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ECOLOG ICAL AND ZOOGEOGRAPHIC RELAT IONSHIPS O F AMPHIBIANS AND REPTILES OF THE CUATRO CIENEGAS BASIN C.J. McCOY Curator Section of Amphibians and Reptiles Carnegie Museum of Natural History Pittsburgh, Pennsylvania 1 521 3 ABSTRACT The herpetofauna of the Cuatro Ci6negas Basin consists of 66 native species (8 anuran amphibians,4 turtles, 23 lizards, 31 snakes) and two introduced species. Ecologically 4L (62%) are desert species, 20% riparian, 9% semiaquatic, and 9% aquatic. Geographic relationships are predominantly with the Chihuahuan Desert (42 species,64To). The Tamaulipan (low- land) fauna consists of 18 species (27%), of which four are endemics. The fauna includes 39% mesic-adapted relict popula- tions, and 10% endemics. The endemics range from poorlydifferentiated subspecies to strongly divergent species, indicating long persistence of the aquatic habitats and their sequential colonization by waves of invading faunal elements. RESUMEN La herpetofauna del Bols-on de Cuatro Ci6negas consiste en 66 especies nativas (8 anfibios anuros,4 tortugas,2Slagar tijas, 31 culebras) y 2 especies introducidas. Ecologicamente 4L (62%) son especies de€rticas, 20% son especies riberefias 9% son semiacu6ticas, y 97o son acu6ticas. Las relaciones geogr-aficas son principalmente con el desicrto chihuahuense (42 especies, 64%). La Fauna tamaulipensc (tierra baja) consiste en 18 especies (27%\,de las cuales cuatxo son end6micas. Dicha fauna incluye 39% poblaciones relictas que est-an adaptadas a condiciones hdmcdas y L0% end6micas, Las endiemicas varian entrc subespecies poco diferenciadas a especies que son muy divergentes. Esto indica persistencia larga de los h6bitats acu6- ticos y su colonizaci6n en succsi-on por oleadas de elementos faunTsticos invasores. INTRODUCTION.-The Cuatro Cidnegas Basin is an inter- (until historic times), elements of the aquatic fauna have montane, closed-drainage valley in the central part of the undergone explosive adaptive radiation and speciation State of Coahuila, Mdxico (approximately 27o N by 1040 resulting in swarms of endemic forms. The groups of W). The Basin is small, roughly 30 by 40 km, and its floor aquatic organisms that have been studied (reviewed in lies at an averag€ 740 m elevation. Surroundingmountains Minckley 1969, 1978, Holsinger and Minckley 1971, and rise to elevations near or exceeding 2,500 m. Geology and papers in this volume) show varying degrees of differentia- physiography of the region were thoroughly reviewed by tion, ranging from strongly differentiated autochthonous Minckley (1969). species or species swarms, to undifferentiated Populations Botfr climatically and geographically the Cuatro of widespread species (outside the basin). This recurring Cidnegas Basin lies within the Chihuahuan Desert, as pattern of differentiation reveals two asPects of aquatic defined by Morafka (1977), and Schmidt (1979). The habitats in thc Cuatro Ci6negas Basin: their persistence dimate is arid, as the highly seasonal precipitation averages through time, and their sequential colonization by waves less than 200 mm annually, and daytime temperatures in of invading faunal elements. summer may exceed 44o C. Vegetation and fauna in the The aquatic herpetofauna includes one strongly dif- Cuatro Cidnegas Basin superficially appear typical of an ferentiated species, several endemic subspecies, several arid intermontane basin at the eastern edge of the plateau relict populations, and others that are continuous with section of the Chihuahuan Desert, but this impression is and indistinguishable from populations in the remainder vastly misleading. From t}le floor of the Cuatro Cidnegas of the Rfo Grande drainage. The overall result is super- Basin a series of major springs arise, and feed a hydrologic position of a rich aquatic and semiaquatic faunal element, system that includes a spectrum of aquatic habitats ranging including both differentiated endemics and relictual popu- from flowing rivers to ponds, lakes, and marshes. More- lations, on an otherwise typical Chihuahuan Desert herpe' ovcr, these aquatic habitats are apparently of griat anti- tofauna. Richness of the herpetofauna is further enhanced quity and permanence. Given combined conditions of by the position of the basin on a major faunal boundary, habitat diversity and permanence, and isolation of the basin that results in juxtaposition or interdigitation of Tamauli' McC,oy, C. J. 19E4. Ecological and Zoogeographic Relationships of Amphibians and Reptiles of the Cuatro Gi-enegas Basin. Journal of the Arizona-Nevada Academy of Sciencc 19:49-59. 50 JOURNAL OF THE ARIZONA-NEVADA ACADEMY OF SCTENCE vol,. 19 pan elements with the Chihuahuan Desert herpetofauna. four turtles, 23 species of lizards, and 3l species ofsnakes These two factors, presence of an aquatic and semiaquatic (Table I ). fauna and overlap with Tamaulipan species, produces Species were categorized ecologically on the basis of remarkable diversity. modal habitat preference of populations in the Cuatro The first biological exploration of the Cuatro Cidnegas Ci6negas Basin, regardless of preferences elsewhere. Four Basin was conducted by E. G. Marsh,Jr., in 1939. Schmidt ecological categories are recognized: (1) Desert, (2) Ripar- and Owens (1944) reported on Marsh's herpetological col- ian, (3) Semiaquatic, and (a) Aquatic. The ecological lections, which included the type-series of the most spec- category to which each species was assigned is indicated in tacular reptilian endemic, the aquatic box turtle Terrapene Table 1. Habitat requirements of species in each category coahuila. For almost 20 years following Marsh's discoveries, are defined as follows: the Cuatro Cidnegas Basin was bypassed by herpetologists Desert (category 1) includes species that are not working in Mdxico, but in 1958 a field party from the limited, in distribution or dispersal potential, to aquatic, University of Kansas visited the basin and initiated the semiaquatic, or riparian habitats. The category admittedly current period of intensive investigation. A member of is broad, and includes many more species than the other that field party, W. L. Minckley, has been the leader in three categories. To establish subdivisions within the desert these studies. Although his interests have been primarily in category, such as grassland, sand, or rock face species, fishes and other aquatic organisms, Minckley and his would obscure the basic faunal unity of included species. students have made significant collections of amphibians Riparian species (category 2) are those which, although and reptiles. My own studies of the herpetofauna date they do not require aquatic or semiaquatic habitats, are from two expeditions to the Basin in 1964. Since then I limited within the Basin ecosystem to areas along water- have made a number of trips to the Basin of 2 to 10 days courses or of high subsurface water. These species were duration, and a longer visit of 6 weeks. Dozens of other categorized by Morafka (1977:f 55) as "facultative riparian herpetologists have also visited the Cuatro Ci6negas Basin associates." Although their dispersal potential is not in the past 25 years, either in search of species of particular limited by stringent aquatic habitat requirements, they interest, or simply lured by prospects of witnessing the tend to be distributed in patterns that reflect drainages adaptational paxadox of an aquatic box turtle. The results and p€rsistent mesic habitats, such as gallery forest and of their research have been published in nearly 100 scien- grassland, Species include the skinks Eumeces obsoletus tific papers and articles (Appendix 1). Nevertheless, despite arrd Eurneces tetragrammus, the snakes Diadophis puncta- this period of intense study, and the collection of thou- tus and Micrurus fulaius, alrd Bufo aalliceps, sands of specimens, some species remain known from only Category three, semiaquatic, includes species that live a handful of specimens and others surely remain to be dis- in marshes, sedge mats, and other such habitats peripheral covered. This analysis, then, must be considered provi- to water courses and lagunas. They are less habitat limited sional, as it is based on an incomplete sample of a complex than aquatic species, may be able to survive seasonal or and elusive fauna. temporary habitat drying, and are capable of dispersal across :ueas of discontinuous aquatic habitat. Among METHODS AND DEFINITIONS.-For this analysis the species in this category are Scincella lateralis, Terrapene herpetofauna is defined as those species that occur natural- co ahuil.a, artd Thamn ophis pro xirnu s ly below 1,000 m elevation in the Cuatro Ci6negas Basin Aquatic species (category 4) require permanent stand- and the annectant Sacramento Basin, to the immediate ing or flowing water. Although some parts of the life east. Above 1,000 m the herpetofauna includes such history (i.e. nesting in oviparous reptiles) may necessitate species as Sceloporus grammicus, Sceloporus ornatus, and emergence, these species live in and disperse only through Gemhonotus liocephalus (among others), that occur permanent aquatic habitats. Examples of species in this throughout the northern Chihuahuan Desert as montane ecological category are the t:,:rtles Trionyx andPseudemys, relicts (Morafka 1977:153). They are not pertinent to and the two species of the snake genv,s Nerodia. It should analysis of the fauna. Within the area defined 68 species of be noted that the riparian herpetofauna characterized by amphibians and reptiles have been collected.

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