The Patagonian Herpetofauna José M. Cei Instituto de Biología Animal Universidad Nacional de Cuyo Casilla Correo 327 Mendoza, Argentina

Reprinted from: Duellman, William E. (ed.). 1979. The South American Herpetofauna: Its origin, evolution, and dispersal. Univ. Kansas Mus. Nat. Hist. MonOgr. 7: 1-485. Copyright © 1979 by The Museum of Natural History, The University of Kansas, Lawrence, Kansas. 13. The Patagonian Herpetofauna José M. Cei Instituto de Biología Animal Universidad Nacional de Cuyo Casilla Correo 327 Mendoza, Argentina

The word Patagonia is derived from the longed erosion. Scattered through the region term “Patagones,” meaning big-legged men, are extensive areas of extrusive basaltic rocks. applied to the tall Tehuelche Indians of The open landscape is dissected by transverse southernmost South America by Ferdinand rivers descending from the snowy Andean Magellan in 1520. Subsequently, this pic­ cordillera; drainage is poor near the Atlantic turesque name came to be applied to a con­ coast. Patagonia is subjected to severe sea­ spicuous continental region and to its biota. sonal drought with about five cold winter Biologically, Patagonia can be defined as months and a cool dry summer, infrequently that region east of the Andes and extending interrupted by irregular rains and floods. southward to the Straits of Magellan and eastward to the Atlantic Ocean. The northern boundary is not so clear cut. Elements of the HISTORY OF THE PATAGONIAN BIOTA Pampean biota penetrate southward along the coast between the Rio Colorado and the Rio In contrast to the present, almost uniform Negro (Fig. 13:1). Also, in the west Pata­ steppe associations in Rio Negro, Chubut, gonian landscapes and biota enter the vol­ and Santa Cruz provinces, during Oligocene canic regions of southern Mendoza, almost and Miocene times tropical and subtropical reaching the Rio Atuel Basin. The Pata­ vegetation occurred along with xerophytic gonian region has a wide ecotonal zone with woodlands with luxuriant mesophytic gallery the Chacoan region (Gallardo, this volume). forests. A comparison of the rich Miocene The monte vegetation (Morello, 1958) with flora of Pichi Leufu, Rio Negro (Berry, 1938) its several formations containing numerous with analagous associations from Mirhoja, subtropical elements extends south to the Chubut; Valcheta, Rio Negro; and Rio Chalia, Peninsula de Valdes; the monte enters the Santa Cruz, shows a mixture of mesic tropical Rio Chubut drainage and extends westward elements (Ficus, Fagara, Nectandra, Tabe- to the Río Neuquén, Río Agrio, and Rio Limay buia, Myristica, Sterculia, tree fems, Erythro- valleys. South of the Rio Negro, the monte xylon, Oreopanax, Maytenus), including associations exist in a system of saline low­ climbers (Buettneria, Banisteria, Bignonia, lands (bajos) and reach irregular spurs of Cissus, Paullinia, Sapindus, Strychnos), to­ the Meseta de Somuncurá, a typical Patagon­ gether with nontropical genera (Araucaria, ian environment (Cei, 1969a,b; Ruiz Leal, Azora, Berberís, Ginkgo, Laurelia, Emboth- 1972). Nevertheless, there is a general, some­ rium, Fitzroya, Libocedrus, Podocarpus, Lo- times remarkable, agreement between the matia, Peumus, Myrceugenia, Drimys). Most phytogeographic boundaries of the Monte- of the latter are characteristic components of Pampean and the Patagonian regions and the the present temperate Valdivian forest. Nev­ distribution patterns of their herpetofaúnas. ertheless, xeric areas in the Middle Tertiary Herein I emphasize the biota of the Cis- of Patagonia are suggested by certain paleo­ Andean steppe to the near exclusion of the floras containing Schinopsis, Schinus, and Cu- Trans-Andean austral forest ecosystems pania. The former is a significant genus of treated by Formas (this volume). trees in the subtropical Chacoan region. Patagonia is a region of sedimentary rocks Nothofagus forests were widespread in and soils, mostly tablelands subjected to pro­ Patagonia in the Eocene and Oligocene, but

309 310 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

Fig. 13:1. Phytogeographic regions of austral South América. Regiones fito geográficas de Sud América austral. 1979 CEI: PATAGONIAN HERPETOFAUNA 311

Fig. 13:2. Paleontological records of the lower Tertiary Patagonian flora and of leptodactylid frogs. Hallazgos paleontológicos de flora patagónica del Terciario inferior y de anuros leptodactylidos. 312 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7 these associations decreased and retreated history of tropical elements in Patagonia. The southward and westward in the Middle and fossil snake Dinilysia patagónica from the Late Tertiary. Chusquea bamboo groves oc­ Upper Cretaceous of Neuquén is related to cur in Cenozoic deposits at Laguna Hunco, boids and aniliids that are widespread in trop­ Chubut. Further evidence of paleoclimatic ical South America. Furthermore, boid snakes conditions in Patagonia is derived from the (Madtsoia), crocodilians (Necrosuchus, Se- extensive paleo-mammal faunas (Báez and becus, Eocaiman), and meiolaniid and pelo- Scillato Yané, this volume) and more limited medusid turtles from Paleocene-Eocene de­ paleo-herpetofaunas ( Báez and Gasparini, posits in Chubut are indicative of tropical this volume). environments (Gasparini and Báez, 1975). Primitive leptodactylid, ceratophrynine, Iguanid (Erichosaurus debilis) and teiid (Di- and bufonid frogs have been recorded by asemosaurus occidentalism lizards lived in Schaeffer (1949), Chaffee (1952), and Casa- southern Santa Cruz in the Miocene. miquela (1963) from the Deseadan, early Oligocene Scarritt Pocket Formation (Can­ quel, Chubut) and from the upper Miocene PATAGONIAN FAUNAL REGIONS of Rio Negro (Fig. 13:2). The living Chilean frog Caudiverbera caudiverbera is almost Two major faunal regions (habitats) can identical to the fossil frogs of the same genus. be defined in Patagonia. These are the north­ A fossil frog from the Oligocene of Chubut ern or ancient region and the southern or referred to Eupsophus by Schaeffer (1949) Santa Cruz region; the border between these has been considered the same as living E. regions is approximately at the Rio Chubut roseus, a species characteristic of the Notho- at 45°S (Fig. 13:3). These habitats corre­ fagus forests of southern Chile ( Bogart, spond to ancient physiographic areas, the 1970). These fossils clearly establish the pres­ Patagonian Massif and the Deseado Massif, ence of telmatobiine frogs in Patagonia in respectively (Figs. 13:4-5). These massifs the Oligocene and Miocene. are ancient structural continental units known as nesocratons (Harrington, 1962). In spite The Oligocene Neoprocoela was provision­ of its less marked subpositive tendency in ally referred to a Batrachophrynus or Telma- comparison with the Pampean Massif, the tobius-iike leptodactylid stock by Schaeffer whole region of the Patagonian Massif has (1949). Tihen (1962) considered it to be a been a site of almost uninterrupted accumu­ species of Bufo in the Palearctic Bufo cala­ lation of continental deposits. More rarely it mita group. The fossil was again associated received shallow marine deposits peripherally with the telmatobiine genera Telmatobufo at times of oceanic transgressions in the and Batrachophrynus by Lynch ( 1971 ). New Eocene-early Oligocene, middle Oligocene, material from the same formation supports and middle Miocene. The smaller Deseado the inclusion of Neoprocoela in Bufo (Estes, Massif had a subpositive tendency even less pers. comm.). The placement of Neoprocoela marked than the Patagonian Massif; accord­ in the Bufo calamita group has interesting ingly, its relief was often depressed, and dur­ biogeographical implications. Serological evi­ ing prolonged subsidences it became a sedi­ dence (Cei, 1977) supports a relationship be­ mentary area like the adjacent pericratonic tween the European Bufo calamita and the basins (Harrington, 1962). small B. variegatus presently restricted to the austral Nothofagus forests of Argentina and The northern or ancient Patagonian re­ Chile (Gallardo, 1962). gion extends through Neuquén, Río Negro, and Chubut provinces (Fig. 13:6). The sub- The presence of a ceratophrynine frog cordilleran area in Neuquén is drained by the (Wawelia gerholdi) in the Miocene provides Río Agrio and Río Neuquén, which flow into herpetological evidence for the southward ex­ the Rio Limay, a tributary of the Rio Negro. tent of tropical elements in the Middle Ter­ Extra-cordilleran mesetas include the large tiary. Reptilian remains substantiate the long Meseta de Somuncurá (1000-1700 m eleva- 1979 CEI: PATAGONIAN HERPETOFAUNA 313

Fig. 13:3. Major herpetofaunal regions of Patagonia. Regiones herpetofaunísticas fundamentales de Patagonia. 314 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

Fig. 13:4. General tectonic structure of South America (after Harrington, 1962). The area in the box is enlarged in figure 5. Estructura geotectónica de Sud América (según Harrington, 1962). El área en el recorte aparece aumentada en la figura 5. 1979 CEI: PATAGONIAN HERPETOFAUNA 315

Fig. 13:5. Location of the southern massifs and their relation to the major Patagonian herpetofaunal regions. Ubicación de los macizos australes y su relación con las regiones herpetofaunísticas fundamentales patagónicas. 316 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7 tion) and the Meseta de Canquel, plus small­ Table 13:1.—Characteristic Types of Vegetation in er mesetas of the same lower and middle the Ancient Patagonian Region. Tertiary age along the Andean front; their Herbs Spiny or Sclerotic Plants small lakes are close to the last eastern patches Stipa Chuquiraga Festuca Nassauvia of Nothofagus and Araucaria forests. Along Poa Ephedra the base of the Andes there is an Austral- Senecio filaginoides Styllingia Patagonian ecotonal zone; however, it is Grindelia chiloensis Verbena Verbena ligustrina Pantacantha much narrower than the Monte-Patagonian Acaena caespitosa Adesmia ecotone extending from the Rio Colorado to Shrubs Austrocactus Valcheta, onto the northern spurs of the Me­ Mulinum spinosurn Hydrophyllic Plants seta de Somuncurá, and southward to near Colliguaja integerrima Juncus Berberis cuneata Carex Bahia Camarones on the Atlantic coast (Ca­ Lycium tenuispinosum Ranunculus brera, 1951). Anarthrophyllum rigidum Hypsela Floristically, the ancient Patagonian re­ Anarthrophyllum Plagiobothrys gion is characterized by a predominantly desideratum Acaena macrostemon Trevoa spinifer Caltha steppe vegetation with scattered green bushes Prosopis patagónica Cortaderia (Mulinum spinosurn), several grasses, low Larrea ameghinoi Azorella herbaceous plants, some spiny plants, and a Halophyllic Plants variety of low bushes; additional kinds of Atriplex Frankenia plants are present in saline environments and Spartina riparian situations (Table 13:1). The most characteristic element of the monte forma­ rugged southernmost Andean cordillera. The tions is the creosote bush (Larrea), which is Andes are commonly bordered by sharp- represented by five sympatric species in the edged basaltic mesetas having elevations of ecotonal zone at Valcheta (Rio Negro). In 1000 to 1500 m. typical Patagonian landscapes only the low Larrea ameghinoi is present. Phytogeographically, the Santa Cruz The cool Patagonian steppes dominated by Faunal Region agrees with Cabrera’s (1951) Mulinum and Stipa, with scattered creeping, Patagonian districts—Patagonico Subandino, cushionlike plants, exist in western Neuquén, Patagonico Central, and Golfo de San Jorge. southwestern Rio Negro, and in most of Chu- The Patagonico Subandino includes the ba­ but, provinces. In these areas the steppes are saltic mesetas (e.g., Meseta Vizcachas, Me- commonly associated with basaltic landscapes seta Asador, Meseta de la Muerte, and the resulting from the rampant Cenozoic volcanic Meseta de Lago del Sello) and the southern activity. The steppes are discontinuous in humid lowlands.1 In these areas open steppe northern Neuquén and Rio Negro provinces, associations of Festuca monticola, Bromus where they occur mostly at elevations of more macranthus, Hordeum comosum, Agropyron than 900 to 1000 m. magellanicum, Poa sp., and Deschampsia sp. The southern or Santa Cruz Faunal Re­ predominate, but some shrubs (Berberis cuneata, Nassauvia aculeata, or Mulinum spi- gion extends from about 45 °S to the Straits nosum) are present. of Magellan (Fig. 13:7). This region encom­ passes some distinct physiographic areas. The Phytogeographic differences between the arid valley of the Río Deseado borders the Sub-Andean district and the central and San northern limits of the large Altiplanicie Cen­ Jorge districts are evident by the monotonous tral, a dead volcanic landscape with scattered grasslands of Stipa humilis in the latter. The clay basins and petrified early Cenozoic trees grasses are interrupted by the broad circular (Auracarites). South of the great plateau the bushes of the blackish “mata negra” (Verbena drainage basins of the ríos Chalía, Santa Cruz, Coyle, and Gallegos provide more moist low­ 1 Although the Sub-Andean District is considered to be a single physiographic unit (Fig. 13:7), for pur­ lands extending to the Straits of Magellan. poses of herpetofaunal analysis, I distinguish the These rivers drain the glacial valleys of the Humid Southern Lowlands. 1979 CEI: PATAGONIAN HERPETOFAUNA 317

Table 13:2.—Comparison of the Herpetofaunas in Ten Districts in Patagonia. (Numbers of species is a given district are in boldface; numbers of species in common to two districts are in Roman, and the italics are Faunal Resemblance Factors [Ni + N2 /2C (Duellman, 1966)].

Sello

Ecotone

Lowlands de

Area

Steppe Lago

Somuncura

Central Highlands

District Southern de del

Monte Monte-Patagonian Patagonian Volcanic Meseta Humid Meseta Coastal Sub-Andean Altiplano Monte 17 17 2 3 1 1 Monte-Patagonian Ecotone ___ 0.69 32 7 7 4 5 4 2 1 Patagonian Steppe ______0.12 0.29 17 12 7 8 7 3 3 Volcanic Highlands______- 0.14 0.25 0.57 25 8 6 5 3 1 _ Meseta de Somtincnrá 0.19 0.50 0.44 11 5 4 2 1 Altiplanicie Central ...... 0.08 0.24 0.62 0.35 0.50 9 8 3 4 Coastal District ...... 0.08 0.20 0.56 0.30 0.42 0.94 8 2 4 Humid Southern Lowlands ___ 0.11 0.26 0.19 0.24 0.40 0.29 6 1 2 Sub-Andean Area ...... 0.06 0.29 0.07 0.13 0.62 0.67 0.20 4 Meseta del Lago de Sello...... — 0.44 3

tridens). Where shrubby formations occur, 13:1). Although lizards are dominant in both dominant plants are Prosopis patagonica, Ly- regions, they comprise a much higher per­ cium ameghinoi, Berberis cuneata, Chuquir- centage of the herpetofauna in the southern aga aurea and avellanedae, Brachyclados cae- region (Fig. 13:8). spitosus, Acantholippia seriphioides, Pleuro- fora patagonica, Ameghinoa sp., and Euphor­ Northern Patagonian Herpetofauna bia sp. The small trees, Trevoa patagonica, For purposes of analysis, the region has around the Golfo de San Jorge and in the arid valley of the Rio Deseado are the most been divided into five ecophysiographic areas conspicuous plants in the southern region. (Fig. 13:6)—1) Monte associations, 2) Monte-Patagonian ecotone, 3) Patagonian steppe, 4) Volcanic highlands, and 5) Meseta COMPOSITION OF THE de Somuncura. The distributions of the spe­ HERPETOFAUNA cies and subspecies of amphibians and rep­ tiles in these five areas are tabulated in Ap­ The herpetofauna of the Patagonian pendix 13:1. The sole Patagonian turtle, Geo- steppe is composed of 60 species, six of which chelone donosobarrosi, and all of the Pata­ have two or more subspecies in Patagonia; gonian anurans are in the northern region, the entire herpetofauna consists of 70 spe­ although one species, Pleurodema bufonina, is cies and subspecies with a noticeable degree widely distributed in the southern region. of endemism. The fauna is made up of 14 Likewise, the single amphisbaenian and all species of anurans (23.3%), one turtle (1.7%), of the snakes are in the northern region, al­ 34 lizards (56.7%), and 11 snakes (18.3%). though Bothrops ammodytoides enters the For purposes of discussion, the herpetofauna southern region. has been divided according to the two major All of the 17 species occurring in the faunal regions. Of the 60 Patagonian species, monte associations are among the 32 species 56 occur in the northern or ancient Pata­ in the Monte-Patagonian ecotone. Included in gonian Region, and 13 occur in the southern these areas are several species characteristic or Santa Cruz Region; nine species are com­ of, or related to species in, the more northern mon to the two regions (Table 13:2, Appendix regions—Pampas and Chaco; this is true of 318 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

Fig. 13:6. Herpetofaunal and physiographic districts of the ancient Patagonian region. Distritos herpetofaunísticos y fisiográficos de la región patagónica antigua. 1979 CEI: PATAGONIAN HERPETOFAUNA 319

Fig. 13:7. Herpetofaunal and physiographic districts of the southern Patagonian region. Distritos herpetofaunísticos y fisiográficos de la región sur-patagónica. 320 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

in rocky areas, some of which also are in­ habited by more cryptic lizards—Liolaemus ceii and L. kriegi. Isolated populations of Phymaturus patagonicus are subspecifically distinct—P. p. patagonicus in the valley of the Rio Chubut and P. p. indistinctus in the Sierra de San Bernardo (Cei and Castro, 1973) (Fig. 13:12). Liolaemus fitzingeri can- queli inhabits the rocky slopes of the Meseta de Canquel and extends eastward through the salt flats to the coast, where it meets L. f. fitzingeri, the subspecies that is common in southern Chubut and Santa Cruz. Diplolae- mus bibronii, more characteristic of the south­ ern region, reaches the northern limits of its distribution at the edge of th$ Meseta de Somuncurá (Cei, 1971b). Clay soils in the region hold water in the form of temporary ponds during the brief rainy season. These ponds and intermittent and permanent streams are the habitats and/or breeding sites for several species of anurans, especially leptodactylids (Barrio, ANCIENT PATAGONIAN SOUTHERN MAJOR MAJOR ANIMAL HABITAT ANIMAL HABITAT 1973; Cei, 1969a,b, 1970b, 1972b; Cei and Roig, 1966, 1968; Gallardo, 1970). Some spe­ Fig. 13:8. Herpetofaunal composition of the cies have restricted ranges; for example, Ate­ Patagonian regions. Composición herpetofaunística de las regiones lognathus solitarius is known only from Ar­ patagónicas. royo Las Bayas, south of Pilcaniyeu, Rio Negro. the anurans Bufo arenarum and Leptodacty- Twenty-four species are known to inhabit lus ocellatus, the lizard Mabuya frenata, and the volcanic plateaus and extra-Andean high­ the snakes Elapomorphus bilineatus, Lystro- lands in western Neuquén and Rio Negro phis semicinctus, and Micrurus frontalis (Fig. provinces. Among them are four species of 13:9). The analysis of latitudinal distribu­ telmatobiine leptodactylid frogs (Fig. 13:3). tions reveals that Geochelone donosobarrosi The aquatic Atelognathus patagonicus is con­ is a true member of the xerophytic scrub asso­ fined to the Laguna Blanca Basin. The semi­ ciation south of the Rio Colorado and only terrestrial Atelognathus praebasalticus is com­ slightly penetrates the neighboring flats of the posed of four geographically isolated sub­ Pampean region. The distributions of the species—A. p. praebasalticus at Laguna Blan­ snake Philodryas patagoniensis and the lizard ca, A. p. agilis at Laguna Casa de Piedra, Proctotretus pectinatus extend eastward into A. p. luisi at Laguna Catan Lil, and A. p. the Pampean region (Figs. 13:10-11). On the dobeslawi at Barda de Santo Tomas (Cei, contrary, the frog Pleurodema bufonina and 1972b). Atelognathus nitoi and Alsodes gár­ the lizards Homonota darwinii and Liolaemus gola gárgola have different distributional bibronii are characteristic Patagonian ele­ traits. In western Rio Negro these frogs oc­ ments and only enter the monte peripherally; cur in small Andean lakes—Laguna Verde these species ascend the Andean slopes north near Cerro Blanco at about 1450 m elevation of the Rio Barrancas. (Barrio, 1973), and Laguna Tonchek and In the Patagonian steppes, extrusive basal­ Laguna Schmoll at 1700 to 1750 m on the tic rocks provide shelter for numerous lizards. slopes of Cerro Catedral near Bariloche (Gal­ Liolaemus elongatus is a conspicuous species lardo, 1970). Together with Alsodes gar gola 1979 CEI: PATAGONIAN HERPETOFAUNA 321

Fig. 13:9. Southern limits of distribution of taxa in the monte formation in northern Patagonia. Areas of sympatry of Pleurodema bufonina and P. thaul are indicated. Límites meridionales de distribución de taxa en la formación del monte en el norte patagónico. Se indi­ ca las áreas de simpatría de Pleurodema bufonina y P. thaul.

neuquensis from the thermal brooks on the Luán. In other ecotonal and subandean areas sandy Meseta Lonco Luán, 1500 m elevation of Neuquén, Pleurodema thaul, Liolaemus in Neuquén (Cei, 1976), they belong to a chilensis, and L. buergeri are found. The lat­ transitional herpetofauna of the austral-Pata- ter occurs sympatrically with the typical Pata­ gonian ecotone. Other transitional species are gonian lizards Liolaemus elongatus and L. the frog Pleurodema thaul and two lizards, kriegi. Likewise, the characteristic lizards of Liolaemus tenuis and Diplolaemus leopardin­ rocky Patagonian communities, Phymaturus as, all characteristic inhabitants of Araucaria palluma and P. patagónicas zapalensis occur forests (Cei, 1970a, 1974b). On the xeric on the basaltic plateaus. Phymaturus palluma Meseta de Lonco Luán, dead patches of extends northward in the Andes at elevations Nothofagus and Chasquea exist near the bor­ of 3000 to 3500 m to La Rioja and San Juan, der of the Araucaria forests. Liolaemus lineo- and P. patagónicas has distinct populations maculatus occurs on the Meseta de Lonco (P. p. payuniae and P. p. nevadoi) to the 322 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

Fig. 13:10. Distribution of Liolaemus darwinii, a characteristic species of the monte formation. The south­ ern edge of this formation is the southern limits of distribution of Geochelone and of colubrid snakes in north­ ern Patagonia. Distribución de Liolaemus darwinii, especie característica de la formación del monte. El límite meridional de esta formación es el límite meridional de Geochelone y de los ofidios colúbridos en la Patagonia septentrional. 1979 CEI: PATAGONIAN HERPETOFAUNA 323

Fig. 13:11. Patterns of distribution of tropidurine iguanid lizards in Patagonia. Distribución de los saurios iguánidos tropidurinos en Patagonia. 324 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

Fig. 13:12. Patterns of distribution of tropidurine iguanid lizards in Patagonia. Distribución de los saurios iguánidos tropidurinos en Patagonia. 1979 CEI: PATAGONIAN HERPETOFAUNA 325

Fig. 13:13. Distribution of extra-Andean telmatobiine frogs in northern Patagonia. Distribución de los anuros telmatobiinos extra-andinos en la Patagonia septentrional.

north in southern Mendoza Province (Cei demic subspecies of lizards. The endemic and Castro, 1973; Cei and Roig, 1975) (Fig. Liolaemus ruizleali inhabits rocky summits of 13:12). the meseta at 1200 to 1700 m. The two en­ The isolated Meseta de Somuncura (150 demic, telmatobiine leptodactylid frogs have X 80 km), with elevations to 1700 m, has unique morphological and ecological traits. some peculiar habitats. Between 800 and The monotypic Somuncuria somuncurensis 1700 m, above the Monte-Patagonian ecotone, lives in streams at 800 to 1000 m issuing from 11 species of amphibians and reptiles are thermal springs on the northeastern slopes known. Six of the species are widespread in of the meseta. Inhabiting the same streams Patagonia—Pleurodema bufonina, Homonota having a temperature of about 18°C is the darwinii, Diplolaemus darwinii, Liolaemus endemic characid fish Gymnochacinus bergi. bibronii, L. boulengeri, and L. rothi. Three Conversely, Atelognathus reverberii, a nearly other species and two subspecies are endemic fossorial frog, inhabits the arid plateau at —Phymaturus patagonicus somuncurensis and elevations of more than 1000 m and breeds in Liolaemus elongatus petrophilus are the en­ small temporary pools. 326 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

Southern Patagonian Herpetofauna narum, and all of the genera of snakes are more northern groups. The distributions of the 11 lizards, one Bufo spinulosus and Alsodes are primarily snake, and one frog are examined with re­ Andean groups. The former enters Patagonia spect to five ecophysiographic areas (Fig. in many disjunct valleys; the species does not 13:7)—1) Altiplano Central, 2) Coastal Dis­ occur on major basaltic mesetas. In Pata­ trict, 3) Humid Southern Lowlands, 4) Sub- gonia, Alsodes occurs only in the volcanic Andean Area, and 5) Meseta de Lago del highlands adjacent to the Andes. Sello. The distributions of the species and Pleurodema is a primitive leptodactyline subspecies of amphibians and reptiles in these that may have originated in, and dispersed five areas are tabulated in Appendix 13:1, from, the austral forests (Duellman and Ve- and the distributional relationships between loso, 1977); the genus has dispersed north­ the areas are analyzed in Table 13:2. ward in nonforested habitats to the Carib­ Nine species occur in the Altiplano Cen­ bean. Two species are peripheral in Pata­ tral; all but the frog Pleurodema bufonina gonia—P. thaul in the austral forest—Pata­ also occur in the coastal district. Only six spe­ gonian steppe ecotone and P. nebulosa in the cies occur in the humid southern lowlands; monte. Pleurodema bufonina is a widespread the southernmost frog, Pleurodema bufonina, species endemic to Patagonian habitats. Al­ is rather rare there. The lizard Liolaemus though P. thaul and bufonina are distinctive magellanicus, the only herpetofaunal species in their morphology and behavior, popula­ on Tierra del Fuego, is common on the south­ tions intermediate between the species exist ern end of the mainland. Only four species in high valleys in western Neuquen. (all lizards) occur in the sub-Andean region. Lynch (1978) provided an hypothesis for The Meseta de Lago del Sello at 47°S is the evolution of lower telmatobiine frogs in nearly 50 km in diameter. On the top of the Patagonia. The peculiar monotypic Somun- plateau, low grasses (Festuca, Poa, and curia is endemic to the Meseta de Somun- Stipa) are dominant with thorny plants or cura, whereas five species of Atelognathus cushion plants (Benthamiella azorella, Ver­ occur in isolated basaltic areas in Patagonia. bena, Senecio, Nassauvia); lichens are abun­ The only extra-Patagonian Atelognathus is dant in rocky areas. Owing to the proximity A. grandisonae from Puerto Eden in extreme of the great Patagonian Ice Field, cold winds southern Chile. whip the plateau, even in summer. Only three Among the iguanid lizards, the monotypic iguanid lizards have been found on the pla­ Vilcunia has characters of both Proctotretus teau (1200-1600 m); the two species of Lio­ and Liolaemus and is endemic to southern laemus are widespread in southern Patagonia, Patagonia (Donoso Barros and Cei, 1971). whereas the monotypic Vilcunia sylvanae is Diplolaemus and Phymaturus are fundamen­ endemic. No amphibians have been found tally austral genera (Figs. 13:11-12). Cten- on the plateau; the widespread Pleurodema oblepharis, Leiosaurus, Proctotretus, and Pris- bufonina ascends the slopes to only 900 m. tidactylus are widely distributed to the north of Patagonia (Cei, 1973c,d). Two species of Proctotretus are distributed in temperate ORIGIN OF THE HERPETOFAUNA areas in southern Brasil, Uruguay, and cen­ tral Argentina; P. pectinatus occurs in the The Patagonian herpetofauna is distinc­ monte-Patagonian ecotone. Pristidactylus fas- tive in the diversity of telmatobiine leptodac- ciatus is primarily Patagonian, but congeners tylid frogs and tropidurine iguanid lizards. occur in disjunct Andean areas (P. scapula- Most other species inhabiting Patagonia are tus) and in isolated extra-Andean massifs of members of groups that are mainly distrib­ central Argentina—Sierra Grande de Cordoba uted to the north of Patagonia. Thus, Geo- (P. achalensis) and Sierra de la Ventana (P. chelone, Cnemidophorus, Mabutja, Homono- casuhatiensis) (Gallardo, 1964, 1968; Cei and ta, Leptodactylus, Odontophrynus, Bufo are­ Castro, 1975). 1979 CEI: PATAGONIAN HERPETOFAUNA 327

Three major points relative to the origin tered relicts, unique witnesses to some of the and evolution of the Patagonian herpetofauna most ancient steps in the history of continental need to be emphasized. vertebrates, whereas the Recent speciation of 1. There has been a radiation of primitive Liolaemus attests to the continued evolu­ leptodactylid frogs, remnants of Gond- tionary activity in the region. wanan elements. Vuilleumier (1968) and Lynch (1971, 1978) noted the aus­ Evolutionary Radiation of tral center of radiation of telmatobiine Patagonian Liolaemus leptodactylids in Patagonia and the austral forests; Formas (this volume) Lizards of the genus Liolaemus are wide­ discussed the biogeography and ecol­ spread in temperate South America. Three ogy of the telmatobiines of the austral species range into southern Rrasil, and several forests—Alsodes, Batrachyla, Caudi- species occur in the Andes, two extending verbera, Eupsophus, Hylorina, Insueto- northward to central Peru (Duellman, this phrynus, and Telmatobufo, some of volume). Twenty-six taxa are Patagonian or which also occur in Argentina (Cei, Andean-Patagonian. 1978). The limited paleontological Four major evolutionary units can be rec­ evidence supports the austral center of ognized among the Patagonian Liolaemus as radiation (Schaeffer, 1949; Chaffee, follows: 1) L. fitzingeri complex, 2) L. elon- 1952; Casamiquela, 1963; Estes and gatus-kriegi complex, 3) L. kingii-archeforus Reig, 1973). complex, and 4) L. magellanicus-lineomacu- 2. An austral South American center of latus complex (Cei, 1971a, 1972a, 1973a,b evolution and adaptive radiation of an 1974a, 1975a,b, 1975d,e; Cei and Scolaro, ancestral stock of iguanid lizards is evi­ 1977; Scolaro and Cei, 1977). The lizards that dent (Cei, 1973c,d, 1975c; Cei and are not members of these groups are pri­ Castro, 1975). Fourteen genera are aus­ marily peripheral to Patagonia and/or are tral in Argentina and Chile; seven of ecotonal elements. Some of these are mem­ these are Patagonian. There are some bers of transcordilleran groups—Liolaemus 20 genera of iguanids in tropical South altissimus, chilensis, cyanogaster, lemniscatus, America and another ten genera in the pictus, and tenuis; L. bibronii is related to Sonoran region of North America. With the Chilean L. fuscus. Liolaemus boulengeri, the exception of Anolis and Sceloporus, darwinii, and gracilis are members of the no other iguanid genus displays such more northern monte fauna and are primarily an impressive adaptive radiation as does peripheral in Patagonia (Fig. 13:10). Liolaemus, the dominant lizards in any Liolaemus fitzingeri complex.—This group Patagonian community. In most of is characterized by 1) patch of enlarged these same communities there exist rep­ scales on posterior surfaces of thighs, espe­ resentatives of the other Patagonian cially well developed in males (Fig. 13:14); iguanid genera—Diplolaemus, Leiosau- 2) high number (52-82) of blunt, slightly rus, Phymaturus, Pristidactylus, Procto- keeled scales around body; 3) high number tretus, and Vilcunia. (7-11) of preanal pores; 4) stout body and 3. An impressive post-Pleistocene adaptive relatively long tail, 1.5 times length of body; radiation has taken place in four groups 5) tendency to have black venters and dark of Patagonian Liolaemus (see following humeral collars; 6) predominate dorsal color section). patterns consisting of wide transverse dark Thus, far from being a totally barren re­ blotches, bordered posteriorly by white, but gion biologically, irregularly colonized by ele­ spotted erythristic, and melanistic variations ments from neighboring biotas, Patagonia has not uncommon. been, and apparently still is, a center of active Content: L. fitzingeri canqueli, L. fitzin­ speciation of several herpetofaunal elements. geri fitzingeri, L. fitzingeri melanops, L. rothi, The old radiations are supported by the scat­ ?L. ruizleali. Liolaemus f. fitzingeri is the 328 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

Fig. 13:14. Morphological traits of lizards of the Liolaemus fitzingeri complex. Características morfológicas de los saurios del conjunto Liolaemus fitzingeri. 1979 CEI: PATAGONIAN HERPETOFAUNA 329

Fig. 13:15. Distribution of the Liolaemus fitzingeri complex in Patagonia. Distribución del conjunto Liolaemus fitzingeri en Patagonia. 330 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7 southernmost member of the group, inhabit­ be considered as a primitive, ecologically gen­ ing sandy Patagonian steppe in Santa Cruz eralized species of the Liolaemus fitzingeri and Chubut; it meets L. f. canqueli in the complex. The poorly known L. ruizleali is vicinity of Trelew and Dos Pozos on the coast morphologically close to L. rothi, except that of Chubut (Cei and Scolaro, 1977). The latter the postfemoral enlarged scales are absent in subspecies is characteristic of the arid vol­ L. ruizleali. canic massif of Canquel south of the Rio Enlarged scales on the posterior surfaces Chubut. Liolaemus f. melanops is distributed of the thighs are characteristic of some other in coastal areas north of the Rio Chubut to the extra-Patagonian Liolaemus. Such is the case Rio Negro and thence inland through Men­ in the Andean L. ornatus and L. mocquardi, doza to San Juan and La Rioja. Liolaemus which morphologically are similar to L. dar­ rothi is endemic to northern Patagonia, as is winii. The character also is present in L. L. ruizleali, known only from the Meseta de wiegmannii in southeastern Brasil, Uruguay, Somuncurá (Fig. 13:15). and the Argentine pampas, and in L. multi- Members of the Liolaemus fitzingeri com­ maculatus from the Atlantic coast of Buenos plex are mostly stout, polymorphic lizards Aires. Because of the many differences dis­ that are psammophilous or fossorial. The played by these two lizards from one another polychromatism in these lizards has been a and from members of the Liolaemus fitzingeri source of confusion. Liolaemus melanops was complex, the enlarged postfemoral scales are considered to be a color variety of L. fitzin­ considered to be independently evolved char­ geri by Donoso-Barros (1966) and Peters and acters in these three lines. Donoso-Barros (1970), and as a northern sub­ Liolaemus elongatus-kriegi complex.—This species of L. fitzingeri by Cei (1975d). How­ group is characterized by 1) no patch of ever, careful analyses of morphological and enlarged scales, but a row of projecting scales serological attributes may suggest that L. on posterior surface of thigh; 2) high number melanops probably is a distinct species. Popu­ (72-120) of acuminate, keeled scales around lations of L. f. melanops near Puerto Madryn body (Fig. 13:16); 3) few (1-4) preanal are highly variable; some individuals are mor­ pores; 4) slender body with very long tail; phologically indistinguishable from L. goet- 5) absence of ventral melanism and dark schi (Cei, 1975a), a monomorphic lizard ex­ nuchal collar; 6) dorsal pattern of blackish tending from the Rio Colorado north to San irregular stripes, not bordered by white, and Juan and La Rioja. Serological analysis confluent into vertebral and lateral bands. shows that populations formerly assigned to Content: L. austromendocinus, L. buer- L. melanops and L. goetschi are conspecific; geri, L. ceii, L. elongatus elongatus, L. elon- thus, only one taxon (L. melanops) is rec­ gatus petrophilus, L. kriegi. Liolaemus elon­ ognized (Cei and Scolaro, 1977). gatus is widespread in rocky habitats in The serological distance between L. dar­ Chubut, Rio Negro, and Neuquén, and north­ winii and members of the Liolaemus fitzin­ ward in the precordillera in Mendoza (Fig. geri complex suggests that L. darwinii di­ 13:17). It is a highly variable species, and verged early from the ancestral stock of that notable serological distances have been found group. Although juveniles and females of L. among scattered, isolated populations (Cei, darwinii and L. boulengeri are strikingly simi­ 1974a); only the population of the Meseta de lar, a noticeable serological distance exists Somuncurá has been recognized taxonomi­ between the species, whereas L. boulengeri cally—L. e. petrophilus. Liolaemus austro­ is serologically closer to L. f. melanops. mendocinus occurs in arid habitats below Liolaemus rothi has morphological char­ 1500 m in volcanic regions in southern Men­ acters that ally it with the Liolaemus fitzingeri doza and in the Río Neuquén and Rio Colo­ group, but serologically it is not so distant rado basins. Liolaemus kriegi occupies ba­ from other Patagonian complexes of Liolae­ saltic areas in Neuquén and Rio Negro, where mus as are the other members of the Liolae­ it occurs sympatrically with L. ceii. Liolae­ mus fitzingeri group. Liolaemus rothi could mus buergeri occurs sympatrically with L. 1979 CEI: PATAGONIAN HERPETOFAUNA 331

Fig. 13:16. Morphological traits of lizards of the Liolaemus elongatus-kriegi complex. Características morfológicas de los saurios del conjunto Liolaemus elongatus-kriegi. 332 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

Fig. 13:17. Distributions of three Liolaemus species complexes in Patagonia. Distribución de tres conjuntos específicos de Liolaemus en Patagonia. 1979 CEI: PATAGONIAN HERPETOFAUNA 333

Fig. 13:18. Morphological traits of lizards of the Liolaemus archeforus-kingii complex. Características morfológicas de los saurios del conjunto Liolaemus archeforus-kingii. 334 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7 elongatus or austromendocinus in Patagonian Content: L. lineomaculatus, L. magellani- associations in southern Mendoza and with cus. The latter occurs in the humid southern L. elongatus or kriegi in valleys in Neuquen. lowlands on the mainland and in isolated Liolaemus buergeri and kriegi occur in Pata­ populations on Tierra del Fuego, whereas L. gonian habitats in Chile; presumably the lineomaculatus is found in the volcanic high­ transcordilleran migration was via Paso Pe- lands in the ancient Patagonian region, where huenche (2500 m, 36°S). it inhabits open formations and Araucaria Several immunological cross-reactions woodlands (Fig. 13:17). Occasional immacu­ among allopatric and sympatric populations late individuals of L. lineomaculatus are of L. austromendocinus and elongatus show known (Cei, 1971a). equally good specific levels of differentiation These last two complexes of Liolaemus are as among L. elongatus, kriegi, and buergeri limited to austral Patagonia and are conserva­ (Cei, 1974a, 1975b). tive in their diversity, as compared to the Liolaemus kingii-archeforus complex.— Liolaemus fitzingeri and elongatus-kriegi This group is characterized by 1) no patch complexes, both of which apparently have of enlarged scales on the posterior surface undergone recent (post-Pleistocene) specia­ of the thigh (Fig. 13:18); 2) moderately high tion. The results of these radiations are nu­ number (58-84) of faintly keeled scales merous morphologically similar species differ­ around body; 3) high number (6-10) of pre- ing from one another biochemically and eco­ anal pores; 4) short legs and relatively short logically. tail, only slightly longer than body; 5) ven­ ter with dark spots; absence of dark nuchal collar; 6) a series of yellowish or whitish ACKNOWLEDGMENTS transverse bars on the dark dorsal ground In acknowledge the special interest and color. efforts of William E. Duellman in rewriting Contents: L. archeforus archeforus, L. the original English draft of this manuscript archeforus sarmientoi, L. kingii. This group is and in his critical analysis of the distribu­ endemic to the southern faunal region (Fig. tional data. I also thank John D. Lynch for 13:17). Liolaemus a. archeforus occurs on information about his research on the mor­ the isolated Meseta de Lago del Sello; it is phology and relationships of telmatobiine replaced by L. a. sarmientoi at lower eleva­ frogs in the Patagonian Region. tions eastward in the Patagonian steppe be­ tween the Rio Coyle and the Rio Gallegos. Liolaemus kingii, which lies at a moderate RESUMEN serological distance from L. archeforus, is a rather stout, apparent ecological generalist Biológicamente, la Patagonia se define inhabiting ravines and open bushy habitats como la región al este de los Andes, extendi­ in most of Santa Cruz. It reaches the Atlantic éndose hasta el Oceano Atlántico, hacia el sur coast, and in the western part of its range is hasta el Estrecho de Magallanes; en el norte broadly sympatric with L. archeforus. hay una zona de transición entre la biota Liolaemus magellanicus-lineomaculatus patagónica y las del norte, entre los ríos complex.—This group is characterized by Negro y Colorado. 1) no patch of enlarged scales on the pos­ La Patagonia es una región de suelos de terior surface of thigh; 2) low number (40- rocas sedimentarias y mesetas de rocas efu­ 70) of large, mucronate, acuminate (dorsally) sivas, presentando severas sequías estacionales scales around body; 3) moderate number con cinco meses de invierno frío, veranos (3-8) of preanal pores; 4) very short limbs usualmente secos y clima fresco. and tail (Fig. 13:19); 5) absence of ventral En contraste con las asociaciones de estepa melanism and dark nuchal collar; 6) dorsum uniformes que existen allí actualmente, una irregularly spotted with black and having vegetación tropical y subtropical occurió al whitish longitudinal lines. mismo tiempo que bosques xerofíticos y bos- 1979 CEI: PATAGONIAN HERPETOFAUNA 335

Fig. 13:19. Morphological traits of lizards of the Liolaemus magellanicus-lineomaculatus com­ plex. Características morfológicas de los saurios del conjunto Liolaemus magellanicus-lineomaculatus.

ques mesofíticos de galería durante el Oligo- En los depósitos del Oligoceno y del Mio­ ceno y el Mioceno. Bosques de Nothofagus ceno telmatóbidos primitivos, ceratofrínidos, existían durante el Eoceno y el Oligoceno. y bufónidos son conocidos, asi como boideos A mediados del Terciario los climas se vol­ primitivos, cocodrilos, y tortugas meiolanidas vieron más secos dando lugar a la expansión y pelomedusidas están representados en los de la vegetación xerofítica. depósitos del Cretáceo superior y del Ceno­ 336 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

zoico inferior. Los fósiles existentes, la pre­ Berry, E. W. 1938. Tertiary flora from the Rio Pichi sente distribución y los patrones de especia- Leufu, Argentina. Geol. Soc. Amer. Spec. Pap. (12):1-149. ción indican que la Patagonia ha sido una Bogart, J. P. 1970. Systematic problems in the am­ región importante para la evolución de la phibians family Leptodactylidae (Anura) as indi­ herpetofauna austral. cated by karyotypic analysis. Cytogenetics 9: 369-383. Dos regiones faunísticas se reconocen en Cabrera, A. L. 1951. Territorios fitogeográficos de la Patagonia—1) la región del norte, o Pata­ la República Argentina. Bol. Soc. Argentina Bot. gonia antigua, 2) la región del sur, o de Santa 4:21-65. Cruz. El límite entre estas dos regiones se Casamiquela, R. M. 1963. Sobre un par de anuros del Mioceno de Rio Negro (Patagonia). Wawelia encuentra aproximadamente en el Río Chu- gerholdi n. gen. et sp. (Ceratophrydidae) y Gi- but, 45°S (Fig. 13:3). gantobatrachus parodi (Leptodactylidae). Ame- La herpetofauna patagónica está compu­ ghiniana 3:141-160. Cei, J. M. 1969a. La meseta basáltica de Somun- esta de 60 especies: 14 anuros, 1 tortuga, 34 curá, Rio Negro. Herpetofauna endémica y sus saurios, 11 ofidios. Hay un grado notable de peculiares equilibrios biocenóticos. Physis 28: endemismo. De las 60 especies, 56 ocurren 257-271. el la región del norte, 13 especies en la región Cei, J. M. 1969b. The Patagonian telmatobiid fauna of the volcanic Somuncura Plateau. J. Herpetol. del sur, y nueve especies están representadas 3:1-18. en ambas regiones. En la región del norte se Cei, J. M. 1970a. Fluctuaciones biocenóticas y re­ encuentra la única tortuga y el único am- lictos herpetológicos de la planicie de Lonco- Luán (Neuquén). Acta Zool. Lilloana 27:193- phisbénido y todas las especies de ofidios y 200. anuros, excepto una especie de rana (Pleuro- Cei, J. M. 1970b. Telmatobius solitarius n. sp., a dema bufonina) y una especie de serpiente new rare telmatobiid frog from the highland (Bothrops ammodytoides) que entran en la Patagonian territories (Rio Negro, Argentina). región del sur. Herpetologica 26:18-23. Cei, J. M. 1971a. Herpetología Patagónica—I. Lio­ El grado de endemismo es alto, especial­ laemus del grupo magellanicus. Características mente en los altiplanos volcánicos, en las taxonómicas y genéticas. Physis 30:417-424. Cei, J. M. 1971b. Herpetología Patagónica—II. estribaciones de los Andes y en las mesetas Notas sobre la distribución geográfica del género aisladas. Los géneros monotípicos Somun- Diplolaemus. Ibid. 30:471-474. curia y Vilcunia son endémicos de las mesetas Cei, J. M. 1972a. Herpetología Patagónica—III. Re­ asi como las especies y/o subespecies de Al- laciones de afinidad seroproteínicas y filéticas en el género Liolaemus. Ibid. 31:411-422. sodes, Atelognathus, Liolaemus, y Phyma- Cei, J. M. 1972b. Herpetología Patagónica—V. Las turus. especies extracordilleranas alto-Patagónicas del En la Patagonia ha habido temprana ex­ género Telmatobius. Ibid. 31:431-449. Cei, J. M. 1973a. Herpetología Patagónica—VI. pansión adaptativa de los telmatóbidos de la Los Liolaemus del grupo fitzingeri en Santa Cruz familia Leptodactylidae. La región también y Chubut (Sauria, Iguanidae). Ibid. 32C:447- fue el centro de evolución de las diversas 458. líneas de iguánidos, y actualmente es un cen­ Cei, J. M. 1973b. Herpetología Patagónica—VII. Notas ecológicas y morfológicas sobre Liolaemus tro de especiación de Liolaemus. Cuatro gru­ bibroni y L. boulengeri. Ibid. 32C: 459-469. pos de especies de Liolaemus tienen su centro Cei, J. M. 1973c. Comentarios sobre algunos géneros de dispersión en la Patagonia. Estos han de iguánidos: Diplolaemus, Leiosaurus, Apero- sufrido especiación post-pleistocenica y suces­ pristis y Cupriguanus. Ibid. 32C:269-276. Cei, J. M. 1973d. Distribución geográfica y carac­ iva dispersión en la región, con el resultado teres poblacionales de Cupriguanus fasciatus de muchas líneas de especies afines bio­ (D’Orbigny) (Sauria, Iguanidae). Ibid. 32C: química y ecológicamente bien definidas. 255-262. Cei, J. M. 1974a. Revision of the Patagonian lizards of the Liolaemus elongatus complex. J. Herpetol. LITERATURE CITED 8:219-229. Cei, J. M. 1974b. Herpetología Patagónica—VIII. La altiplanicie entre Primeros Pinos y Río Kilka, Barrio, A. 1973. Una nueva especie de Telmatobius Neuquén. Physis 33C: 183-185. (Anura, Leptodactylidae) procedente del dominio Cei, J. M. 1975a. Herpetología Patagónica—IX. Lio­ austral cordillerano Argentino. Physis 32C:207- laemus goetschi y el conjunto Liolaemus darwini- 213. boulengeri. Ibid. 34C: 199-202. 1979 CEI: PATAGONIAN HERPETOFAUNA 337

Cei, J. M. 1975b. Herpetología Patagónica—X. El Duellman, W. E., Veloso M., A. 1977. Phylogeny conjunto evolutivo de Liolaemus elongatus: aná­ of Pleurodema (Anura: Leptodactylidae): A lisis serológico. Ibid. 34C: 203-208. biogeographic model. Univ. Kansas Mus. Nat. Cei, J. M. 1975c. Herpetología Patagónica—XI. Hist. Occas. Pap. (64): 1-46. Diferenciación serológica de Diplolaemus dar- Estes, R., Reig, O. A. 1973. The early fossil record wini y Diplolaemus bibroni en poblaciones alo- of frogs: A review of the evidence, pp. 11-63 simpátridas. Ibid. 34C:209-210. in Vial, J. L. (ed.). Evolutionary biology of the Cei, J. M. 1975d. Lialaemus melanops Burmeister anurans: Contemporary research on major prob­ and the subspecific status of the Liolaemus fitz- lems. Univ. Missouri Press, Columbia, 470 p. ingeri group (Sauria-Iguanidae). J. Herpetol. 9: Gallardo, J. M. 1962. A propósito de Bufo varie­ 217-222. gatus (Giinther) sapo del bosque húmedo Ant- Cei, J. M. 1975e. Southern Patagonian lizards of artandico, y las otras especies de Bufo neotropi­ the Liolaemus kingi group. Herpetologica 31: cales. Physis 23:93-102. 109-116. Gallardo, J. M. 1964. Los géneros Urostrophus D. Cei, J. M. 1976. Remarks on some Neotropical am­ & B. y Cupriguanus gen. n. (Sauria, Iguanidae) phibians of the genus Alsodes from southern y sus especies. Neotropica 10:125-136. Argentina (Anura, Leptodactylidae). Atti Soc. Gallardo, J. M. 1968. Dos nuevas especies de Italia Sci. Nat. Mus. Civ. Stor. Nat. Milano 117: Iguanidae (Sauria) de la Argentina. Ibid. 14:1-8. 79-84. Gallardo, J. M. 1970. A propósito de los Telma- Cei, J. M. 1977. Serological relationships of the tobiinae (Anura, Leptodactylidae) patagónicos. Patagonian toad Bufo variegatus (Giinther). Ibid. 16:73-85. Serol. Mus. Bull. 52:2. Gasparini, Z. B., BÁez, A. M. 1975. Aportes al cono­ Cei, J. M. 1979. Amphibians of Argentina. Monit. cimiento de la herpetofauna terciaria de la Argen­ Zool. Italiano Monog. Zool. (in press). tina. Acta I Congr. Argentino Paleontol. Biostrat. Cei, J. M., Castro, L. P. 1973. Taxonomic and sero­ 2:377-413. logic researches on the Phymaturus patagonicus Harrington, H. J. 1962. Paleogeographic develop­ complex. J. Herpetol. 7:237-247. ment of South America. Bull. Amer. Assoc. Petrol. Cei, J. M., Castro, L. P. 1975. A serological con­ Geol. 46:1773-1814. tribution to the taxonomic status of Cupriguanus, Lynch, J. D. 1971. Evolutionary relationships, oste­ a South American genus of iguanid lizards. Serol. ology, and zoogeography of leptodactyloid frogs. Mus. Bull. 51:5-6. Univ. Kansas Mus. Nat. Hist. Mise. Publ. (53): Cei, J. M., Roig, V. G. 1966. Caracteres biocenóticos 1-238. de las lagunas basálticas del oeste de Neuquén. Lynch, J. D. 1978. A re-assessment of the telma- Bol. Est. Geog. Univ. Nac. Cuyo 13:182-201. tobiine leptodactylid frogs of Patagonia. Univ. Cei, J. M., Roig, V. G. 1968. Telmatobiinos de las Kansas Mus. Nat. Hist. Occas. Pap. (72): 1-57. lagunas basálticas de Neuquén (Anura, Lepto­ Morello, J. 1958. La provincia fitogeográfica del dactylidae). Physis 27:265-284. monte. Opera Lilloana 2:5-115. Cei, J. M., Roig, V. G. 1975. A new lizard from the Peters, J. A., Donoso-Barros, R. 1970. Catalogue Sierra del Nevado Mountains, central Argentina. of the Neotropical Squamata II. Lizards and am- J. Herpetol. 9:256. phisbaenians. Bull. U.S. Natl. Mus. (297): 1-293. Cei, J. M., Scolaro, J. A. 1977. Herpetología Pata­ gónica—XIII. La identidad de Liolaemus goet- Ruiz Leal, A. 1972. Los confines boreal y austral schi y de la forma melanops del grupo Liolaemus de las provincias patagónicas y central respectiva­ fitzingeri, en Río Negro y Chubut. Physis 36C: mente. Bol. Soc. Argentina Bot. 13 (Supple­ 225-226. ment): 89-118. Chaffee, R. G. 1952. The Deseadan vertebrate Schaeffer, R. 1949. Anurans from the early Terti­ fauna of Scarritt Pocket, Patagonia. Bull. Amer. ary of Patagonia. Bull. Amer. Mus. Nat. Hist. Mus. Nat. Hist. 98:509-562. 93:47-68. Donoso-Barros, R. 1966. Reptiles de Chile. Ed. Scolaro, J. A., Cei, J. M. 1977. Herpetología Pata­ Univ. Chile, Santiago, 458 p. gónica—XII. Los iguánidos del grupo Liolaemus Donoso-Barros, R., Cei, J. M. 1971. New lizards fitzingeri en Chubut: datos serológicos y posición from Patagonian volcanic tablelands of Argentina. taxonómica. Physis 36C: 219-223. J. Herpetol. 5:89-95. Tihen, J. A. 1962. A review of the New World fos­ Duellman, W. E. 1966. The Central American sil bufonids. Amer. Midi. Nat. 68:1-50. herpetofauna: An ecological perspective. Copeia Vuilleumier, F. 1968. Origin of frogs of Pata­ 1966(4):700—719. gonian forest. Nature 219:87-90. 338 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 7

APPENDIX Appendix 13:1.—Distribution of species and subspecies of amphibians and reptiles in ten districts of the two major faunal regions in Patagonia. Northern Region Southern Region

Sello

Ecotone

Lowlands

de

Area

Taxon Steppe Lago

Somuncura

Central Highlands

District de Southern del

Sub-Andean Volcanic Monte-Patagonian Meseta Humid Monte Patagonian Coastal Meseta Altiplano Anurans Alsodes gárgola gárgola______— — — + — — — — — — Alsodes gárgola neuquensis------— — — + — — — — — — Atelognathus nitoi ______— — — + — — — — — — Atelognathus patagonicus ------— — — + — — — — — — Atelognathus praebasalticus praebasalticus — — — — + — — — — — — Atelognathus praebasalticus agilis ------— — — + — — — — — — Atelognathus praebasalticus dobeslawi ------— — — + — — — — — — Atelognathus praebasalticus luisi ------— — — + — — — — — — Atelognathus reverbera ------— — — — + — — — — — Atelognathus solitarius ------— + — — — — — — — Leptodactylus ocellatus ______— + Odontophrynus occidentalis ______+ + Pleurodema bufonina ______— + + + + + — + + — Pleurodema nebulosa ______+ + Pleurodema thaul ______— — — + — — — — — — Somuncuria somuncuriensis______— — — — + — — — — — Bufo arenarum ______+ + Bufo spinulosus ______— — + + — — — ■ — — — Lizards Homonota darwinii ------— + + + + + + + + — Homonota hórrida ______+ + Ctenoblepharis donosobarrosi ______+ + Diplolaemus bibronii ------— — + — — + + + + — Diplolaemus darwinii ______— — + + + + + — — — Diplolaemus leopardinus ------— — — + — — — — — — Leiosaurus bellii ______— + + — — — — — — — Liolaemus archeforus archeforus ------+ Liolaemus archeforus sarmientoi______— — —. — — — — + — — Liolaemus austromendocinus ______— + Liolaemus bibronii ______— + + + + + + — — — Liolaemus boulengeri ______- + + + + + + — — — Liolaemus buergeri ______— — — + — — — — — — Liolaemus ceii______— — + + — — — — — — Liolaemus chilensis ______— — — + — — — — — — Liolaemus darwinii______+ + Liolaemus elongatus elongatus ______— — + + — — — — — — Liolaemus elongatus petrophilus______— — — — + — — — — — Liolaemus fitzingeri fitzingeri ______- - - - - + + - + — Liolaemus fitzingeri canqueli ______— — + — — — — — — — Liolaemus fitzingeri melanops______+ + Liolaemus gracilis ______+ + Liolaemus kingii______— — — — — + + — + — Liolaemus kriegi ______— — + + — — — — — — Liolaemus lineomaculatus ______- - - + - - - + - + Liolaemus magellanicus ______— — — — — — — + — — Liolaemus rothi ______— — + + + - - - — - 1979 CEI: PATAGONIAN HERPETOFAUNA 339

Appendix 13:1 (Concluded). Northern Region Southern Region Sello

Ecotone

Lowlands

de

Area

Taxon Steppe Lago

Somuncura

Central Highlands

District de Southern del

Monte Volcanic Monte-Patagonian Meseta Sub-Andean Patagonian Altiplano Coastal Humid Meseta Liolaemus ruizleali ______Liolaemus tenuis ______Phymaturus palluma ______Phymaturus patagonicus patagonicus______Phymaturus patagonicus indistinctus ______Phymaturus patagonicus somuncurensis ----- Phymaturus patagonicus zapalensis ------Pristidactylus fasciatus ______+ + + + - _____ Proctotretus pectinatus______+ ______Vilcunia sylvanae ______- - + Mabuya frenata ______+ - ______Cnemidophorus longicauda ______+ + - - - _____ Amphisbaenians Amphisbaena angustifrons______._ + _ _ - _____ Snakes Leptotyphlops australis ______+ + Leptotyphlops borrichiana______+ + Elapomorphus bilineatus ______— + Leimadophis sagittifer ______+ + — + — — — — — — Lystrophis semicinctus ______— + Philodryas burmeisteri ______+ + Philodryas patagoniensis ______— + Philodryas psammophideus ______+ + Pseudotomodon trigonatus______+ + Micrurus frontalis ______— + Bothrops ammodytoides ______4- + + + — + + — — — Turtles Geochelone donosobarrosi ______— + Total Taxa ______17 32 18 29 11 9 8 6 4 3 Total Species 17 32 17 25 11 9 8 6 4 3