Inventory of amphibians and reptiles in SE Senegal, including the Niokola-Koba National Park, with observations on factors influencing diversity

Transcription

1 Tropical Zoology ISSN: (Print) (Online) Journal homepage: Inventory of amphibians and reptiles in SE Senegal, including the Niokola-Koba National Park, with observations on factors influencing diversity U. Joger & M. R.K. Lambert To cite this article: U. Joger & M. R.K. Lambert (2002) Inventory of amphibians and reptiles in SE Senegal, including the Niokola-Koba National Park, with observations on factors influencing diversity, Tropical Zoology, 15:2, , DOI: / To link to this article: Published online: 30 Jul Submit your article to this journal Article views: 423 View related articles Citing articles: 3 View citing articles Full Terms & Conditions of access and use can be found at

2 Tropical Zoology 15: , 2002 Inventory of amphibians and reptiles in SE Senegal, including the Niokola-Koba National Park, with observations on factors influencing diversity U. JOGER 1 and M.R.K. LAMBERT 2,3 1 Zoologische Abteilung, Hessisches Landesmuseum Darmstadt, Friedenplatz 1, D Darmstadt, Germany 2 Natural Resources Institute, University of Greenwich at Medway, Central Avenue, Chatham Maritime, Kent ME4 4TB, United Kingdom Received 6 February 2001, accepted 30 July 2002 Twenty-six Amphibia (plus two possible) and 45 Reptilia have been recorded in SE Senegal; 23 Amphibia (plus three probable) and 39 Reptilia (plus five probable) are listed for the Niokola-Koba National Park. New country records made in October 1996 were Amphibia: Kassina fusca Schiøtz 1967; and Reptilia: Agama sankaranica Chabanaud 1918, Cophoscincopus cf. greeri, Mabuya quinquetaeniata scharica Sternfeld 1917 and Chalcides thierryi pulchellus Mocquard 1906, with extended ranges for Bufo maculatus Hallowell 1854 and Phrynomantis microps Peters Cophoscincopus Mertens 1934 is a genus new to Senegal. Relative density of diurnal woodland lizards was nearly 4-fold that of bushland and thicket lizards of drier northern and western Senegal. Where riparian forest habitat formed within undifferentiated Sudanian woodland, reptile species differed and diversity was lower. Percentage frequency of woodland A. sankaranica was reduced by man-made road structures, while Agama agama Linnaeus 1758 and Hemidactylus brooki complex were only recorded with them. Geckos were not recorded inside the boundary of the Niokola-Koba National Park (baboons and patas monkeys abundant), and diversity was reduced; outside, reptile density was nearly 5-fold, with geckos (Tarentola parvicarinata Joger 1980 and H. brooki complex) predominant. Faunal overlap of lizards in SE Senegal with those of W Mali the same area of endemism was higher than with species in northern and western Senegal. KEY KORDS: Amphibia, Reptilia, SE Senegal, Niokola-Koba National Park, species inventory, herpetofaunal diversity. 3 Present address: Environmental Initiatives, Lydbrook House, Upper Lydbrook, Gloucestershire GL17 9LP, United Kingdom. Published online 30 Jul 2012

3 166 U. Joger and M.R.K. Lambert Introduction Survey methods Locality records Annotated inventory of amphibians and reptiles in SE Senegal (Cercle de Kédougou and Niokola-Koba National Park) Results and discussion Geographical variation Forest vegetation Anthropogenic change Revised check list of the amphibians and reptiles of the Niokola-Koba National Park Biogeographical analysis of amphibian and reptile species in Senegal Acknowledgements References INTRODUCTION The reptiles and amphibians of Senegal were first listed by DE ROCHEBRUNE (1884a, 1884b, respectively), whose area of coverage included part of what is now the Republic of Mali, but his works suffer from inaccuracies due mainly to the large number of nomenclatural changes made subsequently. No species records were included from SE Senegal. A collection of reptiles from West Africa by GRAN- DISON (1956) included material from Senegal, and a further large collection was also made from Senegal by BÖHME (1978); regrettably neither of these authors had the opportunity to visit the south-east of the country. Amphibians were additionally collected in Senegal, as well as reptiles, during West African journeys by JOGER (1981), and these included a visit to Niokola-Koba National Park in SE Senegal by JOGER (1982), who added to the list of reptiles in the Park initially prepared by VIL- LIERS (1956). Amphibians in West Africa have been treated by MERTENS (1938), GUIBÉ & LAMOTTE (1957), LAMOTTE (1966) and SCHIØTZ (1967), and species in Niokola- Koba National Park were listed by LOVERIDGE (1956) and LAMOTTE (1969). The most recent coverage of amphibians in Sahelian Africa is by RÖDEL (2000). Meanwhile, CISSÉ (1974) described the geckos, and later CISSÉ & KARNS (1978) covered all lizard species then known in Senegal. Snakes were collected in Senegal from 1986 by J.-F. TRAPE (pers. comm.), including SE Senegal, Cercle de Kédougou (TRAPE & MANE 1995, TRAPE 1997). Subsequent to preparing an inventory of amphibians and reptiles in neighbouring Mali (JOGER & LAMBERT 1996), and completing a biogeographical analysis based on the distribution of species there (JOGER & LAMBERT 1997), the authors recognised that south-west Mali and south-east Senegal constituted an area of high endemicity. Following the earlier observations made in Niokola-Koba National Park on amphibians (LOVERIDGE 1956, LAMOTTE 1969) and reptiles by VILLIERS (1956) and JOGER (1982), the last was presented with the opportunity to make a further visit to SE Senegal with M.R.K. Lambert in October 1996 (Cercle de Kédougou and Niokola- Koba National Park). Effort was concentrated on lizards, since this reptile group appeared to have received less attention in this part of the country than others. Apart from preliminary observations on herpetofaunal diversity, the main objective of this work was to prepare an inventory of species in SE Senegal (Cercle de Kédougou and Niokola-Koba National Park), using additional records available in literature, and unpublished data from museum collections: Muséum National

4 Amphibians and reptiles in SE Senegal 167 d Histoire Naturelle, Paris (MNHN), the Natural History Museum, London (BMNH) and the Smithsonian Institution, Washington (USNM). Following HOWELL (1992), and as indicated by JOGER & LAMBERT (1996), preparation of regional and national inventories is an important first step in assessing the impact of environmental change on amphibian and reptile species and their habitats in sub-saharan Africa. Inventories which also document the distribution of species additionally enable centres of herpetofaunal richness and endemism to be identified, and, apart from adding new species and extending known ranges, provide the basis for further information to be obtained on habitat and status. Such information is also required for biogeographical studies, and an analysis of data here is included with this work. A check list of amphibians and reptiles, and their distribution in the Republic of Senegal, will also be required at some future date, especially for a distribution atlas, and so it is intended that this work for SE Senegal alone will in due course be extended to provide coverage for the whole of the country. SURVEY METHODS An herpetofaunal survey was conducted at the end of the rainy season from 9-17 October 1996, and the opportunity was taken to make observations on reptile richness and diversity. The survey was chronologically contiguous with one immediately beforehand in northern and western Senegal during September 1996 (LAMBERT 1996). A range of sites was inspected in SE Senegal, and visually encountered species were recorded to establish species richness, and diversity from composition (percentage frequency) and relative density (sighting frequency). With a searching technique common to all, sites were comparably surveyed during daylight hours. Some additional qualitative surveys were conducted after darkness. Time at start and end of searches (together with air temperature and cloud cover) was also recorded at a proportion of the sites, enabling the number of reptiles sighted per man-hour to be calculated as an expression of relative density based on area of coverage corresponding to time spent searching. Distances along roads were travelled through different sectors in a motor vehicle, and, unless under 50 m apart, halts were made at every under-road culvert or tunnel, and associated man-made surrounds, because lizards especially among reptiles, possibly in search of basking and perhaps also feeding sites, tended to aggregate by them. Along woodland tracks without such human artefacts, halts were made in all areas with hard-surfaced open ground in woodland clearings, invariably with flat exposed rock surfaces, again because reptiles for similar reasons tended to aggregate with them. Each site was inspected for reptiles and amphibians, and numbers carefully recorded to provide information on reptile species composition (percentage frequency) and relative density (numbers per man-hr or per site). Sites in the study area were within the bioclimate described as tropical, warm (t > 15 ºC) dry season of long duration sensu UNESCO-FAO (1963), with xerothermic index x (the number of physiologically dry days in a year) 150 < x < 200. Species distribution in the area was also related to major vegetation types sensu WHITE (1983): Forest transitions and mosaics: mosaic of Guineo-Congolian lowland rain forest and secondary grassland [mapping unit 11a]; and Woodland: undifferentiated Sudanian woodland [mapping unit 29a]. In contrast, much of northern and western Senegal, where observations were made by LAMBERT (1996), is made up of Bushland and thicket: Sahel Acacia wooded grassland and deciduous bushland [mapping unit 43] within UNESCO-FAO (1963) subdesertic-dry season shorter bioclimate, with xerothermic index 200 < x < 250. Routine statistical calculations were used to treat the visual encounter survey data obtained from the many sites inspected. They included diversity (Shannon-Wiener index, after

5 168 U. Joger and M.R.K. Lambert MAGURRAN 1988), evenness (after MAGURRAN 1988), faunal overlap between areas (Sorensen s similarity index, after MÜHLENBERG 1989), and resemblance of areas in species dominance (Renkonen s coefficient, after MÜHLENBERG 1989). The representative collection of amphibians and reptiles made was deposited with the collection of the Hessisches Landesmuseum Darmstadt, Germany (HLMD). Animals were mainly caught by hand, some lizards by rod and noose, and were immobilised by placing them inside a polythene collecting bottle, in which there was a pad of chloroform-saturated cotton wool. Specimens were labelled, and preserved in 90% ethanol. LOCALITY RECORDS The number of species recorded at numerous localities during October 1996 provided information on richness, and, where search-time is indicated (0.1+ man-hr), observation intensity (Appendix). Localities (Fig. 1) are enumerated from north to south (in order of ascending mean annual rainfall). ANNOTATED INVENTORY OF AMPHIBIANS AND REPTILES IN SE SENEGAL (CERCLE DE KÉDOUGOU AND NIOKOLA-KOBA NATIONAL PARK) In the following inventory of species in SE Senegal, new country records (***), range extensions (**) and species endemic to Senegal, or having a major part of their range within Senegal (*), are indicated. Only unpublished material in museum collections is listed; accession numbers are included for records of species new to SE Senegal, range extensions and unpublished endemic species. Records of lizards in earlier references are included with CISSÉ & KARNS (1978), and other citations are also given for any previously published records of species in SE Senegal. Ecological notes are added for species recorded in SE Senegal in this work. Locality numbers correspond to those inspected in this work (Appendix). Amphibia A total of 26 species has been recorded. Bufonidae Bufo xeros M. Tandy, S. Tandy, Keith & Duff-Mackay Localities: 1, 3, on open ground by buildings, seeking light-attracted insects during early hours of darkness. **Bufo maculatus Hallowell Localities: 18 [voucher specimen HLMD-RA 2558], 31 [voucher specimen HLMD-RA 2557], 45 (larvae in pool by forest stream), adults collected during early hours of darkness following late afternoon rain, and by gully of forest pocket (locality 31). Bufo regularis Reuss Badi in Niokola-Koba National Park (LOVERIDGE 1956); Niokola-Koba and Dialakota in and by the Niokola-Koba National Park (MERTENS 1938, LAMOTTE 1969). Localities: 26, 27, 40, 41. Hyperoliidae Afrixalis cf. fulvovittatus (Cope 1860). Badi, Tabadian and 10 km S of Diénoun Diala in Niokola-Koba National Park (MERTENS 1938, LAMOTTE 1969). Current

6 Amphibians and reptiles in SE Senegal 169 Fig. 1. Amphibian and reptile observation localities (see Appendix for key to numbers), SE Senegal, 9-17 October 1996.

7 170 U. Joger and M.R.K. Lambert authors agree that there are probably two species in this complex a farmland-bushland species and a savanna species. RÖDEL (2000) differs from SCHIØTZ (1967, 1999) in using the name Afrixalis vittiger (Peters 1876) for the savanna species of fulvovittatus s.l. If this were followed, the Senegalese populations would be vittiger rather than fulvovittatus. However, according to SCHIØTZ (1999), the same savanna species is still unnamed. Afrixalus weidholzi (Mertens 1938). 10 km S of Diénoun Diala (type locality), Badi, Tali Boulou Ba near Niokola-Koba, and near Bangaré in Niokola-Koba National Park (LAMOTTE 1969). Hyperolius nitidulus Peters Badi in Niokola-Koba National Park (LOVERIDGE 1956).?Hyperolius lamottei Laurent Badi by Woulou river, and between Badi and Dar Salam in Niokola-Koba National Park (LAMOTTE 1969). SCHIØTZ (1999) does not mention the occurrence of this species in Senegal. Leptopelis viridis (Günther 1868). Niokola-Koba in Niokola-Koba National Park (LAMOTTE 1969). ***Kassina fusca Schiøtz Locality: 18 [voucher specimen HLMD-RA 2559], collected during early hours of darkness following late afternoon rain. Kassina senegalensis (Duméril & Bibron 1841). Niokola-Koba National Park (MNHN). Ranidae Ptychadena bibroni (Hallowell 1845). Sanyiki near Niokola-Koba, by Doufouroun river between Niokola-Koba and Bangaré, by Niokola-Koba river near Niokola- Koba and Tabadian in Niokola-Koba National Park (LAMOTTE & OHLER 1997). The authors refer to specimens described as Ptychadena maccarthyensis (Anderson 1937), and recorded from Niokola-Koba by MERTENS (1938) and LAMOTTE (1969). Locality: 31, collected during early hours of darkness following late afternoon rain in wet gully of forest pocket. Ptychadena pumilio (Boulenger 1920). Badi, by Woulou river at Badi, by Doufouroun river between Niokola-Koba and Bangaré, Tali Boulou Ba near Niokola-Koba, by Niokola-Koba river at Niokola-Koba, Sanyiki, Dialakoto and Tabadian in and by the Niokola-Koba National Park (MERTENS 1938 as P. m. mascareniensis LAMOTTE 1969). Ptychadena oxyrhynchus (Smith 1884). Niokola-Koba in Niokola-Koba National Park (LAMOTTE 1969). Ptychadena schubotzi (Sternfeld 1917). Tabadienké = Tabadian, Niokola-Koba National Park (MERTENS 1938). Frogs recorded from Badi, Dialakoto and Tabadian in and by the Niokola-Koba National Park, and identified as Ptychadena longirostris Peters 1870 (LAMOTTE 1969), may in fact be P. schubotzi. P. longirostris is a rain forest species, and has often been confused with the similarly patterned but savanna-dwelling P. schubotzi (see RÖDEL 2000). Ptychadena longirostris Peters Locality: 46 (photographic record only), by fast flowing stream. Contrary to the above mentioned specimens from savanna habitat in the Niokola-Koba National Park, our specimen was recorded in a very humid environment, adjacent to the Dinndéfélou waterfall, where the rain forest skink Cophoscincopus cf. greeri was found (see below). This and the fact that M.-O. Rödel, a field expert on West African frogs, identified our photograph as P. longirostris (see RÖDEL 2000), confirms our conclusion that it is not another mistaken P. schubotzi.

8 Amphibians and reptiles in SE Senegal 171?Ptychadena superciliaris (Günther 1858). Badi in Niokola-Koba National Park (LAMOTTE 1969). RÖDEL (2000) considers P. superciliaris as a valid species, but as a forest dweller, and therefore does not include it in his list of amphibians of the West African savanna. Ptychadena tournieri (Guibé & Lamotte 1955). Tali Boulou Ba near Niokola-Koba in Niokola-Koba National Park (LAMOTTE 1969). Ptychadena trinodis Boettger Badi and Dialakoto in and by the Niokola-Koba National Park (MERTENS 1938, LAMOTTE 1969). Ptychadena sp. Locality: 42. Ptychadena floweri (Boulenger 1917). By Sanyiki river near Niokola-Koba in Niokola- Koba National Park (LAMOTTE 1969, as Abrana floweri). Hylarana galamensis (Duméril & Bibron 1841). Badi, Niokola-Koba, Dialakoto and Tabadian in and by the Niokola-Koba National Park (MERTENS 1938, LAMOTTE 1969). Localities: 27, 28, 32, at edge and in water of pools. Hoplobatrachus occipitalis (Günther 1858). Wassa Dou, Niokola-Koba and Dialakoto in and by the Niokola-Koba National Park (MERTENS 1938, LAMOTTE 1969). Localities: 4, 7, 8, 28, 31, 33, 41, 45, 46, in pools. Phrynobatrachus (accraensis) latifrons (Ahl 1924). By Woulou river at Badi, by Niokola-Koba river in Niokola-Koba National Park (LAMOTTE 1969). According to RÖDEL (2000), latifrons might be a separate species, restricted to savanna habitats, whereas typical accraensis are forest dwellers. Phrynobatrachus calcaratus (Peters 1863). Tali Boulou Ba near Niokola-Koba, by Niokola-Koba river at Niokola-Koba, by Doufouroun river between Niokola- Koba and Bangaré, and Saïnti Boulou in Niokola-Koba National Park (LA- MOTTE 1969). Phrynobatrachus francisci Boulenger By the Niokola-Koba river, by the Doufouroun river between Niokola-Koba and Bangaré, by the Sanyiki river near Niokola-Koba, Niokola-Koba, Tabadian and 10 km S of Diénoun Diala in Niokola-Koba National Park (LAMOTTE 1969). MERTENS (1938) mentions P. acridoides (Cope 1867) from Tabadienké (= Tabadian). This is an East African species. LAMOTTE (1969) regards Mertens specimens as belonging to P. francisci, but M.-O. RÖDEL (in litt.) considers the possibility that it might be P. (accraensis) latifrons. Phrynobatrachus sp. Locality: 31, 38, collected during early hours of darkness after rain during late afternoon. Phrynobatrachus natalensis (A. Smith, 1849), which is known from Senegal (RÖDEL 2000) and from Mali (JOGER & LAMBERT 1996), but not definitely recorded from SE Senegal, may be the species in question. Hemisotidae Hemisus marmoratus (Peters 1882). Badi in Niokola-Koba National Park (LAMOTTE 1969). Microhylidae **Phrynomantis microps (Peters 1875). Niokola-Koba National Park (MNHN); locality: 18 [voucher specimen HLMD-RA 2560], collected during early hours of darkness after rain during late afternoon.

9 172 U. Joger and M.R.K. Lambert Reptilia A total of 45 species has been recorded. Chelonia Testudinidae Kinixys belliana nogueyi (Lataste 1886). Oassadou, Badi, Niokola-Koba National Park (VILLIERS 1956); Dialokoto, Gamon (USNM). Trionychidae Cyclanorbis senegalensis (Duméril & Bibron 1835). Niokola-Koba National Park (VILLIERS 1956, MNHN). Pelomedusidae Pelomedusa subrufa (Lacépède 1788). Badi, Niokola-Koba National Park (VILLIERS 1956); localities: 14, 23, 41, in temporary rain or semi-permanent pools. Pelusios subniger subniger (Lacépède 1788). Badi, Niokola-Koba National Park (VIL- LIERS 1956). Crocodylidae Crocodylus niloticus Laurenti Ouolou River, Niokola-Koba National Park (VIL- LIERS 1956). Crocodylus cataphractus Cuvier Koulountou River, Niokola-Koba National Park (VILLIERS 1956). Ostaeolaemus tetraspis Cope Badi, Niokola-Koba National Park (VILLIERS 1956). Sauria Gekkonidae Hemitheconyx caudicinctus (A. Duméril 1851). Kédougou, Niokola-Koba National Park (CISSÉ & KARNS 1978); 5 km SE of Dar Salam in Niokola-Koba National Park (JOGER 1981, 1982); locality: 18, collected during early hours of darkness after late-afternoon rain. Hemidactylus brooki complex. Badi in Niokola-Koba National Park (VILLIERS 1956); Kédougou (CISSÉ & KARNS 1978); 5 km SE of Dar Salam in Niokola-Koba National Park [20 km S of Médina Gounas] (JOGER 1981); localities: 14, 16, 17, 22, 26, 27, 28, 32, 39, 40, 41, recorded on buildings after darkness and in man-made under-road culverts and tunnels. Tarentola ephippiata O Shaughnessy 1875 [Missira, N.-W. of Niokola-Koba National Park (JOGER 1982)]. Tarentola parvicarinata Joger Badi in Niokola-Koba National Park (JOGER 1982); Dialokoto, Gamon (USNM); localities: 1, 2, 15, 20, 24, 27, in man-made under-road culverts and tunnels. Agamidae Agama agama (Linnaeus 1758). Badi in Niokola-Koba National Park (VILLIERS 1956); Kédougou (CISSÉ & KARNS 1978); Dialakoto, Gamon (USNM); localities: 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 20, 21, 22, 26, 27, 28, 29, 32, 36, 38, 39, 40, 44, especially common by man-made structures near woodland.

10 Amphibians and reptiles in SE Senegal 173 ***Agama sankaranica Chabanaud Localities: 4, 8, 14, 25, 27, 28, 30, 31 [voucher specimen HLMD-RA 2568], 32, 34, 35, 36, 38, 39 [voucher specimens HLMD-RA ], 41 [voucher specimen HLMD-RA 2565], on hardsurfaced open ground of woodland clearings. Agama weidholzi Wettstein Niokola-Koba National Park (CISSÉ & KARNS 1978); 5 km SE of Dar Salam in Niokola-Koba National Park [20 km S of Médina Gounas] (JOGER 1981, 1982); 30 km S of Dialokoto by Niokola-Koba National Park (paratype; BMNH ); locality: 9, open ground at edge of woodland. Chameleonidae Chameleo senegalensis Daudin Badi, Niokola-Koba National Park (VILLIERS 1956). Chameleo gracilis gracilis Hallowell Badi in Niokola-Koba National Park, Dialokoto (CISSÉ & KARNS 1978). Scincidae ***,?*Cophoscincopus cf. greeri (Böhme, Schmitz & Ziegler 2000). Locality: 46 [voucher specimens HLMD-RA ], among damp rocks surrounding pool at base of waterfall. Mabuya affinis (Gray 1838). Localities: 31, 45, 46, in riparian forest pockets. Mabuya perroteti (Duméril & Bibron 1839). Badi in Niokola-Koba National Park (VILLIERS 1956); Kédougou (CISSÉ & KARNS 1978); Niokola-Koba National Park (JOGER 1981); localities: 7, 8, 13, 15, 16, 17, 18, 31, 42, 46. An ecologically ubiquitous species. ***Mabuya quinquetaeniata scharica (Sternfeld 1917). Localities: 3 [voucher specimen HLMD-RA 2522], 5, 6, 7, 8, 10, 11, 12, 13, 15, 20, 27 [voucher specimens HLMD-RA 2525, ], 28 [voucher specimens HLMD-RA 2523, 2530], 30 [voucher specimen HLMD-RA 2524], 31, 32, 36, 37, 39 [voucher specimen HLMD-RA 2526], on flat rock surfaces, often man-made, and hard-surfaced open ground of woodland clearings. ***Chalcides thierryi pulchellus Mocquard Locality: 27 [voucher specimens HLMD-RA ; 2534], very local; in soil by fallen trunks on disturbed open ground near woodland. The skink, Chalcides armitagei (Boulenger 1922), an inhabitant of coastal dunes and only known from Gambia (GRUSCHWITZ et al. 1991), is clearly a different species. Varanidae Varanus niloticus (Linnaeus 1758). Badi in Niokola-Koba National Park (VILLIERS 1956). Localities: 11, 12, 21, 31, 33, 42, 43, 45, vicinity of water. Typhlopidae Typhlops punctatus punctatus (Leach 1819). Badi, Niokola-Koba National Park (VIL- LIERS 1956). Typhlops lineolatus lineolatus (Bocage 1891). Badi in Niokola-Koba National Park (MNHN). Leptotyphlopidae Leptotyphlops macrorhynchus (Jan 1862). Sambaïlo, Niokola-Koba National Park (VILLIERS 1956).

11 174 U. Joger and M.R.K. Lambert Boidae Python sebae sebae (Gmelin 1789). Badi, Niokola-Koba National Park (VILLIERS 1956). Atractaspidae Atractaspis dahomeyensis Bocage Youkounkoun, Niokola-Koba National Park (VILLIERS 1956). Colubridae Afronatrix anoscopus (Cope 1861). Bandafassi (TRAPE & MANE 1995). Crotaphopeltis hotamboeia (Laurenti 1768). Sambaïlo, Badi, Youkounkoun, Niokola- Koba National Park (VILLIERS 1956). Haemorrhois (Coluber) dorri (Lataste 1888). Bandafassi (TRAPE 1997); localities: 28, 38, by man-made structures (among wall blocks) adjacent to woodland. Dispholidus typus typus (A. Smith 1829). Badi, Niokola-Koba National Park (VIL- LIERS 1956). Grayia smythi (Leach 1812). Badi, Niokola-Koba National Park (VILLIERS 1956). Lamprophis fuliginosus (Boie 1827). Badi, Niokola-Koba National Park (VILLIERS 1956); locality: 19, collected during early hours of darkness adjacent to woodland. Lamprophis lineatus A.-M.-C. Duméril, A.-H.-A. Duméril & Bibron Badi, Niokola-Koba National Park (VILLIERS 1956). Lycophidion albomaculatum Steindachner Sambaïlo, Niokola-Koba National Park (VILLIERS 1956). Meizodon coronatus (Schlegel 1837). Sambaïlo, Badi (VILLIERS 1956), Badi (MNHN) in Niokola-Koba National Park. Philothamnus irregularis (Leach 1819). Badi, Niokola-Koba National Park (VILLIERS 1956). Prosymna meleagris greigerti Mocquard Badi, Niokola-Koba National Park (VILLIERS 1956). Psammophis elegans (Shaw 1802). Badi, Niokola-Koba National Park (VILLIERS 1956). Psammophis (sibilans) phillipsi (Hallowell 1844). Sambaïlo, Badi, Niokola-Koba National Park (VILLIERS 1956). Rhamphiophis oxyrhynchus oxyrhynchus (Reinhardt 1843). Youkounkoun, Niokola- Koba National Park (VILLIERS 1956). Viperidae Bitis arietans (Merrem 1820). Badi, Youkounkoun, Niokola-Koba National Park as Bitis lachesis (VILLIERS 1956). Causus maculatus (Hallowell 1842). Sambaïlo, Badi, Niokola-Koba National Park as C. rhombeatus (VILLIERS 1956). Echis carinatus. Koundara, Niokola-Koba National Park (VILLIERS 1956). As this specimen from a private collection is not available for redetermination, its specific status remains uncertain. It could be either E. ocellatus Stemmler 1970 or Echis jogeri Cherlin 1990, both of which are known from southwestern Mali (JOGER & LAMBERT 1996). E. carinatus is an Asian species (JOGER 1984).

12 Amphibians and reptiles in SE Senegal 175 Elapidae Naja nigricollis Reinhardt Sambaïlo, Niokola-Koba National Park (VILLIERS 1956). RESULTS AND DISCUSSION Reptile richness and diversity (species composition and relative density) were influenced by a variety of factors in SE Senegal. Species assemblages were a bioindication of bioclimatic, vegetational and other ecological differences, and of anthropogenic change. Geographical variation Relative density of lizards in SE Senegal, based on numbers visually encountered during daylight site searches, was 5.26 per hour (eight sites) within WHITE s (1983) Woodland: undifferentiated Sudanian woodland [29a]. This was in contrast (Table 1i) to 1.43 per hour (five sites), and nearly 4-fold that recorded in drier northern and western Senegal within WHITE s (1983) Bushland and thicket: Sahel Acacia wooded grassland and deciduous bushland [43]; the difference between ranges was significant (Mann-Whitney T = 19, P < 0.05). Conversely, differences between ranges for Agama agama and Varanus niloticus (also recorded in both areas) were not significant. In SE Senegal, the relative density of Agama sankaranica was 3.6 times that of the skink Mabuya quinquetaeniata scharica. The two species, together with Chalcides thierryi pulchellus, were new country records, although all three are known in Mali (JOGER & LAMBERT 1996). Frequency of A. sankaranica was highest in SE Senegal (14 sites), and of Agama agama in northern and western parts (five sites) of the country (Table 1ii). With seven species each (Table 1ii), Shannon-Weiner index of diversity (H ) was apparently a little higher [above 2 is high; below 1 is low] in northern and western Senegal with a lower number of lizards recorded (1.259, evenness 0.647; n = 20) than in wetter SE Senegal (1.178, evenness 0.605; n = 70), but the difference was not significant. Varanus niloticus (in the vicinity of water) and A. agama (except in rocky habitats, JOGER 1979) were widespread in Senegal. Another widespread species, Mabuya perroteti, was not recorded in the time-surveyed sites, although recorded during daylight in other sites elsewhere in SE Senegal (see also Tables 2-4) within WHITE s (1983) undifferentiated Sudanian woodland [29a]. The arboreal Tarentola ephippiata and savannah ground-dwelling Latastia longicaudata and Varanus exanthematicus were not recorded in SE Senegal (Table 1b), nor was the dune-dwelling Acanthodactylus dumerili s.l. Forest vegetation WHITE s (1983) Forest transitions and mosaics: mosaic of Guineo-Congolian lowland rain forest and secondary grassland [11a] made-up an isolated pocket of specialised riparian forest habitat on the north flank of hills bordering Guinea, and formed within adjacent undifferentiated Sudanian woodland [29a]. Reptile diversity

13 176 U. Joger and M.R.K. Lambert Table 1. Variation of reptile diversity in woodland of northern and southern Senegal. Relative density (i) and percentage frequency of lizards (ii) are compared in a. SE Senegal (Cercle de Kédougou), eight sites (41 lizards), and b. northern and western Senegal (from LAMBERT 1996), five sites (20 lizards); frequencies for sites in SE Senegal include data from untimed surveys. WHITE s (1983) major vegetation types for respectively SE and N/W Senegal are undifferentiated Sudanian woodland [29a] and Sahel Acacia wooded grassland and deciduous bushland [43]. Relative densities are from visual encounter surveys (median cumulative no. per hour, site ranges in parentheses) during searches (> 0.1 hr) over air temperature and start-time ranges of a C, 08:42-15:08 hr (10-16.X.96) and b C, 09:41-14:49 hr (14-30.IX.96). Species (i) Relative density (ii) % frequency a. SE Senegal b. N/W Senegal a. SE Senegal b. N/W Senegal (7.8 hr) (14.0 hr) (n = 70) (n = 20) Agama sankaranica 3.72 (0-13.6) 65.7 Mabuya quinquetaeniata scharica 1.03 (0-1.6) 12.9 Agama agama 0.13 (0-1.3) 0.93 (0-3.0) Varanus niloticus 0.13 (0-10) 0.07 (0-0.2) Mabuya perroteti 0.07 (0-1.3) Chalcides thierryi pulchellus 5.7 Mabuya affinis 1.4 Tarentola ephippiata 0.14 (0-0.6) 10.0 Latastia longicaudata 0.07 (0-0.4) 5.0 Varanus exanthematicus 0.07 (0-0.2) 5.0 Acanthodactylus dumerili s.l (0-0.2) 5.0 All lizards 5.26 ( ) 1.43 ( ) in the forest habitat (Table 2a) was significantly lower (t = 2.07; df 24; P < 0.05) than in the woodland surrounding it (Table 2b); faunal overlap was also low (Table 5i). Varanus niloticus (by water) was the only species recorded in both habitats. Mabuya affinis is a forest species found relatively locally in southern Senegal (CISSÉ & KARNS 1978), and with its characteristically fragmented habitat, may be considered a forest bioindicator species. The water skink, Cophoscincopus cf. greeri, was only found by the Cascade de Dinndéfélou, at the base of a cliff among watersplashed rocks surrounding a fall-formed pool. Cophoscincopus greeri has now been described by BÖHME et al. (2000), but in this work, our material has been excluded from the type series because it was found far outside the known range of the species (and genus). Cophoscincopus Mertens 1934 is a genus new to Senegal. Anthropogenic change Man-made structures. Such human artefacts as under-road culverts and tunnels, and associated surrounds, did not apparently greatly affect reptile diversity, for the difference was not significant (Table 3). On the other hand, faunal overlap between these man-made structures and undisturbed sites was low (Table 5ii). The resemblance in species dominance was also low (Table 5ii). In particular, the frequency of Agama sankaranica relative to other species was reduced significantly in man-made structures (c 2 = 44.06, P < 0.001), but not that of the two skinks Mabuya

14 Amphibians and reptiles in SE Senegal 177 Table 2. Influence of vegetation on reptile diversity in SE Senegal (Cercle de Kédougou). Comparison of reptile species composition (%) in adjacent major vegetation types (two) of WHITE (1983): a. mosaic of Guineo-Congolian lowland rain forest and secondary grassland [11a], two sites, which had formed within b. adjacent undifferentiated Sudanian woodland [29a], seven sites. Species were recorded during visual encounter surveys (> 0.1 hr) at air temperature and start-time ranges of a C; 09:30-14:00 hr (11-12.X.96) and b C; 08:45-15:08 hr (10-16.X.96). a. Dinndéfélou area b. between Mako [11a] and Ibel [29a] (n = 13) (n = 30) Lizards Cophoscincopus cf. greeri Mabuya affinis Varanus niloticus Agama sankaranica Agama agama Tarentola parvicarinata Chalcides thierryi pulchellus Mabuya quinquetaeniata scharica Mabuya perroteti Shannon-Wiener index of diversity (H ) Evenness Recorded (a.) during qualitative inspection outside survey period. q. scharica or M. perroteti (Table 3). In addition to elsewhere without man-made structures (Table 2a), the forest skink M. affinis was recorded only at a single forest-fringed locality in this area (Table 3a), but A. agama and Hemidactylus brooki complex in contrast were only recorded adjacent to man-made structures. In light of their popular names, among them house agama and house gecko, respectively, they can indeed be regarded as bioindicator species of anthropogenic change. Varanus niloticus was recorded by man-made structures, adjacent to water, elsewhere in SE Senegal (Tables 1, 4a). Haemorrhois (Coluber) dorri (Table 3b) was first recorded near a man-made structure elsewhere in Senegal (type specimen) by LATASTE (1888). Faunal overlap between similar roadside structures inside and outside the Niokola-Koba National Park (Table 5iii) was much lower than overlap observed between such man-made structures in different areas (Table 5iv, b), whereas resemblance in species dominance had almost the same value. Both faunal overlap and resemblance were rather lower in undisturbed sites than in those with man-made structures in different areas (Table 5iv, a and b). National park protection. Reptile diversity inside Niokola-Koba National Park (Table 4a) was significantly lower than outside the Park boundary (t = 3.13; df 97; P < 0.01); moreover, relative density of reptiles inside the Park (0.57 per site) was a fraction less than 1/4 (0.21) of that outside (2.77 per site). Frequencies of A. agama and M. q. scharica, relative to other species (Table 4), were significantly higher inside than outside the Park (c 2 = 6.85 and 4.07, and P < 0.01 and < 0.05,

15 178 U. Joger and M.R.K. Lambert Table 3. Influence of anthropogenic activity on reptile diversity in SE Senegal (Cercle de Kédougou). Species composition (%) compared in undisturbed sites (n = 11) and those by man-made structures (n = 28) under-road culverts and tunnels, and associated surrounds within WHITE s (1983) undifferentiated Sudanian woodland [29a]. Species were recorded during surveys at air temperature and start-time ranges of a C; 08:42-14:54 hr (12-14.X.96) for visual encounter searches, and b C; 08:45-15:08 hr (14.X.96) for man-made structure inspections. a. Undisturbed sites b. By man-made structures between Saraya and between Saraya and Satadougou-Bafé Kédougou (n = 46) (n = 62) Lizards Agama sankaranica Mabuya quinquetaeniata scharica Agama agama Hemidactylus brooki Mabuya perroteti Mabuya affinis Varanus niloticus Snake Haemorrhois (Coluber) dorri Chelonian Pelomedusa subrufa Shannon-Weiner index of diversity (H ) Evenness respectively), and Park relative densities of respectively 0.29 and 0.21 per site, were fractions a little over 1/3 (0.37 and 0.38) of those outside (0.79 and 0.56). Geckos, in contrast, were totally unrecorded in the Park. And yet one, T. parvicarinata, had the overall highest frequency relative to other species in physiognomically identical habitat outside. The frequency of this species in the area south-east of Niokola-Koba National Park (a road distance of 52-km between Kédougou and the Park boundary) was 27.5% (n = 91) [A. agama and M. q. scharica were at 28.6 and 24.2%, respectively] compared to that north-west of the Park (a road distance of 13-km between the Park boundary and Dar Salaam), where the frequency of T. parvicarinata was 70.6%, with A. agama (the only other species recorded north-west of the Park) at 29.4% (n = 17). Hemidactylus brooki complex was only present southeast of the Park (6.6%). Frequencies of A. sankaranica and M. perroteti were very similar inside and outside the Park. As in the Saraya-Kédougou sector (Table 3b), seven species were recorded in the Kédougou-Dar Salam sector outside the Park boundary (Table 4b), and species composition in both sectors was, not unexpectedly, similar [the snake, H. (C.) dorri, for example, was recorded in each], but T. parvicarinata was absent from the former, as it was from inside the Niokola-Koba National Park. Hemidactylus brooki complex was the only gecko species recorded in the Saraya-Kédougou sector (Table 3b). When M. perroteti was recorded at a site, M. q. scharica was not. This may suggest competition between these two skinks. Large troupes of baboons and patas monkeys are frequently seen in the Niokola-Koba National Park, especially on and at the cleared periphery of the tarmac

16 Amphibians and reptiles in SE Senegal 179 Table 4. Influence of national park protection on reptile diversity in SE Senegal. Reptile species composition (%) compared inside and outside the boundaries of the Niokola-Koba National Park by manmade structures under-road culverts and tunnels, and associated surrounds within WHITE s (1983) undifferentiated Sudanian woodland [29a]. Species were recorded during visual encounter surveys (15.X.96) at a. 32 C (13:22-17:51 hr) and b C (09:33-13:19 hr and 18:02-18:13 hr). a. Niokola-Koba b. Kédougou to Dar Salam, National Park excluding Niokola-Koba National Park (n = 55) (n = 108) Lizards Agama agama Mabuya quinquetaeniata scharica Agama sankaranica Tarentola parvicarinata Hemidactylus brooki Mabuya perroteti Agama weidholzi Varanus niloticus Snake Haemorrhois (Coluber) dorri Shannon-Weiner index of diversity (H ) Evenness No. of sites (road distance in km) 97 (101.0) 39 (65.0) Mean no. of reptiles per site road running through the Park (U. JOGER & M.R.K. LAMBERT pers. obs). The lower relative density of lizards by man-made road structures inside than outside the National Park, and complete absence of geckos from such structures inside the Park, may have been due to the concentration of large numbers of these Park primates, since lizards are very well known to be included with the diet of both species (DART 1963, NAPIER & NAPIER 1967, BUTYNSKI 1982, CHISM et al. 1984). The rather higher relative densities of swift-running Agama and Mabuya species than those of other genera inside than outside the Park may also indicate that they are less readily caught by these quick, deft-fingered vertebrate predators than geckos. Faunal overlap between the inside and outside of the Park may have been somewhat reduced on account of this, notwithstanding that resemblance was still relatively high (Table 5iii). It may also indicate that the Park s primate carrying capacity had been exceeded which may need to be considered in future Park management. REVISED CHECK LIST OF THE AMPHIBIANS AND REPTILES OF THE NIOKOLA-KOBA NATIONAL PARK Check lists of amphibian and reptile species in the Niokola-Koba National Park have been produced previously (LOVERIDGE 1956, VILLIERS 1956), with additional species provided by other authors (LAMOTTE 1969, JOGER 1982). Herewith, the opportunity is provided for an update on these earlier lists. In the list below,

17 180 U. Joger and M.R.K. Lambert Table 5. Statistical comparisons of reptile faunas in different areas. Areas of compared No. of species in Faunal overlap between Resemblance species common/total spp. areas (Sorensen s of dominance similarity index - %) (Renkonen s coefficient) (i) Different vegetation zones 2/9 36 (two) in SE Senegal (Table 2) (ii) Undisturbed sites/man- 3/ made structures (Table 3) (iii) Man-made structures 4/ inside/outside Niokola-Koba National Park (Table 4) (iv) West/East of Kédougou: a. Undisturbed sites 4/ (Tables 2b/3a) b. Man-made structures 6/ (Tables 3b/4b) (v) SE Senegal a. N & W Senegal lizards only 3/ (Table 1) b. W Mali lizards only 11/17 79 (after JOGER & LAMBERT 1996) species in square brackets have not been recorded inside the boundaries of the Niokola-Koba National Park, but in localities adjacent to it, and almost certainly occur also within the Park itself. Amphibia (total 23, plus three probable) Bufonidae: Bufo xeros, [Bufo maculatus], Bufo regularis; Hyperoliidae: Afrixalus cf. fulvovittatus, A. weidholzi, Hyperolius nitidulus,?h. lamottei, Leptopelis viridis, [Kassina fusca new record], K. senegalensis; Ranidae: Ptychadena bibroni, Pt. pumilio, Pt. oxyrhynchus, Pt. schubotzi, Pt. longirostris,?pt. superciliaris, Pt. tournieri, Pt. trinodis, Pt. floweri, Hylarana galamensis, Hoplobatrachus occipitalis, Phrynobatrachus (accraensis) latifrons, Ph. calcaratus, Ph. francisci; Hemisotidae: Hemisus marmoratus; Microhylidae: [Phrynomantis microps]. Reptilia (total 39, plus five probable) Testudinidae: Kinixys belliana nogueyi; Trionychidae: Cyclanorbis senegalensis; Pelomedusidae: Pelomedusa subrufa, Pelusios subniger; Crocodylidae: Crocodylus niloticus, C. cataphractus, Ostaeolaemus tetraspis; Gekkonidae: Hemitheconyx caudicinctus,

18 Amphibians and reptiles in SE Senegal 181 Hemidactylus brooki complex, [Tarentola ephippiata], T. parvicarinata; Agamidae: Agama agama, A. sankaranica new record, A. weidholzi; Chamaeleonidae: Chamaeleo senegalensis, Ch. gracilis gracilis; Scincidae: [Mabuya affinis], M. perroteti, M. quinquetaeniata scharica new record, [Chalcides thierryi pulchellus new record]; Varanidae: Varanus niloticus; Typhlopidae: Typhlops punctatus punctatus, T. lineolatus lineolatus; Leptotyphlopidae: Leptotyphlops macrorhynchus; Boidae: Python sebae sebae; Atractaspidae: Atractaspis dahomeyensis; Colubridae: [Afronatrix anoscopus], Crotaphopeltis hotamboeia, [Haemorrhois (Coluber) dorri], Dispholidus typus typus, Grayia smythi, Lamprophis fuliginosus, L. lineatus, Lycophidion albomaculatum, Meizodon coronatus, Philothamnus irregularis, Prosymna meleagris greigerti, Psammophis elegans, Ps. (sibilans) phillipsi, Rhamphiophis oxyrhynchus oxyrhynchus; Viperidae: Bitis arietans, Causus maculatus, Echis carinatus ; Elapidae: Naja nigricollis. BIOGEOGRAPHICAL ANALYSIS OF AMPHIBIAN AND REPTILE SPECIES IN SENEGAL A comparison of faunal overlap between lizards collected in SE Senegal and northern and western Senegal (Table 5va), with the corresponding index value between lizard faunas of SE Senegal and W Mali (Table 5vb), shows a considerably higher similarity between the latter two. This is consistent with SE Senegal and W Mali being part of a particular area of endemism that also extends into northern Guinea and southern Mauritania (see JOGER & LAMBERT 1997). Lizard species that are endemic to that upper Senegal river region include Agama weidholzi, Agama boulengeri Lataste 1886 and Tarentola parvicarinata; endemic snakes are Rhinoleptus koniagui Villiers 1956 (southern Guinea) and Echis jogeri Cherlin 1990 (southwestern Mali); endemic anurans may be Bufo chudeaui Chabanaud 1919 and Schoutedenella milletihorsini (Angel 1922) (both south-western Mali). The four last mentioned species have not yet been documented from the territory of Senegal. Bufo chudeaui has not been recorded since its description and its existence is doubtful (RÖDEL 2000). Other species, such as Agama sankaranica, Cophoscincopus cf. greeri (the population at the Senegalese/Guinean border may be an endemic taxon), Mabuya quinquetaeniata scharica, Chalcides thierryi pulchellus, Atractaspis dahomeyensis and Haemorrhois (Coluber) dorri, reach their westernmost distribution limit in SE Senegal. Their absence in western and central Senegal can probably be attributed more to ecological than historical reasons. Ecological differences between SE and W Senegal include lack of rocky habitat in western Senegal (except Cap Vert peninsula), differences in vegetation (Sudanian Baobab woodland in western Senegal and dry forest dominating in SE Senegal) and climate (dry season of long duration, but wet season with higher rainfall in SE Senegal than in most of subdesertic western Senegal, except in the Casamance area, which has a maritime wet tropical climate). ACKNOWLEDGEMENTS The study was contiguous with a consultancy by M.R.K. Lambert (10 September to 7 October 1996) for FAO Project GCP/SEN/041/NET (Locustox), Dakar, and we would like to thank Wim C. Mullié for assistance in its implementation. We are indebted to Marc-Oliver Rödel, University of Würzburg, for his help in identifying the anurans collected. Thanks are

19 182 U. Joger and M.R.K. Lambert also due to Wolfgang Böhme and Arne Schiøtz for useful comments on the manuscript, and to Valérie Obsomer (Natural Resources Institute, University of Greenwich, Chatham) for preparing the points map. REFERENCES BÖHME W Zur Herpetofaunistik des Senegal. Bonner Zoologische Beiträge 4 (29): BÖHME W., SCHMITZ A. & ZIEGLER T A review of the West African skink genus Cophoscincopus Mertens (Reptilia: Scincidae: Lygosominae): resurrection of C. simulans (Vaillant, 1884) and description of a new species. Revue Suisse de Zoologie 107: BUTYNSKI T.M Vertebrate predation by primates: a review of hunting patterns and prey. Journal of Human Evolution 11: CHISM J., ROWELL T. & OLSON D Life history patterns of female patas monkeys, pp In: Small M.F., Edit. Female primates. New York: A.R. Liss, 272 pp. CISSÉ M Les geckos du Sénégal. Notes Africaines 144: CISSÉ M. & KARNS D Les sauriens du Sénégal. Bulletin de l Institut Fondamental d Afrique Noire (A) 40: DART R.A The carnivorous propensity of baboons. Symposium of the Zoological Society of London 10: GRANDISON A.G.C On a collection of lizards from West Africa. Bulletin de l Institut Fondamental d Afrique Noire (A) 18: GRUSCHWITZ M., LENZ S. & BÖHME W Zur Kenntnis der Herpetofauna von Gambia (Westafrika). Teil 1: Einführing, Froschlurche (Amphibia, Anura), Schildkröten (Reptilia, Chelonia), Krokodile (Crocodylia) und Echsen (Sauria). Herpetofauna 13 (75): GUIBÉ J. & LAMOTTE M Révision systématique des Ptychadena (Batraciens Anoures Ranidés) d Afrique occidentale. Bulletin de l Institut Fondamental d Afrique Noire (A) 19: HOWELL K.M African Reptile and Amphibian Specialist Group. Species, newsletter of the IUCN Species Survival Commission 18: 51. JOGER U Zur Ökologie und Verbreitung wenig bekannter Agamen Westafrikas (Reptilia: Sauria: Agamidae). Salamandra 15: JOGER U Zur Herpetofaunistik Westafrikas. Bonner Zoologische Beiträge 32: JOGER U Recherches scientifiques dans les Parcs nationaux du Sénégal. XIV. Quatre lézards nouveaux pour la faune du Parc national du Niokolo-Koba. Mémoires de l Institut Fondamental d Afrique Noire 92: JOGER U The venomous snakes of the Near and Middle East. Beihefte Tübinger Atlas des Vorderen Orients, A 12. Wiesbaden: Reichert, 115 pp. JOGER U. & Lambert M.R.K Analysis of the herpetofauna of the Republic of Mali, I. Annotated inventory, with description of a new Uromastyx (Sauria: Agamidae). Journal of African Zoology 110 (1): JOGER U. & LAMBERT M.R.K Analysis of the herpetofauna of the Republic of Mali, II. Species diversity and biogeographical assemblages, pp In: Ulrich H., Edit. Tropical biodiversity and systematics. Proceedings of the International Symposium on Biodiversity and Systematics in Tropical Ecosystems, Bonn Bonn: Zoologisches Forschungsinstitut und Museum Alexander Koenig, 357 pp. LAMBERT M.R.K Assessing the potential of lizards as bioindicators to monitor the environmental impact of pesticides in Mauritania and Sénégal (10 September to 7 October 1996). Chatham (UK): NRI report to FAO Project GCP/SEN/041/NET (Locustox), Dakar, Sénégal (unpublished), V + 71 pp. LAMOTTE M Types de répartition géographiques de quelques batraciens dans l Ouest Africain. Bulletin de l Institut Fondamental d Afrique Noire (A) 28:

20 Amphibians and reptiles in SE Senegal 183 LAMOTTE M III. Amphibiens. Deuxième note. In: Le Parc National du Niokolo-Koba (Sénégal). Mémoires de l Institut Fondamental d Afrique Noire 84: LAMOTTE M. & OHLER A Redécouverte de syntypes de Rana bibroni Hallowell, 1845, désignation d un lectotype et description d une espèce nouvelle de Ptychadena (Amphibia, Anura). Zoosystema 19 (4): LATASTE F Description d un ophidien diacrantérien nouveau (Periops dorri, n. sp.) originaire du Haut-Sénégal. Le Naturaliste (2) 38 (10): LOVERIDGE A VI. Amphibiens. In: Le Parc National de Niokolo-Koba, I. Mémoires de l Institut Fondamental d Afrique Noire 48: MAGURRAN A.E Ecological diversity and its measurement. London: Croom Helm, 179 pp. MERTENS R Über eine Froschsammlung aus Westafrika. Zoologischer Anzeiger 123: MÜHLENBERG M Freilandökologie. Heidelberg: Quelle & Meyer, 214 pp. NAPIER J.R. & NAPIER R.H Handbook of living primates. London and New York: Academic Press, 456 pp. ROCHEBRUNE A.-T. DE 1884a. Reptiles. Faune de la Sénégambie. Paris: Octave Doin, 222 pp., 20 pls. ROCHEBRUNE A.-T. DE 1884b. Amphibiens. Faune de la Sénégambie. Paris: Octave Doin, 50 pp., 10 pls. RÖDEL M.-O Herpetofauna of West Africa. Vol. I. Amphibians of the West African Savanna. Frankfurt am Main: Edition Chimaira, 332 pp. SCHIØTZ A The treefrogs of West Africa. Spolia Zoologica Musei Hauniensis 25: SCHIØTZ A The treefrogs of Africa. Frankfurt am Main: Edition Chimaira, 350 pp. TRAPE J.-F Haemorrhois (= Coluber) dorri (Boie s Racer). Herpetological Review 28: TRAPE J.-F. & MANE Y Afronatrix anoscopus (African Brown Water Snake). Herpetological Review 26: 156. UNESCO-FAO Bioclimatic map of the Mediterranean zone [prepared by L. Emberger, H. Gaussen, M. Kassas & A. de Philippis]. Paris: UNESCO-FAO Arid Zone Research 21, 1 p. VILLIERS A V. Reptiles. In: Le Parc National du Niokolo-Koba (Sénégal). Mémoires de l Institut Fondamental d Afrique Noire 48: WHITE F The vegetation of Africa. Paris: UNESCO, 356 pp. APPENDIX Observation and collection localities, with species recorded to give an indication of richness: (1) Dar Salam ( N, W), 9.X.96, Bufo xeros, Tarentola parvicarinata, Agama agama. (2) 3-5 km SE of Dar Salam ( N, W) (two sites), 16.X.96, T. parvicarinata, A. agama. (3) km SE of Dar Salam, Niokola-Koba National Park ( N, W) (four sites), 16.X.96, B. xeros, A. agama, Mabuya quinquetaeniata scharica. (4) km SE of Dar Salam, Niokola-Koba National Park ( N, W) (two sites), 10, 16.X.96, Hoplobatrachus occipitalis, A. agama, A. sankaranica. (5) km SE of Dar Salam, Niokola-Koba National Park ( N, W) (two sites), 16.X.96, A. agama, M. q. scharica. (6) km NW of Mako, Niokola-Koba National Park ( N, W) (two sites), 16.X.96, A. agama, M. q. scharica. (7) km NW of Mako, Niokola-Koba National Park ( N, W) (five sites), 10, 16.X.96, H. occipitalis, A. agama, Mabuya perroteti, M. q. scharica. (8) km NW of Mako, Niokola-Koba National Park ( N, W) (four sites), 10, 16.X.96, H. occipitalis, A. agama, A. sankaranica, M. perroteti, M. q. scharica.