ECOLOG ICAL AND ZOOGEOGRAPHIC RELAT IONSHIPS O F AMPHIBIANS AND 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 (8 anuran amphibians,4 , 23 , 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 , 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 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 . 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 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. Two of these, Morafka (1977:f 55) includes species in all three categories Hemidactylus turcicus all.d Bufo marinus, are introduced (aquatic, semiaquatic, and riparian) as defined above. exotics. Hemidactylus is well;established in edificarian There were three major geographic categories estab- habitats in the town of Cuatro Ci6negas de Carranza, as it lished for the analysis: Chihuahuan, Tamaulipan, and is elsewhere in northeastem M6xico and southem . ubiquitous. The Chihuahuan fauna is the core desert fauna, Bufo rnarinus was repbrted by Schmidt and Owens (1944) and includes species that have distributions centered in the from the Marsh collection. but has not been taken since. arid northern Mexican Plateau. Tamaulipan species con- A third species, Drymarchon corais, may also represent an stitute the lowland faunal element; distributions of species introduction. It is tentatively included here on the basis of in this group center in the Tamaulipan or Austro-riparian a single specimen. berlandieri taken in the zone east of the Cuatro Ci6negas Basin. Most species in the vicinity of Cuatro Ci6negas de Carranza are regarded as ubiquitous category have distributions that extend from escaped pets (W. L. Minckley, pers. comm.). The native coast to coast (or nearly so) in the United States, south- herpetofauna includes 8 species of anuran amphibians, ward onto the Mexican Plateau. rssuE I,1984 ECOLOGICAL AND ZOOGEOGRAPHIC RELATIONSHIPS OF AMPHIBIANS AND REPTILES 5l

Four further subcategories are recognized in the Chihua- bution of one of these Chihuahuan Dbsert endemics, the huan category. The first includes forms endemic to the gecko Coleonyx breuis, The third subcategory includes Cuatro GiCnegas Basin, but with affinities in the Chihua- species with distributions in both the Chihuahuan Dcsert huan fauna. The second includes species geographically and tlre western Great Plains. Distribution of Eumeces limited to the Chihuahuan Desert. Figure 1 shows distri- obsoletus (fig. 2) illustrates this pattern. The fourth

Table 1. Arnphibians and reptileo of the Cuatro Ci6neg'as Basin, with ecological and geographic category assignments.

Ecological Geographic Ecological Geographic Amphibia: Salientia Categorya Categoryb Reptilia: Sauria (continued) Categorya Categoryb

Scaphiopus couchi 1 A-4 Eumeces tetragramrnus 2 B-2 Bufo speciosas I B-3 Genhonotus lugoi* I A-1 Bufo debilis 1 A4 Cn emid o p h oru s in ornatu s I A-2 Bufo cognatus I A4 Cnemidophorus gularis 2 B-2 * Bufo marinus (introduced) Cn em id o p h oru s s c alaris I A-l Bufo punctatus 1 A4 Cnernidophorus tigris 1 A-4 Bufo oalliceps 2 B-2 Hylactophryne augusti I A-4 Reptilia: Serpentes Rana (pipiens group) 4 A-3 (?) Leptotyphlops dulcis I B-3 Reptilia: Testudines Leptotyphlops humilis I A-4 Arizona elegans 1 L4 Trionyx ater* 4 B-1. Coluber constrictor 2 c Trionyx spiniferus 4 B-3 Dryrnarchon corais 2 B-2 Pseudernys scripta* 4 B-1 Diadophis punctatus , c Terrapene coahuila* 3 B-1 Elaphe obsoleta 2 B-3 Elaphe guttata 2 A-3 Reptilia: Sauria Heterodon nasicus I A-3 Hypsiglena torquata I A4 Coleonyx breais A-2 Lampropeltis alterna I A-2 Coleonyx reticulatus A-2 Larnpropeltis getulus I c Hemidactylus turcicus Lanpropeltis triangulum , c (introduced) Masticop his flage llunt 1 c Crotaphytus collaris ; 1\-4 Masticophis taeniatus I A4 Cro tap hy tu s u isliz enii I A4 Pituop his m elano leucus I A4 Holbrookia macalata I A-4 Rhinocheilus lecontei I A-4 Cophosaurus texonus I A4 Sonora setniannulata 1 L4 Phrynosoma cornutum I A-3 Tantilla atriceps t A-2 Phrynosoma modestum 1 A-2 Nerod,ia erythrogaster 4 B-3 Sceloporus couchi , A-2 Nerod.ia rhombifera 4 B-3 Sceloporus magister 1 A4 Thamnophis cyrtopsis 3 A-4 Sceloporus meniatni I A-2 Thamnophis nzorcianus 3 A-4 Sceloporus oliaaceus I B-3 Thamnophis proximus 3 B-3 Sceloporus poinsetti I A-4 Micrurus fuluius 2 B-2 SceloPorus undulatus I c Sistrurus catenatus 3 B-3 Uta stansburiana I A4 Crotalus atrox A-4 Scincella lateralis* 3 B-1 Crotalus lepidus A4 Eumeces obsoletus 2 A-3 Crotalus molossus L-4 Crotalus scutulatus A-4

a l=Desert, 2=Riparian, 3=Semiaquatic, 4=Aquatic; see test for definitions.

A=Ctrihuahuan (l=Endemic, 2=Chihuahuan Desert only, 3=Chihuahuan Desert and western Great Plains, 4=Widespread, desert and plains); B=Tamaulipan (l=Endemic, 2=Gulf Edge, 3=Plains to desert edge); C=Ubiquitous.

Indicates endemic species or subspecies. 52 JOURNAL OF THE ARIZONA-NEVADA ACADEMY OF SCTENCE vol,. 19

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Figruc l. Distribution of. Coleonyx bretis, a Chihuahrnn l)cscrt cndemic ryecicr. Approximatc locetion of the Cuatro Ci€ncgar Bssitr is indicatcd by thc black equarc.

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Figure 2. Distribution of Eameces obsoletus, a Chihuahuan Des€rt and Great Plains opecies. Approximate location of the Cuatro Ci€negas Bacin is indicated by the bbcl' rquarc. 53 ISSUE l,1984 ECOLOGICAL AND ZOOGEOGRAPHIC RELATIONSHIPS OF AMPHIBIANS AND REPTILES

includes species with wider distributions that include much riparian (Table 2) Twenty-two percent of the lizards, and of the arid southwestem United States and nor*rern the same proportion of snake species are in this category. M6xico, and the western Great Plains. An example of this The only riparian amphibian is Bufo ualliceps. type of distribution is ttlatof. Rhinocheilus lecontei (Fig. 3). The semiaquatic category constitutes 9To of the total In the Tamaulipan category three subcategories are fauna (Table 2). Included species are Scincella lateralis and recognized. First is Basin endemics with affinities in the three species of Thamnophls, although T. marcianus might Tamaulipan fauna. The second includes a series of species as easily be classified as riparian. Sistrurus catenatus is with distributions extending around the Gulf of Mdxico also considered semiaquatic, albeit on scanty evidence and penetrating the desert ula extensions of habitat along (Minckley and Rinne L972). Terrapene coahuila is dif' major river systems. Distribution of Bufo ualliceps (Fig. a) ficult to categoize. Although much of its activity is in illustrates this pattern. The third includes species that are semiaquatic habitats (Brown L974), it is also found in distributed westward across the Great Plains, extending to streams and lagunas (Williams et al. 1960, Webb et al. but not penetrating the Chihuahuan Desert. An example 1963). It is here placed in the semiaquatic category. of this distributional pattem is shown in Figure 5, which Aquatic species also account for 9To of the fauna (Table Many species in the last shows the range of. Bufo speciosus. Table 2. Summary of ecological relationships of the group have wider distributions east of the Great Plains than herpetofauna, by major taxonomic groups. does B. speciosus, but all share a characteristic limit at the edge of the Chihuahuan Desert. Desert Riparian Semi- Aquatic ECOLOGICAL RELATIONSHIPS OF THE HERPETO. aquatic FAUNA.-Forty-one (62V") of the 66 native species eco- logically are characteristic of deserts (Table 2). More than Amphibians 6 I 0 1 half of the snake species (58%) and about three-fourtls of Turtles 0 0 I 3 8 the lizards (74%\ are desert forms. Surprisingly, 6 of the Lizards L7 D 1 0 amphibians from the Basin (Scaphiopus couchi, Hylacto- Snakes 18 1 4 2 phryne augusti, and four species of. Bufo) also are cate- goized, as desert species. Totals 4L 13 66 The next richest fauna in the ecological sense is that of (62%) (20%) (e%\ (e%) riparian zones. Thirteen species (20%) arc classified as

aZ-\ \ lj 1'

NORTH A]"IDRICA Scalo

Figure 3. Distribution of Rhinocbeilus lecontei, a widerpread descrt/plainr cpccies. Approximate location of the Cuatro Gi€ncgas Basin is indicated by the bleck rquarc. 54 JOURNAL OF THE ARTZONA-NEVADA ACADEMY OF SCTENCE vol-. 19

t)\ ,--"\r"

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"z'\ Ai or \ ._.c \

NORTH A]'TDRICA Sca,l e

I I

Figure 4. Dictribution of Bufo oalliceps, a Gulf cdge slrccicr. Approximate location of the Cgatro Ci€negac Basin is indicated by the black square.

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Fignre 5. Distribution of. Bufo speciosus, a species that ranges to the edge of the Ghihuahuan Degert. Approximate location of th€ Cuatro Ci6negas Basin i8 indicatedby the black square. rssuE l, 1984 ECOIOGICAL AND ZOOGEOGRAPIilC REI.A,TIONSHIPS OF AMPHIBIANS AND RETTILES cc

2). Thc Rana ap, (pipiens group) is the only aquatic amphi- transitions, the boundary between the Chihuahuan and bian. Trionyx ater, T. spiniferus, and. Pseudemys scripta Tamaulipan faunal zones is particularly sharp in this area. are the aquatic turtlee. There are no aquatic lizards, and Sixteen (38%) of the Chihuahuan species reach an eastem Nerodia erythrogaster and N. rhombifera are the only range limit at the edge of the Basin. More striking are the aquatic snakes. Tamaulipan species, of which 77% (L4 species) reach their To summarize, the Cuatro Cidnegas herpetofauna is western limit in the Cuatro Cidnegas Basin. Among the composed ecologically of about 60% desert species, and aquatic and semiaquatic species only two (Netodia erythro- 40% aquatic, semiaquatic, and riparian species combined. gaster, Rana sp.) penetrate farther west, beyond Cuatro The presence of extensive aquatic habitats in the Cuabo Ci€negas into the (hihuahuan Desert (Conant 1969, 1978). Ci6negas Basin thus accounts for enhancement of herpe- The abruptness of this faunal boundary is emphasized tofaunal diversity by roughly a third. by the distributions of two pairs of species: Scelo' porus oliaacezs and S. magister, and Cnernidophorus GEOGRAPHIC RELATIONSHIPS OF THE HERPETO. gularis and, C. scalans. The two Tamaulipan species (5. oli' FALINA.-Geographic affinities of the Cuatro Ci6negas uaceus atd C. gularis) reach their westernmost range limits Basin herpetofauna are summarized in Table 3. A pre' at the edge of the Cuatro Cidnegas Basin (Puerto Salado), ponderance of the fauna (42 species, 64%) is classified as and are replaced by the Chihuahuan representatives. In the Chihuahuan. As most of these species are wide-ranging, and case of Cnemidophorus gularis and C. scalaris, secondary not habitat-limited, endemism is rare. Only two forms hybridization occurs where the species are in contact (Genhonotus lugoi, Cnemidophorus scalaris Pallidus) fuom (I{alker 1981). the Chihuahuan fauna are endemic to the Cuatro Ci'enegas Basin. Ten species (7 lizards, 3 snakes) are Chihuahuan SUMMARY.-The herpetofauna of the Cuatro Ci6negas Desert endemics, and 5 species (1 amphibiart, 2 lizards, Basin superficially is typical of the Chihuahuan Desert. 2 snakes) have ranges that encompass the Chihuahuan This is bome out by analysis of both ecological and geo' Desert and western Great Plains. The remaining 26 species graphic relationships of the fauna-62%o arc desert forms, (5 amphibians, 8 lizards, 13 snakes) range widely in plains and. 64% have geographic affinities with the C.hihuahuan and deserts. fauna. Although the Tamaulipan fauna constitutes only majority of the Basin endemics have Table 3. Sunmary of geographic relationrhips of the herpe- 27% of the species, a Cuatro Ci6negas Basin tofauna, by m4ior taxonomic grouPs (number of endemic geographic affinities there. The proportion of relict popula- forms in parentheses). herpetofauna includes a high tions (39%), as noted by Milstead (1960), and Morafka large element is even mote Cftrihuahuan Tamaulipan Ubiquitous (1977:157-158). This relictual remarkable in that 5 species, almost LlVo of. the total fauna, taxonomically recognizable level. Amphibians 6 2 0 are differentiated at some relict amphibians and Turtles 0 4 (3) 0 Patterns of differentiation among Basin are fully as varied and Lizards 18 (2) 4 (1) I reptiles in the Cuatro Ci6negas as fishes, mollusks, and crustaceans. At one Snakes 18 8 5 complicated in end of the spectrum is the highly differentiated Tenapene divergent the that its Totals 42 186 coahuih, a species so within be a subject of contro- (64To) (27%) (e%) phyletic relationships continue to versy (Brown 1971, Milstead 1967). An intermediate level The Tamaulipan or lowland element of the fauna is of differentiation is illustrated by Trionyx ater artd Pseu- smaller (18 species, 27%l bur includes 4 Basin endemics dernys suipta taylori. Although T. ater is distinctive mor- (Trionyx oter, Terrapene coahuila, Pseudemys scripta phologically (Webb 1962) it apparently hybridizes freely in taylori, Scincella lateralis ssp.). Five of the Tamaulipan portions of the Basin with invading populations of T. spin' species have distributions that skirt the arid northern iferus (Webb 1973). Pseudemys scripta taylori intergrades Mexican Plateau around the margin of the Gulf of MCxico, with P. scripta elegans in a narrow zone that begins at and extend to or into the Basin aia the drainage of the Puerto Salado and extends downstream perhaps no farther Ff,o Salado. These are Bufo ttalliceps, Eumeces tetragram' than the Sacramento Basin. Legler (1960) desclibed speci- rnus, Cnemidophorus gularis, Drymtchon corais, and mens from Puerto Salado as intermediate between the two Micrurus fulaius. Nine species are more wiilely distributed, subspecies. Turtles from the eastern end of the Sacramento in the Great Plains and eastward, and characteristically Basin (El Carifro de la Montafra) are "typical" P. s. elegans occur in relict populations located as mesic-adapted isolates (M. A. Ewert, pers. comm.). Conant (1969) noted "some at the desert edge. suggestions of local endemism" in Basin populations of The geographic relationships of the herpetofauna also Nerodia erythrogaster. At the other extreme are relictual, relate to the position of the Cuatro Ci6negas Basin at the but appaxently undifferentiated, populations of such extreme eastern edge of the Chihuahuan Desert Province. species as Eumeces tetragrarnrnus, Coluber constrictor, Because of an abrupt topographic boundary at the eastern Micrurus fulaius, and Sistrurus catenatus. These differing edge of the Basin, and resulting climatic and ecological levels of morphological divergence, as Minckley (1969, 56 JOURNAL OF THE ARIZONA-NEVADA ACADEMY OF SCIENCE voL. l9

1978) observed, indicate a "mosaic of differentiation," Wauer, R. H. and D. Riskind, eds., Transactions of the and result from a long history of successive faunal inva- Symposium on the Biological Regources of the Chihua- sions. Reconstruction of this history, which is essential to huan Desert Region. Trans. Proc. U.S. Natl. Park understanding the complex pattern of differentiation, must Sew.3. await comparative studies of genetic divergence of the MORAFKA, DAVID J. 1977. A biogeographical analysis relictual populations. of the Chihuahuan Desert through its herpetofauna. W.Junk, The Hague. vii + 313 pp. ACKNOWLEDGMENTS.-For specimens or data I thank SCHMIDT, ROBERT H,, JR. 1979. A climatic delineation Garnett R. Brooks, the lateJames A. Peters, T, Paul Maslin, of the 'real' Chihuahuan Desert. J. Arid Environ. D. F. Hoffmeister, R. F. Inger, Stephen F. Hale, Max A. 2:243-250. Nickerson, Arthur C. Hulse, W. S. Brown, Ernest A. Liner, SCHMIDT, KARL P. and DAVID W. OWENS. 1944. and M. J. Fouquette, Jr. Field work was supported by the Amphibians and reptiles of northern Coahuila, . University of Colorado Museum, Carnegie Museum of Zool. Ser. Field Mus. Nat. Hist. 29(6):97-115. Natural History, Sigma Xi, and the U. S. Fish and Wildlife WALKER, J. MARTIN. 1981. Systematics of. Cnernid,o- Service. Assistance in the field was provided by A. V. Bian- phorus gularis. I. Reallocation of populations cur- culli, R. B. Bury, A. C. Hulse, T. P. Maslin, and Harald rently allocated, to Cnernidophorus gularis and Cnemi- Schreiber. The manuscript was reviewed by W. L. Minck- dophorus scalaris in Coahuila, Mexico. Copeia 1981 ley, NormanJ. Scott,Jr., and Robert G. Webb. Finally, my (4):826-8a9. special thanks are due W. L. Minckley and Jose (Pepe) WEBB, ROBERT G. 1962. North American Recent soft- Lugo, for freely sharing their incomparable knowledge of shelled turtles (family ). Univ. Kansas the Cuatro Cidnegas Basin and its fauna. Publ. Mus. Nat. Hist. 13(10):429-6f 1. 1973. Trionyx ater. Cat. Amer. Amphib. LITERATURE CITED Rept.:137.1-137.2. W. L. MINCKLEY and J. E. CRADDOCK. BROWN, WILLIAM S. 1971. Morphometrics of Tenapene 1963. Remarks on the Coahuilan box twtle, Tera- coahuila (Chelonia, ), with comments on its pene coahuila (Testudines, Emydidae). Southwestern evolutionary status. Southwestem Nat. 16(2): L7l-184. Nat.8(2):89-99. L974. Ecology of the aquatic box turtle, WILLIAMS, KENNETH L., HOBART M. SMITH and PETE Tenapene coahuila (Chelonia, Emydidae) in northern S. CHRAPLIWY. 1960. Turtles and lizards from Mexico. Bull. Florida State Mus. f 9(1):1-67. northern Mexico. Trans. Illinois Acad. Sci. 53(1-2): CONANT, ROGER. 1969. A review of the water snakes of 3645. the genus Natrix in Mexico. Bull. Amer. Mus. Nat. Appendix 1. Hist. 142(1):1-140. BIBLIOGRAPHY ON HERPETOLOGY OF THE 1978. Semiaquatic reptiles and amphibians CUATRO CIENEGAS BASIN, COAHUILA, MEXICO of the Chihuahuan Desert and their relationships to drainage pattems of the region . Pp. 455-491ln Wauer, ASHE, VICTOR M., DAVID CHISZAR and HOBART M. R. H., and D. Riskind, eds., Transactions of the Sym- SMITH. L975. Behavior of aquatic and terrestrial posium on the Biological Resources of the Chihuahuan turtles on a visual cliff. Chelonia2(4):3-7. Desert Region. Trans. Proc. U.S. Natl. Park Serv. 3. AUFFENBERG, WALTER. 1958. Fossil turtles of the HOLSINGER, J. R. and W. L. MINCKLEY. 1971. A new gents Terrapene in Florida, Bull. Florida State Mus. genus and two new species of subterranean amphipod 3(21:53-92. crustaceans (Gammaridae) from northern Mexico. AXTELL, RALPH W. 196t, Cnemidophorus inornatus, Proc. Biol. Soc. Washington 83:425444. the valid name for the little striped whiptail lizard, LEGLER, JOHN M. f960. A new subspecies of slider witlr the description of an annectant subspecies. tuttle (Pseudemys scriptal from Coahuila, Mexico. Copeia 1961 (2):fa8-f 58. Univ. Kansas Publ. Mus. Nat. Hist. 13(3):73-84. BEHLER, JOHN L. 1971. 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