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Reptilia: Testudines: Testudinidae REPTILIA: TESTUDINES: TESTUDINIDAE Catalogue of American Amphibians and Reptiles. Ernst. C.H. andT.E.J. Leuteritz. 1999. Geochelot~edenticrrlnra. Geochelone denticulata (Linnaeus) Yellow-footed Tortoise Te.s/~ldodenticrilat~~ Linnaeus 1766:352. Type locality. errone- ously listed as "Virginia." Syntypes, two specimens (one a stuffed juvenile. the other shown in Schoepff 1792:plate 28, fig. 1) in the Muscum de Geer: one of which was kept in Upsala and the other in Stockholm (Andersson 1900, Schoepff 1792). Anderson (1900) lists the holotype as NRM De Greer 21 (not examined by the author). ~estudotabulatae Walbaum 1782: 122. Nome11 illegitimum, see FIGURE 1. Geochelonedenticulott~. Photograph by Roger W. Barbour. Comment. Testudo ressellata Schneider 1792:262. Type locality, not given. Type, not designated. Testudo tabulara Schoepff 179356. Type locality, "In Africa australi?." Type. no1 designated. See Comment. Testudo terrestris hmsiliensis: Schweigger 18 12:445. See Com- ment. ~ Testudo terrestris arnericnna: Schweigger 18 12:445. See Com- ~ ment. Testudo terrestris cayer~rzensis:Schweigger 18 12:445. See Com- ment. Testudo terrestris .rurinatnen.si.s: Schweigger 18 12445. See Comment. Testudo hercules Spix 1824:20. Type locality, "sylvis de flumen Solimoens" [Rio Solimoes], Brazil. Holotype, an adult, now FIGURE 2. Geochelone denriculata, plastron. Photograph by Roger W. Barbour. I lost (Hoogmoed and Gruber 1983). Testirdo sc[rlpfa Spix 1824:21. Type locality, "sylvis juxta flumen Amazonum" [Rio Amazon], Brazil. Syntypes, originally five specimens in the Zoologische Staatssammlung Miinchen, of tebral and pleural areolae; yellowish or orange pigment also which four juveniles ( 273810, a shell; and 275310 A, B, C), occurs at the lower edge of each marginal. The plastron is well fluid preserved specimens, are extant (Hoogmoed and Gruber developed. Its upturned forelobe tapers toward the front and is 1983); Hoogmoed and Gruber ( 1983) designated 275310 A, about as long as, but slightly narrower than, the hindlobe, which the lectotype. bears an anal notch. The average plastral formula is: abd > hum Testudo cagado Spix 1824:23. Type locality, "Cagado in campis > fem > gulxpect x an; the paired gulars are thickened, but et nemoribus campestribus Bahiae" [Salvador], Brazil. Ho- do not extend much, if at all, beyond the carapacial rim. Each lotype, an adult, now lost (Hoogmoed and Gruber 1983). gular scute is longitudinally split on the dorsal side only. The Testudo plunccta Gmelin, in Gray 183 1 :9. Nomen nudum. , bridge is long with a moderately sized axillary and a smaller Testudo foveolata Schinz 1833:40. Nomen nudum. inguinal that barely touches the femoral scute. The plastron is Geochelone (Chelonoidis) rabulata: Fitzinger 1835: 122. yellowish brown with darker pigment along the seams. The Geochelone (Geochelone) denriculata: Fitzinger 1835: 122. head is brown with yellow dorsal scales. It is moderate in size Testudo (Chelonoidis) denticulata: Williams 1 950x22. with a short snout and a slightly hooked upper jaw. The large Chelonoidis denticulata: Froes 1957:9. prefrontal scale is divided medially and is followed by a subdi- Geochelone denticulata: Williams 1960: 10. First use of present vided frontal scale; other head scales are small, and yellow to combination. orange with dark borders; the jaws are dark brown. The ante- Geochelone (Chelonoidis) denticulara: Pritchard 1 967:269. rior surface of each forelimb is covered with large yellow or Geochelone (Chelonoides) denticulata: Auffenberg 197 1: 110. orange, non- or only slightly overlapping scales. No enlarged tubercles occur on the thighs, and the tail lacks a large terminal CONTENT. Geochelone denticulatu is a monotypic species. scale. The karyotype is 2n = 52; 28 macrochromosomes (I 6 meta- DEFINITION. Geochelone denticulota is the largest species centric or submetacentric, 12 telocentric or subtelocentric) and of tortoise (to 82 cm straight carapace length) on the mainland 24 microchromosomes. It differs from that of G. carbonaria by of South America. It has an elongated carapace with a shallow centromere placement of one of the smallest macrochromosomes cervical indentation. The lateral sides of the shell are parallel (Bickham and Baker 1976a, 1976b). or slightly bowed, somewhat flared over the hind limbs, and Adult males average slightly longer than females, but the fe- with slightly scalloped posterior marginals and no cervical scute. males have greater average mass. Males have a concave plas- Vertebrals are broader than long with the I st and 5th laterally tron; carapace expanded over the hindlimbs; low, flattened, elon- expanded. Adults have either weak or no growth rings on the gated profile; gular extensions that usually protrude beyond the vertebral and pleural scutes. Eleven marginals are usually anterior carapacial rim; and a longer, thicker tail. Females have present on each side, and the supracaudal is undivided and a relatively domed, compact carapace; flat plastron; and a shorter downtumed. The carapace is brown with yellow to orange ver- gular and tail. DESCRIPTIONS. Descriptions of adults are in Auffenberg 1975, 1979), Pritchard and Trebbau (1984), Siebenrock (1909), (1971), Boulenger (1889), Castafio-Mora and Lugo-Rugeles and Williams (1960). Juveniles were described by Ernst and (198 I), Ernst and Barbour (1989), Goeldi ( 1905), Gray (1 855 Barbour (1989) and Gray (1 866). Descriptions of eggs are found [I 8561, 1870, 1873b). Hagan ( 1968), Luederwaldt (1926), in Snedigar and Rokosky (1 950). Auffenberg ( 1966) described Medem et al. (1979), Moreira (1991). Miiller (1987), Miiller the wrist and forehand bones, Gray (1869, 1873a) various and Hellmich (1936), Murphy (1997), Pritchard (1964. 1967, skeletal features, and Williams (1950b) the central articula- 691.3 tions of the cervical vertebrae. Sexual dimorphism is de- 197 1; Bour 1980; Castaiio-Mora and Lugo-Rugeles 1981 ;Coto n scribed by Moskovits (1988). Rojas and Acuiia MesCn 1986; Crumly 1988; David 1994; Hoogmoed and Gruber 1983; King 1989; Mertens and Wermuth LLLUSTRATIONS. Color illustrations of adults are in 1955; Sullivan and Riggs 1967; Wermuth and Mertens 1961, Alderton (1988), Boos and Quesnal (1968), Mktrailler and Le 1977; Williams 1950a, 1952,1960), karyotype (Bickham 1976; Gratiet (1996), Murphy (1997), Pritchard (1967, 1979). and Bickham and Baker 1976a,1976b; Dowler and Bickham 1982; Pritchard and Trebbau (1984). Black and white illustrations Sampaio et al. 1971), zoogeography (Pritchard 1979). size of adultsor theshell are in Dowling (1961), Ernst and Barbour (Sajdak and de Barros-Molina 1991), longevity (Dowling 1961, (1989), Hagan (1968), MCtrailler and Le Gratiet (1996), Moreira Oliver 1953), cervical vertebrae (Williams 1950b). eye (1991), Miiller and Hellmich (1936). Oliver (1953), Pough et (Underwood 1970), head muscles (Lakjer 1926), thymus gland al. ( 1998). Paull(1983), Pritchard and Trebbau (1 984), Snedigar (Bockman 1970, Van Bemmelen 1888),parathyroid gland (Van and Rokosky (1950), Sowerby and Lear (1872), Spix (1824). Bemmelen 1888), rostra1 pores (Winokur and Legler 1974), Wermuth and Mertens (l961), and Williams (1960). Juveniles mental glands (Winokur and Legler 1975). breathing (Bellamy or hatchlings are illustrated in color or black and white in and Petersen 1968, Lenfant et al. 1970). cutaneous gas exchange Calmonte (1968), MCtrailler and Le Gratiet (1996), Pritchard (Jackson et al. 1976), metabolism (Benedict 1932),serum pro- and Trebbau (1984), Williams (1960), and a color photograph teins (Newcomer and Crenshaw 1967). hemoglobin (Sullivan of eggs is in Pritchard and Trebbau (1984). Sexually dimor- and Riggs 1967), blood gases (Rahn and Garey 1973), blood phic characters were shown by Snedigar and Rokosky (1 950). buffering (Lenfant et al. 1970, Rahn and Garey 1973), sleep Head scalation is presented in Hagan (1968), MCtrailler and (Monnier 1980, Walker and Berger 1972), courtship Le Gratiet (1996), Mlynarski (1959), and Williams (1960). The (Auffenberg 1964, 1965, 1977; Beltz 1954; Carpenter and skull is illustrated in Auffenberg (197 I), Wermuth and Mertens Ferguson 1977; Pough et al. 1998; Snedigar and Rokosky 1950), (1 961), Gaffney (1979), and Ruckes (1937). Auffenberg (1966) nesting (Dixon and Soini 1977, Ehrenfeld 1979, Medem et al. illustrated the wrist and forehand bones, Gray (1873a, 1873b) 1979. Mowbray 1966, Pough et al. 1998, Snedigar and Rokosky showed the plastral bones, and Hoffmann (1882 [1883]) the 1950). eggs (Castaiio-Mora 1985; Ewert 1979, 1985; Medem pelvic girdle. The karyotype was illustrated by Bickham and 1960; Medem et al. 1979; Moreira 1991; Mowbray 1966; Baker (1976b). Mttrailler and Le Gratiet (1996) presented a Snedigar and Rokosky 1950; Watson 1969; Wilbur and Morin color photograph of habitat and a black and white photograph 1988), hatchlings and juveniles (Calmonte 1968, Medem et of copulation. al. 1979, Watson 1969), social behavior (Auffenberg 1965, 1969, 1970, 1977; Beltz 1954; Carpenter and Ferguson 1977; DISTRIBUTION. Geochelone denticulatu is a denizen of Pough et al. 1998). vocalizations (Auffenberg 1964, 1965; tropical evergreen and deciduous rainforests. It ranges from Campbell and Evans 1967; Devertevil 1858), habitat rn southeastern Venezuela through the Atlantic lowlands of the (Auffenberg and Iverson 1979, Medem 1960, Obst 1986), popu- Guianas to Brazil. where it occurs throughout the Amazon Ba- lations (Moreira 199 I, Moskovits 1988),sex ratio (Auffenberg sin to eastern Ecuador and Colombia. northern and eastern
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