Anuran Surveys in South-Western Nigeria

Home , Amnirana, Leptopelis, Leptopelis viridis, Nectophryne, Phlyctimantis, Phlyctimantis boulengeri

Anuran surveys in south-western Nigeria

SALAMANDRA 45 1 1-14 Rheinbach, 20 February 2009 ISSN 0036-3375

Anuran surveys in south-western Nigeria

Abiodun B. Onadeko & Mark-Oliver Rödel

Abstract. We report on the results of amphibian surveys in three south-western Nigerian states: Lagos, Ogun and Oyo. In total, we recorded between 35 and 38 species. The number of species per state ranged from 22 to 29. Most species were typical farmbush or savanna frogs. Although the overall species richness was comparatively low and only a few species were forest specialists, we observed a surprising number of frogs that could not unambiguously be assigned to described species. This was especially true of the genera Leptopelis and Phrynobatrachus. Whereas some of these taxa (Leptopelis cf. macrotis, Phrynobatrachus cf. liberiensis) seem to indicate closer relationships between south-western Nigeria and the Upper Guinea forests, other ﬁndings (the eastern variation of Phlyctimantis boulengeri, Nectophryne afra) indicate the opposite geographic relation. The position of the study sites between the diversity hotspots of Upper and Lower Guinea would suggest high anuran diversity. We failed to record such a high diversity, but possibly found species from both areas and probably undescribed taxa (Phrynobatrachus sp., Leptopelis sp.). We believe that the high amount of degraded and converted natural forests may have already resulted in an impoverished anuran fauna. Remarkable and even new species may have survived in some of the small remaining forests. Intensive surveys and sustainable conservation of the remaining forests of south-western Nigeria is therefore highly recommended. Key words. Amphibia, distribution, diversity, biology, forest, new records, savanna.

Introduction of West and Central African species alike. So far, about 03 amphibian species, 0 anurans With 30-40 million people, Nigeria is the and two caecilians, have been recorded from most densely populated African country. within Nigeria’s borders (IUCN et al. 2006). The high human population with the result- This is astonishingly few, compared to the ap- ing pressure on natural habitats, as well as the proximate 200 species of neighbouring Cam- country’s position of being the twelfth largest eroon (LeBreton 999, Frétey & Blanc producer of oil, has resulted in a remarkably 2000, IUCN et al. 2006, J.-L. Amiet pers. high proportion of degraded and converted comm.). However, it is questionable whether land (www.en.wikipedia.org/wiki/Nigeria; Cameroon is really that much richer in am- latest accessed 4 May 2007). In contrast, Ni- phibian diversity or if simply Nigeria is less geria also comprises a great variety of ecosys- well explored. tems, ranging from mangrove swamps and Many contributions to the knowledge rainforests along the coast, various types of of the country’s amphibian fauna date back savanna further north, to finally the Jos-Pla- to the sixties of last century (Schiøtz 963, teau in Central Nigeria and the south-eastern 966, 967, Walker 966, 967, 968, 969) or region towards Cameroon offering moun- even earlier (Sanderson 936, Romer 953). tainous habitats. From a biogeographic point Due to long lasting political unrest, with the of view, Nigeria is situated between West and exception of a few papers (Akintola 976, Central Africa, and hence between the bio- Reid 987, Böhme & Nikolaus 989, Reid et diversity hotspots of the Upper and Lower al. 990) not much has been published on Ni- Guinea forest regions (Bakarr et al. 2004). gerian amphibians for the last decades. With- Consequently, Nigeria might be expected to in the last five years several papers on Nige- harbour high biological diversity composed rian amphibians have appeared again, mainly

Fig. 1. Geographic positions of the study sites in south-western Nigeria (compare Appendix 1). dealing with the Cross River region and the ger, Chad and Cameroon. The Federal Re- area of the oil rich Niger Delta, including the public comprises 36 states and the Federal respective environmental problems (Eniang Capital Territory. We investigated the anuran & Luiselli 2002, Lea et al. 2003, 2005, Aka- fauna of three south-western states: Lagos, ni et al. 2004). Ogun and Oyo. Lagos, also comprising the However, still little is known about the sys- homonymous commercial capital, is situat- tematics, distribution and especially biology ed between 6.4-6.7°N and 2.7-4.6°E. Ogun of Nigerian amphibians. Remarkable excep- extends between 6.3-7.8°N and 2.7 4.6°E; tions are the contributions on frog parasites and Oyo extends between 7.-9.2°N and 2.7- by Aisien et al. (e.g. 200, 2004). Nigeria has 4.56°E. Nigeria is characterised by a South- lost the large majority of its rainforests and North gradient of precipitation. This is mir- most of its natural savanna habitats (UICN rored by a gradient of vegetation zones, nat- 996). We know that already minor alteration urally from tropical humid evergreen forests of natural forest ecosystems in West Africa in the South, gradually transforming into dry may result in drastically altered amphibian savannas in the North. assemblages, including the loss of many for- Along the south-western Nigerian coast est specialists (Ernst & Rödel 2005, Ernst there is a belt of mangrove swamps which et al. 2006). Hence, an increased eﬀort to as- naturally do not harbour any amphibians. sess herpetofaunistically unknown parts of However, some species can be recorded just Nigeria seems extremely pressing. The pri- a few metres from the brackish waters, where mary aim of this paper is to present the am- this habitat is replaced by freshwater swamps. phibian results gained throughout several Such areas were located at Ikorodu, Badore, surveys undertaken by one of us (ABO) in Badagry and parts of Epe (all Lagos state). the south-western part of Nigeria. North of these swamps, a forest belt does naturally occur. These forests can be divided into rainforest and mixed deciduous or dry Study areas forest. Unfortunately, these forests have been heavily degraded, fragmented and convert- Nigeria covers an area of 923,768 km² and is ed by many anthropogenic activities, main- bordered by the Atlantic Ocean, Benin, Ni- ly cultivation for agriculture. The remaining

2 Anuran surveys in south-western Nigeria

40 350 40 350

35 300 35 300

30 30 250 250 C) 25 C) 25 200 200 20 20 150 150 15 15 temperature (° temperature (° precipitation (mm) 100 precipitation (mm) 100 10 10

5 50 5 50

0 0 0 0 a Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec b Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

40 450 40 300

35 400 35 250 350 30 30 300 200 C) 25 C) 25 250 20 20 150 200 15 15 temperature (° 150 temperature (° 100 precipitation (mm) precipitation (mm) 10 10 100 50 5 50 5

0 0 0 0 c Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec d Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Fig. 2. Monthly rainfall (black triangles) and monthly average maximum (open squares) and minimum (open circles) temperatures in a) Ijebu-Ode (data from 2003), b) Ikeja (2003), c) Ibadan (2003) and d) Ilorin (2002). forests hence mostly comprise secondary or Climatologically, the three states are charac- tertiary vegetation, characterised by dense terised by a rainy season from April to Oc- undergrowths and an open or broken cano- tober and a dry season from November to py. Farmland, secondary and tertiary forests March. The climate of four south-western Ni- were encountered at Ikorodu, Ogijo, Ishara, gerian towns is summarized in Fig. 2. Ijebu Oru, Ikenne, Onidundun and Fiditi. There only a few pockets of primary or near primary forest were left. Further north, this Material and methods zone is replaced by natural and derived savanna, partly comprising a forest-savanna Our surveys were performed between July mosaic, especially where cultivation is less 2003 and December 2004. The most intense dominant. Examples from this zone were survey time was during the rainy season Onidundu, Fiditi, Oolo, Ibapon Oyewale and which at the same time is the breeding pe- Ogbomosho. The northernmost collection riod of most amphibian species. Between two sites were situated in the moist woodland sa- and four people were engaged in surveying. vanna zone (Guinea savanna), characterised Surveys were normally undertaken during by tall grass with ﬁre resistant trees. Collec- the morning and evening hours. Geographic tion in this zone was done in areas north of positions were taken with a hand-held GPS Ogbomosho, including Gambari and Otte- receiver (Garmin eTrex). A short habitat Egba, an area close to Ilorin. characterisation was noted for each location A list of all study sites including a short investigated and all anurans were registered habitat description is given in Appendix . along with their respective habitats and loca-

3 Abiodun B. Onadeko & Mark-Oliver Rödel

Tab. 1. Amphibian species recorded in Lagos, Ogun and Oyo States and number of sites where the respective species have been recorded (compare Appendix 1). Taxon Lagos Ogun Oyo Pipidae Silurana tropicalis 7, 9, 0 3, 9,  , 2 Xenopus muelleri 4, 8, 9, 0 2, 3,  2, 6, 0 Hemisotidae Hemisus cf. guineensis 9 , 3, 0,  H. marmoratus 2, 3, 7 Bufonidae Amietophrynus maculatus 2, 3, 6-0 -2 -2, 4-8, 0,  A. regularis -4, 3, 4, 6, 8, 0 -3, 5-7, 9-2 2, 3, 5-8, 0,  Nectophryne afra  Dicroglossidae Hoplobatrachus occipitalis , 5, 8- 2-5, 0,  , 2, 5, 6, 8 Ranidae Hydrophylax albolabris 7, 9 2, 8, 0, 2 H. galamensis 2, 6 , 2, 6, 9 Aubria subsigillata 6-8 , 3, 9, 0 2, 3 Ptychadenidae Ptychadena aequiplicata 9,  2 P. bibroni , 3, 5, 8 , 2, 4, 0,  2, 3, 5, 6 P. longirostris 8, 9  P. mascareniensis , 0,  3, 4, 0,  2, 3 P. oxyrhynchus , 6, 0,  , , 2 2, 5 P. pumilio -6, 8,  -4, 6, , 2 2, 3, 5, 6, 8- Phrynobatrachidae Phrynobatrachus accraensis , 2, 6, 8, 0 4, 9,  2, 9, 0 P. francisci 7, 9 3, 7, 9 2, 9, 0 P. cf. liberiensis 7 P. plicatus 2 P. sp. 9 Arthroleptidae Arthroleptis spp. (2-4 species) 7-9 2, 3, 8, 9, , 2 2 Leptopelis cf. macrotis 0 L. cf. occidentalis/boulengeri 6, 0 L. sp. 9 L. viridis 7-9 , 3, 5, 7, 8 3-5, 7, 9 Hyperoliidae Afrixalus dorsalis , 4, 5, 8,  5-7, 9,  2, 5 Hyperolius concolor 3, 4, 8 6, 8, 9, , 2 2, 3 H. fusciventris 3-8,  2, 6, 8-2 2-4, 6 H. guttulatus 4, 0 6,   H. nasutus 5 H. nitidulus 2-4, 7, 9,  Kassina senegalensis 7, 0 Phlyctimantis boulengeri 2

4 Anuran surveys in south-western Nigeria tions. Specimens were located opportunisti- Accounts of selected species cally by thorough visual scanning of the particular environment and careful examination We herein refer to species of particular inter- of suspected hiding places, e.g. under stones, est concerning rarity, distribution or taxon- fallen wood, between leaves, among leaf lit- omy or to interesting observations on habi- ter etc. Most of the specimens were released tat choice and/or biology. For descriptions after determination and recording of their of the species’ general distributions and nat- characters. Measures were taken with a cal- ural history data we refer to Schiøtz (963, liper (± 0. mm). Voucher specimens of all 967, 999), Rödel (2000) and IUCN et al. species were collected, anaesthetized with (2006). chloroform and subsequently preserved in 0 % formalin. Most vouchers are deposited in Xenopus muelleri (Peters, 1844) and the collection of the Department of Zoology, Silurana tropicalis Gray, 1864 University of Lagos. Some vouchers will later These species seem to survive the dry season be deposited in the collection of the Museum by either migrating into streams or other per- of Natural History, Berlin (ZMB). Nomencla- manent waters or by hiding in wet mud or ture follows Frost et al. (2006) and Frost beneath leaves (Schiøtz 963, Rödel 2000). (2007). We recorded Xenopus muelleri as fairly abun- dant at Ijebu Oru (Liberian Refugee Camp), Onidundun and Abowa. The sites at Ijebu Results and discussion Oru and Abowa were located near a forest, while Onidundun was a muddy patch in a In total we recorded between 35 and 38 species rice farm. These sites all dried up during the in the three south-western states (Table ). dry season. In a drying savanna pond at Oolo Within the genus Arthroleptis, we most prob- we collected some X. muelleri from the mud. ably recorded between two and four species. We discovered further X. muelleri in holes, However, because of the lack of acoustic and about half a metre depth, around the drying genetic data, and the morphological similar- up pond. ity of species in this genus, a more precise ﬁg- Most of the registered S. tropicalis were ure and exact species names cannot be given found at Onidundun in a pool not far from a (cf. Rödel & Bangoura 2004). We therefore large stream, close to a forest. However, oth- conservatively count all Arthroleptis records er specimens were collected near Ogbomoso as belonging to one species. In Lagos state in a savanna region. We dug them up on the we recorded 22 species (62.9 %), in Ogun banks of dried up ponds at Onidundun. state we registered 29 species (82.9 %) and in Oyo state we encountered 25 species (7.4 Hemisus marmoratus (Peters, 1854) and %). Without considering the Arthroleptis spe- Hemisus guineensis Cope, 1865 cies, we analysed to what extent the anuran These predominantly fossorial frogs were fauna in the three states was either dominat- collected at various sites, mostly by digging ed by forest or savanna species. As expected soil on farmland. Most likely two species are the northernmost state Oyo had the highest involved, namely the savanna dwelling Hemi- proportion of savanna species, i.e. 9 out of sus marmoratus and the forest species H. 24 (79.2 %). In Ogun state, about half each of guineensis. However, both apparently com- the 28 species were forest (46.4 %) and savan- prise complexes of species and despite the re- na (53.4 %) specialists, respectively. In Lagos vision of the genus by Laurent (972) even state, one third, i.e. 7 of all 22 (3.6 %) species, the distinction between the two names is not preferred forest habitats, while the remaining well understood. Specimens collected from 5 were savanna species. Ijede and Abowa were larger (snout-urostyle-

5 Abiodun B. Onadeko & Mark-Oliver Rödel length – SUL - males: 4.0-4.9 cm, females: 4.5-5.4 cm) than those collected from Fiditi and Ibapon Oyewale (males: 2.2-4.0 cm, females: 3.6-4.2 cm). We tentatively assign the larger Hemisus to H. guineensis, the smaller ones to H. marmoratus (compare Rödel & Ernst 2003). The H. guineensis localities were in forest or former forest.

Nectophryne afra Buchholz & Peters in Peters, 1875 So far the westernmost known locality of this small Central African toad was in the Niger Delta, West of the River Niger (Bouleng- er 93, Perret 966, Schiøtz 966). Our Fig. 3. The tree toad Nectophryne afra from Ogun State. records from a secondary forest with sparse undergrowth close to the Oru River at Ijebu are consumed locally directly by villagers or Oru is hence an expansion of the known dis- smoked and sold on local markets (A.O. and tributional range. This toad is known to breed M.-O.R. unpubl. data). in water-filled tree holes (Scheel 970). We observed it a few metres above ground climb- Ptychadena aequiplicata (Werner, 1898) ing on plants. The SUL was .9 cm. The toad We recorded Ptychadena aequiplicata mostly had a greenish brown back with three trans- from secondary and primary forests that had verse yellow lines, additional yellow spots little undergrowth (cf. Rödel et al. 2002). could be observed on the flanks (Fig. 3). The During the rainy season puddles accumulat- yellow pattern was also present on the hind ed in these parts of the forest. They were col- legs. The webbing between the thumb and lected at Ijebu Oru and Ijede. Schiøtz (963) the first finger was yellow, the remaining part collected this species from Yemoji and Gilli- being orange-brown. The toad only moved gilli but did no report it from south-western slowly in a very clumsy way. Nigeria.

Hoplobatrachus occipitalis (Günther, 1858) Ptychadena bibroni (Hallowell, 1845) Adults and juveniles of this frog were collect- This species was often collected together with ed at most locations though in diﬀerent hab- Ptychadena oxyrhynchus and P. pumilio. We itats. This habitat separation has also been once observed a male P. bibroni in amplexus observed elsewhere (Rödel 2000) and most with a female P. pumilio, but are not aware likely is due to avoidance of intraspeciﬁc pre- if egg deposition had taken place or not. We dation. Adult Hoplobatrachus are known to dug out a few P. bibroni together with X. mu- prey on almost anything smaller than them- elleri at a dried up pond at Oolo. This sup- selves (Rödel 2000), but usually arthropods ports Barbault (972) who stated that these predominate in the prey (Inger & Marx 96, frogs spent the dry season in the mud of dry Lescure 97). We observed an adult H. oc- ponds. cipitalis swallowing a Ptychadena sp. Another adult H. occipitalis (SUL 9.8 cm) tried to de- Ptychadena longirostris (Peters, 1870) vour a Bufo sp. (SUL 5.6 cm), but eventually We collected this species at two locations failed most likely due to the massive produc- only and observed it at another one. Pty- tion of white toxic secretions by the toad. In chadena longirostris inhabited both tempo- Nigeria and other West African countries H. rary and permanent ponds in secondary for- occipitalis, but also many Ptychadena spp., ests. Some were found in puddles on forest

6 Anuran surveys in south-western Nigeria roads. Ptychadena longirostris is usually con- sidered a rainforest dweller where it often re- produces in puddles on forest roads (Rödel & Branch 2002). However, Schiøtz (963) wrote that P. longirostris also invades dry forests in Nigeria.

Phrynobatrachus cf. liberiensis Barbour & Loveridge, 1927 This species is believed to be endemic to the Upper Guinean forest bloc, where it inhabits primary and secondary forests close to small creeks and swamps (Guibé & Lamotte 963, Rödel & Branch 2002, Rödel 2003). Its occurrence in Nigeria hence seems unlike- ly. Nevertheless, we collected a specimen in a secondary forest in Lagos State that mor- phologically exactly ﬁts the Upper Guinean populations. However, as long as genetic and Fig. 4. A subadult Leptopelis cf. macrotis. acoustic comparisons are lacking, aﬃliation of our voucher to P. liberiensis should be taken as tentative.

Phrynobatrachus plicatus (Günther, 1858) This species has a southern West African distribution from Liberia into Nigeria, where it predominantly inhabits open, swampy rainforests (Guibé & Lamotte 963, Rödel et al. 2005a, Leaché et al. 2006, Hillers & Rödel 2007, Segniagbeto et al. 2007). East of the Cross River, Nigeria, it is replaced by the similar Phrynobatrachus auritus (Lamotte & Xavier 966). We recorded the species in a Fig. 5. Leptopelis cf. occidentalis/boulengeri. secondary forest near Ogijo. One larger transverse bar was also present on Phrynobatrachus sp. the lower arms. The venter was white without We collected a few specimens of an unknown any pattern or with small black spots, espe- small Phrynobatrachus (SUL ca. .9-2.0 cm) cially in the pectoral region and the corner of during a light rain from a secondary forest the mouth. The toe tips were slightly enlarged not far from Ijebu-Ode along the Ijebu-Ode- but did not form a disc. The webbing on the Ibadan road (Ogun State). All specimens had feet was well developed. The webbing formu- a dark, almost black, head and eyelids. The la (sensu Rödel & Ernst 2002) was:  (0), 2 back was orange-red and the extremities were (0.5-0), 3 (), 4 (2), 5 (). brown. On the eyelids and the forehead a few Compared to similar sized West African larger warts were discernible. There were Phrynobatrachus these frogs diﬀer as follows: no warts or ridges on the back. The slightly from P. annulatus, P. calcaratus, P. taiensis granular back was almost coloured uniform and P. villiersi by lacking an eyelid cornicle red-brown. The hind legs bear one large dark, (cf. Perret 988); from all these species and transverse bar on thighs and lower legs each. P. gutturosus and P. tokba by the extensive

7 Abiodun B. Onadeko & Mark-Oliver Rödel webbing (Rödel 2000, Rödel et al. 2005b); descriptions of Leptopelis occidentalis and L. from P. annulatus, P. ghanensis, P. taiensis boulengeri. The dorsal coloration best fitted L. and P. villiersi by a different ventral pattern occidentalis. “Real” L. occidentalis are known or colour (Guibé & Lamotte 963, Perret from Liberia to central Ghana and “real” L. 988, Rödel & Ernst 2002); from all other boulengeri have been recorded from the Ni- species, except P. fraterculus (but completely ger Delta south to the Democratic Republic different body shape, see e.g. Fig. 3 in Rödel of Congo. According to Schiøtz (999), sim- & Bangoura 2004), by lacking discernible ilar frogs have been occasionally collected warts or ridges on the back, i.e. in the neck in western Nigeria. Without call recordings region. (different voice in both species) and/or ge- We hence believe these frogs may repre- netic analyses it is currently impossible to as- sent an undescribed species, possibly en- sign these frogs with certainty. L. occidentalis demic to south-western Nigeria, but refrain inhabits almost exclusively primary rainfor- from describing it before its taxonomic po- est in the Upper Guinea region. L. bouleng- sition can be clarified by acoustic and/or ge- eri seems to choose similar habitats (Perret netic data. 966, Amiet 986). Our frog was collected in a rubber plantation that enclosed a small Leptopelis viridis (Günther, 1869) swamp with tertiary vegetation. Farmers reported that Leptopelis viridis were Another juvenile Leptopelis sp. was caught dug up during the dry season while they were about 0 km from Ijebu Ode, sitting on a fern tilling their lands in preparation for the com- in a secondary forest that encircled a swamp. ing rainy season. Subterranean aestivation This small frog (SUL: . cm) had a very broad and cocoon formation were also reported by head, much wider than the body. The dorsal Lamotte (967), Walker (968), Barbault coloration was dark green with conspicu- (972) and Grafe (2000). Schiøtz (967) ous white spots on the heels and the elbows, collected this species from Iperin and four hence not fitting any described West African miles north of the University in Ibadan. Leptopelis (Rödel 2007). Another white spot was situated below the eyes. The hind limbs Leptopelis cf. macrotis Schiøtz, 1967 were fully webbed. We cannot decide wheth- This species is the largest of the West African er this is a juvenile of the above mentioned treefrogs (Schiøtz 967, 999); however, the species or represents another, potentially un- SUL of the sub-adult female that we collected described treefrog. was 5.2 cm only. Our decision to tentatively assign the frog to L. macrotis was due to its Hyperolius fusciventris burtoni coloration (Fig. 4) and the fully webbed feet Schiøtz, 1963 (cf. Schiøtz 967, 999). So far this species Most Hyperolius fusciventris burtoni were col- has been only recorded from rivers in pri- lected from leaves of the oil palm tree, Elais mary forests ranging from Sierra Leone into guineensis. Abandoned farmlands near ponds western Ghana (e.g. Rödel & Branch 2002, or streams also served as habitats. Schiøtz Rödel et al. 2005a, Hillers & Rödel 2007). (963, 967) collected specimens from Iperin, We collected the frog in a secondary forest Ijebu Oru, south of Oyo and 9 km north of beside a river along the Shagamu-Benin ex- University of Ibadan. pressway. Hyperolius concolor ibadanensis Leptopelis cf. occidentalis/boulengeri and Schiøtz, 1967 Leptopelis sp. Hyperolius concolor ibadanensis were collect- This is a medium sized treefrog with a SUL ed mostly in the forest region around ponds of 4.6 cm (Fig. 5). The white spot beneath the at Onidundu, Ikenne, Ijebu Oru and on the eye and the amount of webbing best fitted the campus of the University of Lagos. In the for-

8 Anuran surveys in south-western Nigeria est zone they could be found throughout the geriensis (F), A. vittiger (S), A. weidholzi (S), year in thick vegetation close to water sourc- Hyperolius sylvaticus (F) and Chiromantis es. rufescens (F; for exact localities cf. Schiøtz 963; taxonomy adopted to current names; F Hyperolius guttulatus Günther, 1858 = forest, FB = farmbush, S = savanna). The majority of individuals of the extremely variable Hyperolius guttulatus (cf. Schiøtz 967, Rödel 2000) were observed in swamps Conclusions and recommendations with large stagnant ponds, having dense vegetation, especially floating Pistia sp. in sec- We recorded a surprisingly high number of ondary forests and farmbush habitats. Ac- unexpected frog species, especially of the cording to Schiøtz (967) a typical habitat of treefrog genera Leptopelis (L. cf. macrotis, L. H. guttulatus comprises a large swamp with cf. boulengeri/occidentalis, L. sp.) and Necto- open water surrounded by shrubs but not by phryne, as well as within the leaf litter genus closed forests. We also collected a few speci- Phrynobatrachus (P. cf. liberiensis, P. plicatus, mens from leaves of oil palm trees. P. sp.). Some of these species may be new to science. If our tentative determinations prove Phlyctimantis boulengeri Perret, 1986 to be correct, this would indicate a potentially This treefrog is known from two disjunct ar- high anuran diversity comprising faunal ele- eas: (i) coastal Cameroon (including the type ments from the Upper (e.g. Phrynobatrachus locality) and further south plus easternmost liberiensis) and Lower Guinean (e.g. Nec- Nigeria and (ii) forests from Sierra Leone tophryne afra) forests, as well as potentially into western Ghana (Perret 986, Schiøtz endemic species (Phrynobatrachus sp.). The 999, Rödel et al. 2005a, A. Hillers unpubl. overall amphibian diversity of each state, so data). No records have been reported from far being not only comparatively poor with in-between. It therefore seemed likely that respect to more western localities (cf. Rödel these populations belong to two distinct spe- et al. 2004), but being even below that of par- cies. Our record represents the first record of ticular West African savanna regions (e.g. a frog in the distributional gap. Based on its Lamotte 967, Nago et al. 2006), seems to size (SUL: 5.5 cm) and coloration (dorsum contradict these first findings. A likely expla- beige, hidden part of thighs with traverse nation for these contradictory results is that yellow and black stripes) our frog resem- south-western Nigeria did indeed harbour bles more the eastern populations (Schiøtz a very rich amphibian fauna, but due to the 999). The single specimen originated from a tremendous loss of primary habitats a huge stagnant pond covered by Pistia sp., about 6 number of these species has already been km north of Ibadan. This pond was bordered lost, or at least is now restricted to very iso- by secondary forests and farmland. lated remnants of the former habitats. It there would appear to be extremely important to protect and maintain the remaining natural Further anuran records from or near natural habitats, in particular the for- south-western Nigeria ests (including small fragments) and to survey further forests for those species that so Schiøtz (963, 966) lists a few species for far may have escaped our attention. south-western Nigeria, which have not been recorded by us. In particular these are: Car- dioglossa leucomystax (F), Leptopelis spiritus- Acknowledgements noctis (F), Phrynobatrachus calcaratus (FB), P. gutturosus (S-F), P. alleni (F), P. batesi (F, We thank A. Schiøtz for various comments on determination to be verified), Afrixalus ni- the collected treefrogs, W.O. Odiete for his finan-

9 Abiodun B. Onadeko & Mark-Oliver Rödel cial support, the staff of the Department of Zool- Boulenger, G.A. (93): On the presence of two ogy, University of Lagos for their moral backup, closely allied species of toads of the genus Nec- and J. Penner for compiling figure . tophryne in Cameroon. – Annals and Magazi- ne of natural History, 8é sér., 2: VIII. Eniang, E.A. & L. Luiselli (2002): Ikpan wetland References rainforest: an area of high biodiversity impor- tance in south-eastern Nigeria. – Revue Écolo- Aisien, S.O., F. Ayeni & I. Ilechie (2004): Hel- gie (Terre Vie), 57: 9-28. minth fauna of anurans from the Guinea savanna at New Bussa, Nigeria. – African Zoo- Ernst, R., K.E. Linsenmair & M.-O. Rödel logy, 39: 33-36. (2006): Diversity erosion beyond the species level: Dramatic loss of functional diversity af- Aisien, S.O., A.D. Ugbo, A.N. Ilavbare & O. ter selective logging in two tropical amphibi- Ogunbor (200): Endoparasites of amphibian communities. – Biological Conservation, ans from South-Western Nigeria. – Acta Para- 33:43-55. sitologica, 46: 299-306. Ernst, R. & M.-O. Rödel (2005): Anthropogeni- Akani, G.C., E. Politano & L. Luiselli (2004): cally induced changes of predictability in tro- Amphibians recorded in forest swamp areas of pical anuran assemblages. – Ecology, 86: 3– the River Niger Delta (southeastern Nigeria), 38. and the effects of habitat alteration from oil in- Frètey, T. & C.P. Blanc (2000): Liste des am- dustry development on species richness and phibiens d’Afrique centrale. Cameroun, Con- diversity. – Applied Herpetology, 2: -22. go, Gabon, Guinée-Équatoriale, République Akintola, A.O. (976): Observations on the com- Centrafricaine, République Démocratique du mon African Toad, Bufo regularis at Shagamu Congo, São Tomé et Príncipe. – Les dossiers de Field Station, Nigeria. – Nigerian Field, 4: 23- l’ADIE, Série Biodiversité, N°2: -39. 28. Frost, D.R. (2007): Amphibian species of the Amiet, J.-L. (986): La batrachofaune sylvicole world: an online reference. – Version 5.0. http:// d’un secteur forestier du Cameroun: la région research.amnh.org/herpetology/amphibia/in- de Yaoundé. – Mémoires du Muséum National dex.html [last inquiry date: 3 May 2007]. d’Histoire Naturelle, Série A, 32: 29-42. Frost, D.R., T. Grant, J. Faivovich, R.H. Bain, Assemian, N.E., N.G. Kouamé, B. Tohé, G. A. Haas, C.F.B. Haddad, R.O. De Sá, A. Gourène & M.-O. Rödel (2006): The anur- Channing, M. Wilkinson, S.C. Donnellan, ans of the Banco National Park, Côte d’Ivoire, C.J. Raxworthy, J.A. Campbell, B.L. Blot- a threatened West African rainforest. – Sala- to, P. Moler, R.C. Drewes, R.A. Nussbaum, mandra, 42: 4-5. J.D. Lynch, D.M. Green & W.C. Wheeler Bakarr, M., J.F. Oates, J. Fahr, M. Parren, M.- (2006): The amphibian tree of life. – Bulletin O. Rödel & R. Demey (2004): Guinean forests of the American Museum of Natural History, of West Africa. – pp. 23-30 in: Mittermeier, 297: -370. R.A., P.R. Gil, M. Hoffmann, J. Pilgrim, T. Grafe, T.U. (2000): Leptopelis viridis (West Afri- Brooks, C.G. Mittermeier, J. Lamoreux & can Tree Frog). Cocoon formation. – Herpeto- G.A.B. da Fonseca (eds.): Hotspots revisited: logical Review, 3: 00-0. Earth’s biologically richest and most endange- Guibé, J. & M. Lamotte (963): La réserve natu- red terrestrial ecoregions. – Washington, D.C. relle intégrale du Mont Nimba. XXVIII. Batra- (Conservation International & CEMEX). ciens du genre Phrynobatrachus. – Mémoires Barbault, R. (972): Les peuplements d’amphibiens de l’Institut fondamental d’Afrique noire, 66: des savanes de Lamto (Côte d’Ivoire). – Anna- 60-627. les de l’Université d’Abidjan, Série E, 5: 59–42. Hillers, A. & M.-O. Rödel (2007): The amphi- Böhme, W. & G. Nikolaus (989): Herpetologi- bians of three national forests in Liberia, West cal specimens from the Gotel Mountains and Africa. – Salamandra, 43: -0. Mambilla Plateau, Nigeria. – Tauraco Research Hughes, B. (979): Feeding habits of the frog Report, : 28–30. Aubria subsigillata in Ghana. – Bulletin de

10 Anuran surveys in south-western Nigeria

l’Institut fondamental Afrique noire, Série A, occipitalis (Günther) au Sénégal. – Bulletin de 4: 654-663. l’Institut fondamental d’Afrique noire, Série A, Inger, R. & H. Marx (96): The food of amphi- 33: 446–466. bians. – pp. -86 in: Mission G.F. de Witte: Nago, S.G.A., O. Grell, B. Sinsin & M.-O. Rö- Exploration du Parc National de l’Upemba, In- del (2006): The amphibian fauna of the Pend- stitut des parcs nationaux du Congo et du Ru- jari National Park and surroundings, northern anda-Urundi, 64. Benin. – Salamandra, 42: 93-08. IUCN, Conservation International & Natu- Perret, J.-L. (966): Les amphibiens du Came- reServe (2006): Global Amphibian Assess- roun. – Zoologische Jahrbücher (Systematik), ment. – www.globalamphibians.org [last in- 8: 289–464. quiry date: 3 May 2007]. Perret, J.-L. (986): Considérations sur le genre Knoepffler, L.-P. (976): Food habitats of Aubria Phlyctimantis Laurent et Combaz (Anura, subsigillata in Gabon. – Zoologica Africana, : Hyperoliidae). – Muséum d’Histoire Naturelle 369–37. Genève, 09: 9–26. Lamotte, M. (967): Les batraciens de la régi- Perret, J.-L. (988): Les espèces de Phrynobat- on de Gpakobo (Côte d’Ivoire). – Bulletin de rachus (Anura, Ranidae) à éperon palpébral. l’Institut fondamental d’Afrique noire, Série A, – Archives des Sciences, 4: 275–294. 29: 28-294. Reid, J.C. (987): Notes on the distribution of an- Lamotte, M. & F. Xavier (966): Étude comparée urans among biotopes of the Calabar area of de deux espèces de Phrynobatrachus souvent southeastern Nigeria. – pp. 325-330 in: Van confondues: Phr. plicatus Günther et Phr. au- Gelder, J.J., H. Strijbosch & P.J.M. Bergers ritus Boulenger. – Bulletin de l’Institut fonda- (eds.): Proceedings of the 4th Ordinary General mental d’Afrique noire, Série A, 28: 605–69. Meeting of the Societas Europaea Herpetologi- Laurent, R.F. (972): Tentative revision of the ge- ca, Nijmegen. nus Hemisus Günther. – Musée Royal d’Afrique Reid, J. C., A. Owens & R. Laney (990): Records Centrale, Science Zoologique, 94: –67. of frogs and toads from Akwa Ibom State. – Ni- Lea, J.M., E. Politano & L. Luiselli (2003): gerian Field, 55: 3-28. Changes in the herpetofauna of a fresh water river in southern Nigeria, after 20 years of de- Rödel, M.-O. (2000): Herpetofauna of West Afri- velopment. – Russian Journal of Herpetology, ca. Vol. I. Amphibians of the West African Sa- 0: 9-98. vanna. – Frankfurt/M. (Edition Chimaira). Lea, J.M, L. Luiselli & E. Politano (2005): Are Rödel, M.-O. (2003): The amphibians of Mont there shifts in amphibian faunal composition Sangbé National Park, Ivory Coast. – Sala- in Nigerian landscapes undergoing long-term mandra, 39: 9-0. degradation? A case study from a montane en- Rödel, M.-O. (2007): The identity of Hylambates vironment. – Revue Écologie (Terre Vie), 60: hyloides Boulenger, 906 and description of a 65-76. new small species of Leptopelis from West Afri- Leaché, A.D., M.-O. Rödel, C.W. Linkem, R.E. ca. – Mitteilungen aus dem Museum für Natur- Diaz, A. Hillers & M.K. Fujita (2006): Bio- kunde Berlin, Zoologische Reihe, 83: 90-00. diversity in a forest island: reptiles and amphi- Rödel, M.-O. & M.A. Bangoura (2004): A con- bians of the West African Togo Hills. – Amphi- servation assessment of amphibians in the bian and Reptile Conservation, 4: 22-45. Forêt Classée du Pic de Fon, Simandou Ran- LeBreton, M. (999): A working checklist of the ge, southeastern Republic of Guinea, with the herpetofauna of Cameroon, with localities for description of a new Amnirana species (Am- species occurring in southern Cameroon and phibia Anura Ranidae). – Tropical Zoology, 7: a list of herpetofauna for the Dja Faunal Reser- 20-232. ve. – unpublished report, IUCN Netherlands, Rödel, M.-O., M.A. Bangoura & W. Böhme 39 pp. (2004): The amphibians of south-eastern Re- Lescure, J. (97): L’alimentation du crapaud Bufo public of Guinea (Amphibia: Gymnophiona, regularis Reuss et de la grenouille Dicroglossus Anura). – Herpetozoa, 7: 99-8.

11 Abiodun B. Onadeko & Mark-Oliver Rödel

Rödel, M.-O. & W.R. Branch (2002): Herpetolo- Schiøtz, A. (966): On a collection of amphibia gical survey of the Haute Dodo and Cavally fo- from Nigeria. – Videnskabelige Meddelelser rests, Western Ivory Coast, Part I: Amphibians. fra Dansk Naturhistorisk Forening, 29: 43–48 – Salamandra, 38: 245-268. + 3 plates. Rödel, M.-O. & R. Ernst (2002): A new Phryno- Schiøtz, A. (967): The treefrogs (Rhacophori- batrachus from the Upper Guinean rain forest, dae) of West Africa. – Spolia zoologica Musei West Africa, including a description of a new Haunienses, 25: –346. reproductive mode for the genus. – Journal of Schiøtz, A. (999): Treefrogs of Africa. – Frank- Herpetology, 36: 56-57. furt/M. (Edition Chimaira),. 350 pp. Rödel, M.-O. & R. Ernst (2003): The amphibians Scheel, J.J. (970): Notes on the biology of the Af- of Marahoué and Mont Péko National Parks, rican tree-toad, Nectophryne afra Buchholz & Ivory Coast. – Herpetozoa, 6: 23-39. Peters, 875, (Bufonidae, Anura) from Fern- Rödel, M.-O., M. Gil, A.C. Agyei, A.D. Leaché, ando Poo. – Revue Zoologique Botanique Af- R.E. Diaz, M.K. Fujita & R. Ernst (2005a): ricaine, 8: 225-236. The amphibians of the forested parts of south- Segniagbeto, G.H., J.E. Bowessidjaou, A. Du- western Ghana. – Salamandra, 4: 07-27. bois & A. Ohler (2007): Les Amphibiens du Rödel, M.-O., J. Kosuch, N.G. Kouamé, R. Ernst Togo: état actuel des connaissances. – Alytes, & M. Veith (2005b): Phrynobatrachus alticola 24: 72-90. Guibé & Lamotte, 96 is a junior synonym UICN (996): L’atlas pour la conservation des of Phrynobatrachus tokba (Chabanaud, 92). forêts tropicales d’Afrique. – Paris (Editions – African Journal of Herpetology, 54: 93-98. Jean-Pierre de Monza). Rödel, M.-O., D. Krätz & R. Ernst (2002): The Walker, R.B. (966): An uncommon frog, Pyxice- tadpole of Ptychadena aequiplicata (Werner, phalus adspersus. – Nigerian Field, 3: 77-80. 898) with the description of a new reproductive mode for the genus (Amphibia, Anura, Ra- Walker, R.B. (967): An elusive frog, Rana gala- nidae). – Alytes, 20: -2. mensis. – Nigerian Field, 32: 22-26. Romer, J.D. (953): Reptiles and amphibians coll- Walker, R.B. (968): The amphibians of Zaria, in ected in the Port Harcourt area of Nigeria. – Co- the northern Guinea Savannah, Nigeria. – Co- peia, 953: 2–23. peia, 968: 64–67. Sanderson, I.T. (936): The giant frog problem. Walker, R.B. (969): The burrowing frog: Hemi- – Nigerian Field, 5: 6-70. sus marmoratus (Peters). – Nigerian Field, 34: 72-76. Schiøtz, A. (963): The amphibians of Nigeria. – Videnskabelige Meddelelser fra Dansk Natur- historisk Forening, 25: –92 + 4 plates.

12 Anuran surveys in south-western Nigeria

Appendix 1

Geographic position (see Fig. 1) and short habitat characterisation of collection sites in Lagos, Ogun and Oyo States, Nigeria. F = forest, FB = farmbush, S = savanna. code latitude (N) / site description longitude (E) Lagos state  06°30’064’’ / Badore: swamp with tertiary vegetations and a slow flowing stream, few palm 003°36’276’’ trees, intensive construction work (FB) 2 06°28’74’’ / Badore: dirt road with puddles of various sizes, swamp with dense herb vegeta- 003°24’26’’ tion and shrubs (FB) 3 06°3’074’’ / University of Lagos: pool and fallen tree, few metres from the Lagos Lagoon (FB) 003°24’26’’ 4 06°30’822’’ / University of Lagos: swamp with secondary vegetation, small stream (FB) 003°23’688’’ 5 06°27’736’’ / Lagos-Epe road: small pond, ferns dominating in swamp, many Raphia palms 003°42’347’’ (FB) 6 06°34’996’’ / Epe: close to bridge of Lagos Lagoon, large herbs and grasses, almost permanent 003°56’968’’ swamp forest with Raphia palms (FB) 7 06°39’747’’ / Itokin: about 8 km from Epe, fast flowing stream, banks with sparse vegetation, 003°48’605’’ bordered by secondary forest with sparse under-storey and leaf-litter (F) 8 06°34’072’’ / Ijede: large swamp with slow flowing creek, dense submerse and bank vegetation, 003°35’030’’ overshadowed with bamboo, banana plantation (FB) 9 06°36’77’’ / Ijede: large water-hole without vegetation, supplied by small stream, secondary 003°34’37’’ forest partly degraded, edge of a sugar-cane plantation (F) 0 no GPS data Badagry: large pond in densely vegetated swamp, bank with large herbs; bordered by secondary and degraded forest (FB)  no GPS data Ikorodu: very large densely vegetated pond, Pistia sp., bordered by tall grasses and shrubs; swamp with dense vegetation (FB) Ogun state  06°43’465’’ / Abowa settlement near Ogijo: newly cultivated land bordered by a partly degrad- 003°34’37’’ ed tertiary forest, sloping towards a small stream ( km) (FB) 2 06°44’002’’ / Abowa settlement near Ogijo: small moderately flowing stream without vegeta- 003°29’464’’ tion, banks densely vegetated, in a secondary forest, sparse under-story and nearly closed canopy, fallow 50 m away (F) 3 06°44’403’’ / Erefun settlement near Ogijo: two medium sized temporary pools, sandy soil, 003°29’494’’ dirt road through degraded tertiary forest, Cola acuminata forming canopy, small stream nearby (F) 4 06°46’505’’ / Large and deep pond off the Lagos-Ibadan expressway, water lilies, banks densely 003°25’449’’ vegetated, few banana trees at the perimeter, water filled ditches (FB) 5 06°5’84’’ / large flowing stream, 2 km east of Sagamu, water lilies in stagnant parts, banks 003°38’793’’ densely vegetated, bordered by cultivated land (FB) 6 06°52’059’’ / Ikenne: rubber plantation with low vegetation surrounding a large swamp with 003°4’328’’ densely and sparsely vegetated parts; temporary water bodies present in swamp (F) 7 07°00’27’’ / Ishara: fallow close to a sawmill, sloping towards a small stream; bank with ba- 003°4’55’’ nana trees and herbs (FB)

13 Abiodun B. Onadeko & Mark-Oliver Rödel

code latitude (N) / site description longitude (E) 8 06°45’782’’ / Idowa: tertiary vegetation, mostly low shrubs along the Ikorodu-Ijebu Ode road, 003°5’478’’ bordered by secondary forest (F) 9 06°56’206’’ / about 2 km from Ogun State University along Ijebu Ode road, tertiary vegetation; 003°56’47’’ large mango trees forming canopy, sparse under-story, Ogun river about  km to the east; muddy area with water filled tracks (FB) 0 06°49’487’’ / along Shagamu-Benin expressway: river bordered by secondary / primary forests 003°49’207’’ and farmland (F)  06°56’493’’ / Ijebu Oru: Liberian Refugee Camp Settlement: large swamp with dense vegeta- 003°56’792’’ tion, almost permanent, two large shallow ponds, one being very eutrophic, banana trees, about 50 m from fast flowing with rocky River Oru, river bordered by farmland and tertiary growth (FB) 2 07°05’723’’ / along Lagos-Ibadan expressway: river and cultivated island, partly. dense vegeta- 003°43’796’’ tion and low shrubs, bordered by secondary forest (F) Oyo state  07°36’222’’ / large shallow pond, almost entirely covered by Pistia sp. about 3 km north of 003°5’03’’ Ibadan, bordered by herbs and shrubs, few palm trees (FB) 2 07°37’33’’ / Onidundu: derived savanna, large rice farm in a muddy swamp, bordered by sec- 003°55’258’’ ondary forest and dense herbs and grasses; large stream with overhanging trees and shrubs, Pistia sp., bank with herbs (F) 3 07°43’738’’ / Fiditi: cultivated land bordered by secondary forest sloping towards a valley; small 003°54’939’’ dried up pond overgrown by weeds (F) 4 07°45’90’’ / open degraded savanna about 2 km from Fiditi, bordered by secondary forest (S) 003°55’34’’ 5 07°5’338’’ / large and deep pond in open savanna at the outskirt of Oyo Town, large area with- 003°57’723’’ out vegetation in pond, water lilies, small stream feeding the pond (S) 6 08°0’385’’ / Oolo: small swamp in the savanna region, with a deep water hole present, partly 004°07’435’’ shaded by shrubs and other vegetations (S) 7 08°05’29’’ / maize and millet fields, small gallery forest close to Ibapon Oyewole (S) 004°’32’’ 8 08°08’46’’ / immediate surroundings of Ogomosho, swampy area with puddles, highly pol- 004°5’082’’ luted with mainly domestic wastes (S) 9 08°2’56’’ / very large swamp, partly cultivated, a few kilometres from Ogbomosho, large 004°7’050’’ shallow pond, a partly dried up pond, both surrounded by sparse vegetation, pre- viously cultivated (S) 0 08°7’700’’ / Gbede/Gambari: large pond in a swamp, large area without vegetation, open sa- 004°20’58’’ vanna, small puddles in dried up stream (S)  08°8’778’’ / Otte-Egba: northernmost collection site, savanna, about 0 km from Ilorin, dry- 004°23’209’’ ing up pond with sparse vegetation, few shrubs, large fire-resistant trees (S)

Manuscript received: 12 June 2007 Authors’ addresses: Abiodun B. Onadeko, Department of Zoology, Faculty of Sciences, University of Lagos, Akoka, Yaba, Lagos, Nigeria, E-Mail: [email protected]; Mark-Oliver Rödel, Hum- boldt University Berlin, Museum of Natural History, Herpetology, Invalidenstraße 43, D-10115 Berlin, Germany, E-Mail: [email protected].