92 research outputs found
No ecological opportunity signal on a continental scale?:Diversification and life-history evolution of african true toads (Anura: Bufonidae)
The niche-filling process predicted by the “ecological opportunity” (EO) model is an often-invoked mechanism for generating exceptional diversity in island colonizers. Whether the same process governs lineage accumulation and trait disparity during continental colonization events is less clear. Here, we test this prediction by investigating the rate dynamics and trait evolution of one of Africa's most widespread amphibian colonizers, the true toads (Bufonidae). By reconstructing the most complete molecular phylogeny of African Bufonidae to date, we find that the diversification of lineages in Africa best conforms to a constant rate model throughout time and across subclades, with little support for EO. Evolutionary rates of life-history traits have similarly been constant over time. However, an analysis of generalists and specialists showed a shift toward higher speciation rates associated with habitat specialization. The overall lack of EO signal can be interpreted in a number of ways and we propose several explanations. Firstly, methodological issues might preclude the detection of EO. Secondly, colonizers might not experience true EO conditions and due to the size, ecological heterogeneity and age of landmasses, the diversification processes might be more complex. Thirdly, lower speciation rates of habitat generalists may have affected overall proliferation of lineages
Phrynobatrachus rainerguentheri Rödel, Onadeko, Barej & Sandberger, 2012, sp. nov.
<i>Phrynobatrachus rainerguentheri</i> sp. nov. <p>Figs 1–2</p> <p> <b>Holotype.</b> ZMB 77435, adult female, Nigeria, about 2 km from Ogun State University along Ijebu Ode ─ Ibadan road, Ogun river about 1 km to the East, close to the town of Ijebu Oru, N 06.93906°, E 003.94492°, <100 m a.s.l., 16 July 2004, coll. A.B. Onadeko.</p> <p> <b>Paratypes.</b> ZMB 77436-77437, juveniles, 29 October 2011, type locality, similar location after 6-7 years of vegetation regrowth, other data as holotype; ZMB 77737-77744, all juveniles, Nigeria, Warri (Opara town), near the Niger Delta region, swampy area, N 05.51667°, E 005.73333°, 16 February 2012, coll. A.B. Onadeko.</p> <p> <b>Additional material.</b> DZOOL 211, female, SVL 20.1 mm, 28 September 2003; DZOOL 384, female, 18.4 mm, 14 July 2004; all other data as holotype; DZOOL 544, juvenile, 10.1 mm, Nigeria, town of Warri, near the Niger Delta region, N 05.51667°, E 005.73333°, 18 November 2011, coll. A.B. Onadeko; DZOOL 545, female, 17.6 mm, 19 November 2011, other data as DZOOL 544.</p> <p> <b>Diagnosis.</b> The genetic results show that the new species belongs to the genus <i>Phrynobatrachus.</i> Morphologically this is visible by the body shape; presence of a tarsal tubercle; presence of webbing; lack of a black spot in the tympanal region (as in <i>Arthroleptis</i>); lack of a median dorsal skin raphe (as in <i>Arthroleptis</i>); lack of parallel dorsal ridges (as in <i>Ptychadena</i>); neither finger nor toe tips heart shaped (as in <i>Petropedetes</i>) or expanded to discs.</p> <p>The new species is characterised by a combination of small size; compact body shape; very short and round snout; large eyes, absence of an eyelid cornicle; presence of a conspicuous row of warts with pores on the upper mandible, larger warts on forehead and inter-orbital space; an almost smooth back, a pair of elongated chevron like ridges in the scapular region; an unmarked belly; the absence of discs on finger and toe tips and well developed webbing.</p> <p> <b>Description of holotype</b> (measures in mm). Typical small <i>Phrynobatrachus,</i> female with oval, compact body shape; snout-vent length: 18.8; short snout, rounded in dorsal and lateral view; <i>canthus rostralis</i> indistinct and rounded; loreal region straight to slightly concave; head-width directly behind the eyes: 6.5; large protruding eyes; eye-diameter: 2.6; eyelids, forehead and inter-orbital space with few larger warts; eyelid cornicle absent; distance eye-nostril: 1.8; nares small, round, positioned dorsolaterally; distance nostril-snout tip: 1.3; nares closer to snout than to eye; inter-narial distance: 1.5; inter-orbital distance: 1.5; short supratympanal ridge; tympanum diameter: 1.4, indistinct, much smaller than diameter of eye; distinct row of warts with pores on upper lip; outline of lower mandible mucronate; upper maxilla with minute teeth, hidden by lips; vomerine teeth absent; broad cordiform tongue, tip slightly but broadly notched, median papilla near anterior attachment of tongue; back slightly granular; scapular region almost smooth, without distinct warts or ridges; ventral skin smooth; forelimbs slender; upper arm: 4.9; lower arm to tip of finger III: 8.2; hand with two large, outer round and inner oval carpal tubercles; fingers with small roundish subarticular tubercles, no additional tubercles on hands; relative finger length: IV≤I=II<III; right hand normal, left hand deformed with shortened digit I and II; rudimentary web on bases between fingers; finger tips not broadened; right hand normal; femur: 8.9, shorter than tibia: 10.3; foot including longest toe: 14.9; length of toe I: 2.6; tarsal tubercle present but not conspicuous; larger internal metatarsal tubercle (0.9), narrow and oval; app. 1/3 of length of toe I; external metatarsal tubercle not visible; relative toe length: I<II<V<III<IV; webbing formula I(0), II(0.5-0), III(1), IV(2), V(0.75); toe tips slightly enlarged without forming discs, just slightly exceeding width of subarticular tubercles.</p> <p>Large eggs (largest egg app. 1 mm in diameter), rich in yolk, shining through skin of belly: 17 eggs visible; total egg number estimated to be app. 30, certainly not exceeding 50.</p> <p> <i>Colouration in life</i>: black head and eyelids; black colour extending posteriorly just behind eye and to angle of mouth; tympanum coloured as back; back uniform orange-red; flanks slightly lighter than back; upper arms uniform brown; one dark cross bar on lower arms; fingers dorsally dark cross-bared; vent with black square, enclosed by light band; anterior part of thighs with two (right) and three (left) black cross bars, only one very distinct; posterior half of shanks uniform brown (right) and slightly mottled (left); shanks with three dark cross bars, only one very distinct; foot and toes dark cross-bared; throat, pectoral region and lower parts of flanks with minute brown spots; belly uniform light, almost translucent; ventral parts of feet brownish.</p> <p> <i>Colouration in formalin and ethanol after more than eight years</i>: pattern slightly fainted, black head turned dark brown, reddish back turned to light brown.</p> <p> <b>Variation</b> (measures summarised in Table 1). Snout-vent length in adult females from the type locality were 17.6-20.1 mm (adult males unknown), otherwise identical to holotype. The juvenile paratypes from the type locality are likewise very similar to holotype (Fig. 2); snout-vent length 11.0 mm (ZMB 77436) and 9.6 mm (ZMB 77437, Fig. 2 i); snout in dorsal view round (ZMB 77436) or almost subelliptical (ZMB 77437); distinct larger warts between eyes (ZMB 77437) or this area smooth (ZMB 77436); row of warts on upper lip absent; shoulder with indicated chevron like ridges starting at posterior end of upper eyelid and extending app. to level of forearm bases (ZMB 77437) or smooth (ZMB 77436); back and flanks almost smooth as in holotype (ZMB 77436) or with small flat warts (ZMB 77437); basic coloration, in particular black head and red back as in holotype; lower mandible black; ventral border of flanks with black line (ZMB 77437) or spots (almost fusing, ZMB 77436); basic colour of extremities grey; posterior half of dorsal shanks uniform grey; vent with black square, surrounded by white band; lower arm, thighs and shanks with up to three black cross bars; belly white; sole of feet black; webbing less developed than in holotype or partly destroyed (dried up), the latter being more likely.</p> <p>In contrast to the almost uniform specimens from the type locality, individuals from the second locality (all juveniles), near the town of Warri, exhibited considerable colour and some morphological variation. Only one specimen (ZMB 77741, Fig. 2 j) mirrored exactly the colour pattern and morphology of frogs from the type locality. All others have a distinct pair of elongated chevron like ridges in the scapular region (Fig. 2). The enlarged warts on snout and the interorbital region are sometimes indistinct or missing. In contrast the flanks are warty and the back skin ranges from smooth to warty. The different colour morphs are illustrated in Fig. 2. All specimens from both localities share a white or almost white belly, a white throat with small brown to black dots. The lower mandible is most often dark cross-barred. Some exhibit some large dark spots in the pectoral region and the lateral parts of the belly. Thighs and shanks may carry one or several dark cross bars. However, only one of these bars is particularly broad and distinct on thighs and shanks. In a sitting position the bars on upper and lower leg form one continuous band (Fig. 2).</p> <p> <b>Genetics.</b> <i>Phrynobatrachus rainerguentheri</i> <b>sp. nov.</b> differs between 6.59-22.94% in the investigated part of the 16S rRNA gene from 33 West African and Central African species of the genus <i>Phrynobatrachus</i> (Table 2). The species with the lowest genetic divergence to <i>P. rainerguentheri</i> <b>sp. nov.</b> were <i>P. ghanensis</i> (mean difference 7.06%), <i>P. phyllophilus</i> (7.06%), <i>P. guineensis</i> (7.30%) and <i>P. al le n i</i> (8.55%). The intraspecific genetic variation of nine <i>P. rainerguentheri</i> <b>sp. nov</b>., one from the type locality (ZMB 77437, GenBank JQ730857) and eight from Warri (ZMB 77737-77744, GenBank JQ954860 - JQ954867) was 0-0.27%.</p> <p> <b>Habitat.</b> Five of the 15 known individuals have been collected at the type locality during and towards the end of the rainy season in tertiary farmbush vegetation where large mango trees were forming a “forest” canopy. The understorey was sparsely vegetated and the area was muddy with water filled tracks. There the species seemed to be rare and hard to detect. First surveys, covering app. 400 m ² at the type locality, ranged from June 2003 to December 2004. During that period only three specimens were observed and collected. Although repeatedly revisiting the site, we could not observe the species until September/ November 2011. In late 2007 about 60% of the area was clear-cut during shifting agricultural practices (Fig. 3 a). By the end of 2008 vegetation started to recover and in September 2011 two specimens could be observed and collected. At the type locality all specimens were collected in drier places far away from open water.</p> <p>Species GenBank # voucher p-distance</p> <p> min-max mean ±sd Other anuran species at the type locality where: <i>Silurana tropicalis, Amietophrynus maculatus, A. regularis, Aubria subsigillata, Ptychadena</i> cf. <i>aequiplicata, Phrynobatrachus latifrons, P. francisci, Arthroleptis</i> sp., <i>Leptopelis</i> sp., <i>Afrixalus dorsalis, Hyperolius concolor</i> and <i>H. fusciventris</i> (Onadeko & Rödel 2009). This species composition indicates that the area originally consisted of rainforest. However, the many invasive species, i.e. the two toads and the two other <i>Phrynobatrachus</i> spp., are clearly indicating a heavily altered state of this forest habitat.</p> <p>Most recently (18 & 19 November 2011, 16 February 2012) several additional specimens were recorded near the town of Warri, close to the Niger Delta region (Fig. 3 b-e). Here the frogs, all juveniles, occurred in a dried up swamp in the leaf litter close to a drying pond (about 9 m in diameter) with some water remaining. During the rainy season this entire area is covered with water.</p> <p> <b>Distribution.</b> <i>Phrynobatrachus rainerguentheri</i> <b>sp. nov.</b> is known from the type locality and one further locality app. 250 km south-east from the type locality (Fig. 4).</p> <p> <b>Etymology.</b> The species is dedicated to Dr. sc. Rainer Günther, from 1985-2006 curator of herpetology at the Museum für Naturkunde Berlin, in recognition for his outstanding contributions to herpetology.</p>Published as part of <i>Rödel, Mark-Oliver, Onadeko, Abiodun B., Barej, Michael F. & Sandberger, Laura, 2012, A new polymorphic Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae) from western Nigeria, pp. 55-65 in Zootaxa 3328</i> on pages 56-62, DOI: <a href="http://zenodo.org/record/281218">10.5281/zenodo.281218</a>
Multiple genetic lineages challenge the monospecific status of the West African endemic frog family Odontobatrachidae
BACKGROUND: Correct species identification is crucial in different fields of biology, and in conservation. The endemic West African frog family Odontobatrachidae currently contains a single described species, Odontobatrachus natator. From western Guinea to western Côte d'Ivoire it inhabits forests around waterfalls or cascades. Based on more than 130 specimens from 78 localities, covering the entire distribution, we investigated the molecular diversity of these frogs. - RESULTS: Our analyses included mitochondrial and nuclear genes, with a concatenated alignment of 3527 base pairs. We detected high level of genetic differentiation with five distinct lineages or operational taxonomic units (OTUs). These OTUs were also identified by two different species delimitation approaches, Generalized Mixed Yule Coalescent (GMYC) and cluster algorithm. All OTUs occur in parapatry in the Upper Guinean forests. One OTU, assigned to the “true” Odontobatrachus natator, covers the largest distribution, ranging from the border region of western Sierra Leone-Guinea to eastern Liberia. Two OTUs are restricted to western Guinea (Fouta Djallon and foothills), while two others occur in eastern Guinea and the border region of Guinea-Liberia-Côte d'Ivoire. The OTU representing O. natator consists of two divergent subclades: one restricted to the Freetown Peninsula (Sierra Leone) and the other covering all populations further inland. Environmental niche models indicated that the restricted Freetown Peninsula population is separated by unsuitable habitat from remaining populations. - CONCLUSION: Geographic isolation of OTUs and molecular differences comparable to species level differentiation in other frog families indicate that O. natator contains cryptic species diversity. Respective distribution patterns most probably resulted from repeated changes of forest cover (contraction and expansion) over evolutionary timescales. The survival within forest refugia that have persisted through multiple drier periods and subsequent dispersal during wetter times may best explain the observed geographic distributions of OTUs. According to the IUCN Red List range criteria each OTU should be classified as “Endangered.” If the Freetown Peninsula “natator” population is recognized as a distinct species it would warrant recognition as “Critically Endangered.” The identification of cryptic lineages highlights the urgent need to protect these frogs, all of which are endemic to small areas within the Upper Guinean biodiversity hotspot
Description of two Werneria tadpoles from Cameroon (Amphibia: Anura: Bufonidae)
Hirschfeld, Mareike, Barej, Michael F., Loader, Simon P., Rödel, Mark-Oliver (2012): Description of two Werneria tadpoles from Cameroon (Amphibia: Anura: Bufonidae). Zootaxa 3172 (1): 65-68, DOI: 10.11646/zootaxa.3172.1.5, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.3172.1.
The hairy frog, a curly fighter ? – A novel hypothesis on the function of hairs and claw-like terminal phalanges, including their biological and systematic significance (Anura : Arthroleptidae : Trichobatrachus)
Volume: 117Start Page: 243End Page: 26
A new polymorphic Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae) from western Nigeria
Rödel, Mark-Oliver, Onadeko, Abiodun B., Barej, Michael F., Sandberger, Laura (2012): A new polymorphic Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae) from western Nigeria. Zootaxa 3328: 55-65, DOI: 10.5281/zenodo.28121
Petropedetes johnstoni
Petropedetes johnstoni (Boulenger, 1888 “ 1887 ”) Figs. 1 a, 2 a, 3 a, 4, 5, 7g Cornufer johnstoni Boulenger, 1888 “ 1887 ”, Proc. Zool. Soc. London 1887: 564 Tympanoceros newtonii du Bocage, 1895, J. Sci. Math. Phys. Nat. Lisboa, Ser.2, 3: 270 Petropedetes johnstoni – Boulenger, 1900. Proc. Zool. Soc. London, 1900: 439 Material examined. BM 1947.2. 7.44 (holotype, described as Cornufer johnstoni), subadult male, Cameroon (“ Cameroons ”), Rio del Rey, coll. H.H. Johnston; MSNG 29890 (5 males, 2 females), Equatorial Guinea, Bioko, Musola, app. 500-800 m a.s.l., I-III. 1902, coll. L. Fea; MSNG 50032 (female and tadpole), app. 500- 700 m a.s.l., other data as MSNG 29890; ZFMK 87709 (male), Cameroon, Campo region, Nkoelon, app. 125 m a.s.l., 30.X. 2007, coll. J.A.M. Wurstner & M.F. Barej; ZFMK 87710 (female), 26.X. 2007, other data as for ZFMK 87709. Diagnosis. A medium sized Petropedetes; slender body shape; tympanum round, distinctly smaller than eye diameter (smallest distinct tympanum in the genus); characters of breeding males: tympanic papilla present (papilla slim, compared to other larger Petropedetes), papilla close to the upper border of tympanum; forearm hypertrophy weakly developed; carpal spike present; spinosities on upper arm, throat and breast; few minuscule spines on supratympanal gland; femoral glands brightly orange, prominent and very large; webbing rudimentary. Description. Medium sized Petropedetes with slender body shape; both sexes almost equally large (SUL in males: 28.5–45.6 mm, females: 28.1–47.8 mm); mean head width in males about 41 % of SUL, in females about 38 %; snout rounded to slightly pointed in lateral view; canthus rostralis sharp; loreal region concave; eye diameter about 1.5 times eye-narial distance; nose closer to snout-tip than to eye; distinct round tympanum smaller than eye in both sexes (tympanum / eye in males: 0.49–0.63; in females: 0.45–0.48; Fig. 1 a); tympanic papilla of breeding males close to upper border of tympanum, comparatively slim; supratympanic fold distinct; fingers slender with T-shaped fingertips; relative length of fingers: III> IV> II> I; manual subarticular tubercles single; palmar tubercle and thenar tubercle present; manual webbing absent; forearm hypertrophy weakly developed; carpal spikes present in males; spinosities on upper arm, throat and breast; few minuscule spines on supratympanal gland; back skin granular, beset with small warts; fewer larger warts arranged in more or less longitudinal rows; flanks with few large warts; ventral skin smooth; mean femur length in males 53 % of SUL, in females about 51 %, mean tibia length in both sexes 61 % of SUL; mean foot length in males 79 % of SUL, in females 78 %; pedal subarticular tubercles single; hind limbs slender; femoral glands very large in males, larger and more prominent in males than in females (femoral gland / femur length in males: 0.28–0.46, in females: 0.25–0.27; Fig. 2 a); relative length of toes: IV> III> V> II> I; webbing rudimentary: 1 (1) 2 (1 - 1) 3 (2 - 2) 4 (3 - 3) 5 (2). According to Amiet (1973) the tympanic papilla is shorter than the tympanum diameter and thickened at its tip. This character is hard to verify in preserved specimens, as the papilla is then often shrunken (M.F. Barej pers. obs.). Amiet (1983) remarked that skin on throat and basis of the arms is smooth, showing no trace of spinosities. This observation is not consistent with the MSNG material. The most detailed description referring to “ P. johnstoni ” has been published by Parker (1936). However, this was based on specimens which have later been recognized as a new species, P. parkeri (Amiet 1983). Coloration. Dorsum brownish with small dark spots, marking larger warts; colour on flanks as on back, often with larger spots than on back (Fig. 1 a); iris speckled with gold and silver; throat marbled blackish; belly whitish with minuscule dark speckles (recognizable only under the microscope); upper and lower legs dorsally with large almost bar like dark spots; femoral glands brightly orange in males. Coloration in preservation: as in life, but colours pale. Boulenger (1888) mentioned a dark inter-orbital bar. In the material examined this bar was often incomplete. Tadpole description. The MSNG collection comprises one tadpole (MSNG 50032) collected by L. Fea in March 1902 at Musola (500–700 m) on Bioko. The tadpole (Gosner stage 41) has already fully developed hind limbs with rudimentary webbing. The vent tube is regressed. It had a body length of 11.3 mm (total length 26.3 mm) and long hind legs (18.6 mm), its femoral glands were already 1.8 mm long. As from Bioko, no other Petropedetes species with rudimentary webbing is known (P. cameronensis possesses half-webbed feet) we believe that it is safe to assume that the tadpole is a P. johnstoni. Its body is broad and slightly flattened; flanks not parallel but slightly convex; nostrils closer to eye than to snout tip; tail axis long and very muscular; no tail fins visible; end of tail shrunken, in life probably pointed; keratodont formula: 3 + 3 // 1 + 1: 2; the upper three rows interrupted by the upper jaw sheath (Fig. 3 a). The body shape and tooth formula corresponds to the most advanced P. palmipes tadpoles (Lamotte et al. 1959) and data on P. cameronensis (Schiøtz 1966). Natural history. P. johnstoni inhabits lowland forest and in contrast to other species occurs in areas without flowing water (Amiet 1983; own data). According to Gartshore (1986) this species inhabits littoral forests. In contrast, Fea (in du Bocage 1903) finds the species at a river with rapids, overgrown by vegetation. The call is very characteristic and can be heard from far (du Bocage 1903). Amiet (1989, 1991) finds an egg clutch on a large leaf in the forest, some centimetres above the ground. A male was sitting close to the clutch during daytime. Because of the absence of an open water source he assumed that the tadpoles of P. johnstoni are terrestrial (Amiet 1989, 1991). Lawson (1993) reports Petropedetes clutches in the centre of leaf whorls, deposited at least 75 m away from the next water source. Tadpoles started to hatch after about two weeks. Lawson (1993) assigns these clutches to what he termed “ P. newtonii ” (described as a new species below). However, according to Amiet’s (1989, 1991) and our own observations this behaviour much better fits to the sympatric P. johnstoni. Distribution. An exact type locality of P. johnstoni has not been given by Boulenger (1888). However, due to the title of the publication it has always been regarded as “Rio del Rey, Cameroon ”. Amiet (1983) points out that this region corresponds to a creek of the river Ndian in western Cameroon, close to the Nigerian border. P. johnstoni is a typical lowland species and is known from Victoria (= Limbe) and Bibundi (Werner 1898; Andersson 1907; Amiet 1983) in western Cameroon and further localities in the lowland of the Biafra Bay down to southern Cameroon (Amiet 1983). P. johnstoni has also been reported from Bioko (du Bocage 1895 a, 1895 b, 1903; Boulenger 1906; Mertens 1965; Weinberg 2008; Fig. 4). J.P. Vande weghe photographed a Petropedetes with a tympanum smaller than the eye and a tympanic papilla in LangouĂ© Bai, eastern Gabon (Fig. 7 g). This would be the first record of the species in Gabon. As no voucher is available this record still needs to be confirmed and the taxonomic status of the population requires further study. Genetics. A genetic comparison of a fragment of the mitochondrial 16 S rRNA gene (611 bp, including gaps) using a calculation of the uncorrected p-distances gave the following results for Petropedetes johnstoni: interspecific comparison between P. johnstoni and all other Central African Petropedetes taxa ranged between 3.85 %- 9.03 %, while the intraspecific variation within P. johnstoni was much lower at 0.17 % (N= 2) (Table 4). Systematic remarks. Boulenger (1888) states that the holotype of Cornufer (Petropedetes) johnstoni is a female specimen. However, according to Parker (1936) the determination of the sex of the holotype’s sex is wrong and it indeed is a subadult male of a size, where it is not distinguishable from subadult females. Parker’s correction has been later adopted by Amiet (1983). The holotype, showing the size, the large femoral glands and the lateral aspect of the head is shown in Fig. 5. By the end of the 19 th century Tympanoceros newtonii has been described by du Bocage (1895 a) from the Island Fernando Po (= Bioko). The type of Tympanoceros newtonii deposited in the MusĂ©e Bocage in Lisbon, is lost (Perret 1976). In contrast to the type of Cornufer johnstoni, the holotype of Tympanoceros newtonii must have been a male in breeding condition. The description of a second specimen form Bioko has been supplemented by an illustration of a male with well developed papilla (du Bocage 1895 b). Both descriptions agree, among other characters, in rudimentary webbing, occurrence of a conical papilla on the tongue, and black bars on the limbs; but differ in the tympanum size. While the tympanum of C. johnstoni is given as half the diameter of the eye, it has been stated as equal to three-quarters of the eye for T. newtonii. However, this difference might be due to different methods of measuring the eye diameter. Another difference between the two descriptions concerns the tibiotarsal articulation. It is said to just reach the snout in the description of C. johnstoni (Boulenger 1888), while it is given to exceed the snout by four to five millimetres in T. newtonii (du Bocage 1895 a). Although this difference appears crucial, the examination of this character state in seven specimens of P. “ newtonii ” from Bioko revealed very variable results, ranging from articulation reaching the snout-tip to exceeding the snout-tip by some millimetres. Werner (1898) reports P. johnstoni from Victoria, Cameroon. These frogs possess a small tympanum and their tibiotarsal articulations extend beyond the snout. In 1900 Boulenger synonymised Tympanoceros and Cornufer with Petropedetes and provided the first key to the three species, known at that time. This key was based on the development of webbing and tympanum size. According to Boulenger (1900) the tympanum reaches half the size of the eye in P. johnstoni and more than half the eye diameter in P. newtonii. The differentiation of the species in Boulenger’s (1900) key was based on specimens collected by Mr. Bates on the Benito River (= River Mbini or Woleu River, today Equatorial Guinea) and not from Fernando Po. In this paper Boulenger (1900) describes the tympanum as being even larger than the eye. Subsequently this statement has been adopted by later authors. Amiet (1983) gives a relation of tympanum to eye size of 100.00–120.00% typical for “ P. newtonii ” and 58.18 –75.00% in P. johnstoni. A second diagnostic character presented by Boulenger (1900) is the development of a tympanic papilla in males of “ P. n e w t o n i i ”; a character obviously unknown at that time in P. johnstoni; e.g. Werner (1898) likewise describes males of that species without tympanic papillae. The occurrence of males’ secondary sexual characters like tympanic papillae and the bony carpal spikes in P. johnstoni was first reported by Andersson (1907). Unfortunately Andersson’s (1907) specimens from Bibundi on the foot of Mt. Cameroon, were destroyed during World War II (F. Geller-Grimm, Museum of Wiesbaden, pers. comm.). Another character believed to distinguish “ P. newtonii ” and P. johnstoni was the position of the tympanic papilla (Amiet 1983). According to Amiet the papilla is positioned close to the upper tympanic border in P. parkeri and P. johnstoni, but is positioned more centrally in “ P. newtonii ” (Amiet 1983). However, the papilla of the holotype from Bioko has been described as being close to the upper tympanic border and likewise illustrated for a second specimen of T. newtonii (du Bocage 1895 a, 1895 b). du Bocage’s (1895 b) drawing is thus more similar to P. johnstoni than to “ P. newtonii ” as defined by Amiet (1983). It is hence plausible to assume that the morphological similarities between C. johnstoni and T. newtonii have been simply overseen due to the fact that Boulenger’s (1888) holotype of C. johnstoni does not possess the characteristic seasonal traits of breeding males and consequently never has been associated with du Bocage’ (1895 a) holotype of T. newtonii. Furthermore Boulenger’s (1900) wrong assignment of specimens from the Benito River (Equatorial Guinea) to P. newtonii from Bioko led to the fact that a wrong diagnostic character for P. newtonii (tympanum larger than the eye) has been adopted in subsequent literature. As P. johnstoni from Bioko and the mainland are morphologically indistinguishable, we herein place P. newtonii into the synonymy with the older name P. johnstoni: Tympanoceros newtonii du Bocage, 1895 = Petropedetes johnstoni (Boulenger, 1888 “ 1887 ”) new synonym. Hence, P. “ newtonii ” (sensu Amiet 1983) on the mainland of western Central Africa requires a new scientific name (see below). Bioko is located only 32 km from mainland Africa and is separated by shallow water (<60 m). This island has probably been connected to the mainland during the last ice age (Jones 1994; Rohling et al. 1998). The fauna of Bioko is of continental origin, comprising various faunal elements, like chameleons, lacertids and anurans, being present on the mainland and Bioko, but not occurring on other islands of the Cameroon volcanic line (Jones 1994; Jesus et al. 2005; Weinberg 2008). In contrast, the islands SĂŁo TomĂ© and PrĂncipe in the Gulf of Guinea are seperated by a depth of more than 1800 m and are inhabited by their own endemic fauna with a deviating origin (Loumont 1992; Drewes & Stoelting 2004; Measey et al. 2007). Hence, the occurrence of the same Petropedetes species on Bioko and the mainland seems possible. However, we cannot exclude the possibility that molecular investigations comparing P. johnstoni from Bioko with those from the mainland may reveal differences on the species level. For example Bioko is inhabited by its own subspecies of the Cameroon toad (Amietophrynus camerunensis poensis), with longer extremities, toes and fingers (Perret 1966). On the other hand specimens of Hyperolius ocellatus from Bioko and adjacent mainland are currently regarded as belonging to one subspecies, different to those from south of the Sanaga River (Perret 1975). If future investigations would reveal that specimens from Bioko do belong to a separate evolutionary lineage, the name P. newtonii would be available for the island populations. As herein defined, P. johnstoni is easily differentiated from other Central African congeners by its rudimentary webbing. P. palmipes and P. perreti are both fully webbed, P. c a m e ro n e n s i s and P. juliawurstnerae sp. nov. are half-webbed. The presence of a distinct tympanum and a tympanal papilla distinguishes P. johnstoni from P. cameronensis and P. palmipes, which lack these characters. The small but distinct tympanum and the large femoral glands differentiate P. johnstoni from P. euskircheni sp. nov., P. parkeri and P. v u l p i a e sp. nov., species with reduced webbing. Furthermore P. johnstoni seems to be strictly restricted to lowland localities in coastal areas, whereas other species live in more mountainous areas and further away from the coast. Etymology. This species has been named by Boulenger (1888) to honour Mr. Henry Hamilton Johnston, who collected the type specimen.Published as part of Barej, Michael F., Rödel, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., Böhme, Wolfgang & Schmitz, Andreas, 2010, Review of the genus Petropedetes Reichenow, 1874 in Central Africa with the description of three new species (Amphibia: Anura: Petropedetidae), pp. 1-49 in Zootaxa 2340 on pages 4-13, DOI: 10.5281/zenodo.19328
Petropedetes palmipes Boulenger 1905
Petropedetes palmipes Boulenger, 1905 Figs. 3 d, 4, 8d, 9 d, 16 Petropedetes palmipes Boulenger, 1905, Ann. Mag. Nat. Hist., Ser. 7, 15: 282 Material examined. IRSNB-KBIN 13.840 (male), Gabon, Estuaire Province, Kango Department, 15 km north of Alen-Nkoma, right of route L107, 2004, coll. O.S.G. Pauwels; IRSNB-KBIN 14.936 (female), 14.937-14.939 (3 males), Gabon, Kinguélé, app. 80 m a.s.l., VI. 2001, coll. O.S.G. Pauwels; IRSNB-KBIN 14.940 (male), Gabon, Province du Woleu-Ntem, Song, app. 360 m a.s.l., 25.VI. 2001, coll. O.S.G. Pauwels; MHNG 961.68 (male), Cameroon, Mbikiliki, 18.I. 1956, coll. J.-L. Perret; MHNG 1033.28 - 31 (4 males), 1033.32 - 33 (2 females), Cameroon, Mbikiliki, 1959, coll. J.-L. Perret; MHNG 1033.49 - 50 (2 males), 1033.51 - 52 (2 females), Cameroon, Bipindi, Abiete, 1969, coll. J.-L. Perret; ZFMK 28942 (female), Cameroon, Kribi, Efulen, 1906, coll. G.L. Rosenberg; ZFMK 73210 - 211 (2 females), Gabon, 18 km northeast of Barrage de Kinguélé, app. 165 m a.s.l., 25.- 27.I. 2000, coll. S. Lötters & V. Gossmann; ZMB 73891 Gabon (male), Province du Woleu-Ntem, Tchimbélé, app. 469 m a.s.l., 08.VI. 2001, coll. O.S.G. Pauwels. Diagnosis. Medium to large-sized Petropedetes; robust body shape; tympanum indistinct; breeding males without tympanic papilla; moderate forearm hypertrophy; carpal spike present; spinosities developed on whole body; moderate femoral glands prominent; toes fully webbed. Description. Medium to large-sized Petropedetes with robust body; females growing slightly larger than males (SUL in males: 40.3–57.8 mm, in females: 37.5–54.7 mm); mean head width in males about 41 % of SUL, in females about 37 %; snout more pointed than in other species; canthus rostralis distinct but rounded, loreal region concave; eye diameter about 1.7 times eye-narial distance; nose closer to snout tip than eye; tympanum very small and indistinct (tympanum / eye in males: 0.18–0.23, in females: 0.20–0.25); no tympanic papilla; supratympanic fold distinct; fingers slender, with typically T-shaped fingertips; relative length of fingers: III> IV> II> I; manual subarticular tubercles single; manual webbing absent; palmar tubercle and thenar tubercle present, palmar tubercle sometimes indistinct; forearm hypertrophy moderately developed in males; carpal spike present in males; spinosities on whole body; dorsal surfaces with heterogeneous skin structure, some larger warts on flank and back, otherwise skin texture on flanks and dorsum similar (Fig. 8 d); ventral skin more granular than in other species; mean femur length in males 53 % of SUL, in females 51 %, mean tibia length in males 58 % of SUL, in females 57 %; mean foot length in males 73 % of SUL, in females 72 %; upper hind limbs of moderate size, lower hind limbs slender; femoral glands larger in males than in females (femoral gland / femur length in males: 0.26–0.38, in females: 0.22–0.31; Fig. 9 d); relative length of toes: IV> III> V> II> I; toes fully webbed, on 4 th toe sometimes slightly less developed 4 (0.5 - 0.5). Coloration. Dorsum uniformly dark; green and black with very few minuscule pale spots; throat whitish or dark marbled; upper hind limbs with pale, thin cross bars; coloration in preservation as in life, but pale. According to Lamotte et al. (1959) adults are of a glossy black colour in life, and therefore hard to find between stones. Boulenger (1905) describes them as dark olive above, spotted or marbled dark and light. Limbs bear broad dark bands and narrow light cross-bars, the ventral side is dirty whitish (Boulenger 1905). Tadpole. Lamotte et al. (1959) describe the tadpoles of P. palmipes. Tadpoles were found in high numbers in small rocky pools at the foot of rapids. The general coloration of the tadpoles is grey on dorsum and flanks, a duller colour around eyes and nostrils; venter translucent, intestinal tract being visible (Fig. 3 d); overall body shape flat; large eyes situated in the first third of body; tail almost twice as long as body and tail tip pointed; jaw sheats small, narrow and serrated; keratodont formula in younger stages 4: 3 + 3 // 1 + 1: 5, in older ones 3 + 3 // 1 + 1: 2 (Fig. 3 d). More developed tadpoles are darker than younger stages. Tadpoles do not exceed a total length of 35 mm (25 mm being tail; Lamotte et al. 1959). Natural history. Very little data on the biology of this species has been published. Specimens were caught at night in primary forest among stones in a stream. Clutches were found glued to rocks (Fig. 16). Gossmann et al. (2002) explains that both eggs and larvae occur on submerged rocks. Lamotte et al. (1959) find tadpoles of P. palmipes in stagnant parts of rivers beneath small rapids. Schiøtz (1966) describes similarities in the morphology of advanced larval stages of P. palmipes and P. cameronensis, possessing identical teeth formulae. Schiøtz (1966) interprets the different tadpole mouth development as a change in biology, younger larvae being rheophilous and older ones semiterrestrial, scraping algae off stones. The full webbing of adults points to a more aquatic life history, at least in post-metamorphs. The call of P. palmipes is not known (Amiet 1989). Distribution. Petropedetes palmipes occurs mainly in lowland areas (Fig. 4). It has been found in southern Cameroon (Perret 1966), in Equatorial Guinea (de la Riva 1994; Lasso et al. 2002) and Gabon (Lötters et al. 2001; Pauwels & Rödel 2007). The P. p a l m i p e s record from western Cameroon by Perret & Mertens (1957) is based on a P. perreti specimen (not known at that time). Genetics. The genetic comparison for the uncorrected p-distances of the mitochondrial 16 S rRNAfragment (Table 4) gave the following results for the single genetic voucher of Petropedetes palmipes which we could use for this gentic comparison: interspecific differences to P. palmipes and all other Central African Petropedetes taxa ranged between 7.80 %- 11.76 %. Systematic remarks. Petropedetes palmipes is one of two fully webbed Central African Petropedetes species; the other one being P. p e r re t i. By the amount of webbing it can easily be differentiated from the halfwebbed species P. cameronensis and P. juliawurstnerae sp. nov. and the rudimentary-webbed species P. euskircheni sp. nov., P. johnstoni, P. parkeri and P. vulpiae sp. nov. P. palmipes can be easily distinguished from the fully webbed species P. perreti by the size of the tympanum, which is very small and rather indistinct in both sexes of P. palmipes, while it is large and distinct in P. perreti (Figs. 8 a, 8 d). In addition breeding males of P. p e r re t i possess a tympanic papilla, which is lacking in P. palmipes.Published as part of Barej, Michael F., Rödel, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., Böhme, Wolfgang & Schmitz, Andreas, 2010, Review of the genus Petropedetes Reichenow, 1874 in Central Africa with the description of three new species (Amphibia: Anura: Petropedetidae), pp. 1-49 in Zootaxa 2340 on pages 39-41, DOI: 10.5281/zenodo.19328
Petropedetes
Key to adult Central African <i>Petropedetes</i> species <p>1 toes fully webbed.......................................................................................................................................................... 2</p> <p>1’ toes half-webbed or rudimentary-webbed.................................................................................................................... 3</p> <p> 2 tympanum distinct; in males ¾ of eye diameter or larger, males with tympanic papillae (Fig. 8 a) and carpal spike in breeding season, tympanum in females up to ¾ of eye diameter; femoral glands large (larger in males) <i>...... P. perreti</i></p> <p> 2’ tympanum indistinct in both sexes (Fig. 8 d), small; males without tympanic papillae, but with carpal spike in breeding season; femoral glands large to very large (larger in males)................................................................ <i>P. palmipes</i></p> <p>3 toes half-webbed...........................................................................................................................................................4</p> <p>3’ toes rudimentary-webbed.............................................................................................................................................. 5</p> <p> 4 tympanum distinct; males with tympanic papillae (Fig. 8 c) and carpal spike in breeding season; femoral gland line shaped in both sexes (Fig. 9 c) <i>.............................................................................................. P. juliawurstnerae</i> <b>sp. nov.</b></p> <p> 4’ tympanum indistinct; males without tympanic papillae or carpal spike (Fig. 8 b); femoral gland ovoid (Fig. 9 b), larger in males <i>....................................................................................................................................... P. cameronensis</i></p> <p> 5 tympanum small but distinct, in males smaller or equal to eye diameter, in females tympanum less than half eye diameter (Fig. 1 a); males with tympanic papillae and carpal spike in breeding season; femoral gland large................ <i>..................................................................................................................................................................... P. johnstoni</i></p> <p>5’ tympanum of moderate size or large and distinct, in males usually bigger than the eye, in females half the eye diameter or larger; males with tympanic papillae and carpal spike in breeding season; femoral gland very small to moderate.................................................................................................................................................................................. 6</p> <p> 6 femoral gland of moderate size; tympanum in males usually oval shaped or flattened, males with tympanic papillae closer to the centre than upper border, carpal spike present in breeding season; females larger than males; species of moderate size (males: <43 mm; females: <47 mm)........................................................................ <i>P. vulpiae</i> <b>sp. nov.</b></p> <p>6’ femoral gland small or very small, shifted to the posterior side of the leg; tympanum in males rounded or flattened; males with tympanic papillae closer to upper border than to centre, carpal spike present in breeding season; large species (males:> 60 mm; females:> 50 mm).............................................................................................................. 7</p> <p> 7 femoral gland small but distinct (Fig. 2 c), in males about 22% of femur length; tympanum in males ¾ of eye diameter or larger than eye, occurs in lower altitudes of south-western Cameroon and eastern Nigeria (Fig. 4).... <i>P. parkeri</i></p> <p> 7’ femoral gland very small (smallest in genus Fig. 2 d), in males only about 16% of femur length; tympanum in males as large as eye diameter, occurs in higher altitudes at Mt. Kupe and Mt. Nlonako (Fig. 4) <i>....... P. euskircheni</i> <b>sp. nov.</b></p>Published as part of <i>Barej, Michael F., Rödel, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., Böhme, Wolfgang & Schmitz, Andreas, 2010, Review of the genus Petropedetes Reichenow, 1874 in Central Africa with the description of three new species (Amphibia: Anura: Petropedetidae), pp. 1-49 in Zootaxa 2340</i> on page 41, DOI: <a href="http://zenodo.org/record/193288">10.5281/zenodo.193288</a>
Petropedetes perreti Amiet 1973
<i>Petropedetes perreti</i> Amiet, 1973 <p>Figs. 3 c, 4, 8a, 9a, 10a, 11a, b</p> <p> <i>Petropedetes perreti</i> Amiet, 1973, Bull. I.F.A.N. sér. A., 2: 463</p> <p> <b>Material examined.</b> MHNG 1253.90 (holotype, formerly JLA 71.196) adult male, Cameroon, Mt. Manengouba, Nsoung, 1400-1500 m, 29.III.1971, coll. J.-L. Amiet; MHNG 961.64 (male), Cameroon, Mt. Manengouba, Nsoung, river Medjié, 1957, coll. J.-L. Perret; MHNG 1522.43 (male), Cameroon, Mt. Manengouba, Nsoung, 29.III.1973, coll. J.-L. Perret; SMF 52364 (female), Cameroon, Nkongsamba, 7.II.1955, coll. J.-L. Perret; ZFMK 69212, 69215, 69219, 69221, 69226 (5 males), 69211, 69213-214, 69216- 217, 69223 (6 females), 69232 (tadpole), Cameroon, Mt. Nlonako, above Nguengue, app. 1100–1200 m a.s.l., 23.XI – 6.XII.1998, coll. ALSCO-Expedition; ZFMK 75524, 75526-28 (4 females),75530, 75532 (2 males), Cameroon, Mt. Nlonako, Nguengue, app. 1100 m a.s.l., 18.-21.I.2000, coll. H.-W. Herrmann & A. Schmitz; ZMB 73731 (female), Cameroon, Mt. Manengouba, Manengouba II, app. 1156 m a.s.l., 16.XI.2006, coll. N.L: Gonwouo; ZMB 73732 (male), Cameroon, Mt. Manengouba, Manengouba II, app. 1260 m a.s.l., 17.XI.2006, coll. N.L. Gonwouo; ZMB 73733 (female), Cameroon, Mt. Manengouba, Manengouba II, app. 1260 m a.s.l., 17.XI.2006, coll. N.L. Gonwouo; ZMB 73734 (female), Cameroon, near Nkongsamba, Plot East 0 0 2, app. 1525 m a.s.l., 08.VIII.2008, coll. M.-O. Rödel & N.L. Gonwouo; ZMB 73735 (male), Cameroon, near Nkongsamba, Plot East 0 0 2, app. 1225 m a.s.l., 10.VIII.2008, coll. M.-O. Rödel & N.L. Gonwouo; ZMB 73736 (female), Cameroon, near Nkongsamba, Plot East 0 0 2, app. 1225 m a.s.l., 17.VI.2008, coll. M.-O. Rödel & N.L. Gonwouo; ZMB 73737 (male), Cameroon, Esipa Village, 12.III.2006, coll. N.L. Gonwouo; ZMB 73738 (tadpoles), Cameroon, Mount Manengouba, 10.VIII.2008, mountain stream at 1225 m a.s.l., coll. N.L. Gonwouo & M.-O. Rödel.</p> <p> <b>Diagnosis.</b> Medium sized <i>Petropedetes</i> with slender body; tympanum distinct, round, smaller or almost equal to eye diameter in males, distinctly smaller in females; white conical tubercles close to vent; characters of breeding males: tympanic papilla present (long and slim), situated in the centre of the tympanum (Fig. 8 a); forearm hypertrophy very weakly developed; carpal spike present; spinosities on inner side of arms and forearms (none on throat); femoral glands distinct in both sexes; glands of moderate size, elliptic shape; fully webbed (Fig. 10 a).</p> <p> <b>Description.</b> Medium sized <i>Petropedetes</i> with slender body; females slightly larger than males (SUL in males: 29.4–42.6 mm, in females 32.2–48.5 mm); mean head width in males about 40% of SUL, in females about 38%; snout rounded in lateral view; <i>canthus rostralis</i> distinct but slightly rounded; loreal region concave; eye diameter about 1.7 times distance eye-nose; nose closer to snout tip than to eye; distinct round tympanum; smaller or almost equal to eye in males, smaller in females (tympanum / eye in males: 0.77–1.06, in females: 0.47–0.71); tympanum encircled with minuscule white warts; tympanic papilla positioned in centre of tympanum; supratympanic fold distinct; fingers slender, with typically T-shaped fingertips; relative length of fingers: III> IV> II> I; manual subarticular tubercles single; manual webbing absent; palmar tubercle and thenar tubercle present, palmar tubercle sometimes indistinct; forearm hypertrophy moderate in males; carpal spikes present in males; spinosities in males on inner side of arms and forearms (not on throat); skin texture heterogeneous on dorsum and flanks; ventral skin smooth; mean femur length in males about 53% of SUL, in females about 51%, mean tibia length in males 58% of SUL, in females 56%; mean foot length in males 75% of SUL in females 73%; upper and lower hind limbs moderate; femoral glands distinct, but flat in both sexes; glands larger and slightly more prominent in males (femoral gland / femur length in males: 0.24–0.41, in females: 0.18–0.28; Fig. 9 a); relative length of toes: IV> III> V> II> I; fully webbed (Fig. 10 a; in few specimens webbing absent on posteriormost three fourth on the fourth toe [4 (0.75-0.75)], but then skin folds runs parallel to the last phalange to the tip of the toe).</p> <p> <b>Coloration.</b> Dorsum marbled of brownish or greenish with black parts or more or less uniformly dark; dorsum speckled with white minuscule spots (Fig. 8 a); iris speckled with gold and green; larger warts on flanks can be coloured brighter than surrounding parts; legs darker than dorsum; belly whitish; throat whitish or marbled with dark; femoral glands with dark marbling; femora above with large dark spots (almost bars), continuing on lower legs; coloration in preservation: dorsum mainly uniform brownish or black with very small white spots; belly yellowish.</p> <p> <b>Tadpole.</b> The elongated tadpoles have an ovoid, flat, almost pointed body and a very long and muscular tail that exceed the body length by at least three times (Fig. 3 c); only last third of tail with very narrow hyaline tail fins, the dorsal ones being a bit longer than the ventral ones; eyes very large and positioned almost dorsally (Fig. 11 b); nostrils positioned slightly closer to the eyes than to the snout-tip; mouth positioned anteroventrally; keratodont formulae in stage 28 tadpoles (N= 5; total length appr. 20 mm) is 3+3//2+2:1; the keratodont formula of one tadpole (ZFMK 69232, Gosner stage 41) slightly differed: 3+3 //1+1:2; upper and lower jaw shaped like a narrow V; spiracle small, not visible in dorsal view; overall dorsal coloration consisted of a greenish brown, beset with minute bluish dots; along the vertebral line short reddish stripes discernible; venter almost uniform whitish. Stage 41 larvae (N= 3) had a body length of 12.7 mm (total length 36 mm) and very long hind legs (20.5 mm).</p> <p> <b>Natural history.</b> <i>Petropedetes perreti</i> is a forest species (Herrmann <i>et al</i>. 2005), occurring along mountain streams above 1200 m and reaching altitudes over 1700 m. It is hence an element of Amiet’s orophil fauna (Amiet 1973). Lamotte <i>et al</i>. (1959) report adults adhering with their bellies and limbs to stones in strong currents. This observation has also been made by Amiet (1975), who additionally found specimens on leaves of plants on the river sides. The clutch is deposited on rock surfaces within the splash zone of rapids and waterfalls, where the tadpoles develop (Amiet 1973, 1983, 1989). Males are found at night close to the clutch and appear to defend it (Amiet 1973).</p> <p> We observed <i>P. p e r re t i</i> at various sites on the eastern flanc of Mount Manengouba between 1200 and 1500 m a.s.l. along fast flowing mountain rivers. Some of these sites were already heavily degraded but still possessed at least some forest on the banks. Adults and juveniles could be observed at night sitting on stones amidst the torrent or some meters away from water (1-2 m above water on tree and palm trunks). One male was observed sitting on a clutch of approximately 84 eggs with white embryos. The clutch was deposited between mosses on a rock in the splash zone of a rapid, approximately 50 cm above the water surface. After being disturbed, the male only moved a few cm from the clutch (Fig. 11 a). At another locality we observed tadpoles moving and feeding on wet rocks in the splash zone of a waterfall during daytime (Fig. 11 b). When approaching them, they only jumped away a few centimeters until they were again out of reach. Only when intensively disturbed did they jump into water.</p> <p> <b>Distribution.</b> <i>Petropedetes perreti</i> is endemic to Cameroon. The species is found on the southern slopes of the Bamiléké Plateau, Mt. Manengouba and Mt. Nlonako (Fig. 4), areas which are characterized by a very high precipitation through the rainy season (Amiet 1973; Herrmann <i>et al</i>. 2005).</p> <p> <b>Genetics.</b> The genetic comparison for the uncorrected p-distances of the mitochondrial 16S rRNAfragment (Table 4) gave the following results for <i>Petropedetes perreti</i>: interspecific comparison between <i>P. perreti</i> and all other Central African <i>Petropedetes</i> taxa ranged between 2.84%-10.16%, while there was no intraspecific variation between the included sequences of the <i>P. p e r re t i</i> vouchers (0.00%; N= 6). The lowest interspecific difference has been detected between <i>P. perreti</i> and <i>P. juliawurstnerae</i> <b>sp. nov.</b> ranging between 2.84%-3.59%, but species can be clearly differentiated on the morphological basis alone (see species description of <i>P. juliawurstnerae</i> <b>sp. nov.</b>).</p> <p> <b>Systematic remarks.</b> <i>Petropedetes perreti</i> is one of to the two species of the genus with fully developed webbing. In contrast, <i>P. euskircheni</i> <b>sp. nov.</b>, <i>P. johnstoni</i>, <i>P. parkeri</i> and <i>P. vulpiae</i> <b>sp. nov.</b> show a mere rudiment of web. It can be easily distinguished from <i>P. p a l m i p e s</i> (fully webbed) and <i>P. c a m e ro n e n s i s</i> (halfwebbed) by its bigger and distinct tympanum and the tympanic papilla in males (see Fig. 8 a, 8b, 8d). The most similar species is <i>P. juliawurstnerae</i> <b>sp. nov.</b> (see respective paragraph for distinguishing characters). Lawson (1993) notes the occurrence of <i>P. p e r re t i</i> from the Rumpi Hills in western Cameroon. However, the taxonomic status of these frogs needs confirmation.</p> <p> <b>Etymology.</b> The species has been named by Amiet (1973) to honour the herpetologist and former curator of herpetology at the Natural History Museum in Geneva Dr. Jean-Luc Perret.</p>Published as part of <i>Barej, Michael F., Rödel, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., Böhme, Wolfgang & Schmitz, Andreas, 2010, Review of the genus Petropedetes Reichenow, 1874 in Central Africa with the description of three new species (Amphibia: Anura: Petropedetidae), pp. 1-49 in Zootaxa 2340</i> on pages 28-30, DOI: <a href="http://zenodo.org/record/193288">10.5281/zenodo.193288</a>
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