Venezuelan frogs. V. Gekkonidae). Papeis AVlllos de ZooloKia (Sao P(lIdo) The Venezuelan Guayana. Caribbeall JOllnwl 01 Seiellce 3 1(20),307-343. 4 :411-420. Velasco. A. and Ayarzagüena. J. 1995. Situación aClU al de las Ri vero, J. A. 1964c. The distribution of Venezuelan frogs. VI. poblaciones de baba (Caiman crocodilll s) sometidas a The Llanos and Delta region. Carihhean JOl/mal oI Sei aprovechamiento comercial en los Llanos venezolanos. enee 4:491-495. Pllblicaciolles de la A.HJciaciÓ n de Amigo.\· de DOIj(Jlla Rivero. J. A. 1979. Sobre el origen de la fauna paramera de (Sevilla) 5. 71 p. anfibios venezolanos. p. 165-175 in Salgado-Laboriau, M. Via l. J. L and Saylor. L. 1993. The status of amphibian popula L. (editor). El medio ambiellle páramo. IDEA, Caracas. tions: a compilation and analysis. A repor' 0l,he Dedining Ri vero. J. A. 1982a. Los Elell/herodac/yllls (Amphi bia, Ampltibiall Poplllo/Íons Ta .~k Force (DAPTF) (O /he SSC/ Leptodaclylidae) de los Andes Venezolanos. 1. Especies del IVCN Working Document l . 98 p. páramo. Memorias de la Sociedad de Ciencias Na/li Nd es Wilkinson, M. 1996. Resolution of thc taxonomic stalUs of La Salle (Caracas) 42(11 8):9- 16. Neclocaecilia Iwydee (Roze) and a revised key to th e gen Rivera, J. A. 1982b. Los Eleu/herodacryllls (Amp h ibia. era 01" the Typhlonectidae (Amphibia: Gyrnnophiona). Jour Leptodacty lidae) de los Andes Venezolanos. 11 . Especies l/al of HerpelOluKJI 30(3) :41 3- 41 5. subparameras. Memorias de la Sociedad de Ciellcias Naru YÚ stiz. E. 1996. Aspectos biogeog ráficos de la herpetofauna de rales La Salle (Car(Jcas) 42( 118) :57-132. la cuenca hidrogdfica del Río Turbio (Estado Lara-Ven Ri vera, J. A. 1985. Nuevos eenlrolénidos de Colombia y Ve n ezuela), p. 317-349 in Pél"aur. J. E. (editor). lferpe/oloKía ezuela. Brenesia 23 :335-373. Neorropicol. Unive rsidad de Los Andes . Consejo d e Rive ro. J. A. and Solano, H. 1977. Origen y evolUl.:iÓn de los Publicaciones. Mérida. Elel//herodactyllls (Amphibia: Leptodactyli dae) de los Andes venc701a nos. MC'1II0rill .~ de la Sociedad de Ciencias Naturales La Salle (Caracas) 37:265- 282. Mallllurip/ received : 26-April-1997 Rivero-Blanco. C. and Dixon, J. R. 19 79. Origin and dislribu- A ccepfed: 15-Jafllfary-1998 61 APPENOIX 1: CHECKLlST OF VENEZUELAN AMPHIBIANS ANO REPTILES I zanes C. (:uJllrinl'llsis Péraur 1993 T A x A C. dunni (Rivero 1961) 11 I 1m 11V IVI VI IVII I VIII C. hwni/is Rivero 1978 IAMPHIBIA C. leo/Jar¡/alis Rivero 1976 I ANURA C. mwulelorum (Schmidl 1932) l' I~E C. parkeme Meinhardt and Parmelee 1996 lArdo".. .' . RI"ro C. so/IUCIrS ;S Rivero 1978 lA. I d.a Mar" 1996 C. smlluarlilli Rivera. LaIlJJ;one. and Pri,gioni 1986 lA. . 1 ,"d Manen, 1856) C. ~J1rI?\ ' ei (Rivero 1961) lA. ,Rlvero 1974 D(!/Idmba/es leucnmelas Steindachner 1864 lA. , , 1903 D. rlifll/us Gorzula 1990 lA VillC/IIROi Rivero 1980 Eprpedobares piwlS (fschudi 1938) lA. '""",ro' La Marca 1983 E. IrÍl'il/afl/s (Spix 1824) lA. "'"""O" La Marca~ 1990 MallllO ,/rr\"lIe col/Mis (Boulenger 1912) 11M" ' I! 1 · 1882 M. cordilleriana La Marca 1994 I/J. , S,I, 1824 M. /terminae (Boen2cr 1893) IIJ. , .. "roo .. , SeI", 1799 M. ¡arQ/u/illa Yústlz 1991) IIJ. m{jf¡'ws Linnaeus 1758) M. /wlJ/i lla (fesr 1956) 18. IIl1.\in/S Werner 1903 M. oblillera/a (Rivero 1984) 18. G'nthc~ M. ril'eroi (Donoso-Barros 1965) IIJ. r"/1lroo',,,, , 1758) M. /rini/mis (Garman 1887) , Señar's. . aud Com,l. 1994 M. \'us/izi (La Marca 1989) ''',,"'''0 SeOarl, 1993 Millvolxues Sle)"t' rm(lrki (Rivero 1971) 10. I",b,,' 198í Nephelolxlles alhogl.llllllus (Boulell~e r 1903) 1900 N. dlmm/; (Péfaur 1985) ",,,,, Sefrar',. . a"d Gorzul, 1994 N. Iwwleeae (Rivero 1976) 'I,,,/elrii 1895) N. IIwWlrgai (Rivero 1978 ..,'" '"''',,''' SeO,d,. . a"d Cur""a 1')94 N. meridensis (Dole and Duralll 1972) N. lIIo/inarii (La Marca 1985) rPNlnAF N. oros/onw (Rivero 1976) ',~ N. serruflus (Péfaur 1985) ""d'''''''' (Rlvero 1968) O.. áley' 1882) IIYLlDAE ,o".. la~)992) Aparasphenodorr verrewlanus (Mertens 1950) ! .. osum (Riv~~() 1_9~~ FEee/OIIO/us fr/zxeraldi Parker 1934 , (SeOarl' ,,,,) , , 1993) F. '" '//WCIIS (BoeuRer 1893) 1992) CllSlrutlleca hdemre Dunn 1944 I . 1992) G. lIicefori Caige 1933 , , , ' (GOl" 1963) G. ol'i(cra (Weinland [854) IH. " d,,,,,,,r' (R'vero 1985) G. lralkeri Duellman 1980 111. G. wil/iamsoni Caige 1922 IH. H Y/ll l,lbomem:illC,la Spi;.,: 1824 111 . H. (ll~p,mcww Soix 1824 111 . '1~92) H. alemwri Rivero 1964 111. i I ~I (Rivero 1985) H. emmw/ica AyarzaRúena and Señaris 1993 1/1. 0,;" .. ,,1'.' (Rlvero 1968) H. barlersbvi Rivero 1961 111 . IRlvero 1985) H. Imlile:; I ~ivero 1961 1/1. ""/I"I,,,u (RI"ro 1985) H. bow,~ U/tnaeus 1758) 111. ~er01985) N. ea/camw Troschel 1848 111. H. ere ,i/(I/IS Wied 1824 IH. 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 biogeographical zone. I N. '~) ALLOPIIRYNIDA E I H. ',,,d,,", Rive CROCODYL1A ICA"rOl 'AIf)AE CROCüDYLIDAE C(¡('cili(¡. , , Ro" and Sol'no 1963 CrOClXl\'/u:> ClcullU (Cuvier 1807) I I "'ac"" Shaw 1802 C. illlefmediu,\ Graves 1819 I , , ,Oonn 1942 , , 1 Unn"n' 1758 ALLlGATORIDAE , . , mber (Ro" an~ COilllCllI c/'Owdillu Linnaeus 1758 ji , I r (Mikan Cctimwt lri 'u Linnaeus 1758 J>a/euSllclrus /)(11 ¡ebrosus (Cuvler 1807) IRII 'TIDAE P. lri 'O/la/US (Schneider 1801) , ,(0"'00 1942) I I I Iw,,", AMPlilSBAENIA , """,il 1 . 1882) I I I AMPH ISBAENIDAE 1\111 ,hislx.ella alha Unnaeus 1758 N. , 1834) A llliginosa Linnaeu$ 1758 N. ~ G. seiJ,!/ jei Donoso-Barros 1966 1'I"k'ri seh~m¡.dl.193ZI i (Peters 1862) ~mg G. /(mim' Roze 1963 ¡ 1903) G. "ill(¡HU (Lichlcnslcin 1856 h"b""i La Marca and 1990 Hcmitlm_'f\'lus m{/bOlú(l (Moreau de Jonnes 1818) , , (C"y 1846 )(¡I(lje/,/lltlS Kluge 1969 H . 11 I I Dunn 1 Le úclobleplUlrjs mon/ecanocnsis Markezich and Taphorn 1994 """i Nledcn 191' Ph 'Ilodac/vlu s c/l.'(Olli Ri vero-Blanco and Lancini 1967 ~ , ",i COLUBRIDAE "",oru, [Unnaeu, 1758) Almclus /uuli us (Boie 1827) 1 ~ 1 1 · (~~i~5) A. t/uidellsis Roze 1961 1 f ~ 1 A. elaps (C unther 1BSID 1 ''''"a, (Unnaeu, 758) " ,0mm~M"" 1991 M. remv(w; Roze J 987 I -L -L "phi .. " ' ~Im,,,u, M. E!!xi W~ler 1824 -L -L -L wll/l"",,1 (!lo" 1958) lit surinamensi,) (Cuvicr 1817) 1 1 J , , " "~'bm,,, ",d Ou .."" 1854 , ,,,,,,,,,,,11 (S,u,,' VIPERIDAE Bolltriechis scl ll ~Ii~Bef(hold 1846) M, I 1 ¡ Raddi [820 Borhriopsi.\ bilillca/(¡ (Wied 1821) M, lI"dd"",1 IS"",zeo 1796) lI!ei!usa Stcrnfeld 1920 M, pi,,; 10um" ,Slbcon, aod Oum,," 1854) R iI faelliaw (Wagler 1824) Nillio O/rata (Hallowell 18·1 5) Ho/ llm , .~ as 'er (Gasman 1883) OnDeI;, "''''''' Waglm 24 B, (llrox (Linnacus J 758) O, h,"'d,,, iDa",lIo 180 "",,,,la , ,,758) R bru:ili Hoge 1953 iI cll!0:..d(w Salldner-Montilla 1976 O, 1 "/1 ¡ DumeriL Bibron, and Dumeril 1854 , " Sh CLASS ORDER FAM ILY SPECIES Podoorclllis ex lWISCI AMPHIBIA p, lewwmll· Anura DermochelYidae Al10 hrynidae Dennoche/\,$ corillcell AIIQ JI 'Ile rwhl'('/I¡ Chelonidae Bufonidae Cllrellll curerra Arelo us cllrlxmerell.'ii.l ClrelO/tia 1Il)'das A. cruci er f rellllochd)'.~ imbricllW A. mucuhu'iellsis A. ox~' rh)'nduu Crocod lia A. sorianoj· Crocodylidae Oreo lIn'/leIla IlUberi CrocoddllS llCIIIUS o. maeelme/Ji C. illlemretliu.\ Cenlrolcnidae Alli atoridae Ce'llrole/!f~ altiwd¡,,,,lis· Caillllllr ni er'· C. l/m/ilrus · Hval;IIQfJ(/fr(lc/l;U/II durwui* ·AdditiOI1$ 10 ¡he previolls repon lO I)APT"'SSCtUCN H. /t'I/rolil!Clltwll* Delldrobatidae ColO.tletlws ca lllr¡II/:! II$;S'" C. /wmili s C. leo wrda/is C. ma/Uldorum C. sal/ue/lSis MOn/1O hryne col/aris M. corllillerWlu* N, heloooles albo U1Wlu ~ '"co N. (/u r(lIIli N. /wydec(H! N. mavor tÚ N. lIIeridel l.~il· N. molillll rij* N. oros/oma N. serrll/IIIS Hylidae G(lslrorheCClllice or;- Hyla 'u/mi H. /,m:ill;ll H. meridellsis* H. la -dac/y/a Le todact lidae Cerato hr\'s calcarulll C. co"ulla Le "odacl\'!r1S lab ',iluhiel/s· Elew/ler()(lactrllls ~ille)i E. lauci/úi E. Hlmmem$ Pseudo alutlico/" J!ul/I,,· Caudata Plelhodont idae Boliro ·loHa borlmrata B. ores/es REPTILlA Tesludines Pelomedusidae Pl ate 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 la rgest 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 Ve nezuelan 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.