n REPTILIA: TESTUDINES: TESTUDINIDAE Catalogue of American Amphibians and Reptiles. Ernst, C.H. 1998. Geochelone chilensis. Geochelone chilensis (Gray) Chaco Tortoise Testudo (Gopher) chilensis Gray 1870a: 190. Type locality, "Chili [Chile, South America]. " Restricted to Mendoza. Ar- gentina by Boulenger (1 889) without explanation (see Com- ments). Syntypes, Natural History Museum. London (BMNH), 1947.3.5.8-9, two stuffed juveniles; specimens missing as of August 1998 (fide C.J. McCarthy and C.H. Ernst, see Comments)(not examined by author). Testudo orgentinu Sclater 1870:47 1. See Comments. Testrrdo chilensis: Philippi 1872:68. Testrrrlo (Pamparestrrdo) chilensis: Lindholm 1929:285. Testucin (Chelonoidis) chilensis: Williams 1 950:22. Geochelone chilensis: Williams 1960: 10. First use of combina- tion. Geochelone (Che1onoide.r) chilensis: Auffenberg 197 1 : 1 10. Geochelone donosoharro.si Freiberg 1973533. Type locality, "San Antonio [Oeste], Rio Negro [Province, Argentina]." Ho- lotype, U.S. Natl. Mus. (USNM) 192961, adult male. col- lected by S. Narosky. 22 April 1971 (examined by author). Geochelone petersi Freibeg 197386. Type locality. "Kishka, La Banda. Santiago del Estero [Province, Argentina]." Ho- lotype, USNM 192959. subadult male, collected by J.J. Mar- n cos, 5 May 197 1 (examined by author). Geochelone ootersi: Freiberzm 1973:9 1. E-r errore. Geochelone (Chelonoidis) d1ilensi.s: Auffenberg 1974: 148. MAP. The circle marks the type locality; dots indicate other selected Geochelone ckilensis chilensis: Pritchard 1979:334. records: stars indicate fossil records. Geochelone ckilensis donosoburro.si: Pritchard 1979:335. Chelorroidis chilensis: Bour 1980:546. Geocheloni perersi: Freibeg 1984:30: growth annuli surround the slightly raised vertebral and pleural Chelorioidis donosoharrosi: Cei 1986: 148. areolae. Each side usually has I1 maginals. A single, undi- Chelonoidis perersi: Cei 1986: 148. vided supracaudal scute which is downturned between the pos- terior marginals is present. The well-developed plastron has a CONTENT. No subspecies are recognized. See Comments. deep anal notch. Its forelobe tapers toward the front. and the paired gulars may be slightly divided anteriorly by a notch. The DEFINITION. Adults are 18.0-43.3 cm (greatest known forelobe is longer but slightly narrower than the hindlobe. The length of male, 23.3 cm) in straight carapace length. The oval average plastral formula is: abd > hum >< fem > gul > pect >< carapace is flattened dorsally with descending sides, a cervical an; the paired gulars are thick and do not project much beyond indentation, and a serrated posterior marginal rim. The poste- the anterior rim of the carapace. The bridge is long with an rior marginals are slightly upturned, and no cervical scute is axillary about half as large as the inguinal. The anterior surface present. The first vertebral scute is usually broader than long or of each foreleg is covered with large, angular scales which may its width is at least equal to its length. Other vertebrals are or may not slightly overlap. Several enlarged tubercles occur broader than long, with the fifth laterally flared. Well-defined on each thigh, and the tail ends in an enlarged scale. Like other I;I(;URE. Geoch~,lot~erhilensis (photographs by Jay C. ShaI'lCl-). Geochelone, this species has 52 chromosomes (Bickham and (1982), and Pacheco and Aparicio (1996) for which no speci- Carr 1983). mens are available. Discussions of distribution are in Cei and The carapace may be either totally yellowish-brown or have Castro (1978), Chebez (1994), Laurent and Teran (1 98 I), Lavilla dark-brown or black growth annuli surrounding tan areolae, and and Sirocchi (1991), Ormeiio (1 983), Orrego Arevena (1971, a dark wedge of pigment at the posterior seam of each mar- 1979), Pa~~ll(1983), Richard (1990a, 1990b), Richard et al. ginal. Many Patagonian individuals have dark or dark-ringed (1990), and Scott and Lovett (1975). Distribution maps are in areolae that contrast with the remainder of the cara~acialscutes. Auffenberg (I 969), Buskirk (1993), Cei (1986, 1993), Iverson The plastron varies from uniformly yellow-brown to having a (1986,1992), Pacheco and Aparicio (1 996), Paull(1983), Rich- dark triangular wedge along the seams of each scute. The head ard and de la Fuente (1992b), Walker (1989), Waller (1997), is moderate in size with a short snout and a hooked, bi- or tri- and Waller and Micucci (1 997). Bonin et al. (1996), presum- cuspid upper jaw. The large prefrontal scale is divided longitu- ably by mistake, included a small portion of Brazil. dinally and followed by a large frontal scale which may be ei- ther entire or subdivided; other head scales are small. Head, FOSSIL RECORD. Auffenberg (1971) reported that the limbs, and tail are yellowish-brown. Miocene species Geochelone gringorum (= Testudo gringorum The sexes are difficult to distinguish, but males are smaller Simpson 1942) from the Chubut Valley of Patagonian Argen- and have slightly longer tails. Old males are slate gray with tina is closely related to G. chilensis. He based this on its de- large heads and very enlarged, even columnar antebrachial pressed carapace, which has a round outline when viewed from scales; females and juveniles have apposed, pointed, antebra- above, its slightly bifurcated epiplastron, and its proportionately chial scales (Buskirk 1993). Only male forelimb scales darken long anal scute. Because its holotype (American Museum of and hypertrophy. The interhumeral seam of adult males is more Natural History 3366) can only be separated from G. chilensis than half the length of the interabdominal seam, that of females by its thicker xiphiplastron, Auffenberg (1971) thought may be twice as long. gringorum ancestral to chilensis. He also reported that a group of Plio-Pleistocene tortoise fossils from western Argentina bridge DESCRIPTIONS. General descriptions are in Belmonte the temporal and minor morphological gaps between the Mi- (1991), Boulenger (1889), Buskirk (1 993), Cei (1 986), Chebez ocene G. gringorum and extant G. chilensis, and 1aterAuffenberg (1994), Ernst and Barbour ( 1989), Freiberg (1 938, 197 1, 1973, (1974) referred to these fossils as G. chilensis. De la Fuente 1974, 1981, 1984), Gray (1 870c,d, 1872, 1873), G. Muller and Cabrera ( 1988) reported fossil remains from the Sierra Chica, (1987), Miiller and Hellmich (1936), Paull (1983), Pritchard Cordoba Province, Argentina, and Alderton (1988) reported (1967a, 1979), Siebenrock (1909, 19 12), and Williams (1960). Pleistocene fossils. Other descriptions are as follows: egg and hatchling (Belmonte 199 l), skull (Crumly 1982), shell osteology (Auffenberg 197 1 ), PERTINENT LITERATURE. General accounts are in wrist and forehand bones (Auffenberg 1966), pancreas (Serma Alderton (1988), Auffenberg (1969), Bonin et al. (1 996), Briiner de Esteban 1961), and habitat (Auffenberg and Iverson 1979, (1953), Cei (1986, 1993), Chebez (1994), Emst and Barbour Richard and de la Fuente 1992a). (1989), Fernandez (I 983), Ferrari (l983), Freiberg (1954,1967, 1971, 1972, 1974, 1981, 1984), Gallardo (1974, 1977), ILLUSTRATIONS. Color illustrations of adults are in Bonin Groombridge (1982), Gruss and Waller (1986), Krieg (1951), et al. (1996), Cei (1993), Freiberg (1981), G. Muller (1987), G. Muller ( 1987), N. Muller ( 1987), Orrego Aravena ( 197 I), Pritchard (1967a, 1979), and Rogner (1996). Black and white Pacheco and Aparicio (1996), Philippi (I 872), Pritchard (1967a, photographs or drawings of adults or shells are in Auffenberg 1979), Rogner (1996), Rottman (1969), Rust (1938), Walker (1969), Belmonte (1991), Buskirk (1993), Chebez (1994), (1 989), Waller et al. (1989), and Wermuth (1 967). Other papers Crumly (1989), Ernst and Barbour (1989), Freiberg (1967,1971, are listed by topic as follows: evolution and relationships (Bour 1972, 1973, 1974, 1981, 1984), Muller and Hellmich (1936), 1985; Coto Rojas and Acuiia Mesen 1986; Crumly 1982; Marlow Paull (1983), Rust (1938), Waller (1997), Waller and Micucci and Patton 1981; Williams 1950, 1952, 1960), taxonomy (1 997), Wermuth and Mertens (1 96 l), and Williams (1 960). (Buskirk 1993; Crumly 1988; David 1994; Gray 1870b, 187 1, Hatchlings or juveniles were illustrated in black and white by 1873; Mertens and Wermuth 1955; Miiller and Hellmich 1936; Belmonte (1991), Buskirk (1993), and Freiberg (1971, 1972). Pritchard 1979; Pritchard and Trebbau 1984; Sclater 1870; Eggs were shown by Belmonte (1991) andFreiberg (1981). The Wermuth and Mertens 196 1, 1977), rostra1 pores (Winokur and skull was illustrated in Cei (1986). Bones and scales of the Legler 1974), mental glands (Winokur and Legler 1975), brain shell were illustrated by Cei (1986), Freiberg (1 938, 198 I), chemistry (Juorio 1969), olfaction (Trapani et al. 1977), adre- Siebenrock (I 9 12), and Wermuth and Mertens ( I96 1). Belmonte nal gland (Jones 1957), clitoris (Obst 1986), pancreas (Serma (199 1 ) included a drawing of the foreleg, and Auffenberg (1 966) de Esteban 1961), blood (Rabito et al. 1972; Troiano and Vautier presented a drawing of the wrist and forehand bones. Auffenberg 1986), renin-angiotensin system (Noll y and Fasciolo 1972), (1969) showed the habitat. The burrow or pallet were illus- life history (Wilbur and Morin 1988), reproduction (Buskirk trated by Auffenberg (1969) and Waller and Micucci (1997). 1993, Paull 1983), courtship (Buskirk 1993, Fernandez and Food plants were pictured by Waller and Micucci (1997). Charts Basso 1992), vocalizations (Fernandez and Basso 1992),