36 research outputs found
No ecological opportunity signal on a continental scale?:Diversification and life-history evolution of african true toads (Anura: Bufonidae)
Get PDFThe 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
Second record of the West African hairy bush viper Atheris hirsuta Ernst & Rödel, 2002 (Serpentes: Viperidae)
No full textPenner, Johannes, Gonwouo, Legrand Nono, Rödel, Mark-Oliver (2013): Second record of the West African hairy bush viper Atheris hirsuta Ernst & Rödel, 2002 (Serpentes: Viperidae). Zootaxa 3694 (2): 196-200, DOI: 10.11646/zootaxa.3694.2.
FIGURE 1 in Second record of the West African hairy bush viper Atheris hirsuta Ernst & Rödel, 2002 (Serpentes: Viperidae)
No full textFIGURE 1. Photographs of the second known specimen of Atheris hirsuta showing different aspects from left to right and bottom to top: entire specimen, right lateral side of head, frontal view of head, left lateral side of head showing tongue colouration, midbody dorsal aspects, ventral aspects at midbody, two different views on the fully everted hemipenes of Atheris hirsuta showing the characters described in the text and a map of the two known localities of the species (see text) with the Global Land Cover 2009 as background (dark green depicts lowland rainforest, light green a mosaic of vegetation and croplands).Published as part of <i>Penner, Johannes, Gonwouo, Legrand Nono & Rödel, Mark-Oliver, 2013, Second record of the West African hairy bush viper Atheris hirsuta Ernst & Rödel, 2002 (Serpentes: Viperidae), pp. 196-200 in Zootaxa 3694 (2)</i> on page 198, DOI: 10.11646/zootaxa.3694.2.9, <a href="http://zenodo.org/record/10098932">http://zenodo.org/record/10098932</a>
Petropedetes vulpiae Barej, Rodel, Gonwouo, Pauwels, Bohme and Schmitz 2010
No full textPetropedetes vulpiae Barej, Rödel, Gonwouo, Pauwels, Böhme and Schmitz, 2010 Assignments before official naming of species: in general referred to as P. n e w t o n i concerning specimens on the mainland. Amiet & Perret (1969): P. n e w t o n i corresponds to P. aff. parkeri (locality close to Yaoundé) (see Amiet 1973, 1983) Petropedetes parkeri Amiet, 1983 Assignments before official naming of species: Parker (1936): referred to as P. johnstoni Sanderson (1936): referred to as P. johnstoni Perret (1966): referred to as P. johnstoni Amiet (1973): referred to as P. johnstoniPublished 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 page 48, DOI: 10.5281/zenodo.19328
Petropedetes cameronensis Reichenow 1874
No full text<i>Petropedetes cameronensis</i> Reichenow, 1874 <p>Figs. 3 b, 4, 8b, 9b, 10c, 12, 13</p> <p> <i>Platymantis cameronensis</i> – Peters, 1875, Montasber. Preuss. Akad. Wiss. Berlin, 1875: 211 <i>Hylambates cameronensis</i> – Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 137 <i>Hylambates</i> (<i>Platymantis</i>) <i>cameronensis</i> – Knauer, 1883, Naturgesch. Lurche: 111 <i>Petropedetes cameronensis</i> – Boulenger, 1900, Proc. Zool. Soc. London, 1900: 439 <i>Petropedetes obscurus</i> Ahl, 1924, Zool. Anz., 58: 81-82</p> <p> <b>Material examined.</b> ZMB 8222 (holotype) adult female, 1872-1873, Cameroon, Bimbia, coll. A. Reichenow; MHNG 918.52 (male), Cameroon, Buea, Grand Soppo, 26.IX.1938, coll. R. Mertens & J.-L. Perret; MHNG 1522.32-34 (3 males), 1522.35-37 (3 females), Cameroon, Maholé, Mungo, 28.III.1973, coll. J.-L. Perret; MHNG 1521.40 (male), Cameroon, Buea, 29.XI.1972, coll. J.-L. Perret; MSNG 29891 (male), Cameroon, Mt. Cameroon, Buea, app. 800–1200 m a.s.l., VI.-VII.1902, coll. L. Fea; ZFMK 8798-99 (2 females), Cameroon, Viktoria, II.1973, coll. W. Böhme; ZFMK 14980 (female), Cameroon, Fako, Buea, 20.- 22.III.1974, coll. W. Böhme & W. Hartung; ZFMK 15427 (male), Cameroon, Batoki, 28.II.1954, coll. M. Eisentraut; ZFMK 61343 (female), Cameroon, Ekundu-Kundu, 21.-27.II. l989, coll. M.T. O’Shea; ZFMK 69233 (female), Cameroon, Mt. Nlonako, above Nguengue, app. 1100–1200 m a.s.l., 23.XI.-6.XII.1998, coll. ALSCO-Expedition; ZFMK 78018 (female), Mt. Nlonako: Ekomtolo, app. 500 m a.s.l., XI.2000, coll. H.-W. Herrmann; ZFMK 78355-356 (2 males), 78360 (female), Cameroon, Mt. Nlonako, Ekomtolo, 1/ 5.IV.2001, coll. H.-W. Herrmann <i>et al</i>.; ZFMK 81153 (male), Cameroon, Mt. Nlonako, Ekomtolo, 27.V.2000, coll. M. Solbach; ZFMK 81154 (male), 18.VIII.2000, other data as ZFMK 81153; ZFMK 81555 (female), 81616-617 (2 males), 81614-615 (2 females), Cameroon, Mt. Nlonako: Ekomtolo, 3-7.II.2003, 4.V.2003, 10-11.XII.2002, coll. M. Plath; ZMB 21759, 55222 (2 males, syntypes of <i>Petropedetes obscurus</i> Ahl, 1924), Tscharra, Dana River, coll. Brenner; ZMB 73729-73730 (1 male, 1 female), Nigeria, Rhoko-forest, near Cross River National Park, II.2007, coll. A. Onadeko & M.-O. Rödel; ZMB 73727-72728 (2 females), Nigeria, Mbe Mountains, VI/ VII.2005, coll. A. Onadeko.</p> <p> <b>Problematic material</b>: ZSM 159/1988/1-2 (2 specimens, sex unknown), Cameroon, Victoria, II.1978, coll. F. Sauer.</p> <p> <b>Diagnosis.</b> Small sized <i>Petropedetes</i> with compact body shape; snout short; tympanum very small, rather indistinct; characters of breeding males: tympanic papilla absent; carpal spike absent; forearm hypertrophy not developed; only Central African <i>Petropedetes</i> without spinosity; femoral glands oval shaped; toes approximately half-webbed: 1 (0) 2 (0-0) 3 (0-0) 4 (1.5-1.5) or 4 (2-2) 5 (0) or 5 (0.5) (Fig. 10 c).</p> <p> <b>Description.</b> Small sized <i>Petropedetes</i> with compact body; females distinctly larger than males (SUL in males: 26.8–34.8 mm, in females: 34.2–49.0 mm); mean head width in males about 37% of SUL, in females about 38%; snout in lateral view rounded to slightly pointed; <i>canthus rostralis</i> sharp; loreal region concave; eye diameter about 1.7 times distance eye-nose; nose closer to snout tip than eye; tympanum very small, rather indistinct in both sexes (tympanum / eye in males: 0.31–0.42, in females: 0.28–0.48); no tympanic papilla in breeding males; 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; dorsal skin structure heterogeneous with larger warts on the flanks; few large and longitudinal warts on back; ventral skin smooth; mean femur length in males 51% of SUL, in females 52%; mean tibia length in both sexes 60% of SUL; mean foot length in both sexes 76% of SUL; upper hind limbs of moderate size; lower hind limbs slender; oval femoral glands larger in males than in females (femoral gland / femur in males: 0.30–0.50, in females: 0.22- 0.31, Fig. 9 b); no secondary sexual characters other than femoral gland size known; relative length of toes: IV> III> V> II> I; half-webbed species, toe IV with up to two phalanges free of webbing: 4 (1.5-1.5) or 4 (2- 2); toe V with up to half phalange free of webbing: 5 (0) or 5 (0.5).</p> <p> <b>Coloration.</b> Dorsum with pale spots or light brown colours (Fig. 8 b); one specimen (ZFMK 78355) with dark brown ground colour and small pale dots of about 1 mm; iris golden; flanks can possess white coloured warts; legs coloured similarly as dorsum, but with larger dark bars; throat whitish or dark marbled; belly yellowish grey; coloration in preservation as in life but pale; belly whitish with minuscule speckles, the latter especially dense in the breast region, almost absent on abdomen (recognizable only with dissecting microscope).</p> <p> <b>Tadpole.</b> According to Boulenger’s (1906) figure of the mouth of an advanced <i>P. c a m e ro n e n s i s</i> tadpole the labial-tooth-row formula is 2:1+1//1+1:2 (Fig. 3 b); mouth bordered by papillae; papillae row with a median gap on lower lip. Schiøtz (1966) presents a tooth formula for a still tailed tadpole with fully developed hind limbs as 3+3//1+1:2. The tail length of this specimen is more than 2.5 as long as the body; tail without traces of ventral fin, only a very narrow dorsal fin (Schiøtz 1966).</p> <p> The tadpole assignments of both authors appear reliable, as the species is easily recognizable by the extent of the webbing. Boulenger (1906) even lists another species in his work and Schiøtz (1966) additionally describes the call of adult specimens of <i>P. cameronensis</i>.</p> <p> <b>Natural history.</b> <i>Petropedetes cameronensis</i> occurs mainly in the lowland, but is also known from localities up to 1400 m, where it was found together with <i>P. p e r re t i</i> (Amiet 1973). On Bioko the species is known from mid-altitude, but may also occur in the coastal zone (Boulenger 1906; Mertens 1965). Schiøtz (1963) collects specimens at night on stones in a fast-flowing stream and Amiet (1975) finds the species in rapids and waterfalls. The call of <i>P. cameronensis</i> has been described as a whistle (Schiøtz 1966) or a trill that drowns the noise of torrents (Amiet 1989). Eggs of <i>P. cameronensis</i> are deposited on tree trunks or leaves close to humid rocks, where the saxicolous (living on rocks) tadpoles develop (Amiet 1989). Herrmann <i>et al</i>. (2005) observe males guarding clutches from February to May on Mt. Nlonako. Schiøtz (1966) assumes that the muscular tail without or only narrow fin allows the tadpole to jump on stones.</p> <p> <b>Distribution.</b> In the original description the type locality is given as Bimbia, Cameroon, which has been specified by Perret (1984) as Bimbia, Victoria (= Limbe); and hence the coastal Cameroonian lowland south of Limbe. Schiøtz (1963) calls the type locality “Bimba, Cameroons ”, which is probably a misspelling. This mistake has been further developed as “Bima, Cameroons ” by Frost (2009). <i>P. c a m e ro n e n s i s</i> is also known from moderate altitudes (Lawson 1993), even reaching heights above 1100 m on Mt. Nlonako (Herrmann <i>et al</i>. 2005). The species is known from areas in eastern Nigeria (Schiøtz 1963; Perret 1984) and western Cameroon (e.g. Boulenger 1906; Perret 1966; Fig. 4). Amiet (1973) assumes that the Sanaga River may be the species’ eastern distributional border. <i>P. cameronensis</i> has also been recorded from Bioko (Boulenger 1906; Mertens 1965).</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 cameronensis</i>: interspecific comparison between <i>P. cameronensis</i> and all other Central African <i>Petropedetes</i> taxa ranged between 5.96%-9.25%, while the intraspecific variation within <i>P. cameronensis</i> was much lower at 0.00%-0.50% (N= 4).</p> <p> <b>Systematic remarks.</b> Ahl (1924) describes <i>Petropedetes obscurus</i> from Tscharra, Dana River and distinguishes this half-webbed species from <i>P. cameronensis</i> by the size of its tympanum, which is only one third of the eye diameter in <i>P. obscurus</i> and half the diameter in <i>P. cameronensis</i>. Perret (1984) who studies the validity of <i>P. obscurus,</i> erroneously changes the type locality to Ishiara, Tana River, Kenya. Based on the morphology of the two syntypes (2 males: ZMB 21759, ZMB 55222) he concludes that the diagnostic characters of Ahl (1924) are not sufficient and consequently placed <i>P. obscurus</i> in the synonymy of <i>P. cameronensis</i>. Our examination of the type specimen of <i>P. obscurus</i> confirmed Perret’s (1984) decision. Hence, an affiliation of the two specimens to the East African <i>Arthroleptides</i>, as assumed by Klemens (1998), can be rejected. Both specimens clearly possess vomerine teeth, already mentioned in Ahl’s description (1924, Fig. 12). The placement of the type locality into East Africa must, however be rejected. As such, the type locality remains mysterious. The only Tscharra that we found (www.fallingrain.com) is in Austria (zoogeographically impossible). There are various settlements called Dana in Nigeria, Cameroon, Chad and Central African Republic, however, all in the dry savanna regions.</p> <p> Herrmann (1989) publishes a list of amphibian types in the Zoological Museum, Greifswald (ZIMG) and lists among others, <i>Petropedetes cameronensis</i>. He gives the “ type locality” of this frog as Bonjongo. Our examination of this frog (Acc Cat I – 3411) revealed that it differs from the description of Reichenow (1874).</p> <p> The holotype from ZIMG is bleached out and the femoral glands are hard to identify, but other characters are still recognizable. The tympanum does not possess a papilla, but although being smaller than the diameter of the eye, it is clearly distinct. An even more striking character differentiating this frog from Reichenow’s description is the lack of webbing. These characters are already sufficient to show that this frog cannot be an additional <i>P. cameronensis</i> type. The ZIMG frog is also much smaller than the frog described and figured in Reichenow (1874). Whereas Reichenow (1874) mentions 120 mm, the ZIMG specimen reaches a length of only 82 mm from the tip of the snout to the tip of the toe (Fig. 13). The ZIMG specimen probably belongs to <i>P. johnstoni</i>, which would match with the locality on the foot of Mt. Cameroon. Apparently Herrmann (1989) lists a specimen collected by Buchholz in “ Mai 73 ” (May 1873) from Bonjongo, but misidentifies it. The only identifiable type of <i>P. cameronensis</i>, collected by Reichenow (1874) himself is stored in the ZMB.</p> <p> <i>Petropedetes cameronensis</i> is half-webbed and possesses an indistinct tympanum lacking a tympanic papilla in males. Hence, <i>P. cameronensis</i> is easy to distinguish from the rudimentary-webbed species with a tympanic papilla <i>P. euskircheni</i> <b>sp. nov.</b>, <i>P. johnstoni</i>, <i>P. p a r k e r i</i> and <i>P. vulpiae</i> <b>sp. nov.</b> The species could potentially be mistaken with <i>P. palmipes</i> due to its indistinct tympanum, but differs from <i>P. palmipes</i> by its smaller size, less developed webbing (toes fully webbed in <i>P. p a l m i p e s</i>) and lacking a carpal spike (Amiet 1973). Although having a similar webbing, <i>P. c a m e ro n e n s i s</i> can be unambiguously distinguished from <i>P. juliawurstnerae</i> <b>sp. nov.</b> alike (see the following species description).</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 30-35, DOI: <a href="http://zenodo.org/record/193288">10.5281/zenodo.193288</a>
Petropedetes perreti Amiet 1973
No full text<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>
Petropedetes juliawurstnerae Barej, Rödel, Gonwouo, Pauwels, Böhme & Schmitz, 2010, sp. nov.
No full textPetropedetes juliawurstnerae sp. nov. Figs. 4, 6 c, 7 d, 8 c, 9 c, 10 b, 14, 15 Holotype. ZFMK 67360 (adult male), Cameroon, Mt. Kupe: Nyasoso, 4 ° 49 ’ 42 N, 9 ° 40 ’ 55 E, app. 920 m a.s.l., 19.XII. 1997, coll. O. Euskirchen. Paratypes. IRSNB-KBIN 1989 (female), IRSNB-KBIN 1990 (male), Cameroon, Mt. Kupe, Nyasoso app. 950 m a.s.l., 4 ° 49 ’ 42 N, 9 ° 40 ’ 55 E, 6.VII. 2009, coll. A Schmitz & M.F. Barej; MHNG 2713.12 (male), 5.VII. 2009, other data as IRSNB-KBIN 1989; MHNG 2713.13 (female), same data as IRSNB-KBIN 1989); ZFMK 67357 (female), 2.I. 1998, other data as holotype; ZFMK 67359 (female), 67987 -990, 68170 (4 males), Cameroon, Mt. Kupe, NW-slope, app. 910 m a.s.l., 18.XII. 1997 - 3.I. 1998, coll. A. Schmitz; ZFMK 68131 (male), 68134 - 136 (3 females), app. 950 m a.s.l., other data as ZFMK 67987; ZFMK 75590 (female), 75591 (male), Cameroon, Mt. Kupe: Nyasoso, 4 ° 49 ’ 42 N, 9 ° 40 ’ 55 E, 15.I. 2000, coll. H.-W. Herrmann & A. Schmitz; ZFMK 88866 (female), data as IRSNB-KBIN 1989; ZMB 73694 (male), ZMB (73695) (female), data as IRSNB-KBIN 1989. Additional material. ZFMK 67355 (male), Cameroon, Bakossi Mts., Meked, app. 625 m a.s.l., no date, coll. O. Euskirchen. Diagnosis. Medium sized Petropedetes; slender body shape; tympanum round and distinct, smaller than eye diameter; characters of breeding males: tympanic papilla present, situated in the centre of the tympanum; forearm hypertrophy only weakly developed; carpal spike present; femoral glands long and narrow, not round; a typical dorsal marking of light spots in the form of a flower with five petals recognizable in many specimens (Fig. 14); webbing moderate: 1 (0) 2 (0-0) 3 (1 - 1) 4 (2 - 2) 5 (0.5) (Fig. 10 b). Description of the holotype. The holotype is an adult male with a slender body shape; 32.7 mm SUL; snout in lateral view rounded; head width 13.8 mm; eye diameter 4.8 mm; interorbital distance 3.2 mm; naris closer to snout (1.9 mm) than to eye (3.2 mm); canthus rostralis distinct but rounded; loreal region deeply concave; throat without spinosities (Fig. 6 c); femur length 18.1 mm; tibia length 19.8 mm; foot (incl. longest toe) 24.4 mm; femoral glands large, long and narrow (length x width: 6.8 x 1.9 mm / 6.6 x 2.0 mm); relation femoral gland length to femur length 0.37; spinosity hardly developed on upper forearms; webbing of feet: 1 (0) 2 (0-0) 3 (1 - 0.5) 4 (2 - 2) 5 (0.5); coloration in preservation: overall colour brown-grey; white marking in shape of a 5 -petal flower (indistinct); white warts on flanks, hind limbs and abdomen; upper hind limbs with white transversal bars, continuing on lower legs; ventral side whitish and pale, throat of same coloration with minuscule darker pigmented points; ventral surfaces of femora whitish with darker marbling, which covers in parts the femoral glands; lower leg ventrally with little marbling. Variation. Medium sized Petropedetes with slender body; females being slightly larger (SUL in males: 29.5–38.9 mm, in females: 39.1–42.6 mm); mean head width in males 40 % of SUL in females 39 %; snout in lateral view rounded; canthus rostralis distinct but rounded; loreal region deeply concave; eye diameter about 1.8 times eye-narial distance; nose closer to snout tip than eye; distinct round tympanum, smaller than eye in both sexes (tympanum / eye in males: 0.66–0.8 %, in females: 0.47–0.60); slender tympanic papilla situated in centre of tympanum (Fig. 8 c); 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 weakly developed in males; carpal spike present in males (well visible on X-rays; Fig. 15); spinosities in males weakly developed on upper forearms; dorsal skin heterogeneous with larger warts on flanks; dorsum with prolonged warts, less distinctly developed than in other species; ventral skin smooth; mean femur length in males 54 % of SUL, in females 52 %, mean tibia length in males 60 % of SUL, in females 57 %; mean foot length in males 76 % of SUL, in females 75 %; upper hind limbs of moderate size, lower hind limbs slender; femoral glands line-shaped, long and narrow, of almost similar length in both sexes (femoral gland / femur length in males: 0.36–0.51, in females: 0.32–0.41); relative length of toes: IV> III> V> II> I; half-webbed species, different webbing usually 3 (1 - 1), sometimes 3 (0.5 -0) running along the 3 rd toe, 4 (1.5 - 1.5) up to 4 (2 - 2) and 5 (0) or 5 (0.5); webbing of ZFMK 68170 (paratype) 1 (0) 2 (0-0) 3 (1 -0) 4 (2 - 2) 5 (0). Females of P. juliawurstnerae sp. nov. possess long femoral glands, which may make sex determination difficult. Females are best recognizable by their smaller tympana and more slender femoral glands (Fig. 9 c). Coloration. Dorsum brown with large greenish spots on head, or back dark brown without lighter spots, only light warts (Fig. 8 c); iris golden; flanks with white warts; legs coloured as dorsum with greenish transversal bars; femoral glands pale orange or brownish; coloration in preservation: dorsum predominantly uniform grey or brown with few light, small spots; typical pattern of pale spots, resembling a flower with five petals, in most specimens recognizable; legs with thin white transverse lines. Natural history. The species has been found at a water reservoir and close to small, fast flowing creeks at Mt. Kupe. The streambed was stony and the surrounding vegetation was a transition between primary and secondary rainforest. The frogs were sitting on slippery stones, overgrown by algae, or in the riverine vegetation at heights of 10–20 cm. On Mt. Kupe P. juliawurstnerae sp. nov. occurs in syntopy with P. euskircheni sp. nov. Distribution. At present the species is only known from its type locality at Mt. Kupe and Meked on the adjacent Bakossi Mountains (Fig. 4). Genetics. The genetic comparison for the uncorrected p-distances of the mitochondrial 16 S rRNAfragment (Table 4) gave the following results for Petropedetes juliawurstnerae sp. nov.: interspecific comparison between P. juliawurstnerae sp. nov. and all other Central African Petropedetes taxa ranged between 2.84 %- 12.39 %, while the intraspecific variation within P. juliawurstnerae sp. nov. was much lower at 0.00%- 0.90 % (N= 5). The lowest interspecific difference has been detected between P. p e r re t i and P. juliawurstnerae sp. nov. ranging between 2.84 %- 3.59 %, but species can be clearly differentiated on the morphological basis alone (see below). Systematic remarks. Mostly closely P. juliawurstnerae sp. nov. resembles P. p e r re t i and P. cameronensis. It shares a tympanic papilla in the centre of the tympanum and a similar shape with P. perreti (Figs. 8 a, 8 c), and similar webbing with P. cameronensis (Figs. 10 b, 10 c). From the other species P. juliawurstnerae sp. nov. is distinctly different by its half-webbed toes; i.e. fully webbed in P. palmipes (additionally lacking the tympanal papilla) and rudimentary-webbed in P. euskircheni sp. nov., P. johnstoni, P. parkeri, and P. vulpiae sp. nov. Breeding males of P. juliawurstnerae sp. nov. can be distinguished from other species with a tympanic papilla by the position of the papilla. Only P. juliawurstnerae sp. nov. and P. perreti have papillae in the centre of the tympanum. In all other species this papilla is positioned closer to the dorsal edge of the tympanum. The papilla of P. juliawurstnerae sp. nov. is broad at the end, when not shrivelled after preservation. From the most similar two species, P. p e r re t i and P. cameronensis, P. juliawurstnerae sp. nov. can be distinguished by the following characters: P. juliawurstnerae sp. nov. is smaller than P. p e r re t i (SUL in males: N juliawurstnerae = 12, N perreti = 13, p <0.001; in females: N juliawurstnerae = 10, N perreti = 15, p <0.01). The tympanum diameter / eye diameter ratio is smaller in male P. juliawurstnerae sp. nov. (tympanum diameter / eye diameter in males: N juliawurstnerae = 12, N perreti = 13, p <0.001), but does not differ significantly in females (p = 0.23). The webbing is less developed than in P. p e r re t i (Figs. 10 a, 10 b): 1) webbing deeper notched in P. juliawurstnerae sp. nov., 2) webbing always less developed on the fourth toe, where it usually leaves 1.5–2 phalanges free in P. juliawurstnerae sp. nov.; but only 0 or 1 free phalange in P. perreti. The femoral gland of P. juliawurstnerae sp. nov. is long and narrow (stripe-like) in males and females. The femoral glands of P. perreti are more oval. The gland length / gland width ratio in P. juliawurstnerae sp. nov. is significantly larger (gland length / gland width in males: N juliawurstnerae = 12, N perreti = 13, p <0.001; in females: N juliawurstnerae = 10, N perreti = 15, p <0.001). In contrast, P. c a m e ro n e n s i s and P. juliawurstnerae sp. nov. do not differ in their size (males p= 0.19; females p = 0.29). Petropedetes juliawurstnerae sp. nov. can be distinguished from P. cameronensis by the size of the tympanum which is distinct in P. juliawurstnerae sp. nov., but only rudimentary or even indistinctly developed in P. cameronensis (tympanum diameter / eye diameter in males: N juliawurstnerae = 12, N cameronensis = 16, p <0.001; in females: N juliawurstnerae = 10, N cameronensis = 17, p <0.001). P. c a m e ro n e n s i s males lack the tympanic papillae present in breeding P. juliawurstnerae sp. nov. males. Especially in females, which do not carry a tympanic papilla, the similar webbing makes it difficult distinguishing the new species from P. cameronensis. In this case the shape of the femoral gland is the character best suited for species identification. The femoral glands are oval shaped in P. cameronensis; the gland length / gland width ratio in P. juliawurstnerae sp. nov. is significantly larger (gland length / gland width in males: N juliawurstnerae = 12, N cameronensis = 16, p <0.001; in females: N juliawurstnerae = 10, N cameronensis = 17, p <0.001). Spinosities in examined specimens of P. juliawurstnerae sp. nov. seem not to be fully developed, therefore spines could be developed on the throat and more developed on upper forelimbs as well, as in other species (Fig. 6 c). Etymology. The specific epithet juliawurstnerae is a matronym. The name is dedicated to Mrs. Julia Anna Maria Wurstner (Bad Neuenahr-Ahrweiler, Germany), who accompanied the senior author during fieldwork in 2007 and supported his work from the beginning.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 35-39, DOI: 10.5281/zenodo.19328
Petropedetes euskircheni Barej, Rodel, Gonwouo, Pauwels, Bohme and Schmitz 2010
No full text<i>Petropedetes euskircheni</i> Barej, Rödel, Gonwouo, Pauwels, Böhme and Schmitz, 2010 <p>Assignments before official naming of species:</p> <p> Euskirchen <i>et al</i>. (1999): referred to as <i>P. parkeri</i></p> <p> Herrmann <i>et al</i>. (2000): referred to as <i>P. parkeri</i></p> <p> Plath <i>et al</i>. (2004): referred to as <i>P. parkeri</i></p> <p> Herrmann <i>et al</i>. (2005): referred to as <i>P. parkeri</i></p> <p> <i>Petropedetes perreti</i> Amiet, 1973</p> <p> Assignments before official naming of species, problematic assignment: Perret & Mertens (1957): as <i>P. palmipes</i></p> <p> Lawson (1993): <i>P. p e r re t i</i> from Rumpi Hills, locality unsure</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 48, DOI: <a href="http://zenodo.org/record/193288">10.5281/zenodo.193288</a>
Petropedetes johnstoni
No full textPetropedetes 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 parkeri Amiet 1983
No full text<i>Petropedetes parkeri</i> Amiet, 1983 <p>Figs. 1 c, 2c, 4, 7a, 7b, 7f</p> <p> <b>Material examined.</b> BM 1936.3.4.124 (holotype) adult male, Cameroon, Mamfe region, Atolo, Assumbo, 12.IV.1933, coll. I. Sanderson; BM 1936.3.4.123 (male), Cameroon, Mamfe; BM 1936.3.4.127 (female), 1936.3.4.136 (male), Cameroon, Bachor II; 1936.3.4.128 (female), 1936.3.4.131 (male), Cameroon, Bashau, 1932-1933, coll. I. Sanderson; MHNG 961.66 (male, formerly BM 1936.3.4.133), Cameroon, Mamfe Division, Bashau, 10.III.1933, coll. I. Sanderson; PEM A 9123-9124 (2 males), A 9126 (male), A 9127 (female), Cameroon, Nguti, 15.VII.1996, coll. L. Minter; PEM A 9125 (male), A 9128 (male), 5. & 9.VII.1996, other data as A 9123; ZFMK 61337-338 (2 females), 61342 (male), Cameroon, Ekundu-Kundu, 3-27.II.1989, coll. M.T. O’Shea; ZFMK 87702 (female), Cameroon, Mamfe region, near Amebishu, app. 720 m a.s.l., 14.IX.2007, coll. J.A.M. Wurstner & M.F. Barej; ZMB 73739 (female), Nigeria, Okwangwo, app. 900 m a.s.l., 15.IV.2004, coll. M. Gartshore.</p> <p> <b>Problematic material</b> (see systematic remarks). IRSNB-KBIN 13.842 (female), Gabon, Estuaire Province, Kango Department, 15 north of Alen-Nkoma, right of route L107, 2004, coll. O.S.G. Pauwels; IRSNB-KBIN 14.941 (female), Gabon, Province du Woleu-Ntem, Tchimbélé, app. 469 m a.s.l., 20.VI.2001, coll. O.S.G. Pauwels; MHNG 1521.60 (male), 1521.61 (female), 1521.64-65 (2 females), Cameroon, Bafoussam, Bangwa, 7.IV.1973, coll. J.-L. Perret; MHNG 1521.70-71 (2 males), Cameroon, Yaoundé, Mt. Kala, X.1972, coll. J.-L. Perret; MHNG 1521.78 (male), Cameroon, Dschang, Falaise de Mbos, 25.XI.1972, coll. J.-L. Perret; ZFMK 73209 (female), 73210 (male), Gabon, Barrage de Tchimbélé, coll. V. Gossmann & S. Lötters; ZMB 73892 (female), same data as IRSNB-KBIN 14.941.</p> <p> <b>Diagnosis.</b> Large sized <i>Petropedetes</i>; robust body shape; tympanum round, larger than eye diameter in males, smaller in females; characters of breeding males: tympanic papilla present (broad and fleshy); papilla close to the upper border of the tympanum; forearm hypertrophy strongly developed in males; carpal spike present; tegumental spinosities well developed on throat, forearms, scattered on flanks and dorsum; femoral glands small, coloured similar to limb, shifted slightly to the posterior side of the leg; webbing rudimentary.</p> <p> <b>Description.</b> Large sized <i>Petropedetes</i> with robust body; males distinctly larger than females (SUL in males: 38.0– 74.3 mm, in females: 34.1–61.3 mm); mean head width in males about 44% of SUL, in females about 42%; snout in lateral view short, rounded; <i>canthus rostralis</i> slightly rounded but distinct; loreal region concave; eye diameter about 1.3 times eye-narial distance; nose closer to snout-tip than to eye; distinct tympanum, usually larger than eye in males, always smaller in females (tympanum / eye in males: 0.73-1.39, in females: 0.60–0.89; the lowest value of 0.73 in males measured in the smallest specimen, second smallest value: 0.92; tympanic papilla closer to upper border than centre of tympanum; papilla broad and fleshy with oval basis; supratympanic fold distinct; fingers slender, with typically T-shaped fingertips; relative length of fingers: III> IV> II> I; manual subarticular tubercles single; palmar tubercle and thenar tubercle present, palmar tubercle sometimes indistinct; forearm hypertrophy strongly developed in males; carpal spike present in males; tegumental spinosities well developed on throat, forearms, scattered on flanks and dorsum in males; dorsal tegument with smaller warts than flanks; few larger, elongated warts on dorsum; ventral skin smooth; femora long, mean femur length in males 50% of SUL, in females 51%; mean foot length in both sexes 74% of SUL; upper hind limbs of moderate width, lower hind limbs slender; femoral glands small in both sexes, slightly bigger in males (femoral gland / femur in males: 0.18–0.27, in females: 0.16–0.22); relative length of toes: IV> III> V> II> I; webbing rudimentary: 1 (1) 2 (1-1) or 2 (1.25-1) 3 (2-2) 4 (3-3) 5 (2).</p> <p> <b>Coloration.</b> Dorsum and flanks olive or brownish and dark marbled with diffuse brown-olive spots (Fig. 1 c); throat dirty whitish; belly whitish, slightly translucent; throat can be darker than belly; femora and lower legs with large darker spots, darker spots divided by thin bright coloured transversal bars; ventral surfaces of limbs of the same coloration as belly, pale greenish; iris golden with white-green shades; femoral glands coloured as hind limbs, pale orange or greenish. Coloration in preservation: similar as in life; dorsum uniform brown; belly whitish, with minuscule speckles (less dense than in former species, only recognizable under microscope); throat dirty whitish or pale brown.</p> <p> <b>Natural history.</b> After the first work on the ecology of the species by Sanderson (1936, referred to as <i>P. johnstoni</i>), additional data have been gathered by Amiet. However, some observations by Amiet (e.g. 1975, 1983, 1986) are based on populations of “ <i>P. parkeri</i> ” from Mt. Kala, the Bamiléké Plateau and the region of Nkongsamba. The taxonomic status of these populations at present is not clear (see below). Sanderson (1936) calls this species arboreal, because he found them on lower, broadened leafs, in shade. The breeding season is said to take place in the rainy season (Parker 1936; referred to as <i>P. johnstoni</i>). The species has been found on wet, mossy rocks in the forest; the large rocks were crossed by runlets (Fig. 7 a). According to Sanderson (1936) the adults aggregate in the breeding season on humid rocky surfaces in rough areas with torrent water, while they live in the forest on leaves outside the breeding season.</p> <p> The advertisement call of <i>P. p a r k e r i</i> has been published by Narins <i>et al</i>. (2001). Narins <i>et al</i>. (2001) describe a part of the courtship behaviour, including the female striking the male’s head with her foreleg during amplexus. These authors speculate that this behaviour is connected with secretion of glands in the male tympanal papilla. The small eggs are deposited at the beginning of the rainy season on surfaces of stones within the splash zone. Metamorphosis in <i>P. parkeri</i> is finished before the end of the rains or at the beginning of the dry season (Parker 1936).</p> <p> <i>Petropedetes parkeri</i> shows, in contrast to other members of the genus, a reversed sexual size dimorphism (Amiet 1983). The state that males grow larger than females is rare and occurs only in about 10% of amphibians (Shine 1979). Most often this is connected to some kind of territoriality or male parental care. However, Sanderson (1936) notes the laceration of non-breeding males in the gular region and assumed relics of the previous breeding seasons. This could be regarded as an outcome of territorial behaviour. Further respective data are however lacking. This species appears to live in forests. Some specimens of <i>P</i>. aff. <i>parkeri</i> from Gabon (see below, Fig. 7 f) have been collected during the day in a gallery forest. <i>P</i>. aff. <i>parkeri</i> is called “barking frog” by locals in western Gabon (V. Gossmann <i>pers. comm.</i>).</p> <p> <b>Distribution.</b> We can only assign populations from western Cameroon and eastern Nigeria to <i>P. parkeri</i> with certainty (Fig. 4). At these locations species inhabit altitudes below 1000 m.</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 parkeri</i>: interspecific comparison between <i>P. parkeri</i> and all other Central African <i>Petropedetes</i> taxa ranged between 5.94%-12.39%, while the intraspecific variation within <i>P. p a r k e r i</i> was much lower at 0.00%-0.22% (N= 4).</p> <p> <b>Systematic remarks.</b> <i>Petropedetes parkeri</i> has long been confused with “ <i>P. newtonii</i> ” (described herein as <i>P. v u l p i a e</i> <b>sp. nov.</b>) and <i>P. johnstoni</i> (Gartshore 1994). Amiet (1983) first recognizes that the description of the species’ habitat by Parker (1936) and Sanderson (1936) do not fit the circumstances of the description by Boulenger (1900). He subsequently described this species using distinct morphological characters. <i>P. parkeri</i> and <i>P. euskircheni</i> <b>sp. nov.</b> are the two largest members of the genus. The fleshy papilla in both species has an oval basis, while it is rounder in the remaining species. <i>P. parkeri</i> is only rudimentary-webbed and therefore easily distinguishable from more fully webbed species (<i>P. palmipes</i> and <i>P. p e r re t i</i>, both fully webbed; <i>P. cameronensis</i> and <i>P. juliawurstnerae</i> <b>sp. nov.</b>, both half-webbed). Moreover, the presence of a distinct tympanum and a tympanal papilla seperates <i>P. parkeri</i> from <i>P. cameronensis</i> and <i>P. palmipes</i>, which lack these characters. The size of the femoral gland distinguishes <i>P. parkeri</i> from <i>P. vulpiae</i> <b>sp. nov.</b> and <i>P. johnstoni</i>, which possess much larger ones. Due to a similar appearance, <i>P. p a r k e r i</i> has been confused with a cryptic species in the past. Distinguishing characters between <i>P. p a r k e r i</i> and <i>P. euskircheni</i> <b>sp. nov.</b> are presented in the description of the new species (see below).</p> <p> As holotype, Amiet (1983) has chosen a <i>P. parkeri</i> from Atolo, Mamfe region. The paratypes were from localities in the Mamfe region, but also from the “Falaise de Mbos” and Bangwa. He further investigated material from Kala, Léna (Yaoundé region). He regards the species as vocally and morphologically homogenous throughout its discontinuous range, although he mentioned small deviations in the lengths of the femora between western and eastern populations (Mt. Kala, Yaoundé region; Amiet 1983). The type locality at Atolo is located at app. 425 m a.s.l. The overall altitudinal distribution ranges between 400 and 2300 feet (approximately 120-700 m; Parker 1936; Sanderson 1936). Own and unpublished findings confirm sites below 1000 m. The sites are a hill close to Mukwecha (720 m), Okwangwo (900 m) and material collected by Les Minter from Nkwende Hills near Nguti (400 m). Amiet (1978, 1983) instead lists altitudes between 750– 1400 m from Mbakang and Foto-Ndonchwet and doubts localities in very low altitudes from Sanderson (1936) and Parker (1936). The altitudinal range of <i>P. parkeri</i> may overlap with the altitudinal range of <i>P. euskircheni</i> <b>sp. nov.</b> (type locality at about 900 m; see below). Without genetic comparisons, so far the <i>P. parkeri</i> -like populations in Central Cameroon region around Yaoundé and the northern populations of the Manengouba Mountains cannot be assigned with certainty, neither to <i>P. parkeri</i> nor to <i>P. euskircheni</i> <b>sp. nov.</b></p> <p> Due to the finding of <i>P. euskircheni</i> <b>sp. nov.</b> not all populations can be irrespectively assigned to a particular taxon. We therefore assign unsure localities within Cameroon (e.g. Mt. Kala), but also specimens from Gabon (e.g. Frétey & Blanc 2001; Lötters <i>et al</i>. 2001) and Equatorial Guinea (de la Riva 1994; Lasso <i>et al</i>. 2002) to <i>P.</i> aff. <i>parkeri</i> (Fig. 4).</p> <p> <b>Etymology.</b> The species has been named by Amiet (1983) in honour of Mr. Hampton Wildman Parker, former curator of the Natural History Museum, who conducted a meticulous morphological study on specimens from the Sanderson collection.</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 17-22, DOI: <a href="http://zenodo.org/record/193288">10.5281/zenodo.193288</a>
