(en Scientific Papers Natural History Museum The University of Kansas 3 1 March 2005 N u mber 37: 1 -21 The Herpetofauna of Upper Niger National Park, Guinea, West Africa By UBi-<'^'' Eli Greenbaum' and John L. Carr- ' Division of'Herpctology, Natural Hiftoiy Museum & Biodivcrsiti/ Research Center, and ^^^: r^^. i"~i' Department of Ecology and Evolutio)uiry Biologij, 1345 Jayliawk Bouleimrd, The ilniversiti/ ofKah^n^, tawrence, Kansas 66045-7561, USA. - Department of Biology, and Museum of Natural History, The University of Louisiana at Monroe, Monroe. Loiiisiaiui 71209- 0520, USA. CONTENTS ABSTRACT 1 INTRODUCTION 2 Acknowledgments 2 MATERIALS AND METHODS 3 Stud> Site 3 Inventory Techniques 4 SPECIES ACCOUNTS 5 DISCUSSION 16 LITERATURE CITED 18 ABSTRACT We conducted a preliminary survey of the herpetofauna in Upper Niger National Park at the end of tlie dry season in 2002. Museum holdings and literature records from localities within the park were combined with our data to obtain an overall picture of the herpetofauna. The park contains a mosaic of forest and sa\'anna that straddles the Niger River. Twenty-two species of amphibians and 34 species of reptiles were recorded from the park; four species of amphibians and two of reptiles are the first records from Guinea. Seven species of reptiles are listed as threatened by the lUCN Red List or CITES, but according to the Global Amphibian Assessment, none of the amphibians is threat- ened. Stomach contents of an arboreal colubrid snake included a species of bat (Mops condylurus), which represents the first confirmed record from Guinea. Conservation of the park is paramount because: 1) it is the only protected area containing dry forest in Guinea; 2) gallery forest provides suitable habitat for amphibian, reptile, and mammal spe- cies otherwise restrictecl to tropical rainforest elsewhere in West Africa; and 3) .several species of threatened reptiles endemic to West Africa are in the park. Key Words: Reptiiia, Amphibia, herpetofauna, Africa, Guinea, conservation, distiibution. © Natural History Museum, Tlu' University of Kansas FmSt M iVT Ll'^rarV '^^'^ '^'^' ^^''^■^"'^^^- Museum of Comparative Zoology Harv'srd University Scientific Papers Natural History Museum The University of Kansas 31 Mcirch 2005 Number 37; 1-21 The Herpetofauna of Upper Niger National Park, Guinea, West Africa By UBRAR^ Eli Greenbaum' and John L. Cakr- ' Dii'ision of Hcrpctolos^}/, Nnfiiinl Histoiy Musciiiii t-< Biodiversity Research Center, and ^"^ ,- - i" Y Department of Eeologi/ and Evolutioiuvy Biolo^^i/, 1345 ]ai/hawk Boulevard, The Utiiversity of Ka'hshs,'Eauirence, Kansas b6045-7561, USA. - Department of Biolo^^i/. and Museum of Natural Histor\/. The Universiti/ of Louisiana at Monroe, Monroe, Louisiaiui 71209- 0520, USA'. CONTENTS ABSTRACT 1 INTRODUCTION 2 AckXOVVLEDGMENTS 2 MATERIALS AND METHODS 3 Study Site 3 Inventory Techniques 4 SPECIES ACCOUNTS 5 DISCUSSION 16 LITERATURE CITED 18 ABSTRACT We conducted a preliminoi'v survey of the herpetofauna in Upper Niger National Park at the end of the dry season in 2002. Museum holdings and literature records from localities within the park were combined with our data to obtain an overall picture of the herpetofauna. The park contains a mosaic of forest and savanna that straddles the Niger River. Twenty-two species of amphibians and 34 species of reptiles were recorded from the park; foiu' species of amphibians and two of reptiles are the first records from Guinea. Seven species of reptiles are listed as threatened bv the lUCN Red List or CITES, but according to the Global Amphibian Assessment, none of the amphibians is threat- ened. Stomach contents of an arboreal colubrid snake included a species of bat {Mops coiuii/lurus), which represents the first confirmed record from Guinea. Conservation of the park is paramount because: 1) it is the only protected area containing dry forest in Guinea; 2) gallery forest provides suitable habitat for amphibian, reptile, and mammal spe- cies otherwise restricted to tropical rainforest elsewhere in West Africa; and 3) se\eral species of threatened reptiles endemic to West Africa are in the park. Key Words: Reptilia, Amphibia, herpetofauna, Africa, Giunea, conser\'ation, distribution. © Naturol 1 listory Museum, Tlu' Univursity ot kcins.is Fm nJI BUFFER '^°y^' ] : ZONE1 .Sidakoro [^,,„_,„ •.._ •Park headquarters V BUFFER ■■-. '^^S... ZONE 2 "•" ,^'^ XV' J KOUYA FOREST Faranahj 4 SciENTHif Pahlks, Naiurai. HisioKV MLShLM,Tni: Univi-.rsity Of- Kansas The vegetation of tho park is a complex mosaic of forest and savanna. We used the classification of vegetation formations of Fleur\'-Brugiere and Briigiere (2001) in describing terrestrial habilats. Thev recognized four t\'pes each of forest and savanna: (1 ) closed dry forest; (2) open forest; (3) bamboo forest; (4) gallery forest; (5) woodland sa\'anna; (6) wooded sawmna; (7) shrub savanna; and (8) herbaceous savamia. The forests are dominated by trees, may be multi-layered, and have little or no grass making up the ground layer or understory. Savannas differ by having a nearl)' continuous grounci layer of grasses and varying types, sizes, and densities of woody plants. Our efforts in 2002 were concentrated in the gallery forest around stream courses and in the dry forests (closed anci open). Aquatic habitats were sampled along the Niger River. At Somoria, there is a substantial rocky outcrop that runs perpendicularly across the river bed. During the low l1ow period at the end of the dry season when we visited, the Niger River consists of more or less distinct pools separated by relatively short, narrow runs. One of these runs connects the major pools above anci below Somoria, and the rest of the width of the river bed consists of exposed bedrock, boulders, and isolated, rain-filled pools among the rocks (Fig. 2). The predominant substrate in the river bed was sand. We also sampleci along the Kofing River, which was completely dry during our visit at the end of the dry season in 2002 (the rains were late that year). We concentrated our efforts in the surrounding gallery forest, but we also searched the dry river bed, which had a clay substrate (Fig. 3). Inventory Techniques Our inventory of the park herpetofauna is based on examination of 283 preserved specimens (214 collected bv ULM expetiitions), dozens of photographs, and a review of the literature. We collected amphibians and reptiles in the park from 19 to 29 Mav 2002 as a part of the ULM expedition. During a second ULM expedition, colleagues made a siiort trip to Diaragbela in the buffer zone of the park on 16 and 17 May 2003. In addition, we gathered data from several other sources in order to enhance our taxonomic coverage. The presence of several species was documented photographically based on live material captured and released, either during our visit or at other times in 2002. We also obtained several specimens from the ichthyological team surveying in other parts of the Mafou Forest in 2002. We examined 12 reptile specimens held in the ecomuseum collection at the park headquarters (referred to as PNHN ecomuseum). This is an uncatalogued collection of specimens that are known to have come from Mansiramoribaya, areas near the park headquarters, or other unspecified localities within the park (D. Brugiere and N. Gerhard, pers. comm.). Photographs of specimens representing species ncit otherwise represented by catalogued vouchers have been deposited in the catalogued collection of color transparencies in the Di\'ision of Herpetology of the Natural History Museum, The Uni\'ersity of Kansas (KU CT). Literature records of several species from localities at the margin of the park, i.e., Faranah and Kouroussa, are referred to in the species accounts. Several species are recorded from the park only on the basis of literature records. Mocquard (1902) reported a collection (Museum natii>nale d'Histoire naturelle, MNHN) of 16 species of amphibians and reptiles fre^m the region of Kouroussa, including six snakes not recorded by us. Several anurans (24 specimens) from Kouroussa were examined, as were photos of two turtle specimens (R. Bour, pers. comm.). Table L List of specific localities and the dates sampled ior amphibians and reptiles in LIpper Niger National I\irk in 2002 and 2003. The dominant habitat type(s) at each site is indicated using the system of terrestrial habitat classification tor the park bv Fleury-Brugiere and Brugiere (2001). Elevation tor all localities is approximately 400 m. Locality Coordinates Date(s) Habitat(s) Park headquarters, ca. 1 km S of Sidakom Somoria Kofing River Drift fence # I Drift fence #2 Sulukudjamba, ca. 10 km downstream from Somoria Mafou Bila Diaragbela Un4'40"N, 10"27'41"VV 10"2,S'49"N, 10"26'36"W 10"28'02"N, 10"2fV,"i2"VV 10"27'34"N, 10"26'12"W 10"28'54"N, 10"26'12"W 10"32'36"N, 10"23'56"W 10'31'()4"N, 10'1)8'37"W 10"21'42"N, 09'34'21"VV 19, 20, 28 May 2002 20-28 May 2002 22-27 May 2002 22-29 Mav 2002 23-28 Mav 2002 23 May 2002 27 Ma\- 2002 Kv-17 May 2003 Closed dry forest Ciallerv forest and perennial river (bed and bank) Gallery forest and dry streambed (seasonal river) Gallery forest of Kofing River Open forest near Somoria Galler\- forest and perennial river (bed and bank) Gallery forest and perennial ri\ er (bed and bank) Galler\" forest and perennial river (bed and bank) Hf.rpetofauna of Guinfa These specimens from MNHN include material in addition to that serving as the basis of Mocquard's (1902) report. Because the snakes were not available for examination, we did not include a questionable record from Mocquard's (1902) paper for Psatiiniophif sibilaiis, which might refer to P. elegans already known from the park. A small collection of four anurans from Faranah from the Museum d'histoire naturelle-Geneva (MHNCJ) was examined. Finally, we examined 27 specimens (14 species) from Zoologisches Forschunginstitut und Museum Alexander Koenig (ZFMK) that were collected in the park during an ornithological survey between November 1996 and February 1997 (G. Nikolaus, pers. comm.). Three juvenile Biifo (NLU 70544, ZFMK 64484-85; 12.2, 17.8 and 20.2 mm SVL, respectively) could not be identified to species and were not included in the species accounts. Amphibians and reptiles were collected during the 2002 and 2003 expeditions by means of visual encounter surveys by day and night. Specific localities, dates, and major habitat types sampled are indicated in Table 1. Several snakes were obtained in gill nets of fishermen or the ichthyological team set in the Niger River. Baited turtle traps set in two different pools of the Niger River near Somoria failed to capture specimens. Forested areas near Somciria were sampled with drift fences. Two drift fences made of 60-cm wide black silt fencing were installed within walking distance of the village of Somoria from 22 to 29 May 2002. One fence was installed in the gallery forest of the Kofing River and consisted of three arms of approximately 10 m length each (drift fence #1, Fig. 3). At SPECIES Species are listed in alphabetical order bv family, genus, and species. Snout-vent length (SVL) and tail length (TL) measurements are presented as means ± standard error with the range in parentheses. Carapace length (CL) is the standard measure used for turtles. Information about color pattern in life is based on field notes, color photographs, or color digital photos of live animals. Gender of specimens was cHetermined from secondary sexual characteristics (e.g., vocal slits in bufonids, gular sacs in hyperoliids), presence of eggs in the abdominal cavity, or direct examination of gonads. Where possible, stomach contents from one specimen of each species were examined. ANURA ARTHr Fig. h. Hypewliu^ niluliilii-, (KU 241900), subadult male, collected from damp leaf litter near park headquarters. Photo; E. Greenbaum. ^:^r*: .ga^.* ^j» _u-« ^^^ ^; Fig. II. A\;iuiuni^tima (CNSHB 108), adult temale, collected on a tree trunk in the village ot Somoria. Photo: E. Greenbaum. ♦•-^•-i. Fig. 13. Hcmnlniiyltis hwckii (NLU 70536) collected on the wall of a hut at park headquarters. Photo: E. Greenbaum. 1 ig. 14. Leptofiiiyho^ fcxncos;,-; (KU 291403) collected in leat litter m 1 ig I ^ I araiuif niloticu^ (KU 241925) collected 5 km F ot Soniona, gallery forest adjacent to Kofing River. Photo: E. Greenbaum. Mafou forest. Photo: E. Greenbaum. Fig. 16. Typhlops puiuiatus (NLU 70569) collected in a pitfall trap in gallery forest adjacent to Kofing River. Photo: E. Greenbaum. Fig. 17. I'luliithiiiiiiiii'. hctcwilcnmtf (NLU 70566) coi trap in gallerv forest adjacent to Kofing River. Photo: E ected in funnel- Grccnbaum 8 Scientific Papers, Natural Hisiokv Misilm, Tm. UMvi-Rsn y cji- Kansas The SVL is 46.4 ± 0.62 (40.6-56.3) mm in 25 adult males, 36.2 and 37.8 mm in two subadult males, and 66.9 mm in one adult female. One subadult male was active at night in dry forest. All other males were in the Niger River bed or adjacent gallery forest. Only 2 of 24 males from Somoria were captured during or immediately after rainfall; the others were calling on nights without rainfall 3-7 ciays later. The majority of males were calling from protected locations, either under boulders near the margin of the riverbed, or within the cover of low overhanging branches of small trees lining a small inlet on the south bank of the Niger River. No females were encountered during 13 days of collecting during May of 2002 and 2003. A female from Faranah (MHNG 1011.87) was previously identified as B. pciitoiii by Ferret (1966); however, it is typical of females of B. uiaciilatu^ in Guinea. Stomach contents of CNSHB 120 included ants (Hymenoptera) and true bugs (Heteroptera). Coloration of males in life: Dorsal ground color dull yellow or olive-green with 2 or 3 pairs of dark brown blotches (commonly outlined in black) on the dorsum; flanks bright yellowish green; parotoid and rictal glands brown or yellow; dark brown blotches on the upper lips and transverse bars on the limbs; venter immaculate white or yellow except for blackish throats in all adult males. Males with nuptial excrescences on the thenar tubercle and Digits 1 and 2. Entire dorsum, limbs, and parotoid glands rugose with asperities on tubercles. With the exception of generally having poorly clemarcated parotoid glands and lacking a vertebral stripe on the dorsum, our specimens agree in morphology, size, and color pattern with those described by Rodel (2000). This species has previously been reported from Guinea by Rodel (2000). Biifo regularis Reuss, 1833 Material: JLC 749-750, CNSHB 86-88, 98-99, 110, ll.S-117, 130, 143, KU 291877-78, NLU 70542-43, 70555, Niger River at Somoria; CNSHB 271-74, NLU 70616-18, Diaragbela; MHNG 1011.85-86, Faranah; ZFMK 6448.3, park headquarters near Sidal\vn gender has a SVL of lcS.4 mm. \n preservative the dorsimi is mottled dull vellow and the \-enter is yellowish cream. This species was listed trom Guinea by Riidel (2000). Hyperoliiis iiasutus Ciinther, 1865 M All Ki \i : MNHN l')04.(U40, Koiiroussa, The color pattern ot the single adult female with a SVL of 22.9 mm is faded, but there are no indications of any dark pigment on the body, and the snout is elongate and blunt (Schiotz, 1999:fig. 165). Rodel (2000) noted this species trom Guinea. Hi/pcroliiib iiitidiiiiis Peters, 1875 (Fig. 6) Maiikiai: KU 291900, park headquarters.; MNHN 1904.0442-43. 1999.8148^9, Kouroussa; ZFMK 64486, unknown locality within the park. The SVL is 24.2 + 0.68 (22.6-26.0) mm in fi\ e adult males and 20.1 mm in one subadult male. A subadult male was collected at night as it emerged from forest-tliior leaf litter in closed drv forest during heavy rainfall. Coloration of subadult in life: Dorsum tan with mottled brown pigment forming \ague indications of a dorsolateral stripe; posterior surfaces of thighs and feet reddish orange; venter immaculate cream; iris bronze; spots of brown pigment on underside of jaw, most concentrated toward margin of jaw. A distincti\'e blunt snout and small rectangular gular flap are present. The coloration and morphology are consistent with male Phase A specimens described bv Schiotz (1967). Rodel (2000) noted that subadults of this species are almost unicolor with a beige to brown dc>rsum. The only published record from Guinea is Kouroussa (Laurent, 1951). Kassiiia sciic;fnlciifiis (Dumeril anci Bibron, 1841) Maiikiai: MNHN 1^)04 II44I, Kouroussa. The color pattern and morphology of one adult female with a SVL of 22.6 mm are consistent with the description b\' Riidel (2000), wlw noted this species from Guinea. Lcptopiiis luiidis (Giinther, 1S69) MxiiKiAi: EBG 833, CNSHB 144, Niger River at Somoria; NLU 70570, Somoria. The SVL is 26.2 and 34.0 mm in two adult males and 38.9 mm in one adult female. The smaller male was calling on top of leaves of a plant and the larger male was calling from a cre\ice in a large boulder near the margin of the ri\er bed. The female was on a rock near ground level in the village, no more than about 100 m from the Niger Ri\'er. All specimens were collected at night. The color pattern and morphology are consistent with descriptions bv Schiotz (1967) and Rodel (2000). This species was reported from Guinea bv Rodel (2000). PiPlDAE Siliirann twpicalif Gray, 1864 Maiikiai: .MNHN lM04,n44n-49, kouroussa. Four adults of unknown gender have SVLs of 33.8—14.8 mm (40.5 + 2.49). The color pattern and morphology are like that described b\' Riidel (2000). This species was recorded from Guinea by Arnoult and Lamotte (1968). Rani DAE Aniiiirniin foiiciisi^ Rodel and Bangoura, 2004 Maiikiai: ZFMK 644SI-H2, stream ca. 50 m from road leading to park headquarters Two adult males (60.1 and blA mm SVL) that were calling fmm a stream in November (G. Nikolaus, pers. comm.) ha\'e numerous small, whitish tubercles on the dorsum. These specimens were listed as paratypes in the species description (Riidel and Bangoura, 2004). Aiiniirnna galamciisis (Dumeril and Bibron, 1841) Mmikim: MNHN 1902.0021, 1904.0428-32, Kouroussa. The SVL is 62.2 mm in one adult male, 50.1 + 3.12 (45.2- 3'-*. 2) in four adult females, and 35.9 mm in one subadult of unknown gender. Mocquard (1902) erroneously reported one of these specimens (probably the adult male [MNHN 1902.0021] based on his measurement of (-i3 mm SVL) as Pti/clmdciia loii'^iivstyib. Aiiniiyaua gahiDiciifis has not been repc^rted from Guinea (Riidel, 2000). Hilddmvidtia oniata (Peters, 1878) (Fig. 7) Mam i;i \i : NLU 70h23, Diaragliela. An adult male (54.1 mm SVL) was collected at night on a dirt road. Stomach contents consisted of anuran skin, presumably its own eaten after ecdysis. In life, the dorsum had alternating pale and dark brown longitudinal stripes edged in cream, with green interspaces. The throat was brown with two bright white stripes extending to the posterior of gular region. The venter of the body was immaculate white. This species was not known previously from Guinea. Hoplobntracliiis occipitalis (Giinther, 1858) (Fig. 8) MAitKiAi : NLU 70545, 70578, Niger Ri\ er at Somoria: CNSHB 278, NLU 70624, Diaragbela. An adult male (85.2 mm SVL) and three adult females (74.8, 80.9, and 82.9 mm SVL) were collected at night. Specimens at Somoria were either in or near the edges of shallow pools in the ri\er bed. They dove and swam \'igorously when approached. Stomach contents of NLU 70624 included grasshoppers (Orthoptera). Color in life: Dorsum dark green with numerous lilack blotches on dorsum, flanks, and limbs; brown bars edged in black on jaw, interspaces between bars lime green; lime- green transverse stripe just posterior to eyes on dorsum of head; flanks white with gra\ish and black-mottled 10 SciHNTiFic Pai'hrs, Naiurai. HisioK^ MusLUM.Tiii-; Unimrsii^ ()i Kansas blotches; \'enter mostlv white with some black mottling on hind limbs. Numerous tubercles cover the dorsum and flanks. This species was reported from Guinea by Rtidel (2000). Phn/iiobatrniims accraensis (Ahl, 1925) Material: AHR 20(i, EBG W)3-05, 807, ,S12-13, 815, 846, 848^9, JLC 753, CNSHB 101-104, 106, 129, 145-69, KU 291906, NLU 70551-52, 70562, 70568, 70573-74, 70579-84, Niger River at Somoria; NLU 70563, Somoria; JLC 741, NLU 70557, Sulukudjamba. Four adult females (20.0 + 0.61 [18.5-21.11 mm SVL) and fifty-five unknown gender (15.3 + 0.25 [ 1 1 .6-19.0] mm SVL) were collected during the day and at night; most were among dead leaf piles at the bank of the major river pools, or under rocks. Stomach contents from EBG 813 included reproductive termites (Isoptera). Color in life: KU 291906 — dorsum pale brown with lime-green \ertebral stripe; black lateral stripe from posterior edge of eye to inguinal region; flanks below stripe white. Dorsum smooth or with numerous prominent tubercles; supratympanic ridge barely visible in smaller specimens to distinct in large females; most without enlarged toe tips. The color pattern and morphology are consistent with the virtually indistinguishable taxa Phn/nobafrachiif accnu'iisis and P. latifiviif, but Guibe and Lamotte (1963), Bohme (1994a), and Rodel (2000) noted larger sizes for P. accraensis (ca. 24 mm SVL). Moreover, P. aiX-raciisis has enlarged toe tips and a preference for rainforest or anthropogenic habitats (Rodel, 2000). Phrynobalrachus latifrous reportedly congregates at the edges of rivers during the drv season (Rodel, 2000), which is consistent with our findings at the park. Based on molecular data, Rodel and Agyei (2003) placed P. latifrous into the synonymy of P. accraensis, and we use the latter name for the specimens described herein. This species was reported from Guinea by Rodel (2000). Phnpiobiitnuiiiis cdlcuratus (Peters, 1863) Mathriai : CNSHB 182, drill temi' « 1; NLU 70388, drift fence S2. Two females ( 1 6.3 and 1 3.3 mm SVL) were collected in pitfall traps in galler\' ^^nd open forest. Stomach contents from NLU 70588 consisted entireU' of ants (1 hmenopti'ra: Formicidae). The entire dorsum is tubercul.ik' .ind a supratvmpanic ridge is distinct; eveiid tdrnick-s (sr;;s(/ Rodel, 2000) are present. Color pattern and morphology are consistent with Rodel (2000), who reported this species from CAiinea. I'liryiu'btitnicliiis fraiuisci Btiulengof, 1912 Material: NLU 70537, park headquarters; EBC 806, 823, CNSl IB 89, 105, 118-19, KU 291910, NLU 70585, Niger River at Somoria. Four adult females (21.1 ± 1.65 [18.5-25.9] mm SVL), and five of unknovsn gender (16.3 + 0.55 [ 14.6-17.9] mm SVL) were collected with Phrynobiitniclni^ ai.crnen>i>, the majority of which were among dead leaf piles on the bank of the major pools of the Niger River. One specimen (NLU 70537) was amid leaf litter in closed drv forest. Stomach contents of KU 29! 410 Included reproductixc termites (Isoptera). Cokir in life: KU 291910 — dorsum pale brown with dark brown and cream blotches. Snout short and blunt; supratympanic fold distinct; tympanum distinct in larger individuals; most with prominent tubercles on dorsum; webbing moderate; tarsal tubercle present; toe tips not expanded. This species has not been recorded from Guinea previously. Plin/nobntrnchiis iiatnlciisis (Smith, 1849) Mmiriai: MNHN 1904.0439, Koiiroiissa. One subadult of unknown gender \\ ith a SVL of 16.2 mm is consistent morphologicalK with the description prcn'ided bv Rodel (20(10), who reported this species from Guinea. Pti/chadciia owfrhi/iichus (Smitli, 1849) Matlriai : CNSHB 107, Niger River at Somoria. An adult male (51.0 mm SVL) was found among large boulders near the Niger River at night. The color pattern and morphokigv are cimsistent with that described by Rodel (2000), who reported the species from Guinea. Pti/chadoia piii}iilio (Boiilenger, 1920) Maukial: AHR 158-159, 198, CNSHB 84-85, 92-96, 170, NLU 70540-11, 70564, KU 291911, Niger River at Somoria. Two adult males (23.0 and 29.h mm S\'L), five adult females (23.9 ± 0.74 [22.8-26.8]) and eight unknown gender (25.1 ± 0.51 [23.0-27.0[) were on rocks in shallow pools of the Niger River at night after a heavy rain; one individual (CNSHB 170) was in leaf litter on the bank of the river. Stomach contents from AHR 159 included arachiiids of the famih' Araneidae. Color in life: Dorsum pale brown with numerous dark brown blotches; limbs with brown bars; thin creamv yellow line from posteritir bolder of e\e to inguinal region. This species was reported from Guinea b\- Riidel et al. (2004). Pti/chiuU'iiii k'lliuu (I'eracca, 1904) (Fig. 9) M Ml HI \i : NLU 70625, Diaragbela. An aihill lemale (4S.2 mm S\'L) \\ as in a dr\' rut on the side ot a road at night. Stomach contents oi this indi\ idual included grasshoppers (Ortln)ptera) t\nt.\ reproductive termites (Isopli'ia). color in lite: Dorsum immaculate dull orange, becoming reddish brown on hind limbs; black stripes from rostrum thiough e\'e aiiti tN'mpanum to midbod\ : hidden parts i>t hind limbs \ aiiegatei.1 black and linu- git'en; \enter HERPbTOlAUNA OF GuiNEA 11 ininviculcite white from chin tn midbod\', lime green on underside oi hind limhs. Until receniK, this species was known as Pti/cluidnui schuhotzi, wliicli was placed into tlie s\'nonvmv of P. triliiiii by Largen (2001). Tliis species has not been recorded previously from CJuinea. Ptyclunh'iui trinodis (Boettger, 1881) Materiai : MNl IN ri(l2 I)II20, U)04.l)42S-27, Kouroussa; NLU 70626, Diaragbela. The SVL is 54.4 mm in an adult male and 5b.7 ± 1.21 (54.3-59.0) mm in four adult females. The male was on a drv trail at night. Stomach contents of NLU 7062(1 included grasshoppers (Ortlioptera) and reproductive termites (Isoptera). Color in life: Dorsum grayish brown with scattered small black blotches; yellow vertebral stripe along entire length of dorsum; two lateral vellow stripes from tympanum to midbod\' where tiiey become broken until termination at insertion oi hind limli; hidden parts of thighs mottled black anci \'ellow green; orange stripe above tympanum; venter immaculate white. Mocc]uard (1902) described a specimen from Kouroussa that had a SVL of 65 mm; this exceeds the maximum length of 57 mm given by Rodel (2000). TESTUDINES Pelomedusidae PcloDicdtisa siibnifn (Lacepede, 1788) Mmikiai: KU (. T 11413, ca, 20 km ti of Somona near the Niger Ki\er An apparent juvenile (CL ~ 13 cm) was found on a dirt road just south of the Niger River and photographed in February 2002 during the dry season. The turtle presumabh' must ha\e just emerged from aestivation inasmuch as it is covered by dirt. This species is widespread in sa\'anna regions in sub-Saharan Africa and reportedly spends much of the dry season underground (Loveridge, 1941; Ernst and Barbour, 1989). Although known from the surrounding countries of Sierra Leone, Senegal, Mali, and Cote d'lvoire (Iverson, 1992), this is the first record from Guinea. Pcliisios castaneus (Schweigger, 1812) M ATiRi.M.: MNUN 1904.407A-1^, kouroussa. Photographs of two juveniles have been examined. The smallest (CL = 39.7 mm ) appears ti) have grown minimalh' after hatching and the other (CL - 50.2 mm) has one large growth ring on each scute. Coloration in preservative: Carapace nearly uniform pale to dark brown; plastron with straw-yellow to yellow- brown central area grading into dark brown to black areas at the peripiiery of each scute; one specimen vsith entireh' dark brown to black anal scutes. Carapace with a vertebral keel, particularly prominent on Vertebrals 2-4; no indication of plastral hinge; anal notch shallow and wide. This species is known Irom localities in southern Cuinea and sinrounding countries (Loveridge, 1941; Bour, 1983; Iverson, 1992). Testuiiinioae Kiiiixi/s bclliiiiin Cray, 1831 Mmikiai : KU CT 11923-24, ca. Somoria. Two individuals were found at the margin of savanna and forest during daylight in the vicinity of Somoria (P. DeRidder, pers. comm.) in early April 2002 (dry season) and 24 August 2002 (wet season). Two color photographs of the latter show the coloration of the carapace to be typical for the West African subspecies Kiiiixys belUana nogucyi (Lataste, 1886), in that it has a mostly brown carapace with diffuse, scattered dark brown tci black smudges. The individual appears to be an adult (CL ~ 14-18 cm) with worn growth rings on the carapacial scutes and a well-developed carapacial hinge. This species is widely distributed in savannas and has been reported from several localities in Cuinea (Loveridge and Williams, 1957; Villiens, 1958; Broadley, 1989a; Ernst and Barbour, 1989; Iverson, 1992). Trionychidae Trionyx triioii^uis (Forskal, 1775) M.ATFRIAI : KU CT 11414, unknown localit\ within the park. An adult, dried specimen (carapace only, 25.5 cm long) has lemnants of longitudinally arranged, elongate rugosities on the bony carapace and shriveled posterior periphery; the carapace is dark brown to black. David Brugiere (pers. comm.) reported sighting a large, flat turtle with a black shell basking on a rock in the Niger River; if was almost certainly this species (too large to have been a pelomedusid, and C\/cliiiiorhis sciics^iilcnis has a brown or tan, mottled carapace). This species is known from the Niger River in Mali (Iverson, 1992; Joger and Lambert, 1996), but tiiis is the first record for Guinea. CROCODYLIA Croccidyeidae Cwcodi/liis cntaphrnctus Cuvier, 1825 Mmikiai : kU CT 11412, unknown loealit\- within the park, A dried skin of a jinenile is 121 cm in total length and has a head length of 19 cm and a snout length (from anterior margin of orbit to tip) of 11.5 cm; the snout width at the level of the anterior margins of the orbits is 5.4 cm. This species is found in the Mafou River (13. Brugiere, pers. comm.), and has pre\'iouslv been reported from Guinea (Villiers, 1958; Groombridge, 1982; King and Burke, 1989; Messeletal., 1992). Scientific Papers. Nau km Hisior'i Mi si rM.Tiih UmmiKsii'i oi Kansas Crocodylus suchii^ Geoffrov Stiinl-i lilaire, 1807 (Fig. 10) MATERtAL: KU CT 11915, near Mafou Bil.i; l'\ll\ fKimuM'iini specimen from unknown locality within the park. The following measurements are for the juvenile in the ecomuseum: total length 81 .7 cm, head length 111 cm, snout length (from anterior margin of orbit to tip) 6.0 cm, width of snout at level of anterior margins of orbits 3.8 cm. Individuals attain lengths of up to 5 m (D. Brugiere, pers. comm.). Color in life based on KU CT 11915: Dorsum dark olive-brown, grading into tan on flanks; numerous dark brown, yellowish tan, and black spots on dorsum and flanks; venter immaculate cream. We accept the taxonomic change of Schmitz et al. (2003), who demonstrated that West and Central African Nile crocodiles form a clade distinct from those of East Africa. This species is reported from both the Niger and Mafou Rivers (D. Brugiere, pers. comm.), and has previously been reported from Guinea (Villiers, 1958; Wermuth and Mertens, 1977; King and Burke, 1989; Me.ssel et al., 1992). SAURIA ACAMIDAE As^ivfia n^aiiia (Linnaeus, 1758) (Fig. 11) Mati:i4462-64, Sidakoro; KL' CT 1 19(19, unknown localit)' within the park Two adult males (696 and 759 mm SVL; both tails damaged), one adult female (642 mm SVL; 396 mm TL), and one adult of unknown gender (877 mm SVL; 556 mm TL) have 8 or 9 supralabials (5 and 6 or 4-6 in contact with eve), 11 infralabials, 1 or 2 preoculars, 2 postoculars, 2 anterior temporals, 2 or 3 posterior temporals, 17 dorsal scale rows at midbody, 195-200 (males), 191 (female), and 197 (unknown gender) \'entrals, 159 (female) and 158 (unknown gender) subcaudals, and di\'ided anal plates. This species was reported tn>ni Ciiiinea b\- Chabanaud (1921). Hi RPFTOFAUNA OF GuiNEA 15 Tclcscopii< varii's^iitiis (Reinhardt, 1843) Mocquard (1902) described "Lepiodira Pobcgiiiiii," which is now considered a synonym of T. varic\;atus. The single specimen from Kouroussa (692 mm SVL; 123 mm TL) was reported to ha\'e 19 scale rows, 207 \entrals, a divided anal scale, and 59 suhcaudals. Chippaiix d*-''-''-') mapped a locality from Guinea. Toxicodn/ns blniniiii^^ii (Hallovveil, IS44) Materiai : NLU 70349, Somoria. An adult female (1545 mm SVL; 450 mm TL) was climbing through a tree in the village during the day. Stomach contents included three intact individuals of a molossid bat (Mops coiidi/lnnis) and the hindlegs and tail of a lizard in the genus Chaniaclco. The record of Mops couditliirus adds another species to the dix'erse mammalian fauna knov\n from the park (Ziegler et al., 2002), and pro\ides confirmation t^f its presence in Guinea (listed as "unconfirmed" in the checklist of Barnett and Prangley, 1997). hi life, the dorsum and flanks were pinkish tan w ith 39 brown blotches outlined in a creamy tan border along the flanks. The dorsum of the head is brown; the labials are tan with brown outlines, and the venter is white. Scutellation is: 9 supralabials, 12 infralabials, 2 preoculars, 3 postoculars, 2 anterior temporals, 2 posterior temporals, 23 dorsal scale rows at midbodv, 271 ventrals, 129 subcaudals, and anal plate di\'ided. This species was known previously from Guinea (Chippaux, 1999). El APIPAF Dciidivaspis poh/lcpis Giinther, lSb4 Material: ZFMK M45'3-60, Sidakoro, One subadult male (1085 mm SVL; 318 mm TL) collected in March and one adult (head onh-; 230 cm total length in life [G. Nikolaus, pers. comm.]) collected in December have 8 supralabials (4th in contact with e\'e), 1 1 infralabials, 3 preoculars, 3 postoculars, 3 anterior and 4 posterior temporals, dorsal scales in 21 rows at midbodv, 253 ventrals, 121 suficaudals, and a dixided anal plate. This species was reported pre\iousl\' from Cuunea bv Greenbaum et al. (2003). Dciidwaspis i';r;(/;s (Hallowoll, 1844) Materiai: KU CT II'-MO, iinknnwn localilN' within the park A subadult of unknown gender (464 mm SVL; 146 mm TL) has 9 supralabials (4th in contact with eye), 9 infralabials, 2 preoculars, 4 postoculars, no loreal, 2 anterior and 2 posterior temporals, dorsal scales in 13 rows at midbodv, 215 ventrals, 105 subcaudals, and a divided anal plate. This species was reported previously from Guinea by David and Ineich (1999). Elapsoidt'iJ sciiiiiiuniilnla Bocage, 1882 Materiai : ZFMK (i446l, Sidakoro, An adult female (385 mm SVL; 24 mm TL) preserved specimen with 23 cream bands (1 scale v\ide) that run transversely across the body and tail. Scutellation is: 7 supralabials (3-4 contact eye), 7 infralabials, 1 preocular, 2 postoculars, 1 anterior and 2 posterior temporals, dorsal scales in 13 rows at midbody, 152 ventrals, 18 subcaudals, and anal plate entire. This species was reported previously from Guinea by David and Ineich (1999). Naja n(i>;r;a'//;> Reinhardt, 1843 Mocquard (1902) reported a specimen from Kouroussa. The species is known from Conakry (Maclaud and Mocquard, 1896) and other Guinean localities (Chippaux, 1999). ViPFKlDAE Bitii arietmis (Merrem, 1820) Material: CNSHB 172, gallery forest of Kofing River; ZFMK 64472, Sidakoro; PNHN ecomu^eum specimen from unknou n localitx' \s ithin the park. A subadult of unknown gender (390 mm SVL; 25 mm TL) collected by mammal survey field workers, an adult male (1020 mm SVL; PNHN specimen), and an adult of unknown gender (head only; ZFMK 64472) were examined. In life, the dorsum of the body of the subadult had gray bands containing dark gray and black spots and blotches alternating with crescent-shaped salnmn bands; irregular blotches and spots of white outlined in dark gray contact bcith salmon and gray bands. The dorsum of the head was gray; salmon stripes were present on the side of the head and across the dorsum of the head, and the lower lip was white. Scutellation is: 16 supralabials, 135 (ju\'enile) and 136 (male) ventrals, 17 (juvenile) and 25 (male) subcaudals. This species was reported from Guinea by Da\id and Ineich (1999). Bitis rliiuocL'ws (Schlegel, 1855) Mvlerial: KU CT 1 1*^11 1. unknou n loeaiitN- within the park. An adult of unknown gender (head only). Lenk et al. (1999) elevated this West African subspecies of Bifib ^^abouica to specific status. The species was reported from Guinea by David and Ineich (1999). Causiis niaciihitiis (Hallowell, 1842) MxiLKiAi: CNSHB 111, ea- I km SVV of Scmioria; NLU 70619, Diaragbela; FNHN eeomuseum specimen lioni iinknow n locality within park. An adult of unknown gender (330 mm SVL; 28 mm TL) was dead on a road. An adult female (230 mm SVL; 16 mm TL) was on a path in a recenth' burned area of 16 Scii;ntific Paimrs. Naiirai. History MrsHi m.Tiii. L'mm.rsha oi Kansas grassland early in the morning. An additional specimen of Mocquard (1902) reported specimens of Gjksi/.'; rlioiiibcafiis unknown gender (218 mm SVL; 16 mm TL) was examined from Koiiroussa; however, Hughes (1977) referred this from the ecomuseum. Scutellation data are available t)nl\ record and all other similar specimens from Guinea to C. for CNSHB 111:7 supralabials, 136\entrals, 19siibcaudals. niaciilutii!^. DISCUSSION We recorded 22 species of amphibians and 34 species countries (Table 3). One likely explanation for this finding of reptiles from Upper Niger National Park; four species is incomplete sampling. However, Trioin/x trinii^iiis is of amphibians (Aninimiia j;;alaiiiciisis, Hildcbrandtin uniatn, limited to coastal brackish water and major rivers (Iverson, Plin/iiobatmchiis fmncisci, and Ptychadcua tcllinii) and two 1992), which likely accounts for its absence at other West of reptiles (Pcloiiicdiisn subnifa and Tiioin/x triuii^'^ui^) are African sites. Dciidnmspis poh/lcpis is one of the fastest and the first records for Guinea. Numerous other species of most dangerous snakes in the world, which might explain amphibians and reptiles are likely to occur in the park why previous collectors have not recorded it in other West gi\'en their known habitat preferences for Guinea savanna, African surveys. Given its currently' known distribution in gallery forest, and dry forest in other areas of Guinea, West Africa, this elapid is likely to occur throughout the Mali, and Cote d'lvoire (Joger, 1981; Bohme, 1985; Bohme semiarid and Guinea savanna habitats (Greenbaum et al., et al., 1996; Chippaux, 1999; Rodel, 2000, 2003; Ineich, 2003). The third species is a recently described Aiuiiinvia 2003; Rodel and Ernst, 2003). Thus, our findings should be known onl\' from Guinea (M. -O. Rodel, pers. comm.). considered preliminary, and further ctillecting (especialK' Stuart (1990) suggested that wooded sa\'anna habitats din-ing the rainy season) in the park should document in Guinea should be explored biologically and protected, many additional species. Moreover, some of the taxa Upper Niger National Park protects one of the last large (e.g., Hon/sf/s iiianiiomtKS, Tmcln/lcpis affiiiis, Siiioiiti.'dciu'lliJ blocks of savanna and dry forest in Guinea. Among species poccilouota) discussed herein are in need of additional of international conservation concern that were found to taxonomic work and may represent undescribed taxa (E. inhabit the park are several globalh' threatened species of Greenbaum, pers. obs.; Rodel and Agvei, 2003; Rodel and the Guinean herpetofauna (Table 4). Seven of the dozen Ernst, 2003). non-marine species listed as threatened or endangered Comparisons of the herpetofauna of Upper Niger occur in the park (CITES, 2003; lUCN, 2003), and several National Park to other sites surxeyed in West Africa are additional species from this list are likely to occur in the shown in Tables 2 and 3. Using either the raw percentage park. All anurans currentlv known from the park are similaritv or CBR, PNHN shows the greatest similaritv considered of "Least Concern" in the Global Amphibian to Comoe National Park (Cote d'lvoire), which is in the .Assessment (lUCN etal., 2004). UnfortunateK, poaching is same broad vegetation zone, and the least similaritv to known to be an ongoing problem for man\- large mammals the rainforest and montane forest habitats at Mt. Nimba, in the park core area and liLinling is permitted in the Guinea. Ho\ve\'er, caution shiiuld be used to interpret buffer /ones (Ziegler, 199b; Brugiere and Magassouba, these results, because the sampling of PNHN is relativeh' 2001; Ziegler et al., 2002; Brugiere et al., 2005). It is also incomplete. Although PNHN has more species and known that numerous kinds of amphibians and reptiles habitats (i.e., savanna) in common with Comoe National Park (Cote d'K'oire) than an\' of the other surveyed sites in West Africa, three species that occur in PNHN were not detected in Comoe or an\' ot the other sur\evs in nearbx' that iicciu- throughout West Africa are hunted for skins or human consumption (Messel et al., 1992; Akani et al., 1 998; G. Nikolaus, pers. comm.; P. DeF^idder, pers. comm.). Because man\' ot the species of mammals, birds, reptiles. Tabic 2. Comp.irison ot nunibiT ot spocios in comnmn and C oetlicitMit ot i5iogeot;riiphic ResL'mliLinco (CBR) amons; amphibians and reptiles of Upper Niger Natinnal I'ark (^h species) ,ind ollu'r sites sin'\i.'\'ed in West .'Xtrica. N. 1'. = national park. See lable 3 for sources of data. Abiiko Nuikola-Koba Mt. Nimba, Comoe N.P., Reserve, N. P., Senegal' Guinea Cote d'lvoire Gambia 32/56 34/56 29/56 52/56 (57.1';,) (60.7%) (51.8'';,) (92.9';;.) 0..571 0.535 0.307 0.658 56 71 133 iii: Species in common with PNHN CBR with PNHN Total species known from site 'Pocalitifs in southeastern Senegal near tlii' national park are included Herpbtofalina of Guinua 17 Table 3. List of 56 species of amphibians and reptiles known from Upper Niger National Park, Guinea, and their presence in other areas surveyed in West Africa. ? = presence not clear from citation; N. P. = national park. Ta\on Abuko Reserve, Gambia (Barnettetal., 20(11 Niokola-Koba N.P., Senegal' (Joger and Lambert, 2002) Mt. Nimba, Guinea (Ineich, 2003; Rode! etal., 20t)4) Comoe N.P., Cote d'lvoire- Sclioiili'ili-ihiln poi'ciloiiotii Bufc ninculaluf Hfiiiisufi nwnnoriittis Hypcivliiti coucolor Hiipcwliiis iiiifiitui^ Hypcivlius nitiiiuhni Lcptopclh viriiiis. Siluruiui twpiccilis Atnnlrana foncufis Aiiniiiniin i^nlniiicn^if Hililcbriindtiii ornnta Hoplobntrnclius occipitalis Phn/nobntrnchia iiccrncinii Plui/itohitracluis calcciriitns Plin/nobiititichus fiuiicisci Phn/iiobiilniclm^ nntiilciifif Pti/cliadciiij oxiirln/iuiius Pti/cliiidcnn piimilio Pti/chadcmi tctliiiii Ptycliiuiciiti trincdif Pcloiiicdiiiii fubrufa Pcliisios ca^lijucus Kinixys bctlianci Triom/x triiiu^ui$ Cwcodyliis cntnplunctuf Cwcodyliif buclnis A^aiiui i\^niini Chiiiiiacico ^ll■ncilif Hcnudiictyluf bwokii Hciuillicconyx caudiciiictns Lcptosinplhv to^ociifi> Trncln/lcpis affinis Vijriinnf cxiinthcnmticiif ViiriJiiKS iidoticus Ti/pldops piiiictatiis Python sclnw Crolapliopclttf liotainbociii Dnsypcltif fcnbin' Divniopliis prncor!intu:i^ Gnn/m finithii Mcizodoii coioimtiif^ Nnlricitcrcs varic^atn" Pttilotluiinnuf. Iictcrodcnnus Pbilotluiiituiis inc'fiibirif Psiimiiiopliis cUyaiis Tclcfcopiis i'(!n('5;i7/i(s' Toxicodn/iif blandinf;ii Dciidroniipii^ pol[/lcpii Dciidiviispis viridis EliipMidca scmianiuilntn Nnja iiif^ricollis^ Bitis nrictniis B/fis rlthwccroi Ciiiibus nuiciitntiis Localities in southeastern Senei;al near the park are included -Park species list compiled from Rodel and Crabow, 1993; Rndel et al., 1997, 1999; and Rode! and Spieler, 2IMH1. snce of these la\a in Upper Niger National Park, C.iLJnea is based on lileratiMv records from Mocqiiard, P Th LC preser '»02 18 SciBNiii K Paplrs, Nailiral HisroR'i Mushum,Thh Uni\1;Rsha oi- Kansas Table 4. List of threatened non-mnrine species of amphibians and reptiles in Guinea based on ratings bv CITFS, lUCN Red List, and Global Amphibian Assessment. I'he park occurrence column denotes species known {+), or with potential (?) to occur in Upper \i.;tf National Park. Taxon CITES lUCN Park Occurence Nimbuphn/noidcs occidcntnlis Crocodi/liis cataphractus Crocodylus suchus Oilcolacmus tetraspis Kinixys bcUiana Chamaeleo gracilis Chamadco smcpdensh Varanus exanthcmaticus Varamis niloticua Varanus ornaius Pytlioii regius Python sebae Appendix I Appendix 1 Appendix I Appendix 1 Appendix II Appendix II Appendix II Appendix II Appendix II Appendix II Appendix II Appendix II Critically endangered Data deficient Vulnerable + 7 + + + + 7 7 7 and iimphibians in the park are West African regional endemics and tlireatened, the park should continue to be a high conservation priority for Guinea. Three species from the park, Sihiniiin tivpiculi^, Aiiiiiirmin foiieiifiis, and Bitis rhinoceros, are mostly restricted to rainforest habitats in West Africa, which suggests that PNHN might harbor some species that are adapted to lovvlanci tropica! rainforests. Moreover, several species of mammals tliat occur within the park ha\'e West African distributions that are mainly restricted to trtipical rainforest; these include the Western pied colobus monkey {Colobus polykomos), giant pangolin (Maiiis ^i^aiitca), tree hyrax {Dcndroln/mx dorsalis), giant hog (Hi/lochocrns nu'iucrtzJia^cni), fire-footed rope sc]uirre] {Funisciurus pi/rropiis), Gambian sun scjuirrel {Hcliosciunis ^auibiuiius), hump-nosed mouse (Hi/boim/s planifroiis), and brush-tailed porcupine {Athcniriifi afyiLivnif) (Kingdon, 1997; Ziegleret al., 2002). At Comoe National Park, Cote d'lvoire, gallery forests serve as stepping stone habitats, linking savanna areas with the rainforest zone, thus prt)\'iciing sur\ival possibilities for species that normally do not occur outside of rainforests (M. -O. Rodel, pers. comm.). A similar situation seems to pertain to Upper Niger National Park, which serves to accentuate the conservation importance cif tlie park. LITERATURE CITED Akani, G. 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(Materiaux de la Mission Lamotte au mont Nimba en 1942). 3&me Note. Bulletin du Museum national d'Histoire naturelle, 2&me serie 16:293-294. Angel, R, J. Guibe, and M. Lamotte. 1954a. La reserve naturelle int^grale du Mont Nimba. 21. L<>zards. Memoires de I'lnstifut Frani;ais d'Afriquc Noire (40):371-379. Angel, F., ]. Guibd, M. Lamotte, and K. Roy. I9.S4b. La reserve naturelle intcgralc du Mont Nimba. 22. Serpents. Memoires de I'lnstilut Franijais d'Afrique Noire (4()):.38l^()2. Arnoult, ]., and M. Lamotte. 1968. Les Pipidae de I'Ouest africain et du Cameroun. Bulletin de I'Institut Frani;als d'Afrique Noire, Serie A. 3()A:270-306. Barbault, R. 1975. Place des lezards dans la bioceno.se da Lamto: relations trophiques; production et consommation des populations natu relies. Bulletin de I'Institut Fondamental d'Afriquc .Noire. Serie A, Sciences naturelles 37A:467-514. Barnett, A. A., and M. L. Prangley. 1997. 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New records of amphibians and reptiles from Burkina Faso and Mali. British Herpetological Society Bulletin (56):7-26. Bohme, W., and T. Ziegler. 1997. A taxonomic review of the Viininiif < PolydncilcUuf) niloticus (Linnaeus, 1766) species complex. Herpetological Journal 7:155-162. Bour, R. 1983. Trois populations endemiques du genre Pclnsios (Reptilia, Chelonii, Pelomedusidae) aux iles Seychelles; relations avec les especes africaines et malgaches. Bulletin du Museum National d'Histoire Naturelle, Paris, 4th ser, 5:343-382. Broadlev, D. G. 1989a. Kinixi/s bcUiana: Bell's Hinged Tortoise. Pp. 49-55 /)( I. R. Swingland and M, W, Klemens (eds.). The Conservation Biology of Tortoises. Occasional Papers of the lUCN Species Survival Commission, No. 5. Broadley, D. G. 1989b. A reappraisal of the genus Pnnnspis Cope, with the description of a new species of Li'/ifosiH/'Z/os (Reptilia: Scincidae) from Tanzania. Arnoldia Zimbabwe 9:439^149. Brugiere, D., and B. Magassouba. 2001. 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