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0913 Podocnemis Sextuberculata.Pdf 1 REPTILIA: TESTUDINES: PODOCNEMIDIDAE Podocnemis sextuberculata Catalogue of American Amphibians and Reptiles 913 Iverson, J. B., L. Schneider, and R. C. Vogt. 2017. Podocnemis sextuberculata. Podocnemis sextuberculata Cornalia Six-tubercled River Turtle, Iaçá, Pitiú, or Cupiso Podocnemis sextuberculata Cornalia 1849:13. Figure 1 Mature female Podocnemis sextuber- Type locality: “Fl. Amazonum” [=Ama- culata in Parana da Floresta (a tributary of the zon River], South America. Holotype (a Rio Branco which is a tributary of the Rio Negro), hatchling): Not located; originally in the Roraima, Brazil. Photograph by Richard C. Vogt. Milano Museum (Roger Bour, personal communication). Collected by G. Oscu- CONTENT. No subspecies are recognized. lati (Vanzolini 1977b), date unknown (see Remarks). Not examined by authors. DEFINITION. Podocnemis sextuberculata Podocnemis expansa: Gray 1856:61 (in part). is one of the smaller species of Podocnemis, Podocnemis pitiu Silva Coutinho 1868:150. reaching a carapace length of only 34 cm Type locality: “l’Amazone” [Amazonas, (Vogt 2008a, 2008b, 2008c). The domed cara- Brazil]. Holotype specimen, collector, pace is elliptical and broader behind the cen- and collection date unknown. Not exam- ter. The posterior carapacial rim is serrated ined by authors. in juveniles, but only slightly so or smooth in Bartlettia pitipii Gray 1870:720. Type lo- adults; a cervical indentation may be present. cality: “Lakes of the Upper Amazons.” A blunt medial keel is present on vertebrals Syntypes (Four mentioned by Gray, but V2 and V3. All vertebrals are broader than only three listed by Boulenger [1889] long with V1 and V5 the smallest, and V5 and in the catalog of the British Mu- posteriorly expanded. Surfaces of the carapa- seum of Natural History of London cial scutes are usually smooth in adults and [BMNH]): BMNH 1946.1.22.90 (adult in show few, if any growth annuli. The carapace alcohol), 1947.3.5.82 (adult, stuffed), and is gray to olive brown. The plastron is large, 1947.3.5.83 (adult, skeleton), collected by but does not completely cover the carapacial Edward Bartlett (date unknown). Not ex- opening; the anterior lobe is rounded in front amined by authors. and the posterior lobe is broader posterior- Emys amazonica: Baur 1893:213 (in part). Re- ly. The anal scutes are decidedly tapered and sult of inappropriate synonymy of E. am- much narrower than the femorals; a shallow azonica Spix 1824 (= Podocnemis expan- medial posterior notch is present. A unique sa) with P. sextuberculata Cornalia and character (which gives this turtle both its sci- Bartlettia pitipii Gray. entific and English common names) is the Bartlettia pitipiti: Williams 1954:281. Lapsus. presence of six pairs of prominent tubercles Podecnemis sextuberculata: Medem 1960:347. on the plastron of juveniles. These tubercles Lapsus. occur at the base of the bridge on the pec- Podocnemys sextuberculosa: Voženílek toral and abdominal scutes and at the outer 1981:85. Lapsus. posterior point of the femorals. These protu- Podocnemis sextuber culata: Chegodayev berances disappear with age, although those 2002:82. Lapsus. on the pectoral scutes may persist into adult- 2 ? ? N ? 1000 km ? ? Map 1. Distribution of Podocnemis sextuberculata derived from Ferrara et al. 2017. A single dot may represent multiple adjacent localities; question marks adjacent to dots indicate unverified localities. Figure 2. Mature adult female Podocnemis sex- Figure 3. Mature adult female Podocnemis sex- tuberculata with spotted head pattern. This un- tuberculata. Photograph taken in the Mamirauá usual pattern only occurs in tributaries of the Rio Sustainable Development Reserve, Amazonas, Branco, Roraima, Brazil. Photograph by Richard Brazil by Rafael Bernhard. C. Vogt. 3 Figure 4. Plastron of adult female Podocnemis sextuberculata demonstrating persistent protu- berances on pectoral scutes. Photograph taken in the Mamirauá Sustainable Development Reserve, Amazonas, Brazil by Rafael Bernhard. Figure 5. Plastron views of adult females (top row) and adult males (middle and bottom rows). Females have V-shaped notches in their anal hood. A long intergular separates the gulars. scutes, and adult males have larger U-shaped in- The bridge is not as broad as the width of the dentations in their anal scutes. The plastral color- plastral posterior lobe. Both plastron and ation in males and females can range from solid bridge are yellow to gray or brown. The broad light yellow to cream or mottled brown; the dark- head has a protruding snout and notched er colorations often are stains absorbed from the upper jaw. There is only a single weak ridge water they live in. Photograph taken in the Ma- on the triturating surface of the maxilla. The mirauá Sustainable Development Reserve, Ama- premaxillae separate the maxillae and extend zonas, Brazil by Rafael Bernhard. to the choanal rim. The incisive foramina lie completely within the premaxillae. There is DIAGNOSIS. The body size is small rel- no vomer. The interparietal scale is elongat- ative to other Podocnemis (to only 34 cm ed and widely separates the parietals, which, carapace length; Vogt 2008a, 2008b, 2008c). although also elongated, do not touch be- The species name derives from the presence hind the interparietal. Large subocular scales of three pairs of prominent protuberances are usually present. A deep groove lies be- on the sides of the plastron, and the axillary tween the orbits, and the tympanum is about pair is often still obvious in adults (Williams as broad as the orbit. One or two chin bar- 1954). Several characters can be used to dis- bels are present. The head is olive to reddish tinguish Podocnemis sextuberculata from its brown with cream-colored jaws. The neck is five congeners: the forehead is grooved, the dark gray to olive dorsally but lighter colored masseteric scale does not reach the orbit, the ventrally; limbs are gray to olive. However, interparietal scale usually widely separates the head coloration in the Rio Branco drain- the parietal scales, large subocular scales are age is different: most females retain the juve- present, the jugal bone contacts the parietal nile color pattern of light yellow or light green bone, the vomer is absent, the shell is always markings on the head (see Remarks). Three smooth, the dorsal surface of marginal six is large scales occur on the posterior margin of more than half as wide as long, there are three the hind foot. Males have longer, thicker tails large scales on each hind foot and two chin than do females. barbels are present (Williams 1954). 4 Williams (1954). Cervical vertebrae were de- scribed by Williams (1950), hatchling anato- my by Malvasio et al. (2002), jaw musculature by Schumacher (1954), neural bone variation by Pritchard (1988), plastral anatomy by Ca- dena et al. (2012b) and Gaffney et al. (2006), throat morphology by Magalhães et al. (2014) and Vogt et al. (1998), and comparisons with other species by Müller (1935), Siebenrock (1902), and Williams (1954). The karyotype was described by Ayres et al. (1969), Barros et al. (1976), Bull and Legler (1980), Fantin Figure 6. Normal patterned adult female (left) and dos Santos Monjeló (2011), Gunski et al. and male Podocnemis sextuberculata from Rio (2013), Killebrew (1975), Ortiz et al. (2005), Trombetas, Para, Brazil. Adult males are small- Rhodin et al. (1978), and Ventura et al. (2014). er than adult females. Photograph by Richard C. Vogt. ILLUSTRATIONS. Color photographs were published by Alves et al. (2012), Balensiefer et PHYLOGENETIC RELATIONSHIPS. The al. (2007), Bartlett and Bartlett (2003), Bonin relationships among the six species of Podoc- et al. (1996, 1998, 2006a, 2006b), Castaño-Mo- nemis are still uncertain. A tree based on un- ra (2002), Ceballos et al. (2012), Ferrara et published mitochondrial DNA data was pre- al. (2017), Ferri (1999, 2002), Ferronato and sented by Iverson et al. (2007: erythrocephala Morales (2012), Franklin (2007), Kato (1992), (sextuberculata (lewyana (unifilis (expansa, Koshikawa (1996), Lehr (2002), Nascimen- vogli))))), one based on mitochondrial and to et al. (2012), Obst (1986a, 1986b), Obst nuclear DNA sequences was published by et al. (1984, 1988), Páez (2013), Pritchard Vargas-Ramírez et al. (2008: expansa (vog- (1979), Rogner (1996), Rueda-Almonacid et li (sextuberculata (unifilis (lewyana, eryth- al. (2007), Soini and Cóppula (1995), Vetter rocephala))))), and one based on combined (2005), Vogt (2008a, 2008b, 2008c), and Zhou morphological and molecular data was pub- (2004). Color illustrations were presented lished by Cadena et al. (2012a: expansa (vog- by Bonin et al. (1996, 1998, 2006a, 2006b) li (sextuberculata (erythrocephala (lewyana, and Freiberg (1981). Black-and-white pho- unifilis))))). The molecular phylogeny pub- tographs were published by Carillo de Es- lished by Pereira et al. (2017), based on 13 pinoza (1970), Ernst and Barbour (1989), mitochondrial and nuclear gene loci, is iden- Ferreira (1923), Gaffney et al. (2011), Mitter- tical to that published by Vargas-Ramírez et meier (1975), Neill (1965), Pritchard (1979), al. (2008). and Smith (1979a, 1979b). Black-and-white illustrations were presented by Bonin et PUBLISHED DESCRIPTIONS. General al. (1996, 1998, 2006a, 2006b), Boulenger descriptions were provided by Bonin et al. (1889), Cadena (2011), Castaño Mora (2002), (1996, 1998, 2006a, 2006b), Carvalho et al. Gaffney et al. (2011), Gray (1870), Honegger (2002), Castaño Mora (2002), Ceballos et et al. (1985), King and Burke (1989), Muel- al. (2012), Ernst and Barbour (1989), Hoge ler (1998), Siebenrock (1902), Wermuth and (1952), Honegger et al. (1985), King and Mertens (1961). and Williams (1954). Burke (1989), Mittermeier (1975, 1978), Neill (1965), Pritchard (1967, 1979), Siebenrock DISTRIBUTION. Podocnemis sextubercu- (1902, 1904), Vogt (2008a, 2008b, 2008c), and lata is known in the Amazon River drain- 5 age of northern Brazil, southern Colombia and northeastern Peru (Castro 2007; Iverson A 1986, 1992; Lehr, 2000; Ramo 1982; The IUCN Conservation Monitoring Centre 1988). Dis- tributional information was published for Colombia by Ceballos-Fonseca (2000), Ce- ballos et al. (2012), and Medem (1958, 1960, 1969); for Brazil by Da Silva et al.
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