The Amphibian Fauna of Pendjari National Park and Surroundings, Northern Benin
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The amphibian fauna of Pendjari National Park SALAMANDRA 42 2/3 93-108 Rheinbach, 20 August 2006 ISSN 0036-3375 The amphibian fauna of Pendjari National Park and surroundings, northern Benin S. GILLES A. NAGO, OLAF GRELL, BRICE SINSIN & MARK-OLIVER RÖDEL Abstract. We present the results of the first comprehensive survey of the amphibian fauna of a savannah region in Benin, West Africa. We herein add 17 species to the countries’ amphibian list: Bufo pentoni, Hildebrandtia ornata, Pyxicephalus cf. edulis, Ptychadena bibroni, P. cf. schillukorum, P. tellini, P. tournieri, P. trinodis, Phrynobatrachus accraensis, P. francisci, P. gutturosus, Arthroleptis sp., Kassina cassinoides, K. fusca, K. senegalensis, Leptopelis bufonides and Hyperolius nasutus. We comment on the taxonomy and biology of some rare and/or unusual species. The amphibian species richness of the Pendjari area is among the most diverse of African savannahs. The species assemblage is mainly composed of ‘typical’ West African savannah frogs, especially those that are restricted to drier habitats. However, especially along the mountainous Atakora chain in the South of the area, we also recorded species that are dependent on higher humidity, e.g. Arthroleptis sp., Hyperolius sp. In the future, more emphasis should be given to survey this mountainous area that might prove to have close faunal affini- ties to the Togolese mountains. Key words. Amphibia, Anura, Benin, diversity, savannah, taxonomy, West Africa. Résumé. Nous présentons les résultats du premier aperçu complet des amphibiens d’une région de sa- vane au Bénin, l’Afrique de l’Ouest. Nous ajoutons ainsi 17 espèces à la liste des amphibiens du pays: Bufo pentoni, Hildebrandtia ornata, Pyxicephalus cf. edulis, Ptychadena bibroni, P. cf. schillukorum, P. tellini, P. tournieri, P. trinodis, Phrynobatrachus accraensis, P. francisci, P. gutturosus, Arthroleptis sp., Kassina cassinoides, K. fusca, K. senegalensis, Leptopelis bufonides et Hyperolius nasutus. Nous avons discuté sur la taxonomie et la biologie de quelques espèces rares et/ou peu communes. La rich- esse spécifique des espèces d’amphibiens de la Réserve de Biosphère de la Pendjari, milieu d’étude, fait d’elle l’une des régions africaines savanicoles les plus diversifiées. La communauté d’amphibien se compose principalement d’espèces «typiques» de savane d’Afrique Occidentale, particulièrement ceux qui sont limitées à des habitats secs. Cependant, particulièrement le long de la chaîne montagneuse de l’Atakora dans le Sud du milieu d’étude, nous avons également enregistré les espèces qui dépendent d’une humidité plus élevée, par exemple Arthroleptis sp. et Hyperolius sp. À l’avenir, les efforts devraient être concentrés sur cette chaîne qui pourrait avoir des affinités de faune que la partie togolaise. Mots clés. Amphibia, Anura, Bénin, diversité, savane, taxonomie, Afrique occidentale. Introduction sion of military governments ended in 1972 with the rise to power of MATHIEU KEREKOU Benin is a comparatively small West African whose government had changed the former country (112,622 km²), situated right in the name Dahomey to the People’s Republic of Dahomey Gap, that is a natural gap in the Benin in 1975. In 1991 the first successful coastal rainforest belt of West Africa. Da- transfer of power in Africa from a dictator- homey also was the former country’s name, ship to a democracy resulted in the election originating from the equally named famous of NICEPHORE SOGLO as president. West African kingdom that rose in the 15th Maybe it is for the combination of long century. This territory became a French Col- lasting political unrest and the lack of rain- ony in 1872 and achieved independence in forest, that Benin only rarely was the target 1960, as the Republic of Dahomey. A succes- of zoological surveys (ATACHI et al. 1994). © 2006 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT) http://www.salamandra-journal.com 93 S. GILLES A. NAGO et al. We are aware of only three herpetological these mountains and a few isolated hills, the studies, all unpublished. In 2002, O. GRELL topography of the reserve is generally very started to investigate the Pendjari area in the flat and mostly ranges between 150-200 m North (GRELL 2003), whereas DAOUDA et al. above sea level (DELVINGT et al. 1989). PBR (1998) and ULLENBRUCH (2003) investigated has a Sudanian climate with a seven month one of the few forests in southern Benin. dry period. The mean annual precipitation These contributors found 17 anuran species is 1,000 mm; most of the rain is falling each. Although GRELL (2003) already postu- during a period between late May and early lated the occurrence of many more amphib- October. The mean annual temperature is ian species in the Pendjari area, in general 27 °C (VERSCHUREN 1988). There are a few the available data pointed into the direction large and permanent ponds in PNP. Most of a very poor amphibian fauna for Benin. savannah waters are temporary and form only This is summarized in the current Global after heavy rains in May/June and dry up in Amphibian Assessment (GAA) that lists only October/November (Fig. 2). The vegetation 22 amphibian species for the country (IUCN of the PNP is a mixture of different savannah et al. 2004). However, from other West Af- types, mostly open shrub and tree savannah rican countries it is known that even savan- (Fig. 2). Small island forests are situated in nah habitats may harbour a rich amphibian the areas of former villages. Patches of dry fauna (e.g. LAMOTTE 1967, RÖDEL & SPIELER forests and open gallery forests along water 2000, JOGER & LAMBERT 2002, GREENBAUM courses are present. Huge areas in the North & CARR 2005). We therefore started herpeto- of the reserve are flooded (and thus inacces- logical investigations in one of West Africa’s sible) during the rainy season and only very largest protected areas and its surroundings sparsely vegetated during the dry season. In – the Pendjari biosphere reserve. This paper the periphery of the PNP the landscape is presents the amphibian data of this intensive dominated by fields and fallows. Cultivated survey work. crops are amongst others: rice, yams, maize and cotton, the latter requiring intense use of pesticides (DJIBRIL 2002, CENAGREF 2003). Study site The savannah in the buffer zone is also used for grazing cattle and intensively collecting The Pendjari Biosphere Reserve (PBR) is firewood. So far only the larger mammals situated in the Atakora department, north- of the reserve have attracted considerable western Benin (10°30’ to 11°30 N; 0°50’ scientific interest (VERSCHUREN 1988, SINSIN to 2°00’ E; Fig. 1). It was declared a Game et al. 2000, LAMARQUE 2004). Reserve in 1954 and upgraded to a Nation- al Park in 1961. It nowadays comprises a strictly protected core area (the Pendjari Na- Material and methods tional Park, PNP: 2,660 km²) and a buffer and hunting zone in the West and South of In February 2002 and in October 2003, O. it (1,870 km²). In the North and East, the GRELL started accomplishing first faunal River Pendjari forms a natural border of the surveys for fishes, amphibians, reptiles and PBR that in the North is also the country’s birds in PBR (GRELL 2003). With respect border to Burkina Faso (DELVINGT et al. 1989, to amphibians, he applied visual encounter MAB-UNESCO 1990, BRUCKER 2001). The surveys (VES) and dip-netting (DN). Since River Pendjari is the most important water September 2004, S.G.A. NAGO has regularly course of PNP. In the South the reserve is investigated all possible kinds of waters with- bordered by the Atakora chain (400-513 m in and outside the reserve for amphibians by above sea level), which is directly connected employing VES and acoustic encounter sur- to mountainous northern Togo. Apart from veys (AES) and the box method for tadpoles 94 The amphibian fauna of Pendjari National Park (HEYER et al. 1994, RÖDEL 1998, RÖDEL & 2005 we could recapture all those and add ERNST 2004). The latter investigations are de- several species, resulting in a total of 32 signed to compare the amphibian community species. Based on presence/absence data of composition in pristine and altered savannahs the 19 days of intensive search we estimated (S.G.A. NAGO et al. unpubl. data). During two for the Pendjari region a total of 33.5 ± 2.2 especially intensive searching periods, 1-8 (Chao2) and 35.8 ± 1.7 species (Jack-knife September 2004 and 27 May to 8 June 2005, 1), respectively (Fig. 3). This suggests that S.G.A. NAGO and M.-O. RÖDEL employed we recorded 89.4-95.5 % of the regional VES and AES in search for adult frogs, ac- amphibian fauna. companied by DN for tadpoles. During these periods we tried to find as many species per day as possible to achieve a complete an- Species account uran species list for the region. We thereby tried to investigate all possible habitat types A complete list of recorded amphibian spe- throughout the area. We tested the efficien- cies is summarized in Appendix 1. Most spe- cy of our inventories with the Jack-knife cies are well known West African savannah 1 and Chao2 estimators (software package anurans, already dealt with in detail in RÖDEL EstimateS, COLWELL 1994-2000; cf. VEITH et (2000). Herein we only comment on selected al. 2004). Calculation bases were the daily species of particular interest. presence/absence lists gathered throughout Bufo pentoni ANDERSSON, 1893 lives in the two intensive sampling periods (total 19 xeric habitats south of the Sahara. Known days). To prevent order effects we performed species records range from Mauritania east- 500 random runs of our daily species lists ward to the Red Sea coast of Egypt, Sudan, and calculated mean and standard deviation Eritrea and Djibouti.