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Afrixalus lacustris Greenbaum, Dehling, Kusamba & Portik sp. nov. Great Lakes Spiny Reed Frog urn:lsid:zoobank.org:act: 41D3C41A-4501-4BEC-BFB6-B9C6C1518DC3 Afrixalus laevis (nec Megalixalus laevis Ahl 1930)— Laurent 1950: 24; Laurent 1972: 59 (partim); Laurent 1982: 33 (partim); Schiøtz 1999: 56 (partim); Channing & Howell 2006: 137 (partim); Spawls et al. 2006: 183 (partim); Channing & Rödel 2019: 158 (partim). Holotype. UTEP 20805 (field no. ELI 605), adult male, from the vicinity of Kalundu (03.15552° S, 28.42108° E, 1482 m), South Kivu Province, DRC, collected on 21 December 2010 by Chifundera Kusamba and Félix I. Alonda (Figs. 8D–F, 12A–B). Paratopotype. UTEP 20806 (field no. ELI 606), adult male, collected with the holotype. Referred specimens. Specimens with an asterisk were not included in morphological analyses due to poor quality of preservation, lack of a voucher, or unavailability of specimen(s): UTEP 20807–08 (field nos. ELI 644, 649), vicinity of Kalundu (03.15552° S, 28.42108° E, 1482 m), South Kivu Province, DRC; UTEP 20809, 22422 (field nos. ELI 669–70), Baraka, near shore of Lake Tanganyika (04.10832° S, 29.09705° E, 777 m), South Kivu Province, DRC; UTEP 20810–11 (field nos. EBG 1316, 1319), vicinity of Irangi (01.8873° S, 28.4495° E, 820 m), South Kivu Province, DRC; UTEP 22423 (field no. MUSE 10192), Kahuzi-Biega National Park, Nanwa (02.362138° S, 28.192589° E, 1566 m), South Kivu Province, DRC; UTEP 22424 (field no. MUSE 10137), Itombwe Plateau, Mbandakila (03.47937° S, 28.413049° E, 1135 m), South Kivu Province, DRC; RMCA 77-020 -B-133–136* (four specimens), Itombwe Plateau, Tubutubu (2300 m), South Kivu Province, DRC; RMCA 77-020 -B-89–92 (field nos. MF 14–15 [two specimens]), Kitutu (ca. 03.28° S, 28.11° E, 700 m), South Kivu Province, DRC; RMCA 77-020 - B-0069–71 (1 specimen), Itula (ca. 03.27° S, 28.12° E, 650 m), South Kivu Province, DRC; RMCA 77-020 -B-0081–83, 77-20-B-0086, Kamituga (ca. 03.54° S, 27.69° E, 1050 m), South Kivu Province, DRC; RMCA 77-020 - B-0072–73 (1 specimen), Mwana (ca. 03.15° S, 28.47° E, 1650 m), South Kivu Province, DRC; UTEP 22417 (field no. CRSN 2773), Toyokana (02.02734° N, 030.06653° E, 1294 m), Ituri Province, DRC; (field no. EPLU 395 *, photo and tissue only), ca. 0.5 km E of Epulu, near Mt. Mbiya (01.39594° N, 28.58093° E, 755 m), Ituri Province, DRC; UTEP 22416 (field no. DFH 247), Bwindi Impenetrable National Park, Buhoma (00.99045° S, 29.61884° E, 1523 m), Western Region, Uganda; (field nos. DFH 1102–03 * photos and tissues only), Kibale Forest National Park, Ngogo Research Center (00.49795° N, 30.42262° E, 1363 m), Western Region, Uganda; CAS 202032–36, Bwindi Impenetrable National Park, Buhoma rd., 2 km S (by rd.) of Bizenga River (01.00975° S, 29.620694° E), Western Region, Uganda; CAS 202133, Bwindi Impenetrable National Park, small tributary to Ishasha River (00.905° S, 29.704972° E, 1280 m), Western Region, Uganda; CAS 202109, Bwindi Impenetrable National Park, Bizenga River at Buhoma rd. (00.99275° S, 29.615722° E), Western Region, Uganda; CAS 256035 *, Bwindi Impenetrable National Park, rd. north of Ruhija, Western Region, Uganda; CAS 256128–31 *, Mabira Forest Reserve, marshy area adjacent to unpolluted stream (0.4508° N, 32.9493° E, 1121 m), Mukono District, Central Region, Uganda. Diagnosis. The species is referred to the genus Afrixalus for exhibiting the following characteristics: fingers and toes webbed; tips of fingers and toes enlarged to discs; eye large; pupil vertically elliptical; tympanum indistinct; vomer ridges and teeth absent; dorsal surfaces finely shagreened with minute pointed tubercles; outer metatarsal tubercle distinct; gular gland in males. It is readily distinguished from most other members of the genus by its small size (SVL in males 18.9–22.2 mm, in females 20.8–25.7 mm), being smaller than A. dorsalis (males 25–28 mm, females 26–30 mm), A. fornasini (males 30–38 mm, females 35–40 mm), A. fulvovittatus (males 23–27 mm, females 25–28 mm), A. lacteus (males 22–27 mm, females 25–29 mm), A. leucostictus (males 27–32 mm, females 25–36 mm), A. manengubensis (males to 32 mm), A. nigeriensis (males 28–34 mm, females 32–35 mm), A. osorioi (males 27–31, females 32–35 mm), A. paradorsalis (males 28–34 mm, females 32–35 mm), A. septentrionalis (males 19–21 mm), A. vittiger (males 22–25 mm, females 25–28 mm), and A. wittei (males 27–30 mm, females 29–33 mm); and larger size than A. delicatus (males 15–19 mm, females 17–22 mm) and A. stuhlmanni (males 15–21 mm, females 17–25 mm). Its differs in dorsal coloration and pattern from all species with longitudinal stripes or bands on a yellowish-brown background (A. brachycnemis, A. crotalus, A. delicatus, A. dorsalis, A. enseticola, A. fornasini, A. fulvovittatus, A. knysnae, A. morerei, A. orophilus, A. quadrivittatus, A. schneideri, A. spinifrons, A. stuhlmanni, A. upembae, A. septentrionalis, A. vittiger, and A. wittei); species showing a conspicuous large dark dorsal blotch on a uniform background on anterior dorsum in combination with large blotches on both sides of the hip (A. leucostictus, A. manengubensis, A. osorioi, A. paradorsalis, and A. schneideri) or with longitudinal lateral dark bands (A. equatorialis and A. nigeriensis); species with a uniformly colored dorsum without conspicuous pattern except a dark dorsolateral stripe (A. aureus, A. clarkei, A. delicatus, A. fornasini, A. lacteus, A. leucostictus [dorsum with small white tubercles], A. uluguruensis, and A. weidholzi [might have a black vertebral line and dark brown flanks]); and from all remaining species having either dark stripes running from tip of snout to back, crossing each other and continuing as dorsolateral stripes (A. vibekensis, also males without asperities); anterior part of dorsum yellowish with brown pattern, posterior part of dorsum translucent (A. laevis, also males without asperities); posterior part of dorsum and dorsal sides of limbs semi-translucent with dense small dark white-edged speckles (A. dorsimaculatus); and dark blotches and transverse marks with white speckles (A. sylvaticus). It is most similar to A. phantasma sp. nov. but differs from this species in being smaller (Table 1) and showing an overall darker coloration and more contrasting dorsal pattern that is obvious even in preserved specimens (Figs. 7–8, 12). It is readily distinguished from A. phantasma sp. nov. by the more extensive hand and toe webbing (Fig. 8E–F). Description of holotype. Measurements of the holotype are provided in Table 2. Body very slender, widest at temporal region, slightly tapering to groin (Fig. 8D–F); head small (HL/SVL 0.31, HW/SVL 0.31), about as long as wide (HW/HL 0.99); snout relatively long, rounded in dorsal view and in lateral profile, slightly wider than long; canthus rostralis hardly discernible, slightly concave between eye and nostril in dorsal view; loreal region oblique; nostrils rounded, directed anterolaterally and slightly dorsally; situated much closer to tip of snout and to eye, separated from each other by distance about equal to distance between eye and nostril (IN/EN 1.02); eye directed anterolaterally, strongly protruding, very large (ED/HL 0.39), its diameter shorter than snout (ED/SL 0.90); interorbital distance much wider than upper eyelid (IO/EW 1.61) and wider than internarial distance (IO/IN 1.31); tympanum covered by skin, not visible externally; upper jaw with dentition; teeth on premaxilla larger than teeth on maxilla; choanae small, located far anterolaterally at margins of roof of mouth, its anterior edge covered by maxillary bone, therefore appearing semicircular in ventral view; vomer ridges and teeth absent; tongue heartshaped, longer than broad, bilobed for about one-fourth of its length, free distally for about two-thirds its length; densely covered with minute papillae; median lingual process absent. Dorsal surfaces of head, trunk, and limbs and lateral surfaces of trunk finely shagreened; tiny pointed, widely scattered tubercles on dorsum, more numerous posteriorly than anteriorly; supratympanic fold absent; few tubercles at rear end of mandible without pointed tips; ventral side of head smooth; vocal sac present; gular gland of vocal sac smooth, large and wide, covering about 70 percent of throat width (Fig. 8E); chest smooth, abdomen weakly areolate; ventral side of limbs smooth; short transverse fold above vent. Forelimbs slender; hand large (HaL/SVL 0.28); tips of fingers enlarged into large disks, each with circummarginal groove; relative length of fingers: I I 2-/2 II 2-/3- III 2.25/2 IV; thenar tubercle oval, small and very low, about one-fourth length of metacarpal of Finger I; inner palmar tubercle and outer palmar tubercle very low, small, rounded, almost indiscernible. Hind limbs slender, moderately long; heel reaching to level of eye when legs adpressed forwardly to body; crus moderately long (TibL/SVL 0.44), slightly shorter than thigh; heels not touching each other when knees flexed and thighs held perpendicularly to median plane; foot subequal in length to crus (FoL/TibL 0.94); relative length of toes: I I 1.5/2+ II 1+/2+ III 1+/2+ IV 2/1+ V (Fig. 8E); inner metatarsal tubercle moderately prominent, elongated, about half length of metatarsus of Toe I; outer metatarsal tubercle small, rounded. Coloration in life. During night, skin on dorsal side of head, trunk, forelimbs, crus and tarsus light brown with more or less regular, locally more densely clustered dark brown speckling; distinct dark brown transverse stripe between upper eyelids, another less distinct one in scapula region, and another narrow one at level of anterior end of pelvis; dorsal skin of thigh largely unpigmented with only a broad stripe of widely scattered brown pigmentation; distinct dark brown band along canthus, continuing behind eye on both sides of trunk to level of pelvis; tubercles below eye and in tympanic region at rear end of jaw white; dark brown stripe running diagonally on middle of tibia, forming interrupted, inverted U-shaped band together with pelvic spots when legs folded against body; skin of ventral side of head, trunk and limbs largely unpigmented and translucent; gular gland bright yellow; fingers and toes yellow (Fig. 12). During day, basic dorsal coloration light brown; dark brown dorsal speckles, spots and stripes in more pronounced contrast to basic coloration, more distinctly visible. Coloration in preservative. Dorsal basic coloration largely faded to light brown; darker dorsal pattern elements dark brown, clearly visible; color of gular gland, fingers and toes faded to white (Fig. 8D–F). Variation. The paratypes match the holotype in general appearance and proportions. Coloration and color pattern varies from light-cream to yellowish-orange with a few relatively large and darker blotches and vermiculation, light speckling and indistinct dorsolateral stripe to an almost reticulated pattern of contrasting bright and dark elements with a broad and conspicuous brown dorsolateral band (Fig. 12). Pedal webbing variation is I 1.5[100]/2+[100] II 1+[100]/2+[100] III 1 +[50],1.25[50]/2+[100] IV 2 [100]/1+[100] V. Bioacoustics. The advertisement call and other vocalizations are unknown. Ecology and natural history. We collected males and females in swamps in forest openings and near forest edges (Fig. 11B), from vegetation at the edge of streams (or on vegetation overhanging them) in forest, and from non-forest vegetation near the shore of Lake Tanganyika. Egg deposition has not been observed and tadpoles are unknown. An adult male (CAS 256035) was found in Bwindi Impenetrable National Park (Uganda) ca. 2 m above ground in a tree fern “either calling or moving to a calling site (similar to ‘running’ of Kassina)” (D. Blackburn, in litt., 22 July 2021). Laurent (1955) noted the species from transitional forest. In his paper on amphibians of Virunga National Park, Laurent (1972) noted the species is most commonly found in swamps with herbaceous vegetation in secondary forest, and more rarely, in primary forest. Laurent (1982) noted that, in general, the species is common in the foliage of shrubs in dense forests. He also recorded it from “une mare sombre” (i.e., a dark pond) in syntopy with Chiromantis at Kitutu (DRC), marshes where it reproduces by sticking its eggs under the leaves of fountain grass (Pennisetum, Shabunda, DRC), a shaded bank of a pond in syntopy with Hyperolius “ tuberculatus ” (likely H. hutsebauti sensu Bell et al. 2017) and H. frontalis; the center of this pond had reeds harboring H. kivuensis and Afrixalus orophilus (Shabunda, DRC). Laurent (1983:352) listed the species (along with Phrynobatrachus petropedetoides and Hyperolius frontalis) from “shadowy puddles of the thick transition forest.” Etymology. The species epithet is the Latin adjective “ lacustris,” meaning belonging to or dwelling in lakes; in allusion to the distribution of the new species in the region of the African Great Lakes. Distribution and conservation. The species is distributed from lowland rainforest of the Congo Basin at 460 m (Omaniundu, DRC, Laurent 1982) to transitional forests of the Albertine Rift at 1650 m (Mwana, DRC) and east as far as Mabira Forest, Uganda (1121 m). As shown in Figure 5, the species has a relatively large distribution, including several national parks and protected areas (e.g., Virunga National Park, Bwindi Impenetrable National Park), and thus is not likely to be threatened based on a limited distribution. Using GeoCAT, its extent of occurrence is estimated as 409,238 km ² and its area of occupancy as 160 km ² (Bachman et al. 2011). We therefore classify this species as Least Concern according to the IUCN Red List criteria (IUCN 2021). Four specimens (RMCA 77-020-B-133–136) from Tubutubu, Itombwe Plateau (2300 m) are problematic because they are morphologically consistent with A. lacustris, but they were collected from a substantially higher elevation than all other known specimens. Coordinates for Tubutubu provided by RMCA (4° S, 28.933° E) suggest the locality is east of the highest point of the plateau at an elevation of about 1400 m on Google Earth. Although it is possible that the specimens were collected at the latter elevation, Laurent (1982) mentions bamboo forest as the habitat, which usually occurs at the highest elevations of the Itombwe Plateau (Doumenge 1998). No clarity is offered by Laurent (1964), because he does not mention any Afrixalus in his study of the ecology and distribution of amphibians of the Itombwe Plateau. Given this conflicting information, we do not include the Tubutubu specimens in our current understanding of the elevational distribution of A. lacustris (see gray triangle in Fig. 5)., Published as part of Greenbaum, Eli, Portik, Daniel M., Allen, Kaitlin E., Vaughan, Eugene R., Badjedjea, Gabriel, Barej, Michael F., Behangana, Mathias, Conkey, Nancy, Dumbo, Bonny, Gonwouo, Legrand N., Hirschfeld, Mareike, Hughes, Daniel F., Igunzi, Félix, Kusamba, Chifundera, Lukwago, Wilber, Masudi, Franck M., Penner, Johannes, Reyes, Jesús M., Rödel, Mark-Oliver, Roelke, Corey E., Romero, Soraya & Dehling, J. Maximilian, 2022, Systematics of the Central African Spiny Reed Frog Afrixalus laevis (Anura Hyperoliidae), with the description of two new species from the Albertine Rift, pp. 201-232 in Zootaxa 5174 (3) on pages 218-223, DOI: 10.11646/zootaxa.5174.3.1, http://zenodo.org/record/6986207, {"references":["Ahl, E. (1930) Ueber die afrikanischen Arten der Baumfroschgattung Megalixalus. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1930, 89 - 102.","Laurent, R. (1950) Genres Afrixalus et Hyperolius (Amphibia Salientia). Exploration du Parc National Albert, Mission G. F. De Witte (1933 - 1935), 64, 5 - 120, 7 pls.","Laurent, R. F. (1972) Amphibiens. Exploration du Parc National des Virunga, Deuxieme Serie, 22, 1 - 125, 11 pls.","Laurent, R. F. (1982) Le genre Afrixalus Laurent (Hyperoliidae) en Afrique Centrale. Annales du Musee Royal de l'Afrique Centrale, Serie in Octavo, Science Zoologique, 235, 1 - 93.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Channing, A. & Howell, K. M. (2006) Amphibians of East Africa. Cornell University Press, Ithaca, New York, New York, xi + 418 pp.","Spawls, S., Howell, K. M. & Drewes, R. C. (2006) Pocket Guide to the Reptiles and Amphibians of East Africa. A & C Black Publishers, London, 240 pp.","Channing, A. & Rodel, M. - O. (2019) Field Guide to the Frogs & Other Amphibians of Africa. Struik Nature, Cape Town, 408 pp.","Laurent, R. F. (1955) Apercu de la biogeographie des batraciens et des reptiles de la region des grand lacs. Bulletin de la Societe Zoologique de France, 79, 290 - 310.","Bell, R. C., Parra, J. L., Badjedjea, G., Barej, M. F., Blackburn, D. C., Burger, M., Channing, A., Dehling, J. M., Greenbaum, E., Gvozdik, V., Kielgast, J., Kusamba, C., Lotters, S., McLaughlin, P. J., Nagy, Z. T., Rodel, M. - O., Portik, D. M., Stuart, B. L., VanDerWal, J. & Zamudio, K. R. (2017) Idiosyncratic responses to climate-driven forest fragmentation and marine incursions in reed frogs from Central Africa and the Gulf of Guinea Islands. Molecular Ecology, 26 (19), 5223 - 5244. https: // doi. org / 10.1111 / mec. 14260","Laurent, R. F. (1983) About the herpetofauna of Central African montane forest. In: Rhodin, A. G. J. & Miyata, K. (Eds.), Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. Museum of Comparative Zoology, Cambridge, Massachusetts, pp. 350 - 358.","Bachman, S., Moat, J., Hill, A., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: Geospatial Conservation Assessment Tool. ZooKeys, 150, 117 - 126. https: // doi. org / 10.3897 / zookeys. 150.2109","IUCN (2021) IUCN Red List Categories and Criteria: Version 2021 - 3. Available from: https: // www. iucnredlist. org / (accessed 25 May 2022)","Doumenge, C. (1998) Forest diversity, distribution, and dynamique in the Itombwe Mountains, South-Kivu, Congo Democratic Republic. Mountain Research and Development, 18 (3), 249 - 264. https: // doi. org / 10.2307 / 3674036","Laurent, R. F. (1964) Adaptive modifications in frogs of an isolated highland fauna in Central Africa. Evolution, 18 (3), 458 - 467. https: // doi. org / 10.1111 / j. 1558 - 5646.1964. tb 01622. x"]}

Afrixalus lacustris Greenbaum, Dehling, Kusamba & Portik sp. nov. Great Lakes Spiny Reed Frog urn:lsid:zoobank.org:act: 41D3C41A-4501-4BEC-BFB6-B9C6C1518DC3 Afrixalus laevis (nec Megalixalus laevis Ahl 1930)— Laurent 1950: 24; Laurent 1972: 59 (partim); Laurent 1982: 33 (partim); Schiøtz 1999: 56 (partim); Channing & Howell 2006: 137 (partim); Spawls et al. 2006: 183 (partim); Channing & Rödel 2019: 158 (partim). Holotype. UTEP 20805 (field no. ELI 605), adult male, from the vicinity of Kalundu (03.15552° S, 28.42108° E, 1482 m), South Kivu Province, DRC, collected on 21 December 2010 by Chifundera Kusamba and Félix I. Alonda (Figs. 8D–F, 12A–B). Paratopotype. UTEP 20806 (field no. ELI 606), adult male, collected with the holotype. Referred specimens. Specimens with an asterisk were not included in morphological analyses due to poor quality of preservation, lack of a voucher, or unavailability of specimen(s): UTEP 20807–08 (field nos. ELI 644, 649), vicinity of Kalundu (03.15552° S, 28.42108° E, 1482 m), South Kivu Province, DRC; UTEP 20809, 22422 (field nos. ELI 669–70), Baraka, near shore of Lake Tanganyika (04.10832° S, 29.09705° E, 777 m), South Kivu Province, DRC; UTEP 20810–11 (field nos. EBG 1316, 1319), vicinity of Irangi (01.8873° S, 28.4495° E, 820 m), South Kivu Province, DRC; UTEP 22423 (field no. MUSE 10192), Kahuzi-Biega National Park, Nanwa (02.362138° S, 28.192589° E, 1566 m), South Kivu Province, DRC; UTEP 22424 (field no. MUSE 10137), Itombwe Plateau, Mbandakila (03.47937° S, 28.413049° E, 1135 m), South Kivu Province, DRC; RMCA 77-020 -B-133–136* (four specimens), Itombwe Plateau, Tubutubu (2300 m), South Kivu Province, DRC; RMCA 77-020 -B-89–92 (field nos. MF 14–15 [two specimens]), Kitutu (ca. 03.28° S, 28.11° E, 700 m), South Kivu Province, DRC; RMCA 77-020 - B-0069–71 (1 specimen), Itula (ca. 03.27° S, 28.12° E, 650 m), South Kivu Province, DRC; RMCA 77-020 -B-0081–83, 77-20-B-0086, Kamituga (ca. 03.54° S, 27.69° E, 1050 m), South Kivu Province, DRC; RMCA 77-020 - B-0072–73 (1 specimen), Mwana (ca. 03.15° S, 28.47° E, 1650 m), South Kivu Province, DRC; UTEP 22417 (field no. CRSN 2773), Toyokana (02.02734° N, 030.06653° E, 1294 m), Ituri Province, DRC; (field no. EPLU 395 *, photo and tissue only), ca. 0.5 km E of Epulu, near Mt. Mbiya (01.39594° N, 28.58093° E, 755 m), Ituri Province, DRC; UTEP 22416 (field no. DFH 247), Bwindi Impenetrable National Park, Buhoma (00.99045° S, 29.61884° E, 1523 m), Western Region, Uganda; (field nos. DFH 1102–03 * photos and tissues only), Kibale Forest National Park, Ngogo Research Center (00.49795° N, 30.42262° E, 1363 m), Western Region, Uganda; CAS 202032–36, Bwindi Impenetrable National Park, Buhoma rd., 2 km S (by rd.) of Bizenga River (01.00975° S, 29.620694° E), Western Region, Uganda; CAS 202133, Bwindi Impenetrable National Park, small tributary to Ishasha River (00.905° S, 29.704972° E, 1280 m), Western Region, Uganda; CAS 202109, Bwindi Impenetrable National Park, Bizenga River at Buhoma rd. (00.99275° S, 29.615722° E), Western Region, Uganda; CAS 256035 *, Bwindi Impenetrable National Park, rd. north of Ruhija, Western Region, Uganda; CAS 256128–31 *, Mabira Forest Reserve, marshy area adjacent to unpolluted stream (0.4508° N, 32.9493° E, 1121 m), Mukono District, Central Region, Uganda. Diagnosis. The species is referred to the genus Afrixalus for exhibiting the following characteristics: fingers and toes webbed; tips of fingers and toes enlarged to discs; eye large; pupil vertically elliptical; tympanum indistinct; vomer ridges and teeth absent; dorsal surfaces finely shagreened with minute pointed tubercles; outer metatarsal tubercle distinct; gular gland in males. It is readily distinguished from most other members of the genus by its small size (SVL in males 18.9–22.2 mm, in females 20.8–25.7 mm), being smaller than A. dorsalis (males 25–28 mm, females 26–30 mm), A. fornasini (males 30–38 mm, females 35–40 mm), A. fulvovittatus (males 23–27 mm, females 25–28 mm), A. lacteus (males 22–27 mm, females 25–29 mm), A. leucostictus (males 27–32 mm, females 25–36 mm), A. manengubensis (males to 32 mm), A. nigeriensis (males 28–34 mm, females 32–35 mm), A. osorioi (males 27–31, females 32–35 mm), A. paradorsalis (males 28–34 mm, females 32–35 mm), A. septentrionalis (males 19–21 mm), A. vittiger (males 22–25 mm, females 25–28 mm), and A. wittei (males 27–30 mm, females 29–33 mm); and larger size than A. delicatus (males 15–19 mm, females 17–22 mm) and A. stuhlmanni (males 15–21 mm, females 17–25 mm). Its differs in dorsal coloration and pattern from all species with longitudinal stripes or bands on a yellowish-brown background (A. brachycnemis, A. crotalus, A. delicatus, A. dorsalis, A. enseticola, A. fornasini, A. fulvovittatus, A. knysnae, A. morerei, A. orophilus, A. quadrivittatus, A. schneideri, A. spinifrons, A. stuhlmanni, A. upembae, A. septentrionalis, A. vittiger, and A. wittei); species showing a conspicuous large dark dorsal blotch on a uniform background on anterior dorsum in combination with large blotches on both sides of the hip (A. leucostictus, A. manengubensis, A. osorioi, A. paradorsalis, and A. schneideri) or with longitudinal lateral dark bands (A. equatorialis and A. nigeriensis); species with a uniformly colored dorsum without conspicuous pattern except a dark dorsolateral stripe (A. aureus, A. clarkei, A. delicatus, A. fornasini, A. lacteus, A. leucostictus [dorsum with small white tubercles], A. uluguruensis, and A. weidholzi [might have a black vertebral line and dark brown flanks]); and from all remaining species having either dark stripes running from tip of snout to back, crossing each other and continuing as dorsolateral stripes (A. vibekensis, also males without asperities); anterior part of dorsum yellowish with brown pattern, posterior part of dorsum translucent (A. laevis, also males without asperities); posterior part of dorsum and dorsal sides of limbs semi-translucent with dense small dark white-edged speckles (A. dorsimaculatus); and dark blotches and transverse marks with white speckles (A. sylvaticus). It is most similar to A. phantasma sp. nov. but differs from this species in being smaller (Table 1) and showing an overall darker coloration and more contrasting dorsal pattern that is obvious even in preserved specimens (Figs. 7–8, 12). It is readily distinguished from A. phantasma sp. nov. by the more extensive hand and toe webbing (Fig. 8E–F). Description of holotype. Measurements of the holotype are provided in Table 2. Body very slender, widest at temporal region, slightly tapering to groin (Fig. 8D–F); head small (HL/SVL 0.31, HW/SVL 0.31), about as long as wide (HW/HL 0.99); snout relatively long, rounded in dorsal view and in lateral profile, slightly wider than long; canthus rostralis hardly discernible, slightly concave between eye and nostril in dorsal view; loreal region oblique; nostrils rounded, directed anterolaterally and slightly dorsally; situated much closer to tip of snout and to eye, separated from each other by distance about equal to distance between eye and nostril (IN/EN 1.02); eye directed anterolaterally, strongly protruding, very large (ED/HL 0.39), its diameter shorter than snout (ED/SL 0.90); interorbital distance much wider than upper eyelid (IO/EW 1.61) and wider than internarial distance (IO/IN 1.31); tympanum covered by skin, not visible externally; upper jaw with dentition; teeth on premaxilla larger than teeth on maxilla; choanae small, located far anterolaterally at margins of roof of mouth, its anterior edge covered by maxillary bone, therefore appearing semicircular in ventral view; vomer ridges and teeth absent; tongue heartshaped, longer than broad, bilobed for about one-fourth of its length, free distally for about two-thirds its length; densely covered with minute papillae; median lingual process absent. Dorsal surfaces of head, trunk, and limbs and lateral surfaces of trunk finely shagreened; tiny pointed, widely scattered tubercles on dorsum, more numerous posteriorly than anteriorly; supratympanic fold absent; few tubercles at rear end of mandible without pointed tips; ventral side of head smooth; vocal sac present; gular gland of vocal sac smooth, large and wide, covering about 70 percent of throat width (Fig. 8E); chest smooth, abdomen weakly areolate; ventral side of limbs smooth; short transverse fold above vent. Forelimbs slender; hand large (HaL/SVL 0.28); tips of fingers enlarged into large disks, each with circummarginal groove; relative length of fingers: I I 2-/2 II 2-/3- III 2.25/2 IV; thenar tubercle oval, small and very low, about one-fourth length of metacarpal of Finger I; inner palmar tubercle and outer palmar tubercle very low, small, rounded, almost indiscernible. Hind limbs slender, moderately long; heel reaching to level of eye when legs adpressed forwardly to body; crus moderately long (TibL/SVL 0.44), slightly shorter than thigh; heels not touching each other when knees flexed and thighs held perpendicularly to median plane; foot subequal in length to crus (FoL/TibL 0.94); relative length of toes: I I 1.5/2+ II 1+/2+ III 1+/2+ IV 2/1+ V (Fig. 8E); inner metatarsal tubercle moderately prominent, elongated, about half length of metatarsus of Toe I; outer metatarsal tubercle small, rounded. Coloration in life. During night, skin on dorsal side of head, trunk, forelimbs, crus and tarsus light brown with more or less regular, locally more densely clustered dark brown speckling; distinct dark brown transverse stripe between upper eyelids, another less distinct one in scapula region, and another narrow one at level of anterior end of pelvis; dorsal skin of thigh largely unpigmented with only a broad stripe of widely scattered brown pigmentation; distinct dark brown band along canthus, continuing behind eye on both sides of trunk to level of pelvis; tubercles below eye and in tympanic region at rear end of jaw white; dark brown stripe running diagonally on middle of tibia, forming interrupted, inverted U-shaped band together with pelvic spots when legs folded against body; skin of ventral side of head, trunk and limbs largely unpigmented and translucent; gular gland bright yellow; fingers and toes yellow (Fig. 12). During day, basic dorsal coloration light brown; dark brown dorsal speckles, spots and stripes in more pronounced contrast to basic coloration, more distinctly visible. Coloration in preservative. Dorsal basic coloration largely faded to light brown; darker dorsal pattern elements dark brown, clearly visible; color of gular gland, fingers and toes faded to white (Fig. 8D–F). Variation. The paratypes match the holotype in general appearance and proportions. Coloration and color pattern varies from light-cream to yellowish-orange with a few relatively large and darker blotches and vermiculation, light speckling and indistinct dorsolateral stripe to an almost reticulated pattern of contrasting bright and dark elements with a broad and conspicuous brown dorsolateral band (Fig. 12). Pedal webbing variation is I 1.5[100]/2+[100] II 1+[100]/2+[100] III 1 +[50],1.25[50]/2+[100] IV 2 [100]/1+[100] V. Bioacoustics. The advertisement call and other vocalizations are unknown. Ecology and natural history. We collected males and females in swamps in forest openings and near forest edges (Fig. 11B), from vegetation at the edge of streams (or on vegetation overhanging them) in forest, and from non-forest vegetation near the shore of Lake Tanganyika. Egg deposition has not been observed and tadpoles are unknown. An adult male (CAS 256035) was found in Bwindi Impenetrable National Park (Uganda) ca. 2 m above ground in a tree fern “either calling or moving to a calling site (similar to ‘running’ of Kassina)” (D. Blackburn, in litt., 22 July 2021). Laurent (1955) noted the species from transitional forest. In his paper on amphibians of Virunga National Park, Laurent (1972) noted the species is most commonly found in swamps with herbaceous vegetation in secondary forest, and more rarely, in primary forest. Laurent (1982) noted that, in general, the species is common in the foliage of shrubs in dense forests. He also recorded it from “une mare sombre” (i.e., a dark pond) in syntopy with Chiromantis at Kitutu (DRC), marshes where it reproduces by sticking its eggs under the leaves of fountain grass (Pennisetum, Shabunda, DRC), a shaded bank of a pond in syntopy with Hyperolius “ tuberculatus ” (likely H. hutsebauti sensu Bell et al. 2017) and H. frontalis; the center of this pond had reeds harboring H. kivuensis and Afrixalus orophilus (Shabunda, DRC). Laurent (1983:352) listed the species (along with Phrynobatrachus petropedetoides and Hyperolius frontalis) from “shadowy puddles of the thick transition forest.” Etymology. The species epithet is the Latin adjective “ lacustris,” meaning belonging to or dwelling in lakes; in allusion to the distribution of the new species in the region of the African Great Lakes. Distribution and conservation. The species is distributed from lowland rainforest of the Congo Basin at 460 m (Omaniundu, DRC, Laurent 1982) to transitional forests of the Albertine Rift at 1650 m (Mwana, DRC) and east as far as Mabira Forest, Uganda (1121 m). As shown in Figure 5, the species has a relatively large distribution, including several national parks and protected areas (e.g., Virunga National Park, Bwindi Impenetrable National Park), and thus is not likely to be threatened based on a limited distribution. Using GeoCAT, its extent of occurrence is estimated as 409,238 km ² and its area of occupancy as 160 km ² (Bachman et al. 2011). We therefore classify this species as Least Concern according to the IUCN Red List criteria (IUCN 2021). Four specimens (RMCA 77-020-B-133–136) from Tubutubu, Itombwe Plateau (2300 m) are problematic because they are morphologically consistent with A. lacustris, but they were collected from a substantially higher elevation than all other known specimens. Coordinates for Tubutubu provided by RMCA (4° S, 28.933° E) suggest the locality is east of the highest point of the plateau at an elevation of about 1400 m on Google Earth. Although it is possible that the specimens were collected at the latter elevation, Laurent (1982) mentions bamboo forest as the habitat, which usually occurs at the highest elevations of the Itombwe Plateau (Doumenge 1998). No clarity is offered by Laurent (1964), because he does not mention any Afrixalus in his study of the ecology and distribution of amphibians of the Itombwe Plateau. Given this conflicting information, we do not include the Tubutubu specimens in our current understanding of the elevational distribution of A. lacustris (see gray triangle in Fig. 5).

Afrixalus phantasma Dehling, Greenbaum, Kusamba & Portik sp. nov. Ghost Spiny Reed Frog urn:lsid:zoobank.org:act: 1B2DBF26-5B0A-4DA8-B65E-0BC01704C1B4 Afrixalus laevis (nec Megalixalus laevis Ahl 1930)— Laurent 1972: 59 (partim); Laurent 1982: 33 (partim); Schiøtz 1999: 56 (partim); Channing & Howell 2006: 137 (partim); Spawls et al. 2006: 183 (partim); Channing & Rödel 2019: 158 (partim). Holotype. ZFMK 103454 (field no. JMD 723), adult male, from Gishwati Forest (01.823745° S, 29.360373° E, 2084 m), now part of Gishwati-Mukura National Park, Western Province, Rwanda, collected on 5 April 2011 by J. Maximilian Dehling and Bonny Dumbo (Figs. 7A, 8A–C). Paratypes. ZFMK 103455 (field no. JMD 722), adult female, collected with the holotype; ZFMK 103456 (field no. JMD 2015-30), adult male, ZFMK 103457 (field no. JMD 2015-31), adult female, from the type locality in Gishwati Forest, both collected on 27 September 2015 by J. Maximilian Dehling and Bonny Dumbo; ZFMK 103458–60 (field nos. JMD 677–679), three adult males, collected on 19 March 2011 by J. Maximilian Dehling, ZFMK 103461–62 (field nos. JMD 954–955), two adult males, collected on 16 February 2013 by J. Maximilian Dehling, all from Kamiranzovu Swamp (02.477165° S, 29.158243° E, 1962 m), Nyungwe National Park, Western Province, Rwanda; UTEP 20791–20792 (field nos. EBG 2838, 2843), two adult males, collected on 21 December 2009 by Chifundera Kusamba, Wandege M. Muninga, Mwenebatu M. Aristote, and Maurice Luhumyo from Nyakasanza Swamp near Tshibati (02.22886° S, 28.78017° E, 1979 m), South Kivu Province, DRC. Referred specimens. UTEP 20802 (field no. EBG 1198), forest ca. 4 km NW of Lwiro (02.2226° S, 28.7754° E, 2077 m), South Kivu Province, DRC; UTEP 20803, 22418–20 (field nos. EBG 1232, 1238–40), Kahuzi-Biega National Park, Mugaba (02.2750° S, 28.6631° E, 2298 m), South Kivu Province, DRC; UTEP 20804 (field no. EBG 1283), Kahuzi-Biega National Park, Chinya (02.2671° S, 28.6455° E, 2267 m), South Kivu Province, DRC; UTEP 20793–94 (field nos. EBG 2844–45), Nyakasanza Swamp near Tshibati (02.22886° S, 28.78017° E, 1979 m), South Kivu Province, DRC; UTEP 20795–20800, 22421 (field nos. ELI 414–20), Nyakasanza Swamp near Tshibati (02.22829° S, 28.77972° E, 1979 m), South Kivu Province, DRC; UTEP 20801 (field no. ELI 425), Chanjoka (02.21261° S, 28.77644° E, 2115 m), South Kivu Province, DRC. Diagnosis. The species is referred to the genus Afrixalus by exhibiting the following characteristics: fingers and toes webbed; tips of fingers and toes enlarged to discs; eye large; pupil vertically elliptical; tympanum indistinct; vomer ridges and teeth absent; dorsal surfaces finely shagreened with minute pointed tubercles; outer metatarsal tubercle distinct; gular gland in males. It is readily distinguished from most other members of the genus by its small size (SVL in males 20.1–24.6 mm, in females 22.6–26.4 mm), being smaller than A. dorsalis (males 25–28 mm, females 26–30 mm), A. fornasini (males 30–38 mm, females 35–40 mm), A. lacteus (males 22–27 mm, females 25– 29 mm), A. leucostictus (males 27–32 mm, females 25–36 mm), A. manengubensis (males to 32 mm), A. nigeriensis (males 28–34 mm, females 32–35 mm), A. osorioi (males 27–31 mm, females 32–35 mm), A. paradorsalis (males 28–34 mm, females 32–35 mm), A. septentrionalis (males 19–21 mm), and A. wittei (males 27–30 mm, females 29–33 mm); and larger than A. brachycnemis (males 18–21 mm, females 20–22 mm), A. delicatus (males 15–19 mm, females 17–22 mm), A. spinifrons (males to 20 mm, females to 25 mm), and A. stuhlmanni (males 15–21 mm, females 17–25 mm). Its differs in dorsal coloration and pattern from all species with longitudinal stripes or bands on a yellowish-brown background (A. brachycnemis, A. crotalus, A. delicatus, A. dorsalis, A. enseticola, A. fornasini, A. fulvovittatus, A. knysnae, A. morerei, A. orophilus, A. quadrivittatus, A. schneideri, A. spinifrons, A. stuhlmanni, A. upembae, A. septentrionalis, A. vittiger, and A. wittei); species with a conspicuous large dark dorsal blotch on a uniform background on anterior dorsum in combination with large blotches on both sides of the hip (A. leucostictus, A. manengubensis, A. osorioi, A. paradorsalis, and A. schneideri) or with longitudinal lateral dark bands (A. equatorialis and A. nigeriensis); species with a uniformly colored dorsum without conspicuous pattern except a dark dorsolateral stripe (A. aureus, A. clarkei, A. delicatus, A. fornasini, A. lacteus, A. leucostictus [dorsum with small white tubercles], A. uluguruensis, and A. weidholzi [might have a black vertebral line and dark brown flanks]); and from all remaining species having either dark stripes running from tip of snout to back, crossing each other and continuing as dorsolateral stripes (A. vibekensis, also males without asperities); anterior part of dorsum yellowish with brown pattern, posterior part of dorsum translucent (A. laevis, also males without asperities); posterior part of dorsum and dorsal sides of limbs semi-translucent with dense small dark white-edged speckles (A. dorsimaculatus); and dark blotches and transverse marks with white speckles (A. sylvaticus). The advertisement calls of the following species of Afrixalus consist of a long series of clicks, often initiated by a long buzzing note, and thus differ from the call of A. phantasma: A. aureus, A. brachycnemis, A. clarkei, A. crotalus, A. delicatus, A. dorsalis, A. fornasini, A. fulvovittatus, A. knysnae, A. nigeriensis, A. morerei, A. osorioi, A. quadrivittatus, A. septentrionalis, A. spinifrons, A. vibekensis, A. vittiger, A. weidholzi, and A. wittei. The advertisement calls of the remaining species for which information is available differ from the call of A. phantasma in the following characteristics (in parentheses): A. dorsimaculatus (soft, short buzzing rattle, usually repeated three times), A. equatorialis (series of five notes, repeated at 15–20/s and initiated by a long buzz, energy maximum at 2000–2500 Hz), A. lacteus (8–9 notes, 14–15 pulses,>300 pulses/s), A. paradorsalis (2–3 notes, energy maximum at 2700 Hz), A. stuhlmanni (2–9 notes, energy maximum at 4200–5100 Hz), and A. sylvaticus (2–5 notes, energy maximum at 4000–4500 Hz). The new species is most similar to A. lacustris sp. nov. from which it differs by a number of characteristics (see below). Description of holotype. Measurements of the holotype are provided in Table 2. Body very slender, widest at temporal region, slightly tapering to groin (Fig. 8A–C); head small (HL/SVL 0.31, HW/SVL 0.30), about as long as wide (HW/HL 0.98); snout relatively long, rounded in dorsal view and in lateral profile, slightly longer than wide; canthus rostralis hardly distinct, straight between eye and nostril; loreal region oblique; nostrils rounded, directed anterodorsally and slightly laterally; situated much closer to tip of snout and to eye, separated from each other by distance larger than distance between eye and nostril (IN/EN 1.15); eye directed anterolaterally, strongly protruding, very large (ED/HL 0.41), its diameter shorter than snout (ED/SL 0.84); interorbital distance much wider than upper eyelid and wider than internarial distance (IO/IN 1.21); tympanum covered by skin, not visible externally; upper jaw with dentition; teeth on premaxilla larger than teeth on maxilla; choanae small, located far anterolaterally at margins of roof of mouth, its anterior edge covered by maxillary bone, therefore appearing semicircular in ventral view; vomer ridges and teeth absent; tongue short and moderately broad, bilobed for about one-sixth of its length, free distally for about half its length; densely covered with minute papillae; median lingual process absent. Dorsal surfaces of head, trunk, and limbs and lateral surfaces of trunk finely shagreened; minute pointed, widely scattered tubercles on dorsum, more numerous posteriorly than anteriorly; supratympanic fold absent; small, subcircular, elevated glandular area in tympanic region posterior to eye and rear end of mandible, bearing large tubercles with pointed keratinous tips; ventral side of head smooth; vocal sac present; gular gland of vocal sac smooth, large and wide, covering about 70 percent of throat width (Fig. 8B); chest smooth, abdomen weakly areolate; ventral side of limbs smooth; short transverse fold above vent. Forelimbs slender;hand large (HND/SVL 0.32); tips of fingers enlarged into large disks, each with circummarginal groove; relative length of fingers: I I 2+/2.25 II 2/3+ III 2.75/2.25 IV; thenar tubercle oval, small and low, about one-fourth length of metacarpal of Finger I; palmar tubercles indiscernible. Hind limbs slender, moderately long; heel reaching to level of eye when legs adpressed forwardly to body; crus moderately long (TibL/SVL 0.48), about as long as thigh; heels meeting each other when knees flexed and thighs held perpendicularly to median plane; foot subequal in length to crus (FoL/TibL 0.96); relative length of toes: I I 2/2.25 II 1.5/3- III 1.75/3 IV 3-/1.75 V (Fig. 8C); inner metatarsal tubercle moderately prominent, elongated, about twothirds length of metatarsus of Toe I; outer metatarsal tubercle small, rounded. Coloration in life. During night, skin on dorsal side of head, trunk, forelimbs, crus and tarsus light yellowishbrown with more or less regular dark brown speckling; indistinct darker brown transverse stripe between upper eyelids; large dark brown, irregularly shaped spot in scapula region, two less distinct, somewhat lighter brown, irregularly shaped spots at level of anterior end of pelvis, separated from each other by half their width; cloacal region with dark brown mottling; dorsal skin of thigh largely unpigmented with only a broad stripe of widely scattered brown pigmentation; weak brown stripe along canthus, continuing behind eye on anterior half of trunk on both sides; tubercles below eye, and in tympanic region at rear end of jaw white; indistinct dark brown stripe running diagonally on middle of tibia, forming interrupted, inverted U-shaped band together with pelvic spots when legs folded against body; ventral side of head, trunk and limbs largely unpigmented and translucent; gular gland bright yellow; fingers and toes yellow (Fig. 7A). During day, basic dorsal coloration brighter, very light brown to bright cream-colored; dark brown dorsal speckles, spots and stripes in more pronounced contrast to basic coloration, more distinctly visible (Fig. 7B). Coloration in preservative. Dorsal basic coloration largely faded to yellowish white; darker dorsal pattern elements light to dark brown, clearly visible; yellow color of gular gland, fingers and toes faded to white (Fig. 8A–C). Variation. The paratypes match the holotype in general appearance, proportions, coloration and color pattern. Mensural variation within the species is shown in Table 1. Pedal webbing variation is I 2+[100]/2.5[90],2.25[10] II 2[50],2+[40],2.25[10]/3[10],3+[40],3.5[50] III 1.75 [80],2-[20]/3[100] IV 3 -[40],3[60]/1.75[100] V. Bioacoustics. Series of advertisement calls of six different males were recorded at the following ambient temperatures: 10.9°C (N = 1), 13.6°C (N = 3), and 16.2°C (N = 2). The advertisement call consisted of five to six, rarely four pulse groups (notes) (Fig. 9). Depending on the ambient temperature, notes were repeated at a rate of 7.1–10.0/s (10.9°C), 8.2–11.7/s (13.6°C), and 10.2–11.4/s (16.2°C). The highest repetition rate was always between the first three notes of a series, the lowest at the end of the series. Each note consisted of 10–11 pulses (Fig. 9). Probably due to echo effects, pulsation was often veiled in the waveforms, especially towards the end of the note. Pulse repetition rate varied from 190–210/s at 10.9°C, 222–250/s at 13.6°C, and 277–292/s at 16.2°C, resulting in a note length of 72–94 ms, 55–70 ms, and 36–50 ms, respectively. Amplitude modulation was prominent within individual notes (Fig. 9). Total call length showed linear temperature dependence and varied from 388–397 ms at 16.2°C, 472–499 ms at 13.6°C, and 572–620 ms at 10.9°C for five-note calls (Fig. 10). Energy maximum showed linear temperature dependence, with 3020–3150 Hz at 10.9°C, 3370–3550 Hz at 13.6°C, and 3660–3810 Hz at 16.2°C (Figs. 9–10). There was no marked frequency modulation. Prominent harmonics were at about 6000–7000 Hz and 9000–11000 Hz (Fig. 9). Ecology and natural history. We collected males and females in swamps in forest openings, or at forest edges (Fig. 11A). Males were found calling above standing bodies of water with thick lower vegetation cover. In Gishwati Forest and Kamiranzovu Swamp in Nyungwe Forest, the species was found calling in syntopy with Hyperolius castaneus and H. discodactylus. One adult male (UTEP 20802) from Kahuzi-Biega National Park (DRC) was found 3.5 meters above ground in montane forest. Tadpoles are unknown. Laurent (1955, 1983) listed the species from bushes in montane forest. Laurent (1982) noted that, in general, the species is common in the foliage of shrubs in dense forests. He also recorded it in banana trees in a deep valley (Nyungwe, Rwanda) and “des trous de prospection” (i.e., prospecting holes likely for gold mining, Upper Lubitshako, Kabobo Plateau, DRC). Etymology. The species epithet derives from the Greek noun φάντασμα (phántasma), meaning ghost or phantom, in allusion to the coloration and general appearance of the new species. The epithet is used as an invariable noun in apposition. Distribution and conservation. The occurrence of the species has been confirmed for several locations in western Rwanda (Nyungwe and Gishwati Forest) and eastern DRC (in and near Kahuzi-Biega National Park, Itombwe Plateau and Kabobo Plateau [Laurent 1982], Fig. 5). We expect the species to be eventually found in the AR of southwestern Uganda and northwestern Burundi. So far, the species has been recorded from only a narrow elevation range between 1962 m (Kamiranzovu Swamp, Rwanda) and ca. 2400 m (May ya Moto [and possibly Luemba], Itombwe Plateau, Laurent 1982). Given the relatively limited overall geographic distribution of the species with an estimated extent of occurrence of 19,088 km 2 (Fig. 5), an estimated area of occupancy of 52 km 2 (both calculated with GeoCAT; Bachman et al. 2011), the detection of the amphibian chytrid fungus in one adult male (UTEP 20791, Greenbaum et al. 2015), and the conservation challenges facing natural areas of the AR (Greenbaum 2017; Ayebare et al. 2018), we categorize this species as Vulnerable under the IUCN Red List criteria (IUCN 2021).

Afrixalus phantasma Dehling, Greenbaum, Kusamba & Portik sp. nov. Ghost Spiny Reed Frog urn:lsid:zoobank.org:act: 1B2DBF26-5B0A-4DA8-B65E-0BC01704C1B4 Afrixalus laevis (nec Megalixalus laevis Ahl 1930)— Laurent 1972: 59 (partim); Laurent 1982: 33 (partim); Schiøtz 1999: 56 (partim); Channing & Howell 2006: 137 (partim); Spawls et al. 2006: 183 (partim); Channing & Rödel 2019: 158 (partim). Holotype. ZFMK 103454 (field no. JMD 723), adult male, from Gishwati Forest (01.823745° S, 29.360373° E, 2084 m), now part of Gishwati-Mukura National Park, Western Province, Rwanda, collected on 5 April 2011 by J. Maximilian Dehling and Bonny Dumbo (Figs. 7A, 8A–C). Paratypes. ZFMK 103455 (field no. JMD 722), adult female, collected with the holotype; ZFMK 103456 (field no. JMD 2015-30), adult male, ZFMK 103457 (field no. JMD 2015-31), adult female, from the type locality in Gishwati Forest, both collected on 27 September 2015 by J. Maximilian Dehling and Bonny Dumbo; ZFMK 103458–60 (field nos. JMD 677–679), three adult males, collected on 19 March 2011 by J. Maximilian Dehling, ZFMK 103461–62 (field nos. JMD 954–955), two adult males, collected on 16 February 2013 by J. Maximilian Dehling, all from Kamiranzovu Swamp (02.477165° S, 29.158243° E, 1962 m), Nyungwe National Park, Western Province, Rwanda; UTEP 20791–20792 (field nos. EBG 2838, 2843), two adult males, collected on 21 December 2009 by Chifundera Kusamba, Wandege M. Muninga, Mwenebatu M. Aristote, and Maurice Luhumyo from Nyakasanza Swamp near Tshibati (02.22886° S, 28.78017° E, 1979 m), South Kivu Province, DRC. Referred specimens. UTEP 20802 (field no. EBG 1198), forest ca. 4 km NW of Lwiro (02.2226° S, 28.7754° E, 2077 m), South Kivu Province, DRC; UTEP 20803, 22418–20 (field nos. EBG 1232, 1238–40), Kahuzi-Biega National Park, Mugaba (02.2750° S, 28.6631° E, 2298 m), South Kivu Province, DRC; UTEP 20804 (field no. EBG 1283), Kahuzi-Biega National Park, Chinya (02.2671° S, 28.6455° E, 2267 m), South Kivu Province, DRC; UTEP 20793–94 (field nos. EBG 2844–45), Nyakasanza Swamp near Tshibati (02.22886° S, 28.78017° E, 1979 m), South Kivu Province, DRC; UTEP 20795–20800, 22421 (field nos. ELI 414–20), Nyakasanza Swamp near Tshibati (02.22829° S, 28.77972° E, 1979 m), South Kivu Province, DRC; UTEP 20801 (field no. ELI 425), Chanjoka (02.21261° S, 28.77644° E, 2115 m), South Kivu Province, DRC. Diagnosis. The species is referred to the genus Afrixalus by exhibiting the following characteristics: fingers and toes webbed; tips of fingers and toes enlarged to discs; eye large; pupil vertically elliptical; tympanum indistinct; vomer ridges and teeth absent; dorsal surfaces finely shagreened with minute pointed tubercles; outer metatarsal tubercle distinct; gular gland in males. It is readily distinguished from most other members of the genus by its small size (SVL in males 20.1–24.6 mm, in females 22.6–26.4 mm), being smaller than A. dorsalis (males 25–28 mm, females 26–30 mm), A. fornasini (males 30–38 mm, females 35–40 mm), A. lacteus (males 22–27 mm, females 25– 29 mm), A. leucostictus (males 27–32 mm, females 25–36 mm), A. manengubensis (males to 32 mm), A. nigeriensis (males 28–34 mm, females 32–35 mm), A. osorioi (males 27–31 mm, females 32–35 mm), A. paradorsalis (males 28–34 mm, females 32–35 mm), A. septentrionalis (males 19–21 mm), and A. wittei (males 27–30 mm, females 29–33 mm); and larger than A. brachycnemis (males 18–21 mm, females 20–22 mm), A. delicatus (males 15–19 mm, females 17–22 mm), A. spinifrons (males to 20 mm, females to 25 mm), and A. stuhlmanni (males 15–21 mm, females 17–25 mm). Its differs in dorsal coloration and pattern from all species with longitudinal stripes or bands on a yellowish-brown background (A. brachycnemis, A. crotalus, A. delicatus, A. dorsalis, A. enseticola, A. fornasini, A. fulvovittatus, A. knysnae, A. morerei, A. orophilus, A. quadrivittatus, A. schneideri, A. spinifrons, A. stuhlmanni, A. upembae, A. septentrionalis, A. vittiger, and A. wittei); species with a conspicuous large dark dorsal blotch on a uniform background on anterior dorsum in combination with large blotches on both sides of the hip (A. leucostictus, A. manengubensis, A. osorioi, A. paradorsalis, and A. schneideri) or with longitudinal lateral dark bands (A. equatorialis and A. nigeriensis); species with a uniformly colored dorsum without conspicuous pattern except a dark dorsolateral stripe (A. aureus, A. clarkei, A. delicatus, A. fornasini, A. lacteus, A. leucostictus [dorsum with small white tubercles], A. uluguruensis, and A. weidholzi [might have a black vertebral line and dark brown flanks]); and from all remaining species having either dark stripes running from tip of snout to back, crossing each other and continuing as dorsolateral stripes (A. vibekensis, also males without asperities); anterior part of dorsum yellowish with brown pattern, posterior part of dorsum translucent (A. laevis, also males without asperities); posterior part of dorsum and dorsal sides of limbs semi-translucent with dense small dark white-edged speckles (A. dorsimaculatus); and dark blotches and transverse marks with white speckles (A. sylvaticus). The advertisement calls of the following species of Afrixalus consist of a long series of clicks, often initiated by a long buzzing note, and thus differ from the call of A. phantasma: A. aureus, A. brachycnemis, A. clarkei, A. crotalus, A. delicatus, A. dorsalis, A. fornasini, A. fulvovittatus, A. knysnae, A. nigeriensis, A. morerei, A. osorioi, A. quadrivittatus, A. septentrionalis, A. spinifrons, A. vibekensis, A. vittiger, A. weidholzi, and A. wittei. The advertisement calls of the remaining species for which information is available differ from the call of A. phantasma in the following characteristics (in parentheses): A. dorsimaculatus (soft, short buzzing rattle, usually repeated three times), A. equatorialis (series of five notes, repeated at 15–20/s and initiated by a long buzz, energy maximum at 2000–2500 Hz), A. lacteus (8–9 notes, 14–15 pulses,>300 pulses/s), A. paradorsalis (2–3 notes, energy maximum at 2700 Hz), A. stuhlmanni (2–9 notes, energy maximum at 4200–5100 Hz), and A. sylvaticus (2–5 notes, energy maximum at 4000–4500 Hz). The new species is most similar to A. lacustris sp. nov. from which it differs by a number of characteristics (see below). Description of holotype. Measurements of the holotype are provided in Table 2. Body very slender, widest at temporal region, slightly tapering to groin (Fig. 8A–C); head small (HL/SVL 0.31, HW/SVL 0.30), about as long as wide (HW/HL 0.98); snout relatively long, rounded in dorsal view and in lateral profile, slightly longer than wide; canthus rostralis hardly distinct, straight between eye and nostril; loreal region oblique; nostrils rounded, directed anterodorsally and slightly laterally; situated much closer to tip of snout and to eye, separated from each other by distance larger than distance between eye and nostril (IN/EN 1.15); eye directed anterolaterally, strongly protruding, very large (ED/HL 0.41), its diameter shorter than snout (ED/SL 0.84); interorbital distance much wider than upper eyelid and wider than internarial distance (IO/IN 1.21); tympanum covered by skin, not visible externally; upper jaw with dentition; teeth on premaxilla larger than teeth on maxilla; choanae small, located far anterolaterally at margins of roof of mouth, its anterior edge covered by maxillary bone, therefore appearing semicircular in ventral view; vomer ridges and teeth absent; tongue short and moderately broad, bilobed for about one-sixth of its length, free distally for about half its length; densely covered with minute papillae; median lingual process absent. Dorsal surfaces of head, trunk, and limbs and lateral surfaces of trunk finely shagreened; minute pointed, widely scattered tubercles on dorsum, more numerous posteriorly than anteriorly; supratympanic fold absent; small, subcircular, elevated glandular area in tympanic region posterior to eye and rear end of mandible, bearing large tubercles with pointed keratinous tips; ventral side of head smooth; vocal sac present; gular gland of vocal sac smooth, large and wide, covering about 70 percent of throat width (Fig. 8B); chest smooth, abdomen weakly areolate; ventral side of limbs smooth; short transverse fold above vent. Forelimbs slender;hand large (HND/SVL 0.32); tips of fingers enlarged into large disks, each with circummarginal groove; relative length of fingers: I I 2+/2.25 II 2/3+ III 2.75/2.25 IV; thenar tubercle oval, small and low, about one-fourth length of metacarpal of Finger I; palmar tubercles indiscernible. Hind limbs slender, moderately long; heel reaching to level of eye when legs adpressed forwardly to body; crus moderately long (TibL/SVL 0.48), about as long as thigh; heels meeting each other when knees flexed and thighs held perpendicularly to median plane; foot subequal in length to crus (FoL/TibL 0.96); relative length of toes: I I 2/2.25 II 1.5/3- III 1.75/3 IV 3-/1.75 V (Fig. 8C); inner metatarsal tubercle moderately prominent, elongated, about twothirds length of metatarsus of Toe I; outer metatarsal tubercle small, rounded. Coloration in life. During night, skin on dorsal side of head, trunk, forelimbs, crus and tarsus light yellowishbrown with more or less regular dark brown speckling; indistinct darker brown transverse stripe between upper eyelids; large dark brown, irregularly shaped spot in scapula region, two less distinct, somewhat lighter brown, irregularly shaped spots at level of anterior end of pelvis, separated from each other by half their width; cloacal region with dark brown mottling; dorsal skin of thigh largely unpigmented with only a broad stripe of widely scattered brown pigmentation; weak brown stripe along canthus, continuing behind eye on anterior half of trunk on both sides; tubercles below eye, and in tympanic region at rear end of jaw white; indistinct dark brown stripe running diagonally on middle of tibia, forming interrupted, inverted U-shaped band together with pelvic spots when legs folded against body; ventral side of head, trunk and limbs largely unpigmented and translucent; gular gland bright yellow; fingers and toes yellow (Fig. 7A). During day, basic dorsal coloration brighter, very light brown to bright cream-colored; dark brown dorsal speckles, spots and stripes in more pronounced contrast to basic coloration, more distinctly visible (Fig. 7B). Coloration in preservative. Dorsal basic coloration largely faded to yellowish white; darker dorsal pattern elements light to dark brown, clearly visible; yellow color of gular gland, fingers and toes faded to white (Fig. 8A–C). Variation. The paratypes match the holotype in general appearance, proportions, coloration and color pattern. Mensural variation within the species is shown in Table 1. Pedal webbing variation is I 2+[100]/2.5[90],2.25[10] II 2[50],2+[40],2.25[10]/3[10],3+[40],3.5[50] III 1.75 [80],2-[20]/3[100] IV 3 -[40],3[60]/1.75[100] V. Bioacoustics. Series of advertisement calls of six different males were recorded at the following ambient temperatures: 10.9°C (N = 1), 13.6°C (N = 3), and 16.2°C (N = 2). The advertisement call consisted of five to six, rarely four pulse groups (notes) (Fig. 9). Depending on the ambient temperature, notes were repeated at a rate of 7.1–10.0/s (10.9°C), 8.2–11.7/s (13.6°C), and 10.2–11.4/s (16.2°C). The highest repetition rate was always between the first three notes of a series, the lowest at the end of the series. Each note consisted of 10–11 pulses (Fig. 9). Probably due to echo effects, pulsation was often veiled in the waveforms, especially towards the end of the note. Pulse repetition rate varied from 190–210/s at 10.9°C, 222–250/s at 13.6°C, and 277–292/s at 16.2°C, resulting in a note length of 72–94 ms, 55–70 ms, and 36–50 ms, respectively. Amplitude modulation was prominent within individual notes (Fig. 9). Total call length showed linear temperature dependence and varied from 388–397 ms at 16.2°C, 472–499 ms at 13.6°C, and 572–620 ms at 10.9°C for five-note calls (Fig. 10). Energy maximum showed linear temperature dependence, with 3020–3150 Hz at 10.9°C, 3370–3550 Hz at 13.6°C, and 3660–3810 Hz at 16.2°C (Figs. 9–10). There was no marked frequency modulation. Prominent harmonics were at about 6000–7000 Hz and 9000–11000 Hz (Fig. 9). Ecology and natural history. We collected males and females in swamps in forest openings, or at forest edges (Fig. 11A). Males were found calling above standing bodies of water with thick lower vegetation cover. In Gishwati Forest and Kamiranzovu Swamp in Nyungwe Forest, the species was found calling in syntopy with Hyperolius castaneus and H. discodactylus. One adult male (UTEP 20802) from Kahuzi-Biega National Park (DRC) was found 3.5 meters above ground in montane forest. Tadpoles are unknown. Laurent (1955, 1983) listed the species from bushes in montane forest. Laurent (1982) noted that, in general, the species is common in the foliage of shrubs in dense forests. He also recorded it in banana trees in a deep valley (Nyungwe, Rwanda) and “des trous de prospection” (i.e., prospecting holes likely for gold mining, Upper Lubitshako, Kabobo Plateau, DRC). Etymology. The species epithet derives from the Greek noun φάντασμα (phántasma), meaning ghost or phantom, in allusion to the coloration and general appearance of the new species. The epithet is used as an invariable noun in apposition. Distribution and conservation. The occurrence of the species has been confirmed for several locations in western Rwanda (Nyungwe and Gishwati Forest) and eastern DRC (in and near Kahuzi-Biega National Park, Itombwe Plateau and Kabobo Plateau [Laurent 1982], Fig. 5). We expect the species to be eventually found in the AR of southwestern Uganda and northwestern Burundi. So far, the species has been recorded from only a narrow elevation range between 1962 m (Kamiranzovu Swamp, Rwanda) and ca. 2400 m (May ya Moto [and possibly Luemba], Itombwe Plateau, Laurent 1982). Given the relatively limited overall geographic distribution of the species with an estimated extent of occurrence of 19,088 km 2 (Fig. 5), an estimated area of occupancy of 52 km 2 (both calculated with GeoCAT; Bachman et al. 2011), the detection of the amphibian chytrid fungus in one adult male (UTEP 20791, Greenbaum et al. 2015), and the conservation challenges facing natural areas of the AR (Greenbaum 2017; Ayebare et al. 2018), we categorize this species as Vulnerable under the IUCN Red List criteria (IUCN 2021)., Published as part of Greenbaum, Eli, Portik, Daniel M., Allen, Kaitlin E., Vaughan, Eugene R., Badjedjea, Gabriel, Barej, Michael F., Behangana, Mathias, Conkey, Nancy, Dumbo, Bonny, Gonwouo, Legrand N., Hirschfeld, Mareike, Hughes, Daniel F., Igunzi, Félix, Kusamba, Chifundera, Lukwago, Wilber, Masudi, Franck M., Penner, Johannes, Reyes, Jesús M., Rödel, Mark-Oliver, Roelke, Corey E., Romero, Soraya & Dehling, J. Maximilian, 2022, Systematics of the Central African Spiny Reed Frog Afrixalus laevis (Anura Hyperoliidae), with the description of two new species from the Albertine Rift, pp. 201-232 in Zootaxa 5174 (3) on pages 213-218, DOI: 10.11646/zootaxa.5174.3.1, http://zenodo.org/record/6986207, {"references":["Ahl, E. (1930) Ueber die afrikanischen Arten der Baumfroschgattung Megalixalus. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1930, 89 - 102.","Laurent, R. F. (1972) Amphibiens. Exploration du Parc National des Virunga, Deuxieme Serie, 22, 1 - 125, 11 pls.","Laurent, R. F. (1982) Le genre Afrixalus Laurent (Hyperoliidae) en Afrique Centrale. Annales du Musee Royal de l'Afrique Centrale, Serie in Octavo, Science Zoologique, 235, 1 - 93.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Channing, A. & Howell, K. M. (2006) Amphibians of East Africa. Cornell University Press, Ithaca, New York, New York, xi + 418 pp.","Spawls, S., Howell, K. M. & Drewes, R. C. (2006) Pocket Guide to the Reptiles and Amphibians of East Africa. A & C Black Publishers, London, 240 pp.","Channing, A. & Rodel, M. - O. (2019) Field Guide to the Frogs & Other Amphibians of Africa. Struik Nature, Cape Town, 408 pp.","Laurent, R. F. (1955) Apercu de la biogeographie des batraciens et des reptiles de la region des grand lacs. Bulletin de la Societe Zoologique de France, 79, 290 - 310.","Laurent, R. F. (1983) About the herpetofauna of Central African montane forest. In: Rhodin, A. G. J. & Miyata, K. (Eds.), Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. Museum of Comparative Zoology, Cambridge, Massachusetts, pp. 350 - 358.","Bachman, S., Moat, J., Hill, A., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: Geospatial Conservation Assessment Tool. ZooKeys, 150, 117 - 126. https: // doi. org / 10.3897 / zookeys. 150.2109","Greenbaum, E., Meece, J., Reed, K. & Kusamba, C. (2015) Extensive occurrence of the amphibian chytrid fungus in the Albertine Rift, a Central African amphibian hotspot. The Herpetological Journal, 25 (2), 91 - 100.","Greenbaum, E. (2017) Emerald Labyrinth: A Scientist's Adventures in the Jungles of the Congo. ForeEdge, Lebanon, New Hampshire, xv + 329 pp. https: // doi. org / 10.2307 / j. ctv 1 xx 9 krm","Ayebare, S., Plumptre, A. J., Kujirakwinja, D. & Segan, D. (2018) Conservation of the endemic species of the Albertine Rift under future climate change. Biological Conservation, 220, 67 - 75. https: // doi. org / 10.1016 / j. biocon. 2018.02.001","IUCN (2021) IUCN Red List Categories and Criteria: Version 2021 - 3. Available from: https: // www. iucnredlist. org / (accessed 25 May 2022)"]}

Herein, we describe the tadpoles of six Astylosternus species, A.fallax, A.cf.fallax, A.laurenti, A.montanus, A.perreti, A.ranoides, and Scotoblepsgabonicus, and redescribe the tadpoles of A.batesi, A.diadematus, A.laticephalus, A.occidentalis, A.rheophilus, and Nyctibatescorrugatus. All Astylosternus tadpoles are adapted to torrent currents and share a long, oval body, slightly flattened in lateral view, with very long muscular tails with narrow fins. The jaws are massive, serrated, and often show a tooth-like medial projection (fang) in the upper jaw. Body proportions of Astylosternus tadpoles are extremely similar. The best characters to distinguish species might be life coloration and potentially the shape of labial papillae. The tadpole of Scotoblepsgabonicus is similar to Astylosternus and differs only slightly by a narrower body with a shorter and rounder head. The upper jaw of Scotobleps carries two or three lateral fangs instead of one medial one. The tadpole of Nyctibatescorrugatus is easily distinguishable from the other two genera on the basis of their very long, eel-shaped body and tail and the bluish-black color.

Kpan, Tokouaho Flora, Kouamé, N'Goran Germain, Barej, Michael F., Adeba, Patrick Joël, Emmrich, Mike, Ofori-Boateng, Caleb, Rödel, Mark-Oliver (2018): A new Puddle Frog, genus Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae), from the eastern part of the Upper Guinea biodiversity hotspot, West Africa. Zootaxa 4388 (2): 221-237, DOI: https://doi.org/10.11646/zootaxa.4388.2.5

Phrynobatrachus tanoeensis sp. nov. Figs 2–3 Holotype. ZMB 80870 (field and tissue #: YA 14-03, GenBank #: MG209126), adult male, Ivory Coast, Tanoé-Ehy Swamp Forest, 05°11’181’’N, 02°53’011’’W (date WGS84), swamp forest dominated by Raphia palms, 17–29 July 2010, coll. T.F. Kpan & P.J. Adeba. Paratypes. ZMB 80871 (YA14-01, MG209125), adult male, same data as holotype; ZMB 86953 –86955 (KK 1–3), three adult males, all with damages of dorsal skin, Ivory Coast, Dohouan village forest, Nouamou, 05°11.827'N, 02°53.140'W, 17 m asl, 1–3 October 2014, coll. T.F. Kpan & N.G. Kouamé; ZMB 86956–86960 (KK 4–8; ZMB 86959= MG209122), five adult males, ZMB 86961–86963 (KK 9–11; ZMB 8961= MG209123, ZMB 86963= MK209124), three females, 05°11.718'N, 02°52.854'W, 20 m asl, all other data as in ZMB 86953–86955. Additional material. ZMB 86964 (KK 12), juvenile, 05°11.718'N, 02°52.854'W, 20 m asl, all other data as in ZMB 86953–86955; ZMB 86965 (GAW009), adult male, skin damaged or missing on left flank and right thigh, Ghana, Western Region, Ebonloa, near Essiama, 05°02'0.28’’N, 02°33'05.9W, 16 m asl, peat/swamp forest with slow moving stream near the borders of Ghana and Ivory Coast, coll. C. Ofori-Boateng. Diagnosis. The overall body shape and morphology, i.e. the presence of a tarsal tubercle, as well as the genetic data confirm that the new species is a member of the genus Phrynobatrachus (Phrynobatrachidae). From members of the externally-morphologically similar genus Arthroleptis (Arthroleptidae), the new species differs by: the presence of scapular ridges, pedal webbing and a tarsal tubercle, and the absence of a mid-dorsal skin raphe. From members of the genus Ptychadena (Ptychadenidae; juveniles sometimes morphologically similar to Phrynobatrachus) they differ by a more rounded and shorter snout and the absence of parallel dorsal glandular ridges. The new species can be differentiated from other members of the genus Phrynobatrachus by molecular and acoustic characters (see below), as well as by the combination of the following morphological characters: adult snout–vent length larger than 20 mm but smaller than 30 mm; a dark face mask; absence of a spiny tubercle on the eyelid; narrow and partly indistinct scapular ridges or comma-shaped warts, in particular no long X-shaped pair of dorsal ridges; distinct spinulae on males’ throats and dorsal surfaces; pronounced pedal webbing; finger- and toe tips not enlarged; a black throat in adult males; a white belly with small blackish spotting in both sexes; more than one wide dark cross bar on hind legs; ventral parts of hind legs in both sexes with rosé to reddish colour. In particular the new species is smaller (P. liberiensis and P. plicatus (Günther, 1858); it differs from P. annulatus Perret, 1966, P. calcaratus (Peters, 1863), P. taiensis Perret, 1988, P. villiersi Guibé, 1959 and P. pintoi Hillers, Zimkus & Rödel, 2008 by lacking an eyelid cornicle (spine-like wart on upper eyelid); from all the latter species and P. gutturosus (Chabanaud, 1921), P. hieroglyphicus Rödel, Ohler & Hillers, 2010, and P. tokba (Chabanaud, 1921), the new species differs by more extensive webbing; Phrynobatrachus annulatus, P. fraterculus (Chabanaud, 1921), P. ghanensis, P. maculiventris Guibé & Lamotte, 1958, P. pintoi, P. taiensis and P. villiersi all exhibit very distinct, species-specific patterning on the belly, different to the new species which exhibits many small black spots. In contrast P. latifrons, Ahl, 1924, P. alleni Parker, 1936 (only females, males with yellow belly), P. francisci Boulenger, 1912, P. natalensis (Smith, 1849), and P. rainerguentheri Rödel, Onadeko, Barej & Sandberger, 2012, have predominantly white bellies with at best a few minute black points. Whereas males of the new species have dark grey to black throats, P. alleni and P. latifrons males have yellow throats. These two species, along with all species with an eyelid cornicle, as well as P. francisci, P. gutturosus, P. phyllophilus Rödel & Ernst, 2002, P. guineensis Guibé & Lamotte, 1962, P. natalensis, P. rainerguentheri, P. brongersmai Parker, 1936 have a different body shape and a snout which is rounded in dorsal and lateral view (pointed in lateral view in the new species). In P. alleni the head is also relatively flatter in lateral view. Phrynobatrachus guineensis is smaller (Phrynobatrachus phyllophilus shows some black spots on belly and pectoral region but has a more compact body shape and roundish snout, toe and finger tips which are enlarged to small round discs (straight in the new species) and usually only one wide black cross bar on thigh and lower leg (several in the new species). In West Africa, only P. alleni and P. plicatus have scapular ridges formed like an open X, in P. plicatus this X extends to posterior to half-back. The new species shares a black face mask in West Africa only with P. intermedius, P. plicatus and some P. liberiensis and P. rainerguentheri. However, the face mask is always less distinct than in P. plicatus (see discussion for a more in depth comparison with the genetically most similar species). Description of the holotype (measurements in millimetres mm). Adult male with longish, but compact body shape, short snout and large eyes; snout–vent length 22.1; snout rounded in dorsal, moderately pointed in lateral view; head width 6.9; thigh length 8.5, shorter than crus length, 12.0; foot length including longest toe 15.4; eye diameter 2.8, larger than horizontal diameter of tympanum, 1.6; tympanum distinct, round with round median annuli, diameter 0.5; interorbital distance 2.5; distance eye–nostrils 1.9; distance eye–snout 3.0; nostrils much closer to snout than to eye; canthus rostralis rounded; loreal region straight; distinct, slightly bent supratympanal ridge extending from posterior edge of eye to forearm insertion; dorsal skin fine granular, spinulae on posterior part of back indistinct; spinulae on dorsal surfaces of thighs and lower legs; two pairs of narrow, indistinct, commashaped warts; the anterior pair originating slightly posterior to eyelids, slightly converging and extending to about forearm insertion; the posterior pair being shorter and slightly diverging; a few minute round tubercles on eyelids; throat smooth with parallel skin folds along mandibles; whitish spinulae on throat and anterior part of breast indistinct; skin of other ventral surfaces smooth; palmar surface of hand with large, oval palmar and more slender, longish thenar tubercle; fingers with large, undivided oval subarticular tubercles, additional tubercles on hands absent; finger tips roundish, not enlarged to discs; outer part of thumb with indistinct thin nuptial pad; palmar webbing absent; relative finger length: II = IV ý I Colour of the holotype in life. Dorsal surfaces light beige to yellowish brown; top of snout with Y-shaped dark figure – the single arm pointing towards snout, and further dark spots; margin of snout and upper eyelids with ill-defined light band; upper eyelids dark brown, connected by dark brown inter-orbital band, this band bordered anteriorly and posteriorly by narrow clear bands; the two pairs of narrow comma shaped scapular ridges marked darker, the posterior pair connected by V-shaped black marking; a pair of white spots at the tips of this V; posterior back with a few further blackish markings; dorsal surfaces of thighs (4) and lower legs (3) with black cross bars; arms, hands and feet with less distinct black cross bars; lateral side of snout, upper lip and area below supratympanal ridge black with some minute white spots on lip; supratympanal ridge dorsally bordered by white band; posterior to this band a wide black lateral band, followed by further black spots on whitish base colour; lower mandible, throat and anterior pectoral region almost uniform dark grey; belly white, densely beset with black spots; posterior belly and lower surfaces of thighs and lower legs rosé. Colour of the holotype in preservation (after seven years in 75% ethanol). Back light brown, a few white spots where skin is rubbed off; snout tip and upper eyelids with some black mottling; loreal triangle from nostrils to anterior eye and tympanal area darker; scapular ridges and supratympanal ridge darker; black and light lateral pattern faded; upper arm with two dark cross bars, lower side reddish brown; lower arm with one dark cross bar close to hand; palmar surfaces of hands uniform light reddish brown; lower side of mandibles light blackish grey; throat skin dark blackish grey with minute white points; belly and pectoral region whitish with dark brown spotting, denser towards throat and flanks; vent surrounded by flat black triangle, bordered by narrow light line; posterior part of thighs uniform light brown; dorsal part of thighs banded with 3-4 light and dark transversal bands; lower legs with three dark cross bars; foot and toes brown with indistinct darker pattern; ventral parts of thigh whitish beige with brownish mottling; ventral sides of lower legs with very dense reddish brown spotting; part of foot darker brown than dorsal part. Variation. Paratypes are similar to holotype, but mostly with less contrasting dorsal pattern. Size (SVL) of adult males ranges from 21.1–23.4 mm (mean ± sd: 21.9 ± 0.8 mm; N= 10). They are smaller than females, these ranging from 23.8–26.6 mm (mean ± sd: 25.6 ± 1.5 mm; N= 3; Tables 1–2). Further measures and indices are summarized in Table 1. The dorsal skin may vary from almost smooth to granular. Some small warts may be present on lower flanks and posterior back. In most male paratypes, the posterior half of the back is beset with minute round spinulae. Similar spinulae are present on dorsal surfaces of thigh and lower leg and, less visible, on anterior part of body; the throat and the anterior part of the pectoral region likewise densely beset with whitish, round spinulae. Depending on the preservation the spinulae can be difficult to see. They are best visible when the specimen is slightly dehydrated due to too high ethanol concentration, or after the skin has been carefully towelled. Presumably the distinctiveness of the spinulae may vary as well with the breeding status of the respective male. The spinulae are most pronounced in the male from Ghana (ZMB 86965; Fig. 3). Here they cover the entire back from vent to just behind the eyes, as well as the dorsal surfaces of the hind legs. species SVL HW FL TL FTL ED TD tanoeensis sp. nov. 21.9 ± 0.8 7.5 ± 0.5 9.2 ± 0.9 11.6 ± 0.7 16.8 ± 0.9 2.8 ± 0.4 1.7 ± 0.2 M, N= 10 21.1‒23.4 6.9‒8.3 7.9‒10.3 10.3‒12.2 15.4‒17.9 2.4‒3.5 1.4‒2.0 liberiensis 25.3 ± 1.5 8.1 ± 0.7 12.1 ± 1.1 14.3 ± 1.1 20.2 ± 1.6 3.1 ± 0.3 2.2 ± 0.2 M, N= 15 23.5‒28.9 6.2‒9.1 10.8‒14.4 12.8‒17.0 18.1‒24.2 2.6‒3.5 1.8‒2.7 tanoeensis sp. nov. 25.6 ± 1.5 8.9 ± 0.4 9.6 ± 0.9 13.4 ± 0.9 19.8 ± 1.5 3.1 ± 0.1 2.0 ± 0.4 F, N= 3 23.8‒26.6 8.5‒9.2 8.6‒10.2 12.4‒14.1 18.1‒21.1 3.0‒3.2 1.8‒2.4 intermedius 27.2 7.8 14.4 16.5 20.3 3.4 2.3 F, N= 1 liberiensis 31.7 ± 2.4 10.1 ± 0.8 14.7 ± 1.2 17.7 ± 1.1 24.0 ± 1.7 3.3 ± 0.4 2.4 ± 0.3 F, N= 31 25.5‒35.2 8.0‒11.2 11.9‒17.2 13.9‒19.3 19.1‒26.5 2.7‒4.0 1.9‒3.1 continued. species IOD EN ES HW/ SUL FL/ SUL TL/ SUL EN/ES tanoeensis sp. nov. 2.3 ± 0.3 2.0 ± 0.2 3.1 ± 0.4 0.3 ± 0 0.4 ± 0 0.5 ± 0 0.6 ± 0.1 M, N= 10 1.9‒2.6 1.7‒2.4 2.7‒4.0 0.3‒04 0.4‒05 0.5‒0.6 0.5‒0.8 liberiensis 2.1 ± 0.2 1.9 ± 0.2 2.9 ± 0.3 0.3 ± 0 0.5 ± 0 0.6 ± 0 0.6 ± 0.1 M, N= 15 1.8‒2.7 1.6‒2.2 2.4‒3.7 0.2‒0.4 0.4‒0.5 0.5‒0.6 0.6‒0.8 The dorsal warts/ridges may vary from very distinct to almost absent (in live and preserved specimens); often only the anterior pair is present and then longer than in the holotype (e.g. Figs. 2e, g). Dorsal and lateral colour varies from dark grey or light to dark brown to almost black, with or without lighter patterning. Sometimes a greenish tinge may be present on the back (Fig. 2d). Females exhibit more uniform dorsal coloration than males (Fig. 2g). Black lateral bands and white lines may be present (Figs. 2a, c-e) or absent. The dark face mask may be more or less pronounced. The juvenile specimen and some adults exhibit a pair of light dorsal spots posterior to the dorsal ridges (Figs. 2a, d). The Throats of female vary from light grey (Fig. 2h) to brownish dark with white spots. Further greyish or blackish belly spots are particularly dense close to the forearm insertion and towards flanks. The central parts of female bellies are almost uniform white. The ventral surfaces of female hind legs are greyish to orange, in males the colour is reddish beige or rosé. The pedal webbing may vary slightly. In particular toes III and IV may show slightly less or more webbing than the holotype (in the range of 0.5 phalanx). liberiensis 2.8 ± 0.4 2.3 ± 0.3 3.5 ± 0.3 0.3 ± 0 0.5 ± 0 0.6 ± 0 0.7 ± 0.1 F, N= 31 2.1‒3.6 1.7‒3.0 2.8‒4.0 0.3 0.4‒0.5 0.5‒0.6 0.5‒0.8 Genetics. We compared parts (up to 538 bp) of the 16S RNA gene from five individuals of the new species, with two sequences of P. intermedius and P. tokba, respectively and, considering the erroneous initial field identification, 27 individuals of P. liberiensis. The P. liberiensis samples covered almost the entire known range of that species from eastern Sierra Leone, Liberia, south-eastern Guinea, western and eastern Ivory Coast and western Ghana (Fig. 1). Phrynobatrachus liberiensis showed some clear intraspecific genetic structure, i.e. a western clade comprising frogs from the Ivorian Banco National Park to Sierra Leone and a Ghanaian clade, comprising samples from Kakum and Ankasa National Parks. The intraspecific variation in P. liberiensis ranged from 0–4.42% (mean ± sd: 2.02 ± 1.35%, N= 351 comparisons). This high genetic variation may indicate further cryptic diversity, i.e. a further undescribed taxon from Ghana. The two included P. tokba sequences differed by 1.36%, while intraspecific variation in P. intermedius was zero. In P. tanoeensis sp. nov. the intraspecific variation ranged from 0–0.19% (mean ± sd: 0.08 ± 0.10%, N= 10 comparisons). Phrynobatrachus tanoeensis sp. nov. differed from P. intermedius by 6.39–6.70% (mean ± sd: 6.64 ± 0.13%, N= 10), and from P. liberiensis by 5.25–7.03% (mean ± sd: 5.95 ± 0.44%, N= 135), and from P. tokba by 6.60– 7.27% (mean ± sd: 6.97 ± 0.30%, N= 22). These differences are well within the range of what has been accepted as genetic distances between other species of the genus (see Discussion). Acoustic. The recorded advertisement calls of two males of the new species comprise an amplitude modulated, short note, with few pulses that are difficult to tell apart (no full amplitude modulation = no complete silence between pulses; Fig. 4a, b). Call duration and frequency varied and showed some overlap to P. liberiensis calls, recorded in Banco and Taï National Parks, the latter calls as well showing some variation (Fig. 4, Table 3). However, a comparison of the spectrogram and oscillogram between the calls of both taxa, shows distinct differences (Fig. 4). The advertisement calls of P. tanoeensis sp. nov. are much shorter and less pulsed than the calls of P. liberiensis. These differences are also audible to the human ear. Whereas P. liberiensis calls can be described like a loud, hammer like “ackk,” the calls of the new species sound like a deep, short “tick.” Both species call predominantly during the day, in swamps or from the banks of small forest creeks. Distribution. The new species is so far only known from two swamp forests, the Tanoé-Ehy Swamp Forest region in south-eastern Ivory Coast and a forest near Ebonloa, south-western Ghana. These localities are only about 40 km apart. Natural history. Both known sites of the new species are predominantly swamp forests (Fig. 5). The mean annual temperature of the Tanoé-Ehy forest is 26°C, the mean annual precipitation is about 2000 mm. This region is characterized by a longer dry season (December to March), followed by the period with high precipitation (March to July). A minor rainy season extends from October to November (Avenard et al. 1971). The Tanoé-Ehy forest mainly consists of moist, partly primary forests on predominantly sandy soil, with vegetation typical for south-eastern Ivory Coast (Béligné 1994). Whereas all collected males from the Tanoé-Ehy forest were encountered well concealed in areas thickly covered with Raphia palms (Fig. 5b), the females were seen on muddy ground in open patches of the forest. The canopy of the sites where P. tanoeensis sp. nov. was found had gaps (Fig. 5a), and traces of recent human activity were visible. For instance, Raphia palms were used by the local population for palm wine extraction and as construction material. An unpaved road, leading to Dohouan and Nouamou villages, crossed the area. Due to frequent traffic with motorbikes the Dohouan site was quite noisy. The road was only eight meters from the calling site of one male. This swamp forest was surrounded by heavily degraded forest and plantations for subsistence farming (i.e. cassava plantation associated with coconut trees). The Dohouan River, which crosses the Tanoé-Ehy forest, is polluted due to the activity of launderers and people washing their cars and motorbikes. In Ghana, the species was collected in small patches of land within an inundated landscape (Fig. 5c). The dominant vegetation on these ‘islands’ are trees with stilt-like roots and Raphia palms. Etymology. The specific name refers to the type locality, the Tanoé-Ehy Swamp Forest. We suggest Tanoé Puddle Frog as common name.

Phrynobatrachus tanoeensis sp. nov. Figs 2���3 Holotype. ZMB 80870 (field and tissue #: YA 14-03, GenBank #: MG209126), adult male, Ivory Coast, Tano��-Ehy Swamp Forest, 05��11���181������N, 02��53���011������W (date WGS84), swamp forest dominated by Raphia palms, 17���29 July 2010, coll. T.F. Kpan & P.J. Adeba. Paratypes. ZMB 80871 (YA14-01, MG209125), adult male, same data as holotype; ZMB 86953 ���86955 (KK 1���3), three adult males, all with damages of dorsal skin, Ivory Coast, Dohouan village forest, Nouamou, 05��11.827'N, 02��53.140'W, 17 m asl, 1���3 October 2014, coll. T.F. Kpan & N.G. Kouam��; ZMB 86956���86960 (KK 4���8; ZMB 86959= MG209122), five adult males, ZMB 86961���86963 (KK 9���11; ZMB 8961= MG209123, ZMB 86963= MK209124), three females, 05��11.718'N, 02��52.854'W, 20 m asl, all other data as in ZMB 86953���86955. Additional material. ZMB 86964 (KK 12), juvenile, 05��11.718'N, 02��52.854'W, 20 m asl, all other data as in ZMB 86953���86955; ZMB 86965 (GAW009), adult male, skin damaged or missing on left flank and right thigh, Ghana, Western Region, Ebonloa, near Essiama, 05��02'0.28������N, 02��33'05.9W, 16 m asl, peat/swamp forest with slow moving stream near the borders of Ghana and Ivory Coast, coll. C. Ofori-Boateng. Diagnosis. The overall body shape and morphology, i.e. the presence of a tarsal tubercle, as well as the genetic data confirm that the new species is a member of the genus Phrynobatrachus (Phrynobatrachidae). From members of the externally-morphologically similar genus Arthroleptis (Arthroleptidae), the new species differs by: the presence of scapular ridges, pedal webbing and a tarsal tubercle, and the absence of a mid-dorsal skin raphe. From members of the genus Ptychadena (Ptychadenidae; juveniles sometimes morphologically similar to Phrynobatrachus) they differ by a more rounded and shorter snout and the absence of parallel dorsal glandular ridges. The new species can be differentiated from other members of the genus Phrynobatrachus by molecular and acoustic characters (see below), as well as by the combination of the following morphological characters: adult snout���vent length larger than 20 mm but smaller than 30 mm; a dark face mask; absence of a spiny tubercle on the eyelid; narrow and partly indistinct scapular ridges or comma-shaped warts, in particular no long X-shaped pair of dorsal ridges; distinct spinulae on males��� throats and dorsal surfaces; pronounced pedal webbing; finger- and toe tips not enlarged; a black throat in adult males; a white belly with small blackish spotting in both sexes; more than one wide dark cross bar on hind legs; ventral parts of hind legs in both sexes with ros�� to reddish colour. In particular the new species is smaller (P. liberiensis and P. plicatus (G��nther, 1858); it differs from P. annulatus Perret, 1966, P. calcaratus (Peters, 1863), P. taiensis Perret, 1988, P. villiersi Guib��, 1959 and P. pintoi Hillers, Zimkus & R��del, 2008 by lacking an eyelid cornicle (spine-like wart on upper eyelid); from all the latter species and P. gutturosus (Chabanaud, 1921), P. hieroglyphicus R��del, Ohler & Hillers, 2010, and P. tokba (Chabanaud, 1921), the new species differs by more extensive webbing; Phrynobatrachus annulatus, P. fraterculus (Chabanaud, 1921), P. ghanensis, P. maculiventris Guib�� & Lamotte, 1958, P. pintoi, P. taiensis and P. villiersi all exhibit very distinct, species-specific patterning on the belly, different to the new species which exhibits many small black spots. In contrast P. latifrons, Ahl, 1924, P. alleni Parker, 1936 (only females, males with yellow belly), P. francisci Boulenger, 1912, P. natalensis (Smith, 1849), and P. rainerguentheri R��del, Onadeko, Barej & Sandberger, 2012, have predominantly white bellies with at best a few minute black points. Whereas males of the new species have dark grey to black throats, P. alleni and P. latifrons males have yellow throats. These two species, along with all species with an eyelid cornicle, as well as P. francisci, P. gutturosus, P. phyllophilus R��del & Ernst, 2002, P. guineensis Guib�� & Lamotte, 1962, P. natalensis, P. rainerguentheri, P. brongersmai Parker, 1936 have a different body shape and a snout which is rounded in dorsal and lateral view (pointed in lateral view in the new species). In P. alleni the head is also relatively flatter in lateral view. Phrynobatrachus guineensis is smaller (Phrynobatrachus phyllophilus shows some black spots on belly and pectoral region but has a more compact body shape and roundish snout, toe and finger tips which are enlarged to small round discs (straight in the new species) and usually only one wide black cross bar on thigh and lower leg (several in the new species). In West Africa, only P. alleni and P. plicatus have scapular ridges formed like an open X, in P. plicatus this X extends to posterior to half-back. The new species shares a black face mask in West Africa only with P. intermedius, P. plicatus and some P. liberiensis and P. rainerguentheri. However, the face mask is always less distinct than in P. plicatus (see discussion for a more in depth comparison with the genetically most similar species). Description of the holotype (measurements in millimetres mm). Adult male with longish, but compact body shape, short snout and large eyes; snout���vent length 22.1; snout rounded in dorsal, moderately pointed in lateral view; head width 6.9; thigh length 8.5, shorter than crus length, 12.0; foot length including longest toe 15.4; eye diameter 2.8, larger than horizontal diameter of tympanum, 1.6; tympanum distinct, round with round median annuli, diameter 0.5; interorbital distance 2.5; distance eye���nostrils 1.9; distance eye���snout 3.0; nostrils much closer to snout than to eye; canthus rostralis rounded; loreal region straight; distinct, slightly bent supratympanal ridge extending from posterior edge of eye to forearm insertion; dorsal skin fine granular, spinulae on posterior part of back indistinct; spinulae on dorsal surfaces of thighs and lower legs; two pairs of narrow, indistinct, commashaped warts; the anterior pair originating slightly posterior to eyelids, slightly converging and extending to about forearm insertion; the posterior pair being shorter and slightly diverging; a few minute round tubercles on eyelids; throat smooth with parallel skin folds along mandibles; whitish spinulae on throat and anterior part of breast indistinct; skin of other ventral surfaces smooth; palmar surface of hand with large, oval palmar and more slender, longish thenar tubercle; fingers with large, undivided oval subarticular tubercles, additional tubercles on hands absent; finger tips roundish, not enlarged to discs; outer part of thumb with indistinct thin nuptial pad; palmar webbing absent; relative finger length: II = IV �� I Colour of the holotype in life. Dorsal surfaces light beige to yellowish brown; top of snout with Y-shaped dark figure ��� the single arm pointing towards snout, and further dark spots; margin of snout and upper eyelids with ill-defined light band; upper eyelids dark brown, connected by dark brown inter-orbital band, this band bordered anteriorly and posteriorly by narrow clear bands; the two pairs of narrow comma shaped scapular ridges marked darker, the posterior pair connected by V-shaped black marking; a pair of white spots at the tips of this V; posterior back with a few further blackish markings; dorsal surfaces of thighs (4) and lower legs (3) with black cross bars; arms, hands and feet with less distinct black cross bars; lateral side of snout, upper lip and area below supratympanal ridge black with some minute white spots on lip; supratympanal ridge dorsally bordered by white band; posterior to this band a wide black lateral band, followed by further black spots on whitish base colour; lower mandible, throat and anterior pectoral region almost uniform dark grey; belly white, densely beset with black spots; posterior belly and lower surfaces of thighs and lower legs ros��. Colour of the holotype in preservation (after seven years in 75% ethanol). Back light brown, a few white spots where skin is rubbed off; snout tip and upper eyelids with some black mottling; loreal triangle from nostrils to anterior eye and tympanal area darker; scapular ridges and supratympanal ridge darker; black and light lateral pattern faded; upper arm with two dark cross bars, lower side reddish brown; lower arm with one dark cross bar close to hand; palmar surfaces of hands uniform light reddish brown; lower side of mandibles light blackish grey; throat skin dark blackish grey with minute white points; belly and pectoral region whitish with dark brown spotting, denser towards throat and flanks; vent surrounded by flat black triangle, bordered by narrow light line; posterior part of thighs uniform light brown; dorsal part of thighs banded with 3-4 light and dark transversal bands; lower legs with three dark cross bars; foot and toes brown with indistinct darker pattern; ventral parts of thigh whitish beige with brownish mottling; ventral sides of lower legs with very dense reddish brown spotting; part of foot darker brown than dorsal part. Variation. Paratypes are similar to holotype, but mostly with less contrasting dorsal pattern. Size (SVL) of adult males ranges from 21.1���23.4 mm (mean �� sd: 21.9 �� 0.8 mm; N= 10). They are smaller than females, these ranging from 23.8���26.6 mm (mean �� sd: 25.6 �� 1.5 mm; N= 3; Tables 1���2). Further measures and indices are summarized in Table 1. The dorsal skin may vary from almost smooth to granular. Some small warts may be present on lower flanks and posterior back. In most male paratypes, the posterior half of the back is beset with minute round spinulae. Similar spinulae are present on dorsal surfaces of thigh and lower leg and, less visible, on anterior part of body; the throat and the anterior part of the pectoral region likewise densely beset with whitish, round spinulae. Depending on the preservation the spinulae can be difficult to see. They are best visible when the specimen is slightly dehydrated due to too high ethanol concentration, or after the skin has been carefully towelled. Presumably the distinctiveness of the spinulae may vary as well with the breeding status of the respective male. The spinulae are most pronounced in the male from Ghana (ZMB 86965; Fig. 3). Here they cover the entire back from vent to just behind the eyes, as well as the dorsal surfaces of the hind legs. species SVL HW FL TL FTL ED TD tanoeensis sp. nov. 21.9 �� 0.8 7.5 �� 0.5 9.2 �� 0.9 11.6 �� 0.7 16.8 �� 0.9 2.8 �� 0.4 1.7 �� 0.2 M, N= 10 21.1���23.4 6.9���8.3 7.9���10.3 10.3���12.2 15.4���17.9 2.4���3.5 1.4���2.0 liberiensis 25.3 �� 1.5 8.1 �� 0.7 12.1 �� 1.1 14.3 �� 1.1 20.2 �� 1.6 3.1 �� 0.3 2.2 �� 0.2 M, N= 15 23.5���28.9 6.2���9.1 10.8���14.4 12.8���17.0 18.1���24.2 2.6���3.5 1.8���2.7 tanoeensis sp. nov. 25.6 �� 1.5 8.9 �� 0.4 9.6 �� 0.9 13.4 �� 0.9 19.8 �� 1.5 3.1 �� 0.1 2.0 �� 0.4 F, N= 3 23.8���26.6 8.5���9.2 8.6���10.2 12.4���14.1 18.1���21.1 3.0���3.2 1.8���2.4 intermedius 27.2 7.8 14.4 16.5 20.3 3.4 2.3 F, N= 1 liberiensis 31.7 �� 2.4 10.1 �� 0.8 14.7 �� 1.2 17.7 �� 1.1 24.0 �� 1.7 3.3 �� 0.4 2.4 �� 0.3 F, N= 31 25.5���35.2 8.0���11.2 11.9���17.2 13.9���19.3 19.1���26.5 2.7���4.0 1.9���3.1 continued. species IOD EN ES HW/ SUL FL/ SUL TL/ SUL EN/ES tanoeensis sp. nov. 2.3 �� 0.3 2.0 �� 0.2 3.1 �� 0.4 0.3 �� 0 0.4 �� 0 0.5 �� 0 0.6 �� 0.1 M, N= 10 1.9���2.6 1.7���2.4 2.7���4.0 0.3���04 0.4���05 0.5���0.6 0.5���0.8 liberiensis 2.1 �� 0.2 1.9 �� 0.2 2.9 �� 0.3 0.3 �� 0 0.5 �� 0 0.6 �� 0 0.6 �� 0.1 M, N= 15 1.8���2.7 1.6���2.2 2.4���3.7 0.2���0.4 0.4���0.5 0.5���0.6 0.6���0.8 The dorsal warts/ridges may vary from very distinct to almost absent (in live and preserved specimens); often only the anterior pair is present and then longer than in the holotype (e.g. Figs. 2e, g). Dorsal and lateral colour varies from dark grey or light to dark brown to almost black, with or without lighter patterning. Sometimes a greenish tinge may be present on the back (Fig. 2d). Females exhibit more uniform dorsal coloration than males (Fig. 2g). Black lateral bands and white lines may be present (Figs. 2a, c-e) or absent. The dark face mask may be more or less pronounced. The juvenile specimen and some adults exhibit a pair of light dorsal spots posterior to the dorsal ridges (Figs. 2a, d). The Throats of female vary from light grey (Fig. 2h) to brownish dark with white spots. Further greyish or blackish belly spots are particularly dense close to the forearm insertion and towards flanks. The central parts of female bellies are almost uniform white. The ventral surfaces of female hind legs are greyish to orange, in males the colour is reddish beige or ros��. The pedal webbing may vary slightly. In particular toes III and IV may show slightly less or more webbing than the holotype (in the range of 0.5 phalanx). liberiensis 2.8 �� 0.4 2.3 �� 0.3 3.5 �� 0.3 0.3 �� 0 0.5 �� 0 0.6 �� 0 0.7 �� 0.1 F, N= 31 2.1���3.6 1.7���3.0 2.8���4.0 0.3 0.4���0.5 0.5���0.6 0.5���0.8 Genetics. We compared parts (up to 538 bp) of the 16S RNA gene from five individuals of the new species, with two sequences of P. intermedius and P. tokba, respectively and, considering the erroneous initial field identification, 27 individuals of P. liberiensis. The P. liberiensis samples covered almost the entire known range of that species from eastern Sierra Leone, Liberia, south-eastern Guinea, western and eastern Ivory Coast and western Ghana (Fig. 1). Phrynobatrachus liberiensis showed some clear intraspecific genetic structure, i.e. a western clade comprising frogs from the Ivorian Banco National Park to Sierra Leone and a Ghanaian clade, comprising samples from Kakum and Ankasa National Parks. The intraspecific variation in P. liberiensis ranged from 0���4.42% (mean �� sd: 2.02 �� 1.35%, N= 351 comparisons). This high genetic variation may indicate further cryptic diversity, i.e. a further undescribed taxon from Ghana. The two included P. tokba sequences differed by 1.36%, while intraspecific variation in P. intermedius was zero. In P. tanoeensis sp. nov. the intraspecific variation ranged from 0���0.19% (mean �� sd: 0.08 �� 0.10%, N= 10 comparisons). Phrynobatrachus tanoeensis sp. nov. differed from P. intermedius by 6.39���6.70% (mean �� sd: 6.64 �� 0.13%, N= 10), and from P. liberiensis by 5.25���7.03% (mean �� sd: 5.95 �� 0.44%, N= 135), and from P. tokba by 6.60��� 7.27% (mean �� sd: 6.97 �� 0.30%, N= 22). These differences are well within the range of what has been accepted as genetic distances between other species of the genus (see Discussion). Acoustic. The recorded advertisement calls of two males of the new species comprise an amplitude modulated, short note, with few pulses that are difficult to tell apart (no full amplitude modulation = no complete silence between pulses; Fig. 4a, b). Call duration and frequency varied and showed some overlap to P. liberiensis calls, recorded in Banco and Ta�� National Parks, the latter calls as well showing some variation (Fig. 4, Table 3). However, a comparison of the spectrogram and oscillogram between the calls of both taxa, shows distinct differences (Fig. 4). The advertisement calls of P. tanoeensis sp. nov. are much shorter and less pulsed than the calls of P. liberiensis. These differences are also audible to the human ear. Whereas P. liberiensis calls can be described like a loud, hammer like ���ackk,��� the calls of the new species sound like a deep, short ���tick.��� Both species call predominantly during the day, in swamps or from the banks of small forest creeks. Distribution. The new species is so far only known from two swamp forests, the Tano��-Ehy Swamp Forest region in south-eastern Ivory Coast and a forest near Ebonloa, south-western Ghana. These localities are only about 40 km apart. Natural history. Both known sites of the new species are predominantly swamp forests (Fig. 5). The mean annual temperature of the Tano��-Ehy forest is 26��C, the mean annual precipitation is about 2000 mm. This region is characterized by a longer dry season (December to March), followed by the period with high precipitation (March to July). A minor rainy season extends from October to November (Avenard et al. 1971). The Tano��-Ehy forest mainly consists of moist, partly primary forests on predominantly sandy soil, with vegetation typical for south-eastern Ivory Coast (B��lign�� 1994). Whereas all collected males from the Tano��-Ehy forest were encountered well concealed in areas thickly covered with Raphia palms (Fig. 5b), the females were seen on muddy ground in open patches of the forest. The canopy of the sites where P. tanoeensis sp. nov. was found had gaps (Fig. 5a), and traces of recent human activity were visible. For instance, Raphia palms were used by the local population for palm wine extraction and as construction material. An unpaved road, leading to Dohouan and Nouamou villages, crossed the area. Due to frequent traffic with motorbikes the Dohouan site was quite noisy. The road was only eight meters from the calling site of one male. This swamp forest was surrounded by heavily degraded forest and plantations for subsistence farming (i.e. cassava plantation associated with coconut trees). The Dohouan River, which crosses the Tano��-Ehy forest, is polluted due to the activity of launderers and people washing their cars and motorbikes. In Ghana, the species was collected in small patches of land within an inundated landscape (Fig. 5c). The dominant vegetation on these ���islands��� are trees with stilt-like roots and Raphia palms. Etymology. The specific name refers to the type locality, the Tano��-Ehy Swamp Forest. We suggest Tano�� Puddle Frog as common name., Published as part of Kpan, Tokouaho Flora, Kouam��, N'Goran Germain, Barej, Michael F., Adeba, Patrick Jo��l, Emmrich, Mike, Ofori-Boateng, Caleb & R��del, Mark-Oliver, 2018, A new Puddle Frog, genus Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae), from the eastern part of the Upper Guinea biodiversity hotspot, West Africa, pp. 221-237 in Zootaxa 4388 (2) on pages 224-233, DOI: 10.11646/zootaxa.4388.2.5, http://zenodo.org/record/1187953, {"references":["Perret, J. - L. (1988) Les especes de Phrynobatrachus (Anura, Ranidae) a eperon palpebral. Archives des Sciences Geneve, 41, 275 - 294.","Hillers, A., Zimkus, B. & Rodel, M. - O. (2008) A new species of Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae) from north-western Guinea, West Africa. Zootaxa, 1815, 43 - 50.","Rodel, M. - O., Ohler, A. & Hillers, A. (2010) A new extraordinary Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae) from West Africa. Zoosystematics and Evolution, 86, 257 - 261. https: // doi. org / 10.1002 / zoos. 201000008","Rodel, M. - O., Barej, M. F., Hillers, A., Leache, A. D., Kouame, N. G., Ofori-Boateng, C., Assemian, N. E., Tohe, B., Penner, J., Hirschfeld, M., Doumbia, J., Gonwouo, L. N., Nopper, J., Brede, C., Diaz, R., Fujita, M., Gil, M., Segniagbeto, G. H., Ernst, R. & Sandberger, L. (2012) The genus Astylosternus in the Upper Guinea rainforests, West Africa, with the description of a new species (Amphibia: Anura: Arthroleptidae). Zootaxa, 3245, 1 - 29.","Rodel, M. - O. & Ernst, R. (2002) A new Phrynobatrachus from the Upper Guinean rain forest, West Africa, including a description of a new reproductive mode for the genus. Journal of Herpetology, 36, 561 - 571. https: // doi. org / 10.1670 / 0022 - 1511 (2002) 036 [0561: ANPFTU] 2.0. CO; 2","Avenard, J. - M., Eldin, M., Girard, G., Sircoulon, J., Touchebeuf, P., Guillaumet, J. - L. & Adjanohoun, E. (1971) Le milieu naturel de la Cote d'Ivoire. ORSTOM, Paris, 391 pp.","Beligne, V. (1994) Etude de l'etat du milieu naturel du Parc National du Banco: recommandation pour sa sauvegarde et son amenagement. Rapport du Fonds Mondial pour la Nature (WWF), Abidjan, 48 pp."]}

Afroedura gorongosa sp. nov. (Figs. 4���5) Afroedura transvaalica (part) Broadley 1966, p. 111. Holotype. ZMB 83293 (GNP 484), adult male, western flank of Mount Gorongosa, Gorongosa National Park, Sofala Province, Mozambique (1038 m a.s.l., 18˚28���04.3���S, 34˚02���53.2���E), collected by M.- O. R��del and M.F. Barej, 11 December 2015. Paratypes (three specimens). ZMB 83290 (GNP 438), adult female, with incision in left thigh; PEM R22220 (GNP 439, previously ZMB 83291), adult female, with incision in left thigh; ZMB 83292 (GNP 440; will later be deposited in the zoological collection of the E.O. Wilson Laboratory in Gorongosa National Park), adult male, with everted hemipenes and incision in left thigh; collected on 22 May and 21 July 2015 by J.A. Guyton and P. Naskrecki from gallery forest near Murombodzi Waterfall, Mount Gorongosa, Gorongosa National Park, Sofala Province, Mozambique (842 m a.s.l; 18��29'0.1''S, 34��2'34.6''E). Additional material (two specimens). ZMB 83288 (GNP 433), adult male, ZMB 83289 (GNP 434), adult female; both collected in April 2015 by H. Farooq, P. Naskrecki and J.A. Guyton from the top of Bunga Inselberg, Gorongosa National Park, Sofala Province, Mozambique (approx. 200 m a.s.l., 18˚35���58.1���S, 34˚20���34.8���E). Etymology. The specific name refers to Mount Gorongosa and Gorongosa National Park in Central Mozambique, to which the species is endemic. We suggest Gorongosa Flat Gecko is a suitable common name. Afroedura, based on Oedura, is feminine and the specific epithet is treated as a noun in apposition. Diagnosis. A member of the A. transvaalica -complex in possessing two pairs of enlarged scansors per digit and a strongly verticillate and flattened tail (Jacobsen et al. 2014). Differs from other members of the complex by having 97���102 midbody scale rows (less than 95 in A. bogerti- complex, 113���120 (���= 117) in A. loveridgei and 102���119 in A. transvaalica (102���118, ���= 109 in South Africa���Jacobsen et al. 2014; 108���119, ���= 114 in Zimbabwe adjacent to Mount Gorongosa); by the rostral bordering the nostril (nostril excluded from rostral in A. loveridgei); by the anterior nasals being separated by 1���3 granules (always in broad contact in A. loveridgei; usually in broad contact in A. transvaalica - A. loveridgei; 17���20 in South African A. transvaalica, and 15���19 in Zimbabwe populations adjacent to Mozambique); and in having a higher average number of precloacal pores in males (8���13, ���= 10; 6���10, ���= 8 in A. transvaalica; 8���10, ���= 9 in A. loveridgei). Holotype description. Morphology. Adult male; SVL 50.8 mm; tail 52.2 mm (partly regenerated), with a small incision in the left thigh for the removal of muscle tissue. Head and body dorsoventrally depressed; HL 12.3 mm, HW 9.0 mm, broadest at mid-eye and 1.37x longer than wide. Eye large (3.7 mm wide), pupil vertical with indented margins; circumorbital scales small and granular, becoming elongate, almost ciliate, at upper posterior margin. Snout rounded, 4.9 mm long, longer than distance between eye and ear openings (3.6 mm). Scales on top of snout granular, rounded and convex, the largest being 2/3rds the width of the scales on the back, which are granular and juxtaposed, with no intervening minute granules. Scales on snout almost twice as large as those on crown and throat. Scales on belly flattened, imbricate, more-or-less ovate at mid-ventrum, and twice the size of lateral granules and 1.5x those along backbone. Ear opening deep, oblique and roughly oval, only half as high as wide. Limbs well developed, hindlimbs slightly longer than forelimbs, both without obvious mite pockets at posterior margin of limb insertions. All digits with a large pair of distal scansors, separated by a large, curved claw, and followed after a large gap (twice length of terminal scansor) by a smaller pair of scansors; infero-median row of digital scales enlarged transversely, particularly towards the scansors, where the terminal scale adjoining the first pair of scansors may be medially constricted, swollen and scansor-like; 12 scale rows under 4th toe. Nostril pierced between rostral, 1 st supralabial and three nasal scales; the supranasal being much larger than the subequal postnasals and separated by a granule bordering the rostral, followed behind by two smaller granules. Rostral roughly rectangular but with its upper edges elongated due to extensions into the nostril. Nine supralabials on each side, the labial margin flexing dorsad at the rictus, with 1���2 minute scales proximal to the flexure. Nine infralabials on either side, with a small scale proximad to the flexure. At the lip, mental slightly narrower than adjacent infralabial, only 2/3rds the width of rostral, and in contact with two distinctly elongate postmental scales. Approximately 20 scales across crown at level of front of eye; 13 scales between nostril and front of eye; 23 scales from front of ear to back of eye; 99 scales around midbody. A roughly V-shaped row of 13 precloacal pores occurs eight scale rows anterior to the cloacal lip, with these scales more imbricate and triangular adjacent to the precloacal pores. Original portion of tail slightly dorsoventrally flattened and indistinctly verticillate, without obvious lateral constrictions; each verticil comprising 6���7 imbricate rows of scales dorsally and 5 imbricate scale rows ventrally, and with ventral scales approximately twice the size of those on dorsal surface. Hemipenal pouches well-developed, with an angular, ventro-lateral series of 3 enlarged tubercular scales on each side (largest nearest cloaca), and with two openings of the cloacal sacs on the posterior lip of the cloaca. Colour in life (from photographs, Fig. 4a and unpublished). Dark black-brown above with a series of five white dorsal spots (4���5 scales across) between the fore- and hind-limb insertions, that are surrounded by a chocolate brown zone that encompasses a series of 2���4 laterally arranged smaller pale spots (2���3 scales wide), and posteriorly by a diffuse pale region that forms an irregular band across the mid dorsum. Top of head, flanks and upper surfaces of limbs with small, scattered pale spots. Head with a dark brown band across the posterior edge of crown encompassing a small central pale spot; a vague, thin pale canthal stripe, extends on both sides from the nasal region to front of eye; upper and lower labials grey-brown with diffuse yellowish bars; iris reddish brown with a heavy dark brown reticulation, and pupil with crenulated edge. Original segment of tail mottled in dark brown and pale blotches; regenerated tail grey with vague darker blotches. Ventrum mottled grey-cream. Colour in preservative. Dark black-brown above with irregular markings on back, comprising four equally spaced dark bars, each with a central white spot 4���5 scales wide, and an irregular, paler brown bar behind; top and sides of head, flanks and upper surfaces of limbs with small, irregular pale spots; ventral surface of throat, neck, belly, limbs and tail uniform dark grey. Side of head and labials dark brown, with a few scattered pale spots on upper labials, and vague pale edges to lower labials. Variation in paratypes and additional material (Figs. 4b, 4c, 5; Tables 3���4). Morphology. SVL from 54.6 mm (ZMB 83289) to 61.2 mm (PEM 22220); head length 1.24���1.38 times head width (max. in paratypes 1.38); snout only 1.5 times diameter of eye in one specimen. The supranasals always separated by granules, usually with a single large granule in contact with the rostral between the supranasals, followed by 2���3 smaller granules in lateral contact; the first upper labial and rostral always enters the nostril, and the width of the rostral at the lip margin is always wider than that of the mental (except in ZMB 83290 were they are subequal); always two postmental scales; supralabials more than in holotype (10���11), infralabials (9���10); scales between anterior edge of eye 19���21; scales between nostril and anterior edge of orbit 12���13; scales between anterior edge of ear and rear margin of orbit 21���24; scales around mid-body 97���101; subdigital lamella beneath 4th toe 10���11; dorsal scales per tail verticel 7���8; ventral scales per tail verticel 5. In one female (ZMB 83290) the terminal, medially-constricted subdigital lamella of the 3rd toe of left foot is swollen and scansor-like, bearing small setae. Precloacal pores 8���13 in males, females without dimples in precloacal scales (as in A. transvaalica, but unlike A. loveridgei where five females had 5���9 dimpled precloacal scales, that lacked secretory activity, and only one had none). Colour in life (based on female paratype PEM R22229). Boldly patterned, with six dorsal dark transverse bands, posteriorly edged with cream, the first on the neck, the second and sixth at the fore- and hindlimb insertions, respectively, and with three thin bands equally spaced between them on the body; background colour of body, head and upper surfaces of limbs flecked on light brown and cream with occasional pale spots 2���4 scales wide; a dark canthal stripe runs through the rostral to the front of the eye, with a paler zone above; the scales of the anteriordorsal region of the orbital rim are bright yellow, with a faint yellowish infusion of the crown scales between the eyes; the original tail is boldly barred dorsally with seven, subequally-spaced, dark brown-black bars with a yellow posterior edge; belly uniform cream with scattered yellowish scales, greyer on ventral surfaces of limbs and tail. Colour in preservative. Similar to holotype, but dorsum pale brown to grey; larger dorsal markings usually reticulated, occasionally forming irregular transverse bands (6���7 from occiput to between hindlimbs); smaller dorsal markings vary from vague to distinct. A few specimens have irregular pale blotches along the posterior edges of the darker dorsal patches/bands, and also scattered on the dorsal surfaces of the limbs and back. Dorsal surface of regenerated tails usually with irregular pale and dark blotches that may form wavy bands, and which become flecks or grey blotches below. Size. Largest male (ZMB 83288)���SVL 56.5 mm, tail (partially regenerated) 55.7 mm; largest female (PEM R22220)���SVL 61.2 mm, tail (original) 64.0 mm. It is thus intermediate in size between A. transvaalica (malemax. SVL 72 mm, female���max. SVL 64 mm) and A. loveridgei (male���max. SVL 59 mm, female���max. SVL 58 mm). Only one of six specimens had an original tail (105% SVL). Hemipenis (based on ZMB 83292). Approximately 5 mm long, with a simple, smooth, possibly longitudinally flounced pedicel; capped with two large cups covered with fine calyculate ornamentation; sulcus simple and draining into the conjoined base of the distal cups. Distribution. Known from only three localities in the Mount Gorongosa region (Fig. 2). The holotype and paratypes, plus various non-collected individuals, have been found on the western flanks of Mount Gorongosa between about 900 and 1100 m a.s.l. (Figs. 6a���c). Access to further potential areas on the mountain, i.e. rock faces at the eastern slopes, was not possible due to security concerns. The additional material and various non-collected specimens have been observed at a lower altitude on the Bunga Inselberg (about 212 m a.s.l.) in the north-western corner of the National Park, approximately 40 km from the type locality (Fig. 6d). No tissue samples from the Bunga individuals were available, and the assignment of these individuals to the new species was thus based on morphological similarity. Natural history. The Gorongosa National Park is situated in Central Mozambique approx. 100 km south-west of the Zambezi River, and is in the southern part of the great African Rift Valley. Mount Gorongosa is an isolated granite mountain massif rising to 1863 m a.s.l., north-west of the National Park���s plains. At its highest elevations the massif���s plateau is covered with montane grasslands and forest patches dominated by cypresses (Widdringtonia nodiflora); the upper slopes of the mountain (900���1600 m) support a belt of moist evergreen forest. The type series was collected in an area between 900 and 1100 m on the western flanks of the mountain (Figs. 6a���c, 7). The paratypes were collected in a small alcove beneath a boulder pile in the closed-canopy riverine forest beside the Murombodzi Waterfall, and thus in comparatively cool and moist microclimates (Figs. 6c, 7). Some rocks with lizards present were located within the splash zone of the waterfall. Other specimens were observed but not collected at this site. The holotype was collected from a deep narrow crack in a large granite rock (i.e. 8 m ��) in highly degraded grassland below the edge of the rainforest and almost fully exposed to the sun (Fig. 6b). It was the only specimen seen at that site. At both sites the new species lived in syntopy with the cordylid Smaug mossambicus (FitzSimons, 1958). The additional non-type material comes from the Bunga Inselberg (Fig. 6d), a partly eroded inselberg composed mostly of trachyte in the north-western corner of the national park, covered and surrounded by dense Miombo woodland at a relatively low altitude (approx. 212 m a.s.l.). Here the geckos were observed during the day and at night in several places in deep cracks between very large boulders. No geckos were observed outside of these cracks. Syntopic lizards were Smaug mossambicus, the skink Trachylepis margaritifera (Peters, 1854) and the gecko Hemidactylus platycephalus Peters, 1854. Further sympatric, but not syntopic geckos were: Lygodactylus cf. capensis (Smith, 1849), Pachydactylus punctatus Peters, 1854 and Chondrodactylus turneri (Gray, 1864). The new species was observed regularly between April and December 2015. Like A. transvaalica, it is tolerant of conspecifics and frequently several specimens, and sometimes as many as over 20, were observed sheltering in deep rock crevices. One adult female (ZMB 83290) contained two eggs of approximately 11.5 x 7.5 mm. Conservation Status. The population of the new gecko present on the lower slopes of Mount Gorongosa is threatened by illegal deforestation taking place within the riverine forest adjacent to Murombodzi Waterfall. Slashand-burn removal of tall, old growth trees around the rocks may cause changes to the humidity and water availability in these currently very moist, shaded microhabitats that seem to be preferred by the lizards. Reforestation efforts led by Gorongosa National Park and introduction of shade-grown coffee as an alternative to slash-and-burn agriculture still practiced around the mountain will hopefully slow or halt the loss of the native riverine and evergreen vegetation in that area. On Bunga Inselberg the population of A. gorongosa is currently protected within the core area of the National Park and no immediate threat to its survival exists, though deforestation for agriculture is slowly encroaching on the area. However, based on the existence of only a few known sites and small area of occupancy, i.e. the presumed restriction to a fragmented range on Mount Gorongosa and potentially some rocky areas surrounding the mountain, as well as the ongoing high rate of habitat degradation and conversion on the mountain, the species may be considered highly threatened, potentially justifying an assessment as Endangered or even Critically Endangered. Further surveys and formal IUCN Red List assessment are required to confirm this., Published as part of William R. Branch, Jennifer A. Guyton, Andreas Schmitz, Michael F. Barej, Piotr Naskrecki, Harith Farooq, Luke Verburgt & Mark-Oliver R��del, 2017, Description of a new flat gecko (Squamata: Gekkonidae: Afroedura) from Mount Gorongosa, Mozambique, pp. 142-160 in Zootaxa 4324 (1) on pages 150-155, DOI: 10.11646/zootaxa.4324.1.8, http://zenodo.org/record/997117, {"references":["Broadley, D. G. (1966) The herpetology of south-east Africa. Unpublished Ph. D. thesis, University of Natal, Pietermaritzburg, 680 pp.","Jacobsen, N. H. G., Kuhn, A. L., Jackman, T. R. & Bauer, A. M. (2014) A phylogenetic analysis of the southern African gecko genus Afroedura Loveridge (Squamata: Gekkonidae), with the description of nine new species from Limpopo and Mpumalanga provinces of South Africa. Zootaxa, 3846 (4), 451 - 501. https: // doi. org / 10.11646 / zootaxa. 3846.4.1"]}

Key to the Afroedura transvaalica- group 1. Midbody scale rows more than 95........................................................................2 - Midbody scale rows less than 95; Angola.................................................. A. bogerti (sensu lato) 2. Rostral bordering nostril................................................................................3 - Rostral excluded from nostril................................................................... A. loveridgei 3. Anterior nasals in contact (very rarely separated); scales around midbody-South Africa 102-118 (= 109), northern Zimbabwe 108-119 (= 114)........................................................................... A. transvaalica - Anterior nasals separated by 1-3 granules; scales around midbody 99-101 (= 100)................ A. gorongosa sp. nov.

Key to the Afroedura transvaalica- group 1. Midbody scale rows more than 95........................................................................2 - Midbody scale rows less than 95; Angola.................................................. A. bogerti (sensu lato) 2. Rostral bordering nostril................................................................................3 - Rostral excluded from nostril................................................................... A. loveridgei 3. Anterior nasals in contact (very rarely separated); scales around midbody-South Africa 102-118 (���= 109), northern Zimbabwe 108-119 (���= 114)........................................................................... A. transvaalica - Anterior nasals separated by 1-3 granules; scales around midbody 99-101 (���= 100)................ A. gorongosa sp. nov., Published as part of William R. Branch, Jennifer A. Guyton, Andreas Schmitz, Michael F. Barej, Piotr Naskrecki, Harith Farooq, Luke Verburgt & Mark-Oliver R��del, 2017, Description of a new flat gecko (Squamata: Gekkonidae: Afroedura) from Mount Gorongosa, Mozambique, pp. 142-160 in Zootaxa 4324 (1) on page 156, DOI: 10.11646/zootaxa.4324.1.8, http://zenodo.org/record/997117

Afroedura gorongosa sp. nov. (Figs. 4–5) Afroedura transvaalica (part) Broadley 1966, p. 111. Holotype. ZMB 83293 (GNP 484), adult male, western flank of Mount Gorongosa, Gorongosa National Park, Sofala Province, Mozambique (1038 m a.s.l., 18˚28’04.3”S, 34˚02’53.2”E), collected by M.- O. Rödel and M.F. Barej, 11 December 2015. Paratypes (three specimens). ZMB 83290 (GNP 438), adult female, with incision in left thigh; PEM R22220 (GNP 439, previously ZMB 83291), adult female, with incision in left thigh; ZMB 83292 (GNP 440; will later be deposited in the zoological collection of the E.O. Wilson Laboratory in Gorongosa National Park), adult male, with everted hemipenes and incision in left thigh; collected on 22 May and 21 July 2015 by J.A. Guyton and P. Naskrecki from gallery forest near Murombodzi Waterfall, Mount Gorongosa, Gorongosa National Park, Sofala Province, Mozambique (842 m a.s.l; 18°29'0.1''S, 34°2'34.6''E). Additional material (two specimens). ZMB 83288 (GNP 433), adult male, ZMB 83289 (GNP 434), adult female; both collected in April 2015 by H. Farooq, P. Naskrecki and J.A. Guyton from the top of Bunga Inselberg, Gorongosa National Park, Sofala Province, Mozambique (approx. 200 m a.s.l., 18˚35’58.1”S, 34˚20’34.8”E). Etymology. The specific name refers to Mount Gorongosa and Gorongosa National Park in Central Mozambique, to which the species is endemic. We suggest Gorongosa Flat Gecko is a suitable common name. Afroedura, based on Oedura, is feminine and the specific epithet is treated as a noun in apposition. Diagnosis. A member of the A. transvaalica -complex in possessing two pairs of enlarged scansors per digit and a strongly verticillate and flattened tail (Jacobsen et al. 2014). Differs from other members of the complex by having 97–102 midbody scale rows (less than 95 in A. bogerti- complex, 113–120 (= 117) in A. loveridgei and 102–119 in A. transvaalica (102–118, = 109 in South Africa—Jacobsen et al. 2014; 108–119, = 114 in Zimbabwe adjacent to Mount Gorongosa); by the rostral bordering the nostril (nostril excluded from rostral in A. loveridgei); by the anterior nasals being separated by 1–3 granules (always in broad contact in A. loveridgei; usually in broad contact in A. transvaalica - A. loveridgei; 17–20 in South African A. transvaalica, and 15–19 in Zimbabwe populations adjacent to Mozambique); and in having a higher average number of precloacal pores in males (8–13, = 10; 6–10, = 8 in A. transvaalica; 8–10, = 9 in A. loveridgei). Holotype description. Morphology. Adult male; SVL 50.8 mm; tail 52.2 mm (partly regenerated), with a small incision in the left thigh for the removal of muscle tissue. Head and body dorsoventrally depressed; HL 12.3 mm, HW 9.0 mm, broadest at mid-eye and 1.37x longer than wide. Eye large (3.7 mm wide), pupil vertical with indented margins; circumorbital scales small and granular, becoming elongate, almost ciliate, at upper posterior margin. Snout rounded, 4.9 mm long, longer than distance between eye and ear openings (3.6 mm). Scales on top of snout granular, rounded and convex, the largest being 2/3rds the width of the scales on the back, which are granular and juxtaposed, with no intervening minute granules. Scales on snout almost twice as large as those on crown and throat. Scales on belly flattened, imbricate, more-or-less ovate at mid-ventrum, and twice the size of lateral granules and 1.5x those along backbone. Ear opening deep, oblique and roughly oval, only half as high as wide. Limbs well developed, hindlimbs slightly longer than forelimbs, both without obvious mite pockets at posterior margin of limb insertions. All digits with a large pair of distal scansors, separated by a large, curved claw, and followed after a large gap (twice length of terminal scansor) by a smaller pair of scansors; infero-median row of digital scales enlarged transversely, particularly towards the scansors, where the terminal scale adjoining the first pair of scansors may be medially constricted, swollen and scansor-like; 12 scale rows under 4th toe. Nostril pierced between rostral, 1 st supralabial and three nasal scales; the supranasal being much larger than the subequal postnasals and separated by a granule bordering the rostral, followed behind by two smaller granules. Rostral roughly rectangular but with its upper edges elongated due to extensions into the nostril. Nine supralabials on each side, the labial margin flexing dorsad at the rictus, with 1–2 minute scales proximal to the flexure. Nine infralabials on either side, with a small scale proximad to the flexure. At the lip, mental slightly narrower than adjacent infralabial, only 2/3rds the width of rostral, and in contact with two distinctly elongate postmental scales. Approximately 20 scales across crown at level of front of eye; 13 scales between nostril and front of eye; 23 scales from front of ear to back of eye; 99 scales around midbody. A roughly V-shaped row of 13 precloacal pores occurs eight scale rows anterior to the cloacal lip, with these scales more imbricate and triangular adjacent to the precloacal pores. Original portion of tail slightly dorsoventrally flattened and indistinctly verticillate, without obvious lateral constrictions; each verticil comprising 6–7 imbricate rows of scales dorsally and 5 imbricate scale rows ventrally, and with ventral scales approximately twice the size of those on dorsal surface. Hemipenal pouches well-developed, with an angular, ventro-lateral series of 3 enlarged tubercular scales on each side (largest nearest cloaca), and with two openings of the cloacal sacs on the posterior lip of the cloaca. Colour in life (from photographs, Fig. 4a and unpublished). Dark black-brown above with a series of five white dorsal spots (4–5 scales across) between the fore- and hind-limb insertions, that are surrounded by a chocolate brown zone that encompasses a series of 2–4 laterally arranged smaller pale spots (2–3 scales wide), and posteriorly by a diffuse pale region that forms an irregular band across the mid dorsum. Top of head, flanks and upper surfaces of limbs with small, scattered pale spots. Head with a dark brown band across the posterior edge of crown encompassing a small central pale spot; a vague, thin pale canthal stripe, extends on both sides from the nasal region to front of eye; upper and lower labials grey-brown with diffuse yellowish bars; iris reddish brown with a heavy dark brown reticulation, and pupil with crenulated edge. Original segment of tail mottled in dark brown and pale blotches; regenerated tail grey with vague darker blotches. Ventrum mottled grey-cream. Colour in preservative. Dark black-brown above with irregular markings on back, comprising four equally spaced dark bars, each with a central white spot 4–5 scales wide, and an irregular, paler brown bar behind; top and sides of head, flanks and upper surfaces of limbs with small, irregular pale spots; ventral surface of throat, neck, belly, limbs and tail uniform dark grey. Side of head and labials dark brown, with a few scattered pale spots on upper labials, and vague pale edges to lower labials. Variation in paratypes and additional material (Figs. 4b, 4c, 5; Tables 3–4). Morphology. SVL from 54.6 mm (ZMB 83289) to 61.2 mm (PEM 22220); head length 1.24–1.38 times head width (max. in paratypes 1.38); snout only 1.5 times diameter of eye in one specimen. The supranasals always separated by granules, usually with a single large granule in contact with the rostral between the supranasals, followed by 2–3 smaller granules in lateral contact; the first upper labial and rostral always enters the nostril, and the width of the rostral at the lip margin is always wider than that of the mental (except in ZMB 83290 were they are subequal); always two postmental scales; supralabials more than in holotype (10–11), infralabials (9–10); scales between anterior edge of eye 19–21; scales between nostril and anterior edge of orbit 12–13; scales between anterior edge of ear and rear margin of orbit 21–24; scales around mid-body 97–101; subdigital lamella beneath 4th toe 10–11; dorsal scales per tail verticel 7–8; ventral scales per tail verticel 5. In one female (ZMB 83290) the terminal, medially-constricted subdigital lamella of the 3rd toe of left foot is swollen and scansor-like, bearing small setae. Precloacal pores 8–13 in males, females without dimples in precloacal scales (as in A. transvaalica, but unlike A. loveridgei where five females had 5–9 dimpled precloacal scales, that lacked secretory activity, and only one had none). Colour in life (based on female paratype PEM R22229). Boldly patterned, with six dorsal dark transverse bands, posteriorly edged with cream, the first on the neck, the second and sixth at the fore- and hindlimb insertions, respectively, and with three thin bands equally spaced between them on the body; background colour of body, head and upper surfaces of limbs flecked on light brown and cream with occasional pale spots 2–4 scales wide; a dark canthal stripe runs through the rostral to the front of the eye, with a paler zone above; the scales of the anteriordorsal region of the orbital rim are bright yellow, with a faint yellowish infusion of the crown scales between the eyes; the original tail is boldly barred dorsally with seven, subequally-spaced, dark brown-black bars with a yellow posterior edge; belly uniform cream with scattered yellowish scales, greyer on ventral surfaces of limbs and tail. Colour in preservative. Similar to holotype, but dorsum pale brown to grey; larger dorsal markings usually reticulated, occasionally forming irregular transverse bands (6–7 from occiput to between hindlimbs); smaller dorsal markings vary from vague to distinct. A few specimens have irregular pale blotches along the posterior edges of the darker dorsal patches/bands, and also scattered on the dorsal surfaces of the limbs and back. Dorsal surface of regenerated tails usually with irregular pale and dark blotches that may form wavy bands, and which become flecks or grey blotches below. Size. Largest male (ZMB 83288)—SVL 56.5 mm, tail (partially regenerated) 55.7 mm; largest female (PEM R22220)—SVL 61.2 mm, tail (original) 64.0 mm. It is thus intermediate in size between A. transvaalica (malemax. SVL 72 mm, female—max. SVL 64 mm) and A. loveridgei (male—max. SVL 59 mm, female—max. SVL 58 mm). Only one of six specimens had an original tail (105% SVL). Hemipenis (based on ZMB 83292). Approximately 5 mm long, with a simple, smooth, possibly longitudinally flounced pedicel; capped with two large cups covered with fine calyculate ornamentation; sulcus simple and draining into the conjoined base of the distal cups. Distribution. Known from only three localities in the Mount Gorongosa region (Fig. 2). The holotype and paratypes, plus various non-collected individuals, have been found on the western flanks of Mount Gorongosa between about 900 and 1100 m a.s.l. (Figs. 6a–c). Access to further potential areas on the mountain, i.e. rock faces at the eastern slopes, was not possible due to security concerns. The additional material and various non-collected specimens have been observed at a lower altitude on the Bunga Inselberg (about 212 m a.s.l.) in the north-western corner of the National Park, approximately 40 km from the type locality (Fig. 6d). No tissue samples from the Bunga individuals were available, and the assignment of these individuals to the new species was thus based on morphological similarity. Natural history. The Gorongosa National Park is situated in Central Mozambique approx. 100 km south-west of the Zambezi River, and is in the southern part of the great African Rift Valley. Mount Gorongosa is an isolated granite mountain massif rising to 1863 m a.s.l., north-west of the National Park’s plains. At its highest elevations the massif’s plateau is covered with montane grasslands and forest patches dominated by cypresses (Widdringtonia nodiflora); the upper slopes of the mountain (900–1600 m) support a belt of moist evergreen forest. The type series was collected in an area between 900 and 1100 m on the western flanks of the mountain (Figs. 6a–c, 7). The paratypes were collected in a small alcove beneath a boulder pile in the closed-canopy riverine forest beside the Murombodzi Waterfall, and thus in comparatively cool and moist microclimates (Figs. 6c, 7). Some rocks with lizards present were located within the splash zone of the waterfall. Other specimens were observed but not collected at this site. The holotype was collected from a deep narrow crack in a large granite rock (i.e. 8 m ³) in highly degraded grassland below the edge of the rainforest and almost fully exposed to the sun (Fig. 6b). It was the only specimen seen at that site. At both sites the new species lived in syntopy with the cordylid Smaug mossambicus (FitzSimons, 1958). The additional non-type material comes from the Bunga Inselberg (Fig. 6d), a partly eroded inselberg composed mostly of trachyte in the north-western corner of the national park, covered and surrounded by dense Miombo woodland at a relatively low altitude (approx. 212 m a.s.l.). Here the geckos were observed during the day and at night in several places in deep cracks between very large boulders. No geckos were observed outside of these cracks. Syntopic lizards were Smaug mossambicus, the skink Trachylepis margaritifera (Peters, 1854) and the gecko Hemidactylus platycephalus Peters, 1854. Further sympatric, but not syntopic geckos were: Lygodactylus cf. capensis (Smith, 1849), Pachydactylus punctatus Peters, 1854 and Chondrodactylus turneri (Gray, 1864). The new species was observed regularly between April and December 2015. Like A. transvaalica, it is tolerant of conspecifics and frequently several specimens, and sometimes as many as over 20, were observed sheltering in deep rock crevices. One adult female (ZMB 83290) contained two eggs of approximately 11.5 x 7.5 mm. Conservation Status. The population of the new gecko present on the lower slopes of Mount Gorongosa is threatened by illegal deforestation taking place within the riverine forest adjacent to Murombodzi Waterfall. Slashand-burn removal of tall, old growth trees around the rocks may cause changes to the humidity and water availability in these currently very moist, shaded microhabitats that seem to be preferred by the lizards. Reforestation efforts led by Gorongosa National Park and introduction of shade-grown coffee as an alternative to slash-and-burn agriculture still practiced around the mountain will hopefully slow or halt the loss of the native riverine and evergreen vegetation in that area. On Bunga Inselberg the population of A. gorongosa is currently protected within the core area of the National Park and no immediate threat to its survival exists, though deforestation for agriculture is slowly encroaching on the area. However, based on the existence of only a few known sites and small area of occupancy, i.e. the presumed restriction to a fragmented range on Mount Gorongosa and potentially some rocky areas surrounding the mountain, as well as the ongoing high rate of habitat degradation and conversion on the mountain, the species may be considered highly threatened, potentially justifying an assessment as Endangered or even Critically Endangered. Further surveys and formal IUCN Red List assessment are required to confirm this.

Phrynobatrachus horsti sp. nov. Figs. 1���2 Holotype. ZMB 81698 (adult male; field # MBUR 3237), Republic of the Congo, Niari Department, 02�� 12 ��� 47.8 ������ S, 12 �� 50 ���06.0������ E, 28 September 2012, leg. M. Burger & A.G. Zassi-Boulou. Paratypes (all from Republic of the Congo, all collected by M. Burger & A.G. Zassi-Boulou). ZMB 81699 (adult male; field # MBUR 3098), Kouilou Department, Yombo River, 04�� 23 ��� 26.1 ������ S, 11 �� 40 ���26.0������ E, 27 July 2012; ZMB 81700 (adult male; field # MBUR 3122), ZMB 81701 (adult female; field # MBUR 3123), Kouilou Department, Yombo River, 04�� 23 ��� 26.1 ������ S, 11 �� 40 ���26.0������ E, 29 July 2012; ZMB 81702 (adult male; field # MBUR 3148), ZMB 81703 (adult female; field # MBUR 3149), Kouilou Department, Yombo River, 04�� 23 ��� 26.1 ������ S, 11 �� 40 ���26.0������ E, 31 July 2012; ZMB 81704 (adult female; field # MBUR 3236), Niari Department, 02�� 12 ��� 47.8 ������ S, 12 �� 50 ���06.0������ E, 28 September 2012; ZMB 81705 (adult male; field # MBUR 3286), Niari Department, 02�� 12 ��� 47.8 ������ S, 12 �� 50 ���06.0������ E, 30 September 2012; ZMB 81706 (adult male; field # MBUR 3357), Niari Department, 02�� 12 ��� 47.8 ������ S, 12 �� 50 ���06.0������ E, 4 October 2012; ZMB 81707 (adult female; field # MBUR 3541), Niari Department, 02�� 19 ��� 36.2 ������ S, 12 �� 47 ��� 32.7 ������ E, 26 October 2013; ZMB 81708 (adult male; field # MBUR 3591), Niari Department, 02�� 12 ��� 38.3 ������ S, 12 �� 49 ��� 56.6 ������ E, 19 October 2013; ZMB 81709 (adult female; field # MBUR 3610), ZMB 81710 (adult male; field # MBUR 03611), Niari Department, 02�� 19 ��� 36.2 ������ S, 12 �� 47 ��� 32.7 ������ E, 21 October 2013; ZMB 81711 (adult female; field # MBUR 3612), ZMB 81712 (adult male; field # MBUR 3613), Niari Department, 02�� 12 ��� 38.3 ������ S, 12 �� 49 ��� 56.6 ������ E, 20 October 2013; ZMB 81713 (adult male; field # MBUR 3641), Niari Department, 02�� 12 ��� 38.3 ������ S, 12 �� 49 ��� 56.6 ������ E, 21 October 2013; ZMB 81714 (adult female; field # MBUR 3634), ZMB 81715 (adult male; field # MBUR 3652), ZMB 81716 (adult female; field # MBUR 3653), ZMB 81717 (adult female; field # MBUR 3677), ZMB 81718 (adult male; field # MBUR 3678), Niari Department, 02�� 19 ��� 36.2 ������ S, 12 �� 47 ��� 32.7 ������ E, 26 October 2013; ZMB 81719 (adult male; field # MBUR 3681), ZMB 81720 (adult male; field # MBUR 3682), ZMB 81721 (adult male; field # MBUR 3683), Niari Department, 02�� 12 ��� 38.3 ������ S, 12 �� 49 ��� 56.6 ������ E, 23 October 2013; ZMB 81722 (adult female; field # MBUR 3696), ZMB 81723 (adult female; field # MBUR 3697), ZMB 81724 (adult male; field # MBUR 3698), ZMB 81725 (adult male; field # MBUR 3699), ZMB 81726 (adult female; field # MBUR 3732), ZMB 81727 (adult male; field # MBUR 3742), Niari Department, 02�� 12 ��� 38.3 ������ S, 12 �� 49 ��� 56.6 ������ E, 23 October 2013; ZMB 81728 (adult male; field # MBUR 3770), ZMB 81729 (adult male; field # MBUR 3771), Niari Department, 02�� 24 ���08.8������ S, 12 �� 59 ���08.0������ E, 14 November 2013; ZMB 81730 (adult male; field # MBUR 4005), ZMB 81731 (adult male; field # MBUR 4006), Niari Department, 02�� 24 ���08.8������ S, 12 �� 59 ���08.0������ E, 26 November 2013. Additional material deposited in IRSEN (all from the Republic of the Congo, all collected by M. Burger & A.G. Zassi-Boulou). MBUR 3028���3031, Kouilou Department, Yombo River, 04�� 23 ��� 26.1 ������ S, 11 �� 40 ���26.0������ E, 22 July 1012; MBUR 3038���3039, Kouilou Department, Yombo River, 04�� 23 ��� 33.1 ������ S, 11 �� 40 ���16.0������ E, 21 July 2012; MBUR 3063, 3099, 3103���3106, Kouilou Department, Yombo River, 04�� 23 ��� 26.1 ������ S, 11 �� 40 ���26.0������ E, 23, 26 and 27 July 2012; MBUR 3641, Niari Department, 02�� 12 ��� 38.3 ������ S, 12 �� 49 ��� 56.6 ������ E, 21 October 2013; MBUR 4005, Niari Department, 02�� 24 ���08.8������ S, 12 �� 59 ���08.0������ E, 26 November 2013. Diagnosis. The genetic data (16 S rRNA) identifies the new species as a member of the genus Phrynobatrachus. Morphologically this is confirmed by: the general body shape; the presence of a tarsal tubercle; and the presence of pedal webbing. From the morphologically similar Arthroleptis they differ by lacking a median dorsal skin raphe and a black tympanic spot. From juvenile Ptychadena they can be distinguished by the lack of parallel dorsal ridges. Compared to other Phrynobatrachus the new species can be identified by the combination of the following characters: having snout-vent lengths smaller than 20 mm; males lacking tooth like projections in the lower jaw; males having black throats with spines; lacking a spiny eyelid tubercle; possessing a distinct lateral black facial mask; lacking X-shaped dorsal ridges; possessing comma-shaped dorsal ridges; lacking distinct markings on the whitish coloured belly; having yellowish ventral colouration on hind limbs; showing reduced pedal webbing and small but distinct discs on fingers and toes; and by a unique 16 S rRNA sequence. Differential diagnosis. The new species differs from other western African Phrynobatrachus species by: a small compact body (P. acridoides, P. africanus, P. auritus, P. batesii, P. brongersmai, P. cornutus, P. cricogaster, P. danko, P. francisci, P. giorgii, P. intermedius, P. jimzimkusi, P. latifrons, P. liberiensis, P. plicatus, P. natalensis, P. njiomock, P. sandersoni, P. schioetzi, P. steindachneri); lacking tooth like projections in lower jaw of males (present in P. africanus and P. sandersoni); a black lateral face mask originating from at least the level of eyes and extending to below the supratympanic ridge (black face mask only shared with: P. auritus, P. batesii, P. chukuchuku, P. intermedius, P. plicatus, P. ruthbeateae; less distinct brown face masks: P. werneri; some P. rainerguentheri have entirely black heads); small, but distinct tympanum (tympanum absent according to original description: P. ogoensis; indistinct but visible in BMNH 1947.2.6.83); black axillary spots of varying size, but generally being small and irregular shaped (larger distinct axillary and/or lateral spots and bands: P. dispar, P. fraterculus, P. gutturosus, P. maculiventris, P. pintoi, P. ruthbeateae); males with a black throat (yellow in P. acridoides, P. alleni, P. batesii, P. fraterculus, P. latifrons; throat with large black or brown spots: P. brongersmai, P. cornutus, P. giorgii; white or whitish greyish: P. annulatus, P. dispar, P. hieroglyphicus), the entire males��� throat���but in particular the centre���beset with black spines (black spines on throat shared with: P. chukuchuku, P. danko, P. jimzimkusi, P. ruthbeateae, P. steindachneri; spines hyaline or white: P. hylaios, P. schioetzi; P. werneri; spines mostly along throat margins: P. leveleve, P. danko); belly of males dirty white, dark grey towards flanks (belly at least partly yellow in P. alleni, P. batesii; P. manengoubensis, P. werneri; orange to yellow: P. danko; black: P. chukuchuku; white or bluish with black pattern: P. villiersi; white with blackish spots or dots: P. annulatus, P. ghanensis, P. hylaios, P. maculiventris, P. pintoi; a black circle and median spot on white or yellow background���bull���s eye pattern: P. cricogaster); females with yellowish grey throat, belly dirty greyish (throat and belly with brown blotches: P. intermedius, P. jimzimkusi, P. njiomock, P. steindachneri); lower parts of hind limbs in both sexes uniform yellow (yellow colouration shared with: P. alleni, P. batesii, P. danko, P. manengoubensis, possibly also with P. sternfeldi; greyish/brownish to yellowish in P. ruthbeateae, P. schioetzi; vaguely dark blotched in P. hylaios); absence of spiny eyelid tubercles (spiny eyelid tubercle present in P. annulatus, P. calcaratus, P. cornutus, P. pintoi, P. taiensis, P. villiersi); comma-shaped scapular ridges converging towards vent (scapular ridges long and converging towards mid-body, almost X-shaped: P. alleni, P. auritus, P. brongersmai, P. jimzimkusi, P. plicatus, P. steindachneri; scapular ridges indistinct or absent: P. hieroglyphicus, P. intermedius, P. liberiensis, P. maculiventris, P. manengoubensis, P. tokba); short dorsolateral ridges often distinct (short dorsolateral ridges shared with: P. r u t h be a t e a e); dorsal skin granular (smooth: P. fraterculus, P. hieroglyphicus, P. maculiventris, most P. rainerguentheri, P. tokba; tuberculate: P. annulatus, P. calcaratus, P. cornutus, P. cricogaster, P. guineensis, P. sternfeldi, most P. guttursous, P. jimzimkusi, P. steindachneri, P. sternfeldi, P. taiensis, P. villiersi; the new species described below); ventral skin smooth in both sexes (ventral skin with asperities in both sexes: P. chukuchuku); absence of a blue band along the interior half of ventral shanks (present in P. auritus); absence of nuptial pads in males (nuptial pads or swollen thumb in males present: P. b at e s i i, P. e l b e r t i, P. guineensis, P. hieroglyphicus, P. maculiventris, P. ogoensis, P. werneri); distinct roundish discs on finger and toe tips (discs absent: most other species e.g. P. hieroglyphicus, P. intermedius, P. liberiensis, P. maculiventris, P. ogoensis, P. plicatus, P. rainerguentheri, P. tokba, P. werneri; discs heart-shaped: P. sandersoni); carpal gland in males lacking (carpal gland present: P. africanus); femoral glands absent (present in males: P. calcaratus, P. cornutus, P. fraterculus, P. leveleve, P. maculiventris, P. ogoensis, P. phyllophilus; present in both sexes: P. sandersoni); rudimentary but distinct webbing (webbing indistinct or absent: P. annulatus, P. calcaratus, P. chukuchuku, P. cornutus, P. dispar, P. fraterculus, P. gutturosus, P. hieroglyphicus, P. maculiventris, P. manengoubensis, P. tokba, P. villiersi; webbing more developed: P. acridoides, P. africanus, P. alleni, P. auritus, P. brongersmai, P. cricogaster, P. danko, P. elberti, P. francisci, P. giorgii, P. intermedius, P. jimzimkusi, P. latifrons, P. liberiensis, P. natalensis, P. njiomock, P. ogoensis, P. plicatus, P. steindachneri, P. sternfeldi); small round tarsal tubercle (numerous tarsal tubercles fused to tarsal ridge: P. cricogaster). Morphologically the new species can be only mistaken for P. ruthbeateae and P. ba t es i i. A detailed comparison with these two taxa is presented in the discussion. A comparison with the second new species described herein, is included in the diagnosis of the latter species. For further character states of western African Phrynobatrachus species see: Peters (1870); Boulenger (1906 a, b); Nieden (1910); Sternfeld (1917; Arthroleptis taeniatus; replacement name by Ahl 1924: A. sternfeldi; a Phrynobatrachus species according to Fr��tey 2008); de Witte (1921); Parker (1936); Angel (1940); Perret (1957, 1959, 1966); Guib�� (1959); Guib�� & Lamotte (1963); Lamotte & Xavier (1966 a, b); R��del (2000); R��del & Ernst (2002 b); R��del et al. (2005, 2009a, b, 2010, 2011, 2012 a, b); Uyeda et al. (2007); Zimkus (2009); Blackburn (2010); Blackburn & R��del (2011); and Zimkus & Gvožd��k (2012). Description of holotype [measures in mm]. Typical, small adult male Phrynobatrachus with short oval, compact body shape; snout-vent length: 14; short snout, rounded to slightly truncate in dorsal and rounded to slightly protruding in lateral view; canthus rostralis rounded; loreal region straight to very slightly convex; headwidth directly behind the eyes: 5; medium sized eyes positioned laterally; eye-diameter: 2.1; distance anterior corner of eye to nostril: 1.5; nares small and roundish to slightly vertically elliptical, positioned laterally; straight distance nostril-snout tip: 1.1; nostril closer to snout than to eye; inter-narial distance: 2.4, larger than inter-orbital distance: 1.8; pupil round; supratympanic ridge curving down from posterior corner of eye to forearm base; tympanum diameter: 1.1, with distinct tympanic annulus, annulus round with central round papilla; tympanum much smaller than diameter of eye; upper and lower maxillae and praemaxillae with minute teeth, hidden by lips; vomerine teeth absent; elongated, almost rectangular tongue, tip with distinct v-shaped notch, median conical papilla near anterior attachment of tongue; tongue densely beset with small round papillae; small round choanae positioned at lateral edges of mouth roof. Forelimbs slender; upper arm: 3.4; lower arm: 3.6; hand to tip of finger III: 4.6; hand with two large tubercles, small and oval thenar tubercle, large and round palmar tubercle; carpal gland absent; fingers with round, nondivided, protruding subarticular tubercles, no further tubercles on hand and fingers; relative finger length: IVColouration in ethanol after 1.5 years. Back and dorsal part of flanks more or less uniform brown; snout slightly clearer, dorsal with dark median spot, snout tip including anterior part of upper lip and loreal region light brown; clear spot between anterior part of eyelids; posterior part of inter-orbital space with dark bar; eyelids dark grey; lateral part of upper lip, sub-orbital region, supratympanic ridge and temporal area including tympanum dark brown to black, extending into a similarly coloured blotch on the anterior part of the upper arm; flanks posterior to forearm bases with large dark axillary blotch; brown flanks fainting ventrally into whitish belly; arms coloured as back, lower arm towards hand with two dark transversal bars; hands dorsally lighter coloured with dark crossbands; lower legs brown as back; thighs with 3-4 faintly indicated dark cross-bars; anterior part of thigh with dark longitudinal stripe, being the border to whitish ventral parts; lower leg dorsally with three (right) and four (left) dark cross-bars; feet and toes with dark cross-bands; dark blotch around vent, dorsally bordered by narrow yellowish line; lower parts of feet and hands greyish brown; remaining ventral parts of limbs and posterior part of belly uniform whitish; lateroventral parts of venter, anterior part of belly and pectoral region whitish but densely beset with minute black spots; throat and lower mandibles dark grey to black; lower mandibles with few small clear spots. Colouration in life (Fig. 1). Basic colour of dorsal surface of head, upper part of flank, back and dorsal surfaces of extremities light brown; snout, loreal region and anterior part of eyelids orange brown; snout with median longitudinal black spot; black inter-orbital bar; back with irregular darker patterning; lower arms, thighs, lower legs and feet with dark crossbars; anterior part of upper mandible brown with a dark bar; a V-shaped darker figure in scapular region, pointing anteriorly; black lateral face mask extending from below the eye and supratympanic ridge to forearm bases, covering tympanum; continuing below arms as a lateroventral black patch to axillar region; eye posterodorsally bordered by fine blue line; black pupil; iris copper red; central belly dirty white; belly laterally and pectoral area dark grey, no distinct black ventrolateral line or band; ventral part of forelimbs and hands dark grey; groin area and lower surfaces of thighs, lower leg and tarsus yellowish to orange; lower part of lower legs bordered by blackish spots; ventral surfaces of toes blackish. Variation (Figs. 1���2). Snout-vent length in males with spiny black throats (presumably indicating maturity) 13.5���15.8 mm and 13.1���19.7 mm in females (smallest one presumably not adult); body dimensions generally as in holotype, for small deviations see Table 1. The scapular ridges may differ in lengths and distinctness, but are always discernible and comma-shaped. Short dorsolateral ridges, composed of a very dense row of small warts, are visible in most specimens. Scapular and dorsolateral ridges are usually coloured dark. The webbing formulae showed some slight variation (Table 1). The black face mask may cover either the entire lateral parts of the head (sometimes with minute white spots; Figs. 2 c, d), extending from snout-tip to dorsal of the forearm insertion, or the parts anterior to the eye are coloured in clearer brown (Figs. 1, 2), sometimes with darker spots. Individuals with clearer coloured loreal regions also have dark brown upper mandible with black bars instead of entirely black mandibles. The back might be uniform dark or reddish brown, show some darker patterning, or carry a broad central dark band (Fig. 2 a). A light inter-orbital stripe is usually present. The iris colour varies from golden to copper red. The flank might be either uniform brown, grey or mottled with darker dots and lines (Fig. 2 c). The black axillary spot may be distinct and extending as far caudally as to about mid-body. Usually, the edges of this spot are irregularly shaped. In other individuals it is nothing but a rather indistinct mottling of blackish spots with clearer dots. We neither observed in males, nor in females distinct black bordering of belly and lower shanks as in P. ruthbeateae. The inguinal region, Published as part of R��del, Mark-Oliver, Burger, Marius, Zassi-Boulou, Ange Ghislain, Emmrich, Mike, Penner, Johannes & Barej, Michael F., 2015, Two new Phrynobatrachus species (Amphibia: Anura: Phrynobatrachidae) from the Republic of the Congo, pp. 55-80 in Zootaxa 4032 (1) on pages 56-66, DOI: 10.11646/zootaxa.4032.1.3, http://zenodo.org/record/289765, {"references":["Peters, W. (1870) Uber neue Amphibien (Hemidactylus, Urosaura, Tropidolepisma, Geophis, Uriechis, Scaphiophis, Hoplocephalus, Rana, Entomoglossus, Cystignathus, Hylodes, Arthroleptis, Phyllobates, Cophomantis) des Koniglich zoologischen Museums. Monatsberichte der Koniglichen Akademie der Wissenschaften zu Berlin, 1870 (August), 641 - 652 + pls. 1 - 2.","Boulenger, G. A. (1906 a) Report on the batrachians collected by the late L. Fea in West Africa. Annali del Museo Civico di Storia Naturale di Genova, 42, 157 - 172 + 2 plates.","Boulenger, G. A. (1906 b) Descriptions of new batrachians discovered by Mr. G. L. Bates in South Cameroon. Annals and Magazine of Natural History, 17 (17), 317 - 323.","Sternfeld, R. (1917) Lieferung 11, Reptilia und Amphibia. In: Schubotz, H. (Ed.), Ergebnisse der zweiten deutschen Zentral- Afrika-Expedition 1910 - 1911 unter Fuhrung Adolf Friedrichs, Herzog zu Mecklenburg. Band I. Zoologie. Verlag Klinkhardt & Biermann, Leipzig, pp. 407 - 502 + pls. XXII-XXIV.","Ahl, E. (1924) Zur Kenntnis der Froschfauna Afrikas. Zoologischer Anzeiger, 58, 81 - 82.","Fretey, T. (2008) Revue des genres Africains Arthroleptis Smith, 1849 et Phrynobatrachus Gunther, 1862 (Amphibia, Anura). Alytes, 25, 99 - 172.","Witte, de G. F. (1921) Description de batraciens nouveaux du Congo Belge. Revue Zoologique Africaine, 9, 1 - 21 + pls. I - V.","Parker, H. W. (1936) Amphibians from Liberia and the Gold Coast. Zoologische Mededelingen, 19, 87 - 102.","Angel, F. (1940) Description de trois amphibiens nouveaux du Cameroun, materiaux de la Mission P. Lepesme, R. Paulian et A. Villiers (2 e note). Bulletin du Museum national d'Histoire naturelle, Serie 2, 12, 238 - 243.","Perret, J. - L. (1957) Un nouveau Phrynobatrachus du Cameroun. Revue Suisse de Zoologie, 64, 527 - 531. http: // dx. doi. org / 10.5962 / bhl. part. 75500","Guibe, J. (1959) Description d'un batracien nouveau de Cote d'Ivoire: Phrynobatrachus villiersi n. sp. Bulletin du Museum National d'Histoire Naturelle, Serie A, 31, 134 - 136.","Guibe, J. & Lamotte, M. (1963) La reserve naturelle integrale du Mont Nimba. XXVIII. Batraciens du genre Phrynobatrachus. Memoires d'Institute fondamental d'Afrique noire, 66, 601 - 627.","Lamotte, M. & Xavier, F. (1966 a) Phrynobatrachus natalensis (Smith) et Phrynobatrachus francisci (Boulenger): deux especes de l'Ouest Africain difficiles a distinguer. Bulletin du Museum National d'Histoire Naturelle, Serie A, 28, 344 - 361.","Lamotte, M. & Xavier, F. (1966 b) Etude comparee de deux especes de Phrynobatrachus souvent confondues: Phr. plicatus Gunther et Phr. auritus Boulenger. Bulletin du Museum National d'Histoire Naturelle, Serie A, 28, 1605 - 1619.","Rodel, M. - O. (2000) Herpetofauna of West Africa. Vol. I. Amphibians of the West African savanna. Edition Chimaira, Frankfurt / M., 335 pp.","Rodel, M. - O. & Ernst, R. (2002 b) A new Phrynobatrachus from the Upper Guinean rain forest, West Africa, including a description of a new reproductive mode for the genus. Journal of Herpetology 36, 561 - 571.","Uyeda, J. C., Drewes, R. C. & Zimkus, B. M. (2007) The California Academy of Sciences Gulf of Guinea Expeditions (2001, 2006) VI. A new species of Phrynobatrachus from the Gulf of Guinea Islands and a reanalysis of Phrynobatrachus dispar and P. feae (Anura: Phrynobatrachidea). Proceedings of the California Academy of Sciences, 58 (18), 367 - 385.","Zimkus, B. (2009) Biogeographical analysis of Cameroonian puddle frogs and description of a new species of Phrynobatrachus (Anura: Phrynobatrachidae) endemic to Mount Oku, Cameroon. Zoological Journal of the Linnean Society, 157, 795 - 813. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.2009.00579. x","Blackburn, D. C. (2010) A new puddle frog (Phrynobatrachidae: Phrynobatrachus) from the Mambilla Plateau in eastern Nigeria. African Journal of Herpetology, 59, 33 - 52. http: // dx. doi. org / 10.1080 / 04416651003742160","Blackburn, D. C. & Rodel, M. - O. (2011) A new puddle frog (Phrynobatrachidae: Phrynobatrachus) from the Obudu Plateau in eastern Nigeria. Herpetologica, 67, 271 - 287. http: // dx. doi. org / 10.1655 / HERPETOLOGICA-D- 10 - 00046.1","Zimkus, B. M. & Gvozdik, V. (2013) Sky Islands of the Cameroon Volcanic Line: a diversification hot spot for puddle frogs (Phrynobatrachidae: Phrynobatrachus). Zoologica Scripta, 42, 591 - 611. http: // dx. doi. org / 10.1111 / zsc. 12029","Zimkus, B. M. & Larson, J. G. (2013 a) Phrynobatrachus ruthbeatae (Ruth-Beate's Puddle Frog). Gabon: Haut Ogooe Province: Bateke Plateau National Park. Herpetological Review, 44, 104.","Zimkus, B. M. & Larson, J. G. (2013 b) Assessment of the amphibians of Bateke Plateau National Park, Gabon, including results of chytrid pathogen tests. Salamandra, 49, 159 - 170.","Burger, M., Branch, W. R. & Channing, A. (2004) Amphibians and reptiles of Monts Doudou, Gabon: species turnover along an elevational gradient. In: Fisher, B. L. (Ed.), Monts Doudou, Gabon: a floral and faunal inventory with reference to elevational variation, California Academy of Sciences Memoir, 28, pp. 145 - 186.","Rodel, M. - O., Doherty-Bone, T., Talla Kouete, M., Janzen, P., Garrett, K., Browne, R., Gonwouo, N. L., Barej, M. F. & Sandberger, L. (2012 a) A new small Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae) from southern Cameroon. Zootaxa, 3431, 54 - 68.","Rodel, M. - O., Rudolf, V. H. W., Frohschammer, S. & Linsenmair, K. E. (2004) Life history of a West African tree-hole breeding frog, Phrynobatrachus guineensis Guibe & Lamotte, 1961 (Amphibia: Anura: Petropedetidae). In: Lehtinen, R. M. (Ed.), Ecology and evolution of phytotelm-breeding anurans, Miscellaneous Publications of the Museum of Zoology, University of Michigan, No. 193, pp. 31 - 44,","Channing, A. & Howell, K. M. (2006) Amphibians of East Africa. Cornell University Press, New York, 418 pp.","Kouame, N. G., Tohe, B., Assemian, N. E., Gourene, G. & Rodel, M. - O. (2008) Prey composition of two syntopic Phrynobatrachus species in the swamp forest of Banco National Park, Ivory Coast. Salamandra, 44, 177 - 186."]}

Phrynobatrachus mayokoensis sp. nov. Fig. 6 Holotype. ZMB 81732 (female; field # MBUR 3833), Republic of the Congo, Niari Department, Lekoumou, about 23 km E of Tsinguidi, about 4 km SE of Mayoko Post��, start of path to site 20, 740 m a.s.l., 02�� 26 ���05.6������ S, 12 �� 56 ��� 16.9 ������ E, 18 November 2013, coll. M. Burger & A.G. Zassi-Boulou. Paratype. ZMB 81733 (female, field # MKO 001), Republic of the Congo, Niari Department, Mount Lekoumou, about 4 km south-east of Mayoko Post��, 704 m a.s.l., 02�� 19 ���04.5������ S, 12 �� 49 ��� 34.5 ������ E, 31 March 2012, coll. A.G. Zassi-Boulou & P. Moukala. Diagnosis. The genetic data (16 S rRNA) characterize the new species as a member of the genus Phrynobatrachus. Morphologically this is confirmed by: the general body shape and presence of a tarsal tubercle. From the morphologically similar Arthroleptis they differ by lacking a median dorsal skin raphe and a black tympanic spot; from juvenile Ptychadena they can be distinguished by lacking parallel dorsal ridges and pedal webbing. The new species can be distinguished from other Phrynobatrachus by the following combination of characters: small size; compact body shape; short round snout; absence of a black lateral mask, absence of a spiny eyelid tubercle; females with conspicuous ventral pattern with a white throat carrying the tip of a dark figure originating in the pectoral region, belly white; very conspicuous red blotch on anterior craniad bases of thighs, bordered by black line; short comma-shaped scapular ridges diverging posteriorly; dorsal skin tuberculate, warts particularly abundant and spiny on rear of back; flanks with some larger warts; absence of discs on finger and toe tips; no webbing and a unique 16 S rRNA sequence. Differential diagnosis. The new species differs from other western African Phrynobatrachus species by: body size being distinctly smaller than 20 mm (species with adults larger than 20 mm: P. acridoides, P. africanus, P. auritus, P. batesii, P. brongersmai, P. cornutus, P. cricogaster, P. danko, P. francisci, P. giorgii, P. intermedius, P. jimzimkusi, P. latifrons, P. liberiensis, P. plicatus, P. natalensis, P. njiomock, P. sandersoni, P. schioetzi, P. steindachneri); the absence of a spiny eyelid tubercles (spiny eyelid tubercle present in P. annulatus, P. calcaratus, P. cornutus, P. pintoi, P. taiensis, P. villiersi); absence of a dark lateral face mask (black face mask present: P. auritus, P. batesii, P. chukuchuku, P. horsti sp. nov., P. intermedius, P. plicatus, P. ruthbeateae; less distinct brown face mask: P. w e r ne r i); tympanum present���although not very distinct (tympanum absent according to original description: P. ogoensis; indistinct but visible in BMNH 1947.2.6.83); distinct short, chevron shaped pair of scapular ridges (scapular ridges long and converging towards mid-body, almost X-shaped: P. alleni, P. auritus, P. brongersmai, P. jimzimkusi, P. plicatus, P. steindachneri; scapular ridges indistinct or absent: P. hieroglyphicus, P. intermedius, P. liberiensis, P. maculiventris, P. manengoubensis, P. tokba); very tuberculate dorsal skin (smooth skin: P. fraterculus, P. hieroglyphicus, P. maculiventris, P. manengoubensis, P. tokba; almost smooth, scapular ridges present: P. rainerguentheri, P. schioetzi); ventral skin smooth (ventral skin with asperities: P. chukuchuku); absence of a very distinct large black axillary blotch or black band on flanks���anterior flank only with indistinct black pattern (distinct black blotches and/or bands on flank: P. dispar, P. fraterculus, P. gutturosus, P. maculiventris, P. pintoi, P. ruthbeateae; some P. cornutus); venter white with a dark triangle originating in pectoral region and tip extending to posterior part of throat (belly at least partly yellow in P. batesii, P. werneri; orange to yellow: P. danko; white or bluish with black pattern: P. villiersi; white with blackish spots or dots: P. annulatus, P. ghanensis, P. hylaios, P. maculiventris, P. pintoi; a black circle and median spot on white or yellow backgroundbull���s eye pattern: P. cricogaster; white with brown mottling: P. intermedius, P. jimzimkusi, P. njiomock, P. steindachneri); lower parts of hind limbs white with some dark patterning at the edges (yellow: P. alleni, P. batesii, P. danko, P. manengoubensis; greyish/brownish to yellowish in P. ruthbeateae, P. schioetzi; vaguely dark blotched in P. hylaios); absence of a blue band along the interior half of ventral shanks (present in P. auritus); presence of a distinct red spot at the anterio-dorsal bases of thighs, bordered by black line (not known in any other Phrynobatrachus); absence of pedal webbing (webbing well developed: P. acridoides, P. africanus, P. alleni, P. auritus, P. brongersmai, P. cricogaster, P. danko, P. elberti, P. francisci, P. giorgii, P. intermedius, P. jimzimkusi, P. latifrons, P. liberiensis, P. natalensis, P. njiomock, P. ogoensis, P. plicatus, P. rainerguentheri, P. steindachneri, P. sternfeldi); no discs on toe and finger tips (discs present, roundish: P. cornutus, P. guineensis, P. horsti, P. leveleve, P. phyllophilus, P. ruthbeateae, P. sternfeldi; discs heart shaped: P. sandersoni); small round tarsal tubercle (numerous tarsal tubercles fused to tarsal ridge: P. cricogaster). A detailed comparison with the genetically most similar species is given in the discussion. For further character states of western African Phrynobatrachus species see the literature cited at the end of the differential diagnosis paragraph for P. horsti sp. nov. Description of holotype [measures in mm]. Typical, small female Phrynobatrachus with short roundish, compact body shape; snout-vent length: 15.9; short snout, rounded in dorsal and lateral view; canthus rostralis rounded; loreal region straight; head-width directly behind the eyes: 5.5; large eyes positioned laterally; eyediameter: 2.1; distance anterior corner of eye to nostril: 1.5; nares small and round, positioned laterally; straight distance nostril-snout tip: 0.9; nostril much closer to snout than to eye; inter-narial distance: 2.0, much larger than inter-orbital distance (distance between eyelids): 1.1; pupil round; supratympanic ridge curving down from slightly ventrad of posterior corner of eye towards forearm bases (in preservation indistinct and vanishing just posterior to tympanum); tympanum indistinct, diameter: 0.9, no tympanic annulus; tympanum much smaller than diameter of eye; upper maxillae with minute teeth, hidden by lips; vomerine teeth absent; broad, almost heart shaped tongue, tip broad and almost straight, median conical papilla near tongue tip; tongue smooth to slightly granular; small round choanae positioned at anterior-lateral edges of mouth roof. Forelimbs long and slender; upper arm: 3.9; lower arm: 3.7; hand to tip of finger III: 3.8; hand with two large tubercles, oval thenar tubercle, large and round palmar tubercle; fingers with round, non-divided, protruding subarticular tubercles, no further tubercles on hand and fingers; relative finger length: IV���II=IColouration in ethanol after one year. Back and dorsal part of flanks and extremities light brown; snout almost uniform light brown; upper eyelids dark grey; interorbital area mottled dark brown; back posterior to scapular region with dark brown triangle, pointing towards head; posterior third of back slightly darker, large warts almost black; remaining back and flanks with few smaller and irregular shaped darker patches; lower arms with two indistinct dark cross-bars; anterio-dorsal part of femur���bordering groin area���with lighter area (red in life, see below), bordered by black line; femur and tibia with one distinct and 1-2 indistinct darker crossbars; foot and toes with dark crossbars; upper lip with three dark blotches on either side; tympanum carrying dark blotch; lower lip white with four dark blotches on either side; throat white with a black triangular figure pointing towards mouth tip, originating on level with forearm bases; belly white, anterior half with small round dark spots (two on right, three on left side); lower parts of extremities white, edges with greyish specking; lower part of thigh with some dark mottling. Colouration in life (Fig. 6). Basic colour of head, anterior two thirds of back, flanks and anterior extremities reddish brown; anterior part of eyelids and interorbital space with dark pattern; posterior parts of the two chevron like scapular ridges enclose a medial dark figure on the back; anterior part of flanks darker brown; upper lip with three dark patches on each side; tympanum with dark spot; upper two-thirds of iris brownish-golden, lower third greyish; bases of forearms almost white; elbow and lower arm with dark mottling. Basic colour of posterior third of back and dorsal parts of hind limbs dark brown; thighs, tibiae and feet with dark crossbars; warts on posterior part of back and lower legs reddish brown; dorso-craniad bases of thighs with large red blotch, bordered by distinct black line. Ventral colouration as in preservation. Variation. Paratype in bad state of preservation, flattened; right foot missing, right thumb short and fat (presumably healed injury); otherwise almost identical to holotype; back with slightly more dark patterning, crossbaring on extremities more pronounced; ventral pattern as in holotype. Measures are summarized in Table 4. Males are currently unknown. Genetics. Compared to 40 West African and Central African species of the genus Phrynobatrachus, P. mayokoensis sp. nov. exhibited a mean difference between 1.34���16.98 % in the sampled region of the 16 S rRNA gene (Table 5). The genetically most similar sample was a frog determined as P. ogoensis by Larson & Zimkus (2015; MCZ A- 149217, GenBank: KP 247505; the second most similar species being P. dispar with a mean difference of 5.27 %). However, a comparison of the new species with two P. ogoensis syntypes (BMNH 1947.2.6.83- 84) revealed that P. ogoensis is not conspecific with P. mayokoensis sp. nov. (see discussion). It thus seemed, that the Gabonese frog is either conspecific with P. mayokoensis sp. nov. or represents a further undescribed taxon. The intraspecific difference between the two P. mayokoensis specimens was 0.20 %. From the morphologically most similar species, P. cornutus, P. mayokoensis sp. nov. differed by 7.33���7.53 %. Genetically, Phrynobatrachus mayokoensis sp. nov. is most similar to various West and Central Africa species of the clade B sensu Zimkus et al. (2010). Distribution. Phrynobatrachus mayokoensis sp. nov. is only known from two localities in the Mayoko District of the Niari Department, south-western Republic of the Congo, close to the border with Gabon (Fig. 4). It might also occur in the Ogoou��-Ivindo Province of Gabon (Larson & Zimkus 2015; compare above). Habitat and natural history. The mid altitude habitat (740 m a.s.l.) of the holoytpe is shown in Figure 7 a. There it was encountered during the day (3: 15 pm), being active on the forest floor, within 20 m of a road. The nearest wetland was a stream about 180 m away at an altitude of 673 m. The habitat of the paratype (Fig. 7 b) likewise consisted of mid-altitude rainforest (704 m a.s.l.) on a slope. There were no wetlands within 100 m. At both sites the forest habitat seemed relatively dry compared to forest in the surrounding region. It seems possible that the species is reproducing independently of water or in small water bodies like tree holes and water filled snail shells (compare e.g. R��del & Ernst 2002 a, b; R��del et al. 2004). Etymology. The species��� name refers to the type locality situated in the Mayoko District of the Niari Department, Republic of the Congo. The specific epithet is considered an invariable noun in apposition to the generic name. The suggested English name for this new species is Mayoko Puddle Frog., Published as part of R��del, Mark-Oliver, Burger, Marius, Zassi-Boulou, Ange Ghislain, Emmrich, Mike, Penner, Johannes & Barej, Michael F., 2015, Two new Phrynobatrachus species (Amphibia: Anura: Phrynobatrachidae) from the Republic of the Congo, pp. 55-80 in Zootaxa 4032 (1) on pages 67-72, DOI: 10.11646/zootaxa.4032.1.3, http://zenodo.org/record/289765, {"references":["Larson, J. C. & Zimkus, B. M. (2015) Phrynobatrachus ogoensis (Ogooue River Frog). Gabon: Ogooue-Ivindo Province. Herpetological Review, 46, 57.","Rodel, M. - O. & Ernst, R. (2002 a) A new reproductive mode for the genus Phrynobatrachus: Phrynobatrachus alticola has nonfeeding, nonhatching tadpoles. Journal of Herpetology, 36, 121 - 125.","Rodel, M. - O. & Ernst, R. (2002 b) A new Phrynobatrachus from the Upper Guinean rain forest, West Africa, including a description of a new reproductive mode for the genus. Journal of Herpetology 36, 561 - 571.","Rodel, M. - O., Rudolf, V. H. W., Frohschammer, S. & Linsenmair, K. E. (2004) Life history of a West African tree-hole breeding frog, Phrynobatrachus guineensis Guibe & Lamotte, 1961 (Amphibia: Anura: Petropedetidae). In: Lehtinen, R. M. (Ed.), Ecology and evolution of phytotelm-breeding anurans, Miscellaneous Publications of the Museum of Zoology, University of Michigan, No. 193, pp. 31 - 44,"]}

Rödel, Mark-Oliver, Burger, Marius, Zassi-Boulou, Ange Ghislain, Emmrich, Mike, Penner, Johannes, Barej, Michael F. (2015): Two new Phrynobatrachus species (Amphibia: Anura: Phrynobatrachidae) from the Republic of the Congo. Zootaxa 4032 (1): 55-80, DOI: http://dx.doi.org/10.11646/zootaxa.4032.1.3

Phrynobatrachus mayokoensis sp. nov. Fig. 6 Holotype. ZMB 81732 (female; field # MBUR 3833), Republic of the Congo, Niari Department, Lekoumou, about 23 km E of Tsinguidi, about 4 km SE of Mayoko Posté, start of path to site 20, 740 m a.s.l., 02° 26 ’05.6’’ S, 12 ° 56 ’ 16.9 ’’ E, 18 November 2013, coll. M. Burger & A.G. Zassi-Boulou. Paratype. ZMB 81733 (female, field # MKO 001), Republic of the Congo, Niari Department, Mount Lekoumou, about 4 km south-east of Mayoko Posté, 704 m a.s.l., 02° 19 ’04.5’’ S, 12 ° 49 ’ 34.5 ’’ E, 31 March 2012, coll. A.G. Zassi-Boulou & P. Moukala. Diagnosis. The genetic data (16 S rRNA) characterize the new species as a member of the genus Phrynobatrachus. Morphologically this is confirmed by: the general body shape and presence of a tarsal tubercle. From the morphologically similar Arthroleptis they differ by lacking a median dorsal skin raphe and a black tympanic spot; from juvenile Ptychadena they can be distinguished by lacking parallel dorsal ridges and pedal webbing. The new species can be distinguished from other Phrynobatrachus by the following combination of characters: small size; compact body shape; short round snout; absence of a black lateral mask, absence of a spiny eyelid tubercle; females with conspicuous ventral pattern with a white throat carrying the tip of a dark figure originating in the pectoral region, belly white; very conspicuous red blotch on anterior craniad bases of thighs, bordered by black line; short comma-shaped scapular ridges diverging posteriorly; dorsal skin tuberculate, warts particularly abundant and spiny on rear of back; flanks with some larger warts; absence of discs on finger and toe tips; no webbing and a unique 16 S rRNA sequence. Differential diagnosis. The new species differs from other western African Phrynobatrachus species by: body size being distinctly smaller than 20 mm (species with adults larger than 20 mm: P. acridoides, P. africanus, P. auritus, P. batesii, P. brongersmai, P. cornutus, P. cricogaster, P. danko, P. francisci, P. giorgii, P. intermedius, P. jimzimkusi, P. latifrons, P. liberiensis, P. plicatus, P. natalensis, P. njiomock, P. sandersoni, P. schioetzi, P. steindachneri); the absence of a spiny eyelid tubercles (spiny eyelid tubercle present in P. annulatus, P. calcaratus, P. cornutus, P. pintoi, P. taiensis, P. villiersi); absence of a dark lateral face mask (black face mask present: P. auritus, P. batesii, P. chukuchuku, P. horsti sp. nov., P. intermedius, P. plicatus, P. ruthbeateae; less distinct brown face mask: P. w e r ne r i); tympanum present—although not very distinct (tympanum absent according to original description: P. ogoensis; indistinct but visible in BMNH 1947.2.6.83); distinct short, chevron shaped pair of scapular ridges (scapular ridges long and converging towards mid-body, almost X-shaped: P. alleni, P. auritus, P. brongersmai, P. jimzimkusi, P. plicatus, P. steindachneri; scapular ridges indistinct or absent: P. hieroglyphicus, P. intermedius, P. liberiensis, P. maculiventris, P. manengoubensis, P. tokba); very tuberculate dorsal skin (smooth skin: P. fraterculus, P. hieroglyphicus, P. maculiventris, P. manengoubensis, P. tokba; almost smooth, scapular ridges present: P. rainerguentheri, P. schioetzi); ventral skin smooth (ventral skin with asperities: P. chukuchuku); absence of a very distinct large black axillary blotch or black band on flanks—anterior flank only with indistinct black pattern (distinct black blotches and/or bands on flank: P. dispar, P. fraterculus, P. gutturosus, P. maculiventris, P. pintoi, P. ruthbeateae; some P. cornutus); venter white with a dark triangle originating in pectoral region and tip extending to posterior part of throat (belly at least partly yellow in P. batesii, P. werneri; orange to yellow: P. danko; white or bluish with black pattern: P. villiersi; white with blackish spots or dots: P. annulatus, P. ghanensis, P. hylaios, P. maculiventris, P. pintoi; a black circle and median spot on white or yellow backgroundbull’s eye pattern: P. cricogaster; white with brown mottling: P. intermedius, P. jimzimkusi, P. njiomock, P. steindachneri); lower parts of hind limbs white with some dark patterning at the edges (yellow: P. alleni, P. batesii, P. danko, P. manengoubensis; greyish/brownish to yellowish in P. ruthbeateae, P. schioetzi; vaguely dark blotched in P. hylaios); absence of a blue band along the interior half of ventral shanks (present in P. auritus); presence of a distinct red spot at the anterio-dorsal bases of thighs, bordered by black line (not known in any other Phrynobatrachus); absence of pedal webbing (webbing well developed: P. acridoides, P. africanus, P. alleni, P. auritus, P. brongersmai, P. cricogaster, P. danko, P. elberti, P. francisci, P. giorgii, P. intermedius, P. jimzimkusi, P. latifrons, P. liberiensis, P. natalensis, P. njiomock, P. ogoensis, P. plicatus, P. rainerguentheri, P. steindachneri, P. sternfeldi); no discs on toe and finger tips (discs present, roundish: P. cornutus, P. guineensis, P. horsti, P. leveleve, P. phyllophilus, P. ruthbeateae, P. sternfeldi; discs heart shaped: P. sandersoni); small round tarsal tubercle (numerous tarsal tubercles fused to tarsal ridge: P. cricogaster). A detailed comparison with the genetically most similar species is given in the discussion. For further character states of western African Phrynobatrachus species see the literature cited at the end of the differential diagnosis paragraph for P. horsti sp. nov. Description of holotype [measures in mm]. Typical, small female Phrynobatrachus with short roundish, compact body shape; snout-vent length: 15.9; short snout, rounded in dorsal and lateral view; canthus rostralis rounded; loreal region straight; head-width directly behind the eyes: 5.5; large eyes positioned laterally; eyediameter: 2.1; distance anterior corner of eye to nostril: 1.5; nares small and round, positioned laterally; straight distance nostril-snout tip: 0.9; nostril much closer to snout than to eye; inter-narial distance: 2.0, much larger than inter-orbital distance (distance between eyelids): 1.1; pupil round; supratympanic ridge curving down from slightly ventrad of posterior corner of eye towards forearm bases (in preservation indistinct and vanishing just posterior to tympanum); tympanum indistinct, diameter: 0.9, no tympanic annulus; tympanum much smaller than diameter of eye; upper maxillae with minute teeth, hidden by lips; vomerine teeth absent; broad, almost heart shaped tongue, tip broad and almost straight, median conical papilla near tongue tip; tongue smooth to slightly granular; small round choanae positioned at anterior-lateral edges of mouth roof. Forelimbs long and slender; upper arm: 3.9; lower arm: 3.7; hand to tip of finger III: 3.8; hand with two large tubercles, oval thenar tubercle, large and round palmar tubercle; fingers with round, non-divided, protruding subarticular tubercles, no further tubercles on hand and fingers; relative finger length: IV≤II=I Dorsal skin tuberculate (in life) to mostly smooth (in preservation); a pair of chevron shaped warts on neck, the anterior parts of these warts shorter, parallel to body and to each other, the posterior parts of the warts longer, extending towards flanks; flanks and anterior part of back with smaller and larger flat warts (not visible in preservation); posterior third of back with distinct and pointed warts (in preservation: round); eyelid, interorbital space, dorsal parts of thighs towards knees and lower legs with many distinct and pointed warts (in preservation round: eyelids, or invisible: extremities); spiny eyelid tubercle absent; skin on lower parts of head, limbs, belly and pectoral region smooth. Sex determined via dissection: few small, white ovarian eggs, presumably not fully developed. Colouration in ethanol after one year. Back and dorsal part of flanks and extremities light brown; snout almost uniform light brown; upper eyelids dark grey; interorbital area mottled dark brown; back posterior to scapular region with dark brown triangle, pointing towards head; posterior third of back slightly darker, large warts almost black; remaining back and flanks with few smaller and irregular shaped darker patches; lower arms with two indistinct dark cross-bars; anterio-dorsal part of femur—bordering groin area—with lighter area (red in life, see below), bordered by black line; femur and tibia with one distinct and 1-2 indistinct darker crossbars; foot and toes with dark crossbars; upper lip with three dark blotches on either side; tympanum carrying dark blotch; lower lip white with four dark blotches on either side; throat white with a black triangular figure pointing towards mouth tip, originating on level with forearm bases; belly white, anterior half with small round dark spots (two on right, three on left side); lower parts of extremities white, edges with greyish specking; lower part of thigh with some dark mottling. Colouration in life (Fig. 6). Basic colour of head, anterior two thirds of back, flanks and anterior extremities reddish brown; anterior part of eyelids and interorbital space with dark pattern; posterior parts of the two chevron like scapular ridges enclose a medial dark figure on the back; anterior part of flanks darker brown; upper lip with three dark patches on each side; tympanum with dark spot; upper two-thirds of iris brownish-golden, lower third greyish; bases of forearms almost white; elbow and lower arm with dark mottling. Basic colour of posterior third of back and dorsal parts of hind limbs dark brown; thighs, tibiae and feet with dark crossbars; warts on posterior part of back and lower legs reddish brown; dorso-craniad bases of thighs with large red blotch, bordered by distinct black line. Ventral colouration as in preservation. Variation. Paratype in bad state of preservation, flattened; right foot missing, right thumb short and fat (presumably healed injury); otherwise almost identical to holotype; back with slightly more dark patterning, crossbaring on extremities more pronounced; ventral pattern as in holotype. Measures are summarized in Table 4. Males are currently unknown. Genetics. Compared to 40 West African and Central African species of the genus Phrynobatrachus, P. mayokoensis sp. nov. exhibited a mean difference between 1.34–16.98 % in the sampled region of the 16 S rRNA gene (Table 5). The genetically most similar sample was a frog determined as P. ogoensis by Larson & Zimkus (2015; MCZ A- 149217, GenBank: KP 247505; the second most similar species being P. dispar with a mean difference of 5.27 %). However, a comparison of the new species with two P. ogoensis syntypes (BMNH 1947.2.6.83- 84) revealed that P. ogoensis is not conspecific with P. mayokoensis sp. nov. (see discussion). It thus seemed, that the Gabonese frog is either conspecific with P. mayokoensis sp. nov. or represents a further undescribed taxon. The intraspecific difference between the two P. mayokoensis specimens was 0.20 %. From the morphologically most similar species, P. cornutus, P. mayokoensis sp. nov. differed by 7.33–7.53 %. Genetically, Phrynobatrachus mayokoensis sp. nov. is most similar to various West and Central Africa species of the clade B sensu Zimkus et al. (2010). Distribution. Phrynobatrachus mayokoensis sp. nov. is only known from two localities in the Mayoko District of the Niari Department, south-western Republic of the Congo, close to the border with Gabon (Fig. 4). It might also occur in the Ogooué-Ivindo Province of Gabon (Larson & Zimkus 2015; compare above). Habitat and natural history. The mid altitude habitat (740 m a.s.l.) of the holoytpe is shown in Figure 7 a. There it was encountered during the day (3: 15 pm), being active on the forest floor, within 20 m of a road. The nearest wetland was a stream about 180 m away at an altitude of 673 m. The habitat of the paratype (Fig. 7 b) likewise consisted of mid-altitude rainforest (704 m a.s.l.) on a slope. There were no wetlands within 100 m. At both sites the forest habitat seemed relatively dry compared to forest in the surrounding region. It seems possible that the species is reproducing independently of water or in small water bodies like tree holes and water filled snail shells (compare e.g. Rödel & Ernst 2002 a, b; Rödel et al. 2004). Etymology. The species’ name refers to the type locality situated in the Mayoko District of the Niari Department, Republic of the Congo. The specific epithet is considered an invariable noun in apposition to the generic name. The suggested English name for this new species is Mayoko Puddle Frog.

Phrynobatrachus horsti sp. nov. Figs. 1–2 Holotype. ZMB 81698 (adult male; field # MBUR 3237), Republic of the Congo, Niari Department, 02° 12 ’ 47.8 ’’ S, 12 ° 50 ’06.0’’ E, 28 September 2012, leg. M. Burger & A.G. Zassi-Boulou. Paratypes (all from Republic of the Congo, all collected by M. Burger & A.G. Zassi-Boulou). ZMB 81699 (adult male; field # MBUR 3098), Kouilou Department, Yombo River, 04° 23 ’ 26.1 ’’ S, 11 ° 40 ’26.0’’ E, 27 July 2012; ZMB 81700 (adult male; field # MBUR 3122), ZMB 81701 (adult female; field # MBUR 3123), Kouilou Department, Yombo River, 04° 23 ’ 26.1 ’’ S, 11 ° 40 ’26.0’’ E, 29 July 2012; ZMB 81702 (adult male; field # MBUR 3148), ZMB 81703 (adult female; field # MBUR 3149), Kouilou Department, Yombo River, 04° 23 ’ 26.1 ’’ S, 11 ° 40 ’26.0’’ E, 31 July 2012; ZMB 81704 (adult female; field # MBUR 3236), Niari Department, 02° 12 ’ 47.8 ’’ S, 12 ° 50 ’06.0’’ E, 28 September 2012; ZMB 81705 (adult male; field # MBUR 3286), Niari Department, 02° 12 ’ 47.8 ’’ S, 12 ° 50 ’06.0’’ E, 30 September 2012; ZMB 81706 (adult male; field # MBUR 3357), Niari Department, 02° 12 ’ 47.8 ’’ S, 12 ° 50 ’06.0’’ E, 4 October 2012; ZMB 81707 (adult female; field # MBUR 3541), Niari Department, 02° 19 ’ 36.2 ’’ S, 12 ° 47 ’ 32.7 ’’ E, 26 October 2013; ZMB 81708 (adult male; field # MBUR 3591), Niari Department, 02° 12 ’ 38.3 ’’ S, 12 ° 49 ’ 56.6 ’’ E, 19 October 2013; ZMB 81709 (adult female; field # MBUR 3610), ZMB 81710 (adult male; field # MBUR 03611), Niari Department, 02° 19 ’ 36.2 ’’ S, 12 ° 47 ’ 32.7 ’’ E, 21 October 2013; ZMB 81711 (adult female; field # MBUR 3612), ZMB 81712 (adult male; field # MBUR 3613), Niari Department, 02° 12 ’ 38.3 ’’ S, 12 ° 49 ’ 56.6 ’’ E, 20 October 2013; ZMB 81713 (adult male; field # MBUR 3641), Niari Department, 02° 12 ’ 38.3 ’’ S, 12 ° 49 ’ 56.6 ’’ E, 21 October 2013; ZMB 81714 (adult female; field # MBUR 3634), ZMB 81715 (adult male; field # MBUR 3652), ZMB 81716 (adult female; field # MBUR 3653), ZMB 81717 (adult female; field # MBUR 3677), ZMB 81718 (adult male; field # MBUR 3678), Niari Department, 02° 19 ’ 36.2 ’’ S, 12 ° 47 ’ 32.7 ’’ E, 26 October 2013; ZMB 81719 (adult male; field # MBUR 3681), ZMB 81720 (adult male; field # MBUR 3682), ZMB 81721 (adult male; field # MBUR 3683), Niari Department, 02° 12 ’ 38.3 ’’ S, 12 ° 49 ’ 56.6 ’’ E, 23 October 2013; ZMB 81722 (adult female; field # MBUR 3696), ZMB 81723 (adult female; field # MBUR 3697), ZMB 81724 (adult male; field # MBUR 3698), ZMB 81725 (adult male; field # MBUR 3699), ZMB 81726 (adult female; field # MBUR 3732), ZMB 81727 (adult male; field # MBUR 3742), Niari Department, 02° 12 ’ 38.3 ’’ S, 12 ° 49 ’ 56.6 ’’ E, 23 October 2013; ZMB 81728 (adult male; field # MBUR 3770), ZMB 81729 (adult male; field # MBUR 3771), Niari Department, 02° 24 ’08.8’’ S, 12 ° 59 ’08.0’’ E, 14 November 2013; ZMB 81730 (adult male; field # MBUR 4005), ZMB 81731 (adult male; field # MBUR 4006), Niari Department, 02° 24 ’08.8’’ S, 12 ° 59 ’08.0’’ E, 26 November 2013. Additional material deposited in IRSEN (all from the Republic of the Congo, all collected by M. Burger & A.G. Zassi-Boulou). MBUR 3028–3031, Kouilou Department, Yombo River, 04° 23 ’ 26.1 ’’ S, 11 ° 40 ’26.0’’ E, 22 July 1012; MBUR 3038–3039, Kouilou Department, Yombo River, 04° 23 ’ 33.1 ’’ S, 11 ° 40 ’16.0’’ E, 21 July 2012; MBUR 3063, 3099, 3103–3106, Kouilou Department, Yombo River, 04° 23 ’ 26.1 ’’ S, 11 ° 40 ’26.0’’ E, 23, 26 and 27 July 2012; MBUR 3641, Niari Department, 02° 12 ’ 38.3 ’’ S, 12 ° 49 ’ 56.6 ’’ E, 21 October 2013; MBUR 4005, Niari Department, 02° 24 ’08.8’’ S, 12 ° 59 ’08.0’’ E, 26 November 2013. Diagnosis. The genetic data (16 S rRNA) identifies the new species as a member of the genus Phrynobatrachus. Morphologically this is confirmed by: the general body shape; the presence of a tarsal tubercle; and the presence of pedal webbing. From the morphologically similar Arthroleptis they differ by lacking a median dorsal skin raphe and a black tympanic spot. From juvenile Ptychadena they can be distinguished by the lack of parallel dorsal ridges. Compared to other Phrynobatrachus the new species can be identified by the combination of the following characters: having snout-vent lengths smaller than 20 mm; males lacking tooth like projections in the lower jaw; males having black throats with spines; lacking a spiny eyelid tubercle; possessing a distinct lateral black facial mask; lacking X-shaped dorsal ridges; possessing comma-shaped dorsal ridges; lacking distinct markings on the whitish coloured belly; having yellowish ventral colouration on hind limbs; showing reduced pedal webbing and small but distinct discs on fingers and toes; and by a unique 16 S rRNA sequence. Differential diagnosis. The new species differs from other western African Phrynobatrachus species by: a small compact body (P. acridoides, P. africanus, P. auritus, P. batesii, P. brongersmai, P. cornutus, P. cricogaster, P. danko, P. francisci, P. giorgii, P. intermedius, P. jimzimkusi, P. latifrons, P. liberiensis, P. plicatus, P. natalensis, P. njiomock, P. sandersoni, P. schioetzi, P. steindachneri); lacking tooth like projections in lower jaw of males (present in P. africanus and P. sandersoni); a black lateral face mask originating from at least the level of eyes and extending to below the supratympanic ridge (black face mask only shared with: P. auritus, P. batesii, P. chukuchuku, P. intermedius, P. plicatus, P. ruthbeateae; less distinct brown face masks: P. werneri; some P. rainerguentheri have entirely black heads); small, but distinct tympanum (tympanum absent according to original description: P. ogoensis; indistinct but visible in BMNH 1947.2.6.83); black axillary spots of varying size, but generally being small and irregular shaped (larger distinct axillary and/or lateral spots and bands: P. dispar, P. fraterculus, P. gutturosus, P. maculiventris, P. pintoi, P. ruthbeateae); males with a black throat (yellow in P. acridoides, P. alleni, P. batesii, P. fraterculus, P. latifrons; throat with large black or brown spots: P. brongersmai, P. cornutus, P. giorgii; white or whitish greyish: P. annulatus, P. dispar, P. hieroglyphicus), the entire males’ throat—but in particular the centre—beset with black spines (black spines on throat shared with: P. chukuchuku, P. danko, P. jimzimkusi, P. ruthbeateae, P. steindachneri; spines hyaline or white: P. hylaios, P. schioetzi; P. werneri; spines mostly along throat margins: P. leveleve, P. danko); belly of males dirty white, dark grey towards flanks (belly at least partly yellow in P. alleni, P. batesii; P. manengoubensis, P. werneri; orange to yellow: P. danko; black: P. chukuchuku; white or bluish with black pattern: P. villiersi; white with blackish spots or dots: P. annulatus, P. ghanensis, P. hylaios, P. maculiventris, P. pintoi; a black circle and median spot on white or yellow background—bull’s eye pattern: P. cricogaster); females with yellowish grey throat, belly dirty greyish (throat and belly with brown blotches: P. intermedius, P. jimzimkusi, P. njiomock, P. steindachneri); lower parts of hind limbs in both sexes uniform yellow (yellow colouration shared with: P. alleni, P. batesii, P. danko, P. manengoubensis, possibly also with P. sternfeldi; greyish/brownish to yellowish in P. ruthbeateae, P. schioetzi; vaguely dark blotched in P. hylaios); absence of spiny eyelid tubercles (spiny eyelid tubercle present in P. annulatus, P. calcaratus, P. cornutus, P. pintoi, P. taiensis, P. villiersi); comma-shaped scapular ridges converging towards vent (scapular ridges long and converging towards mid-body, almost X-shaped: P. alleni, P. auritus, P. brongersmai, P. jimzimkusi, P. plicatus, P. steindachneri; scapular ridges indistinct or absent: P. hieroglyphicus, P. intermedius, P. liberiensis, P. maculiventris, P. manengoubensis, P. tokba); short dorsolateral ridges often distinct (short dorsolateral ridges shared with: P. r u t h be a t e a e); dorsal skin granular (smooth: P. fraterculus, P. hieroglyphicus, P. maculiventris, most P. rainerguentheri, P. tokba; tuberculate: P. annulatus, P. calcaratus, P. cornutus, P. cricogaster, P. guineensis, P. sternfeldi, most P. guttursous, P. jimzimkusi, P. steindachneri, P. sternfeldi, P. taiensis, P. villiersi; the new species described below); ventral skin smooth in both sexes (ventral skin with asperities in both sexes: P. chukuchuku); absence of a blue band along the interior half of ventral shanks (present in P. auritus); absence of nuptial pads in males (nuptial pads or swollen thumb in males present: P. b at e s i i, P. e l b e r t i, P. guineensis, P. hieroglyphicus, P. maculiventris, P. ogoensis, P. werneri); distinct roundish discs on finger and toe tips (discs absent: most other species e.g. P. hieroglyphicus, P. intermedius, P. liberiensis, P. maculiventris, P. ogoensis, P. plicatus, P. rainerguentheri, P. tokba, P. werneri; discs heart-shaped: P. sandersoni); carpal gland in males lacking (carpal gland present: P. africanus); femoral glands absent (present in males: P. calcaratus, P. cornutus, P. fraterculus, P. leveleve, P. maculiventris, P. ogoensis, P. phyllophilus; present in both sexes: P. sandersoni); rudimentary but distinct webbing (webbing indistinct or absent: P. annulatus, P. calcaratus, P. chukuchuku, P. cornutus, P. dispar, P. fraterculus, P. gutturosus, P. hieroglyphicus, P. maculiventris, P. manengoubensis, P. tokba, P. villiersi; webbing more developed: P. acridoides, P. africanus, P. alleni, P. auritus, P. brongersmai, P. cricogaster, P. danko, P. elberti, P. francisci, P. giorgii, P. intermedius, P. jimzimkusi, P. latifrons, P. liberiensis, P. natalensis, P. njiomock, P. ogoensis, P. plicatus, P. steindachneri, P. sternfeldi); small round tarsal tubercle (numerous tarsal tubercles fused to tarsal ridge: P. cricogaster). Morphologically the new species can be only mistaken for P. ruthbeateae and P. ba t es i i. A detailed comparison with these two taxa is presented in the discussion. A comparison with the second new species described herein, is included in the diagnosis of the latter species. For further character states of western African Phrynobatrachus species see: Peters (1870); Boulenger (1906 a, b); Nieden (1910); Sternfeld (1917; Arthroleptis taeniatus; replacement name by Ahl 1924: A. sternfeldi; a Phrynobatrachus species according to Frétey 2008); de Witte (1921); Parker (1936); Angel (1940); Perret (1957, 1959, 1966); Guibé (1959); Guibé & Lamotte (1963); Lamotte & Xavier (1966 a, b); Rödel (2000); Rödel & Ernst (2002 b); Rödel et al. (2005, 2009a, b, 2010, 2011, 2012 a, b); Uyeda et al. (2007); Zimkus (2009); Blackburn (2010); Blackburn & Rödel (2011); and Zimkus & Gvoždík (2012). Description of holotype [measures in mm]. Typical, small adult male Phrynobatrachus with short oval, compact body shape; snout-vent length: 14; short snout, rounded to slightly truncate in dorsal and rounded to slightly protruding in lateral view; canthus rostralis rounded; loreal region straight to very slightly convex; headwidth directly behind the eyes: 5; medium sized eyes positioned laterally; eye-diameter: 2.1; distance anterior corner of eye to nostril: 1.5; nares small and roundish to slightly vertically elliptical, positioned laterally; straight distance nostril-snout tip: 1.1; nostril closer to snout than to eye; inter-narial distance: 2.4, larger than inter-orbital distance: 1.8; pupil round; supratympanic ridge curving down from posterior corner of eye to forearm base; tympanum diameter: 1.1, with distinct tympanic annulus, annulus round with central round papilla; tympanum much smaller than diameter of eye; upper and lower maxillae and praemaxillae with minute teeth, hidden by lips; vomerine teeth absent; elongated, almost rectangular tongue, tip with distinct v-shaped notch, median conical papilla near anterior attachment of tongue; tongue densely beset with small round papillae; small round choanae positioned at lateral edges of mouth roof. Forelimbs slender; upper arm: 3.4; lower arm: 3.6; hand to tip of finger III: 4.6; hand with two large tubercles, small and oval thenar tubercle, large and round palmar tubercle; carpal gland absent; fingers with round, nondivided, protruding subarticular tubercles, no further tubercles on hand and fingers; relative finger length: IV Dorsal skin fine granular (in life) to smooth (in preservation); a pair of long comma-shaped scapular ridges, ranging from about posterior corner of upper eyelid to slightly beyond level of forearm bases, converging towards vent; dorsolateral ridges faintly indicated; posterior edge of eyelid with few flat round warts, spiny eyelid tubercle absent; no white spines or warts on flanks; skin on outer parts of thighs and dorsal part of lower legs granular; skin on lower parts of limbs, belly and pectoral region smooth; throat without skin folds running parallel to mandibles; anterior third of lower mandible and anterior third of throat densely beset with rounded spines. Colouration in ethanol after 1.5 years. Back and dorsal part of flanks more or less uniform brown; snout slightly clearer, dorsal with dark median spot, snout tip including anterior part of upper lip and loreal region light brown; clear spot between anterior part of eyelids; posterior part of inter-orbital space with dark bar; eyelids dark grey; lateral part of upper lip, sub-orbital region, supratympanic ridge and temporal area including tympanum dark brown to black, extending into a similarly coloured blotch on the anterior part of the upper arm; flanks posterior to forearm bases with large dark axillary blotch; brown flanks fainting ventrally into whitish belly; arms coloured as back, lower arm towards hand with two dark transversal bars; hands dorsally lighter coloured with dark crossbands; lower legs brown as back; thighs with 3-4 faintly indicated dark cross-bars; anterior part of thigh with dark longitudinal stripe, being the border to whitish ventral parts; lower leg dorsally with three (right) and four (left) dark cross-bars; feet and toes with dark cross-bands; dark blotch around vent, dorsally bordered by narrow yellowish line; lower parts of feet and hands greyish brown; remaining ventral parts of limbs and posterior part of belly uniform whitish; lateroventral parts of venter, anterior part of belly and pectoral region whitish but densely beset with minute black spots; throat and lower mandibles dark grey to black; lower mandibles with few small clear spots. Colouration in life (Fig. 1). Basic colour of dorsal surface of head, upper part of flank, back and dorsal surfaces of extremities light brown; snout, loreal region and anterior part of eyelids orange brown; snout with median longitudinal black spot; black inter-orbital bar; back with irregular darker patterning; lower arms, thighs, lower legs and feet with dark crossbars; anterior part of upper mandible brown with a dark bar; a V-shaped darker figure in scapular region, pointing anteriorly; black lateral face mask extending from below the eye and supratympanic ridge to forearm bases, covering tympanum; continuing below arms as a lateroventral black patch to axillar region; eye posterodorsally bordered by fine blue line; black pupil; iris copper red; central belly dirty white; belly laterally and pectoral area dark grey, no distinct black ventrolateral line or band; ventral part of forelimbs and hands dark grey; groin area and lower surfaces of thighs, lower leg and tarsus yellowish to orange; lower part of lower legs bordered by blackish spots; ventral surfaces of toes blackish. Variation (Figs. 1–2). Snout-vent length in males with spiny black throats (presumably indicating maturity) 13.5–15.8 mm and 13.1–19.7 mm in females (smallest one presumably not adult); body dimensions generally as in holotype, for small deviations see Table 1. The scapular ridges may differ in lengths and distinctness, but are always discernible and comma-shaped. Short dorsolateral ridges, composed of a very dense row of small warts, are visible in most specimens. Scapular and dorsolateral ridges are usually coloured dark. The webbing formulae showed some slight variation (Table 1). The black face mask may cover either the entire lateral parts of the head (sometimes with minute white spots; Figs. 2 c, d), extending from snout-tip to dorsal of the forearm insertion, or the parts anterior to the eye are coloured in clearer brown (Figs. 1, 2), sometimes with darker spots. Individuals with clearer coloured loreal regions also have dark brown upper mandible with black bars instead of entirely black mandibles. The back might be uniform dark or reddish brown, show some darker patterning, or carry a broad central dark band (Fig. 2 a). A light inter-orbital stripe is usually present. The iris colour varies from golden to copper red. The flank might be either uniform brown, grey or mottled with darker dots and lines (Fig. 2 c). The black axillary spot may be distinct and extending as far caudally as to about mid-body. Usually, the edges of this spot are irregularly shaped. In other individuals it is nothing but a rather indistinct mottling of blackish spots with clearer dots. We neither observed in males, nor in females distinct black bordering of belly and lower shanks as in P. ruthbeateae. The inguinal region

Leptodactylodon erythrogaster Amiet, 1971 We examined 11 tadpoles that all originated from Mount Manengouba: ZMB 78479 (one tadpole, Gosner stage 25, Caldera, 5 �� 1 ��� 50.5 ���N, 9 �� 48 ��� 38.5 ���E, 1959 m, 30 September 2010); ZMB 78481 (two tadpoles, Gosner stage 25, Caldera, 5 ��0��� 51.78 ���N, 9 �� 49 ��� 24.24 ���E, 2114 m, 8 November 2010); ZMB 78482 (seven tadpoles, Gosner stage 25, Caldera, 5 �� 1 ��� 50.5 ���N, 9 �� 48 ��� 38.5 ���E, 1959 m, 31 August 2011); ZMB 78483 (one tadpole, Gosner stage 36, near summit, 5 �� 2 ��� 24 ���N, 9 �� 51 ��� 39.3 ���E, 2012 m, 26 September 2011). All individuals were captured in small forest streams and rivers between 1959 m and 2114 m altitude (Fig. 2). The description is based on two genotyped tadpoles (ZMB 78481 ��� 82) at Gosner stage 25. Body length/total length ratio and description of tail tips was based on non-genotyped specimens of the same developmental stage. Morphology. Long slender tadpole with narrow and long muscular tail; body elliptical in dorsal and depressed in lateral view (Fig. 7 a, b); body length 28.8 �� 1.7 % (N= 8) of total length; body height 39.6���46.5 % (N= 2) of body length; body width 50.0��� 57.4 % (N= 2) of body length; maximum body width on level of spiracle insertion; snout nearly rounded in dorsal view; nostrils oval and situated laterally; nostrils equidistant from eye and snout tip; eye diameter 8.3���11.9 % (N= 2) of body length; inter-orbital distance slightly exceeds inter-nostril distance; tail fins narrow; dorsal fin originates slightly anterior (1.1 mm in ZMB 78481) or posterior (0.2 mm in ZMB 78482) to tail base, reaching deepest point at mid-tail or posterior to it, almost as deep or slightly narrower than ventral fin; narrow ventral fin originates at level of tail base, runs almost parallel to tail axis, reaches deepest point slightly anterior to tail tip; tail tip rounded; tail axis broad and muscular; body height 90.5 ���94.0% (N= 2) of total tail height; maximum height of tail axis 61.9 ���62.0% (N= 2) of maximum tail height; vent tube dextral; lateral sacs present, extending from spiracle to end of body, covering lower two thirds of flanks; short sinister spiracle, translucent, opening lateral, not visible in dorsal view, originating slightly posterior to mid-body; mouth opening frontal; labial tooth row formula 0/0; both jaw sheaths fully keratinized; upper jaw of narrow almost rectangular shape, distinctly and uniformly serrated; lower jaw more massive, depressed U-shaped; distinctly serrated, with a lateral pair of caniniform projections (fangs), median part with four to seven needle-like cusps (Fig. 7 d); two distinct serrations abaxial to fangs; posterior lip large and semicircular to kidney shaped, covered with 20 papillae; arrangement of papillae symmetrical along vertical body axis; papillae arranged in two semicircular rows (Fig. 7 c); short almost straight skin fold on lower lip, just posterior to lower jaw sheath; oral disc width 31.3���51.5 % (N= 2) of body length; mouth width 18.0���25.0 (N= 2) of oral disc width. The largest tadpole at Gosner stage 25 (ZMB 78481) measured 40.7 mm total length (body length: 11.2 mm; tail length: 29.5 mm). The most developed individual (ZMB 78483, Gosner stage 36) had a body length of 13.2 mm (tail incomplete). Amiet (1980) measured metamorphs, still possessing 2���18 mm tails, with 13.5���15 mm SVL. Coloration in preservation. Pale brown, dark brown mottling irregularly spread over dorsal body parts and tail; snout may be darker or lighter in some individuals; tail axis with black line in anterior two thirds; ventral parts and last fourth of ventral fin with some darker speckling; dorsal fins darker spotted, otherwise translucent creamwhite., Published as part of Mapouyat, Lissa, Hirschfeld, Mareike, R��del, Mark-Oliver, Liedtke, H. Christoph, Loader, Simon P., Gonwouo, L. Nono, Dahmen, Matthias, Doherty-Bone, Thomas M. & Barej, Michael F., 2014, The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae), pp. 29-53 in Zootaxa 3765 (1) on pages 36-37, DOI: 10.11646/zootaxa.3765.1.2, http://zenodo.org/record/285560, {"references":["Amiet, J. - L. (1980) Revision du genre Leptodactylodon Andersson (Amphibia, Anura, Astylosterninae). Annales de la Faculte des Sciences de Yaounde, 27, 69 - 224."]}

Leptodactylodon ovatus orientalis Amiet, 1971 Nine tadpoles where examined. All were found in Ebo Forest at the Bekob and Njuma sites: ZMB 78534 (five tadpoles, Gosner stages 25 & 26, Ebo Forest Bekob, 4 �� 21 ��� 51.96 ���N, 10 �� 25 ��� 10.26 ���E, 893 m, 30 September 2011); ZMB 78535 (one tadpole, Gosner stage 25, Ebo Forest Bekob, 4 �� 21 ��� 51.96 ���N, 10 �� 25 ��� 10.26 ���E, 893 m, 10 September 2011); ZMB 78540 (one tadpole, Gosner stage 37, Ebo Forest Bekob, 4 �� 21 ��� 55.68 ���N, 10 �� 25 ��� 18.48 ���E, 825 m, 10 January 2011); ZMB 78541 (two tadpoles, Gosner stage 25, Ebo Forest Njuma, 4 �� 20 ��� 59.34 ���N, 10 �� 14 ��� 15.72 ���E, 311 m, 5 August 2011). All tadpoles were found in mid-sized rivers within primary and secondary forests, at 311 to 893 m. The description is based on two genotyped individuals (ZMB 78534 ��� 35) of Gosner stage 25���26. Body length/ total length ratio and description of tail tips was based on non-genotyped specimens of Gosner stage 25. Morphology. Long slender tadpole with narrow and long muscular tail; body depressed in lateral and elliptical in dorsal view (Fig. 10 a, b); body length 28.3 �� 0.6 % (N= 4) of total length; body height 43.1 ���43.9.0% (N= 2) of body length; body width 52.9���55.3 % (N= 2) of body length; maximum body width on level of spiracle insertion; snout truncate in dorsal view; nostrils oval, situated laterally; nostrils equidistant from eye and snout tip; eye diameter 11.1���11.4 % (N= 2) of body length; inter-orbital distance almost equals inter-nostril distance; tail fins narrow; dorsal fin originates distinctly posterior to the tail base, broader than ventral fin, reaching deepest point in last third of tail; ventral fin narrow, originates at level of tail base, runs almost parallel to tail axis, deepest point close to tail tip (Channing et al. 2012 illustrate the tadpole with a very narrow dorsal fin, originating at about 25 % of the tail, the ventral fin being distinctly deeper than the dorsal one); tail tip pointed to rounded; tail axis broad, converges in last third towards tail tip; vent tube dextral; lateral sacs present, extending from spiracle to end of body, covering lower two thirds of flanks; short sinister spiracle, translucent, opening lateral, not visible in dorsal view, originating slightly posterior to mid-body; mouth opening frontal; labial tooth row formula 0/0; both jaw sheaths completely keratinized, serrated and with a lateral pair of caniniform projections (fangs); median part of lower jaw with six to seven needle-like cusps (Fig. 10 d); one distinct serration abaxial to fangs; upper jaw sheath narrow and almost straight; lower jaw sheath heavier, slightly bent, serration more pronounced; semicircular to bilobate posterior lip, covered with 20 large papillae (Channing et al. 2012: 18) and numerous small papillae; arrangement of large papillae symmetric along vertical body axis, arranged in two semicircular rows; small papillae arranged along edge of posterior lip (Fig. 10 c); short, straight to depressed w-shaped skin fold on lower lip, just posterior to lower jaw sheath; oral disc width 41.8 �� 7.1 % (N= 4) of body length; mouth width 24.5 �� 4.6 % (N= 4) of oral disc width. The largest known individual (ZMB 78534, Gosner stage 25) measured 51.3 mm (body length. 14.3 mm; tail length: 37.0 mm). The most developed tadpole had a body length of 13.4 mm (ZMB 78540, Gosner stage 37, tail incomplete). Amiet (1980) reports a juvenile with 17 mm body length, still having a 10 mm tail. Coloration in preservation. Dark brown body and tail, densely and irregularly mottled with dark brown spots; some individuals lighter colored; venter slightly lighter; tail fins mottled, dorsal one brownish with dark spots, ventral fin translucent with few spots., Published as part of Mapouyat, Lissa, Hirschfeld, Mareike, R��del, Mark-Oliver, Liedtke, H. Christoph, Loader, Simon P., Gonwouo, L. Nono, Dahmen, Matthias, Doherty-Bone, Thomas M. & Barej, Michael F., 2014, The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae), pp. 29-53 in Zootaxa 3765 (1) on pages 40-41, DOI: 10.11646/zootaxa.3765.1.2, http://zenodo.org/record/285560, {"references":["Channing, A., Rodel, M. - O. & Channing, J. (2012) Tadpoles of Africa - The biology and identification of all known tadpoles in sub-Saharan Africa. Edition Chimaira, Frankfurt am Main, 404 pp.","Amiet, J. - L. (1980) Revision du genre Leptodactylodon Andersson (Amphibia, Anura, Astylosterninae). Annales de la Faculte des Sciences de Yaounde, 27, 69 - 224."]}

Leptodactylodon bicolor Amiet, 1971 We examined nine tadpoles from several sites on Mount Manengouba and the South-West Province of Cameroon: ZMB 78452 (four tadpoles, Gosner stages 25���26, Ebonemin, 5 �� 1 ��� 27.6 ���N, 9 �� 45 ��� 53.2 ���E, 1417 m, 15 August 2011); ZMB 78453 (one tadpole, Gosner stage 26, Nkikoh, 5 �� 5 ��� 26.34 ���N, 9 �� 48 ��� 24.36 ���E, 1319 m, 5 December 2010); ZMB 78457 (one tadpole Gosner stage 25, Pola, 5 �� 3 ��� 26.6 ���N, 9 �� 50 ��� 10.2 ���E, 1788 m, 20 October 2011); ZMB 78458 (two tadpoles, Gosner stages 25���26, Pola, 5 �� 3 ��� 27.9 ���N, 9 �� 49 ��� 39 ���E, 1719 m, 20 October 2011); ZMB 78461 (one tadpole, Gosner stage 35, Fotabong, 5 �� 28 ��� 46.44 ���N, 9 �� 55 ��� 53.22 ���E, 1221 m, 1 November 2011). The tadpoles were all found in small to medium-sized streams with farmbush vegetation on river banks (Fig. 2). They inhabited slowflowing parts of fast flowing streams with rapids in 1221���1788 m The description of L. bicolor is predominantely based on three (ZMB 78452, 78453, 78458) genotyped individuals of Gosner stage 26. Body length/total length ratio and description of tail tips was based on nongenotyped specimens of the same developmental stage. Morphology. Long slender tadpole with narrow and long muscular tail; body elliptical in dorsal view, slightly depressed in lateral view (Fig. 4 a, b); body shape in lateral view either slightly convex, extending level of tail axis (Fig. 4 a) or flat, in line with tail axis (Channing et al. 2012); body length 28.1 �� 2.1 % (N= 7) of total length; body height 42.6 �� 4.2 % (N= 3) of body length; body width 54.0 �� 4.3 % (N= 3) of body length; maximum body width on the level of spiracle insertion; snout nearly rounded to truncate in dorsal view; nostrils oval, positioned laterally, opening slightly pointed ventrally; nostrils equidistant from eye and snout tip; eye diameter 13.1 �� 1.8 % (N= 3) of body length; inter-orbital distance slightly exceeds inter-nostril distance; tail fins narrow; dorsal fin originating posterior to tail base (1.7 �� 0.3 mm; N= 3; Fig. 102 in Channing et al. 2012 and ZMB 78461: at level with tail base), equal to slightly deeper than ventral fin, reaching deepest point at about mid-tail, fin edge almost parallel to tail axis; ventral fin originating on level with tail base, narrow and almost parallel to tail axis; tail axis muscular, only slowly converging towards tail tip; tail tip rounded; body height 78.1���100 % (N= 3; flatter individuals possibly due to preservation artifact) of total tail height; maximum height of tail axis 62.5 ���75.0% (N= 3) of total tail height; vent tube dextral; lateral sacs present, extending from spiracle to end of body, covering lower two thirds of flanks; short spiracle, sinistral, translucent, opening lateral, not visible in dorsal view, originating slightly posterior to midbody; mouth opening frontal; labial tooth formula 0/0; both jaw sheaths fully keratinized and of narrow almost rectangular shape; upper jaw distinctly serrated, serrations uniform; lower jaw U-shaped, serrated, lateral pair of caniniform projections (���fangs���; Fig. 4 d), median part with six to seven needle-like cusps (Channing et al. 2012: 8); three distinct serrations abaxial to fangs; large almost semicircular posterior lip covered with 20 papillae; papillae symmetrically arranged along body axis; most papillae arranged in two semicircular rows, one near the lower jaw sheath, the second closer to posterior margin of lower lip (Fig. 4 c); almost straight skin fold on lower lip, just posterior to lower jaw sheath; oral disc width 33.9���45.4 % (N= 3) of body length; mouth width 22.2���33.3 % (N= 3) of oral disc width. The largest tadpole at Gosner stage 25 (ZMB 78462) measured 48.8 mm total length (body length: 11.9 mm; tail length: 38.9 mm). The most developed individual (ZMB 78461, Gosner stage 35) measured 45.6 mm total length (body length: 12.8 mm; tail length: 32.8 mm). Amiet (1980) reports tailed metamorphs measuring 13.5 mm SVL. Coloration in preservation. Dorsal body parts and tail axis irregularly speckled dark brown on brown ground; mottling on tail axis strong, less pattern towards tail tip; anterior part of tail with longitudinal black line; ventrum clearer with very little dark speckling; dorsal fin brownish with darker spots or translucent; ventral fin predominantly translucent cream-white with few darker spots. Coloration in life. Light brown with dark brown speckling, some dots being lighter; venter light grayishreddish; dark intestinal loops shining through skin; some individuals with cream-white snout., Published as part of Mapouyat, Lissa, Hirschfeld, Mareike, R��del, Mark-Oliver, Liedtke, H. Christoph, Loader, Simon P., Gonwouo, L. Nono, Dahmen, Matthias, Doherty-Bone, Thomas M. & Barej, Michael F., 2014, The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae), pp. 29-53 in Zootaxa 3765 (1) on pages 32-34, DOI: 10.11646/zootaxa.3765.1.2, http://zenodo.org/record/285560, {"references":["Channing, A., Rodel, M. - O. & Channing, J. (2012) Tadpoles of Africa - The biology and identification of all known tadpoles in sub-Saharan Africa. Edition Chimaira, Frankfurt am Main, 404 pp.","Amiet, J. - L. (1980) Revision du genre Leptodactylodon Andersson (Amphibia, Anura, Astylosterninae). Annales de la Faculte des Sciences de Yaounde, 27, 69 - 224."]}

Leptodactylodon ornatus ornatus Amiet, 1971 Two L. ornatus ornatus tadpoles were found on Mount Manengouba: ZMB 78502 (two tadpoles, Gosner stages 25 & 32, Ebonemin, 5 �� 1 ��� 6.2 ���N, 9 �� 46 ��� 16.8 ���E, 1447 m, 9 October 2011). The two individuals were found in a small river surrounded by farmbush vegetation at 1447 m (Fig. 2). The description is based on the genotyped individual (ZMB 78502) at Gosner stage 32. Body length/total length ratio and description of tail tips was based on the non-genotyped specimen. Morphology. Long slender tadpole with narrow and long muscular tail; body elliptical to almost parallel in dorsal and depressed in lateral view (Fig. 9 a, b); body length 26.0% of total length; body height 44.6 % of body length; body width 56.2 % of body length; maximum body width on level of spiracle insertion; snout truncate in dorsal view; nostrils nearly rounded, situated lateroventrally; nostrils equidistant from eye and snout tip; eye diameter 13.1 % of body length; inter-orbital distance slightly exceeds inter-nostril distance; tail fins narrow; dorsal fin originating posterior (5.7 mm) to tail base, slightly deeper than ventral fin, reaching deepest point slightly posterior to mid-tail; ventral fin narrow, originates on level with tail base, reaches deepest point near tail tip; tail tip rounded; tail axis broad and muscular; body height 82.9 % of total tail height; maximum height of tail axis 61.4 % of total tail height; vent tube dextral; lateral sacs present, extending from spiracle to end of body, covering lower two thirds of flanks; short sinister spiracle, translucent, opening lateral, not visible in dorsal view, originating slightly posterior to mid-body; mouth opening frontal; labial tooth formula 0/0; both jaw sheaths completely keratinized; upper jaw sheath distinctly and strongly serrated, narrow, almost rectangular, edges with large, caniniform projections; lower jaw distinctly but less serrated, more massive than upper jaw, slightly bent, a lateral pair of caniniform projections, median part without needle-like cusps (Fig. 9 d); four distinct serrations abaxial to fangs; posterior lip large and broad bilobate, covered with 21 large and numerous small papillae; small papillae arranged along edge of the lower lip; large papillae arranged in two semicircular rows (Fig. 9 c); short, straight to depressed w-shaped skin fold on lower lip, just posterior to lower jaw sheath; oral disc width 41.5 % of body length; mouth width 37.0% of oral disc width. The tadpole at Gosner stage 25 measured 45.7 mm total length (body length: 11.9 mm; tail length: 33.8 mm). The more developed individual (Gosner stage 32) had a body length of 13.0 mm (tail incomplete). Amiet (1980) gives the size of three metamorphs as 14���16 mm. Coloration in preservation. Light to dark brown; dorsal parts of body and in particular tail axis mottled with larger dark dots and densely covered with smaller dark speckles, lighter towards tail tip; anterior half of tail axis with median, longitudinal dark line; venter lighter; dorsal fin and posterior-most part of ventral fin mottled with dark spots, otherwise, particularly anterior part of ventral tail fin translucent cream-white., Published as part of Mapouyat, Lissa, Hirschfeld, Mareike, R��del, Mark-Oliver, Liedtke, H. Christoph, Loader, Simon P., Gonwouo, L. Nono, Dahmen, Matthias, Doherty-Bone, Thomas M. & Barej, Michael F., 2014, The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae), pp. 29-53 in Zootaxa 3765 (1) on page 39, DOI: 10.11646/zootaxa.3765.1.2, http://zenodo.org/record/285560, {"references":["Amiet, J. - L. (1980) Revision du genre Leptodactylodon Andersson (Amphibia, Anura, Astylosterninae). Annales de la Faculte des Sciences de Yaounde, 27, 69 - 224."]}

Leptodactylodon mertensi Perret, 1959 We examined 48 tadpoles, all from Mount Manengouba: ZMB 78499 (two tadpoles, Gosner stages 25 & 40, Ebonemin, 5 �� 1 ��� 33.42 ��� N, 9 �� 45 ��� 46.74 ���E, 1356 m, 13 December 2010); ZMB 78500 (eight tadpoles, Gosner stages 25 & 42, Ebonemin, 5 ��0��� 46.5 ���N, 9 �� 45 ��� 46.74 ���E, 1372 m, 16 December 2010); ZMB 78501 (one tadpole, Gosner stage 25, Ebonemin, 5 ��0��� 46.5 ���N, 9 �� 46 ��� 5.8 ���E, 1372 m, 7 October 2011); ZMB 78503 (one tadpole Gosner stage 25, Ebonemin, 5 �� 1 ��� 6.2 ���N, 9 �� 46 ��� 16.8 ���E, 1447 m, 9 October 2011); ZMB 78504 (two tadpoles, Gosner stages 25 & 27, Caldera, 5 �� 2 ��� 26.2 ���N, 9 �� 48 ��� 30 ���E, 1889 m, 2 September 2011); ZMB 78505 (two tadpoles, Gosner stage 25 & 28, Caldera, 5 �� 2 ��� 26.2 ���N, 9 �� 48 ��� 30 ���E, 1889 m, 17 October 2011); ZMB 78507 (three tadpoles, Gosner stage 25, Manengouba village, 4 �� 56 ��� 50.3 ���N, 9 �� 52 ��� 45.8 ���E, 996 m, 28 June 2011); ZMB 78508 ���09 (six tadpoles and one tadpole, respectively, Gosner stage 25, M���Bouroukou, 5 �� 3 ��� 48.2 ���N, 9 �� 52 ��� 0.09 ���E, 1518 m, 22 September 2011); ZMB 78511 (one tadpole, Gosner stage 25, Nkack, 5 �� 2 ��� 17.4 ���N, 9 �� 46 ��� 27.3 ���E, 1466 m, 12 October 2011); ZMB 78513 (five tadpoles, Gosner stage 25, Pastoral Nkongsamba, 4 �� 59 ��� 13.26 ���N 9 �� 52 ��� 45.9 ���E, 1459 m, 18 January 2011); ZMB 78514 ��� 16 (two tadpoles, eight tadpoles and six tadpoles, respectively; Gosner stages 25 & 26, Pastoral Nkongsamba, 4 �� 58 ��� 47.1 ���N, 9 �� 53 ��� 41.9 ���E, 1225 m, 29 September 2011). The tadpoles occurred in small to mid-sized rivers between 996 m and 1889 m altitude. These riverbanks were covered with forest and farmbush vegetation (Fig. 2). The description is based on ten genotyped tadpoles (ZMB 78500 ���01, 7850 ���304, 78507���09, 78511, 78513, 78515) at Gosner stage 25. Body length/total length ratio and description of tail tips was based on non-genotyped specimens of the same developmental stage. Morphology. Long but robust tadpole with long tail and very broad muscular tail axis; body elliptical in dorsal and robust and pointed in lateral view (Fig. 8 a, b); body length 28.3 �� 2.4 % (N= 31) of total length; body height 42.2 �� 3.9 % (N= 10) of body length; body width 54.0 �� 4.2 % (N= 10) of body length, maximum body width on level of spiracle insertion; snout truncate to nearly rounded in dorsal view; nostrils oval and situated laterally; nostrils equidistant from eye and snout tip; eye diameter 12.7 �� 0.9 % (N= 10) of body length; inter-orbital distance slightly smaller than inter-nostril; tail fins narrow; dorsal fin mostly originating posterior to tail base (exceptionally fin insertions at or slightly anterior to tail base were observed); narrow dorsal fins, reaching deepest point posterior to mid-tail, slightly deeper than ventral fin; ventral fin originates at level with tail base, runs almost parallel to tail axis; tail tip rounded; tail axis very broad and muscular in dorsal and lateral view; body height 91.0 �� 4.1 % (N= 10) of total tail height; maximum height of tail axis 69.1 �� 5.9 % (N= 9) of maximum tail height; vent tube dextral; lateral sacs present, extending from spiracle to end of body, covering lower two thirds of flanks; short sinister spiracle, translucent, opening lateral, not visible in dorsal view, originating at mid-body; mouth opening frontal; labial tooth row formula 0/0; both jaw sheaths completely keratinized; upper jaw narrow, almost rectangular, slightly bent, distinctly and uniformly serrated; lower jaw U-shaped, distinctly serrated with a lateral pair of caniniform projections (fangs), median part with six to seven needle-like cusps (Fig. 8 d; Channing et al. 2012: 8); two distinct serrations abaxial to fangs; posterior lip huge semicircular to oval bean shaped, covered with 21 papillae (Channing et al. 2012: 20); arrangement of papillae symmetric to vertical body axis; papillae arranged in two semicircular rows (Fig. 8 c); straight to semicircular skin fold on lower lip, just posterior to lower jaw sheath; oral disc width 34.0 �� 2.6 % (N= 10) of body length; mouth width 26.2 �� 3.2 % (N= 10) of oral disc width. The most advanced individuals (ZMB 78499, Gosner stage 40; ZMB 78500, three tadpoles of Gosner stages 40���42) measured 43.3���46.2 mm (body length. 11.3���13.9 mm; tail length: 29.7���32.6 mm). A few tadpoles of Gosner stages 25���26 even reached 46.1���49.5 mm total length. Channing et al. (2012) reported 52 mm total length. Amiet (1980) measured metamorphs, still possessing 4���5 mm tails, with 13.5���15.5 mm SVL. Coloration in preservation. Pale brown body and tail; dorsal parts of body and entire tail axis heavily mottled with little dark spots; ventral parts lighter; fins with very few dark markings, translucent cream white; some individuals with cream-white snout., Published as part of Mapouyat, Lissa, Hirschfeld, Mareike, R��del, Mark-Oliver, Liedtke, H. Christoph, Loader, Simon P., Gonwouo, L. Nono, Dahmen, Matthias, Doherty-Bone, Thomas M. & Barej, Michael F., 2014, The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae), pp. 29-53 in Zootaxa 3765 (1) on pages 37-39, DOI: 10.11646/zootaxa.3765.1.2, http://zenodo.org/record/285560, {"references":["Channing, A., Rodel, M. - O. & Channing, J. (2012) Tadpoles of Africa - The biology and identification of all known tadpoles in sub-Saharan Africa. Edition Chimaira, Frankfurt am Main, 404 pp.","Amiet, J. - L. (1980) Revision du genre Leptodactylodon Andersson (Amphibia, Anura, Astylosterninae). Annales de la Faculte des Sciences de Yaounde, 27, 69 - 224."]}

Leptodactylodon perreti Amiet, 1971 Three tadpoles of L. perreti were examined and genotyped. All were found on Mount Oku in the Emfveh-mi forest, near Elak. ZMB 79084 ��� 86 (Gosner stages 25, 35, 37, respectively; Mount Oku, 6 �� 13 ��� 42.4 ���N, 10 �� 31 ��� 17.4 ���E, 2269 m, 23 August and 2 September 2012). Tadpoles were collected in a stream in montane forest, ca. 300 m from the farm-forest boundary. The description is based on all three tadpoles. Morphology. Long and slender tadpoles with long, muscular tail; body elliptical in dorsal and narrow elliptical lateral view (see Fig. 3 in Cruz et al. 2013); body length 36.2���39.3 % (N= 3) of total length; body width 47.2��� 57.9 % (N= 3) of body length; body widest at level of spiracle; snout rounded in dorsal view; nostrils oval and situated lateroventrally; nostrils equidistant from eye and snout tip (closer to eye Cruz et al. 2013); eye diameter 11.7���12.4 % (N= 3) of body length; inter-orbital distance slightly exceeds inter-nostril distance; tail fins narrow; narrow dorsal fin originates distinctly posterior to tail base (3.1���3.5 mm), running almost parallel to muscular tail axis; ventral fin originates at tail base, only slightly narrower than dorsal fin, runs parallel to tail axis; tail tip roundish to slightly pointed; tail axis broad, continuously converging towards tail tip; body height equal to or slightly less than total tail height; maximum height of tail axis 67.2���70.7 % (N= 3) of total tail height; vent tube dextral; lateral sacs present, extending from spiracle to end of body, covering lower two thirds of flanks; short spiracle, sinistral, translucent, opening lateral, not visible in dorsal view, originating slightly anterior (N= 1) or posterior (N= 2) of mid-body; mouth opening frontal; labial tooth row formula 0/0; both jaw sheath completely keratinized, serrated, almost straight and narrow; two small caniniform projections (fangs) towards edges of lower sheath; lower jaw slightly bent; median part of lower jaw with indistinct needle-like cusps (Fig. 11 b; Channing et al. 2012: 8); four distinct serrations abaxial to fangs; broad semicircular posterior lip covered with 21 large papillae (Fig. 11 a; Channing et al. 2012: 30); arrangement of papillae symmetrical to vertical body axis; papillae arranged in two semicircular rows; short, very depressed w-shaped skin fold on lower lip, just posterior to lower jaw sheath; oral disc width 29.8���33.3 % (N= 3) of body length; mouth width 22.2���28.6 % (N= 2) of oral disc width. ZMB 79084 (Gosner stage 25) had a total length of 38.3 mm (body length: 10.8 mm; tail length: 27.5 mm). Our most advanced tadpole (ZMB 79086, Gosner stage 37) measured 48.2 mm total length (body length: 12.8 mm; tail length: 35.4 mm). Cruz et al. (2013) report total lengths of 45.1 and 51.6 mm in Gosner stage 25 tadpoles. Metamorphs with completely resorbed tails measure 15.5���17.5 mm SVL (Amiet 1980). Coloration in preservation. Pale to deep dark brown dorsal parts of body and tail, dense irregular dark mottling; venter slightly lighter; fins with irregular dark mottling, more pronounced in dorsal parts, otherwise translucent cream-white; fins lighter than in other species., Published as part of Mapouyat, Lissa, Hirschfeld, Mareike, R��del, Mark-Oliver, Liedtke, H. Christoph, Loader, Simon P., Gonwouo, L. Nono, Dahmen, Matthias, Doherty-Bone, Thomas M. & Barej, Michael F., 2014, The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae), pp. 29-53 in Zootaxa 3765 (1) on pages 41-42, DOI: 10.11646/zootaxa.3765.1.2, http://zenodo.org/record/285560, {"references":["Cruz, D., Dawson, A. L. & Blackburn, D. C. (2013) Description of the tadpoles of two Cameroonian frogs, Leptodactylodon axillaris and L. perreti (Anura: Arthroleptidae). African Journal of Herpetology, 62, 28 - 39. http: // dx. doi. org / 10.1080 / 21564574.2013.777370","Channing, A., Rodel, M. - O. & Channing, J. (2012) Tadpoles of Africa - The biology and identification of all known tadpoles in sub-Saharan Africa. Edition Chimaira, Frankfurt am Main, 404 pp.","Amiet, J. - L. (1980) Revision du genre Leptodactylodon Andersson (Amphibia, Anura, Astylosterninae). Annales de la Faculte des Sciences de Yaounde, 27, 69 - 224."]}

Mapouyat, Lissa, Hirschfeld, Mareike, Rödel, Mark-Oliver, Liedtke, H. Christoph, Loader, Simon P., Gonwouo, L. Nono, Dahmen, Matthias, Doherty-Bone, Thomas M., Barej, Michael F. (2014): The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae). Zootaxa 3765 (1): 29-53, DOI: http://dx.doi.org/10.11646/zootaxa.3765.1.2

Leptodactylodon cf. polyacanthus punctiventris Amiet, 1971 Three tadpoles from Fotabong were tentatively assigned to L. polyacanthus punctiventris based on known amphibian assemblages from that area (e.g. Amiet 1980). One specimen was genotyped: ZMB 78551 (three tadpoles, Gosner stage 25, Fotabong, 5 �� 28 ��� 54 ���N, 9 �� 55 ��� 26.58 ���E, 1197 m, 1 November 2011). The tadpoles were collected from shallow, sandy streams at 1997 m. The description is based on the genotyped individual (Gosner stage 25). Body length/total length ratio and description of tail tips was based on non-genotyped specimens of the same developmental stage. Morphology. Long and slender tadpole with long, muscular tail (non-genotyped tadpoles not as slender as drawn in Fig. 12 a, b); body elliptical in dorsal and depressed in lateral view; body length 27.5���29.3 % (N= 2) of total length; body height 37.2 % of body length, body width 46.3 % of body length, maximum body width on level of spiracle insertion; snout nearly rounded in dorsal view; nostrils oval and situated laterally; nostrils equidistant from eye and snout tip; eye diameter 9.9 % of body length; inter-orbital distance equals inter-nostril distance; tail fins narrow, dorsal fin originating 4.6 mm anterior to tail base, reaching deepest point in first third of tail, slightly deeper than ventral fin; ventral fin originates at tail base, runs parallel to tail axis; tail tip rounded to slightly pointed; tail axis broad, continuously converging towards tail tip; maximum height of tail axis 57.0��� 60.5 % (N= 2) of total tail height; vent tube dextral; lateral sacs present, extending from spiracle to end of body, covering lower two thirds of flanks; very short spiracle, sinistral, translucent, opening lateral, not visible in dorsal view, originating at mid-body; mouth opening frontal; labial tooth row formula 0/0; both jaw sheaths completely keratinized, strongly serrated, each with lateral pair of caniniform projections (fangs); median part of lower jaw with six to seven needle-like cusps (Fig. 12 d); three distinct serrations abaxial to fangs; upper jaw narrow and long, lower jaw shorter; large, broad bilobate posterior lip covered with 21 large papillae (Fig. 12 c); papillae arranged symmetrically along in vertical body axis; papillae arranged in two semicircular rows; almost straight skin fold on lower lip, just posterior to lower jaw sheath; oral disc width 39.7 % of body length; mouth width 25.0% of oral disc width. The two individuals with intact tails (both Gosner stage 25) had total lengths of 39.6 and 43.6 mm, respectively (body length: 11.6 & 12.0 mm; tail length: 28.0 & 31.6 mm). Metamorphs measure about 14 mm SVL (Amiet 1980). Coloration in preservation. Body and tail light brown, mottled with bigger and smaller dark spots and speckles on dorsal part of body and tail; anterior half of tail axis with median, longitudinal dark line (not visible in life); venter lighter with fewer dark patterns; fins translucent cream-white. Coloration in life. Light brown with light to dark brown speckles; dark line from snout tip to anterior corner of eye; anterior part of tail axis with white spots, posterior part with black dots; fins light with dark spots, posterior part almost white; venter lighter (Fig. 6)., Published as part of Mapouyat, Lissa, Hirschfeld, Mareike, R��del, Mark-Oliver, Liedtke, H. Christoph, Loader, Simon P., Gonwouo, L. Nono, Dahmen, Matthias, Doherty-Bone, Thomas M. & Barej, Michael F., 2014, The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae), pp. 29-53 in Zootaxa 3765 (1) on pages 42-43, DOI: 10.11646/zootaxa.3765.1.2, http://zenodo.org/record/285560, {"references":["Amiet, J. - L. (1980) Revision du genre Leptodactylodon Andersson (Amphibia, Anura, Astylosterninae). Annales de la Faculte des Sciences de Yaounde, 27, 69 - 224."]}

Leptodactylodon ventrimarmoratus (Boulenger, 1904) Fifteen tadpoles of L. ventrimarmoratus were found near Mount Kala: ZMB 78552 ��� 53 (one tadpole and seven tadpoles, respectively; all Gosner stage 25, 3�� 50 ���24,72���N, 11 �� 21 ��� 6.66 ���E, 858 m); ZMB 78554 (seven tadpoles, Gosner stage 25, 3�� 50 ��� 23.46 ���N, 11 �� 21 ��� 6.54 ���E, 901 m). All tadpoles were found on 8 November 2011 near Kala village in small, rocky rivers, mostly beneath leaf litter, at 858 and 901 m altitude. The description is based on three genotyped individuals (ZMB 78552, 78553, 78554) at Gosner stage 25. Body length/total length ratio and description of tail tips was based on non-genotyped specimens of the same developmental stage. Morphology. Long and slender tadpole with long, muscular tail; body elliptical to almost parallel in dorsal and depressed long ovoid in lateral view (Fig. 13 a, b); body length 30.8 �� 2.5 % (N= 12) of total length; body height 29.9���48.2 % of body length (high value probably a preservation artifact; Amiet 1970 reports the body length to be 1.7 to 2 times the body widths); body width 48.2���56.1 % of body length; snout truncate in dorsal view; nostrils oval and situated laterally; nostrils equidistant from eye and snout tip; eye diameter 10.5���12.3 % of body length; interorbital distance equal to inter-nostril distance; tail fins narrow; dorsal fin originating (1.3���2.5 mm, N= 3) anterior to tail base, narrow, reaching deepest point at or posterior to mid-tail, slightly deeper or as deep as ventral fin; ventral fin originating at tail base, parallel to tail axis to about mid-tail, then broader (Amiet 1970 reports the dorsal fin originating distinctly posterior to tail base and a more robust tail axis); tail tip rounded to slightly pointed; tail axis broad, continuously converging towards tail tip; body height 96.6 % of total tail height; maximum height of tail axis 64.9 % of total tail height; vent tube dextral; lateral sacs present, extending from spiracle to end of body, covering lower two thirds of flanks; short spiracle, sinistral, translucent, opening directed lateral, not visible in dorsal view, originating at mid-body; mouth opening frontal; labial tooth row formula 0/0; both jaw sheath completely keratinized, serrated, with a lateral pair of caniniform projections (fangs); upper jaw almost straight; lower jaw Ushaped, median part of lower jaw edge with six to seven needle-like cusps (Fig. 13 d); two distinct serrations abaxial to fangs; posterior lip large and broadly bilobate, covered with 21 large papillae and numerous small papillae; large papillae arranged symmetrical and in two semicircular rows along vertical body axis; small papillae along edge of lower lip (Fig. 13 c); depressed w-shaped skin fold on lower lip, just posterior to lower jaw sheath; oral disc width 34.7���57.9 % of body length (lower value due to preservation artifact, lip not completely enfolded); mouth width 24.2���32.6 % of oral disc width (see above). The two largest known individuals (ZMB 78553 & 78554, Gosner stage 25) measured 41.6 and 42.8 mm total length, respectively (body length: 12.2 & 12.4 mm; tail length: 29.2 & 30.6 mm). Amiet (1970) reports that a total length of 60 mm may be reached. Metamorphs measure 17���21 mm (Amiet 1980). Coloration in preservation. Dorsal parts of body and tail light brown, densely and irregularly covered with dark dots; anterior half of tail axis with median, longitudinal dark line; pattern on tail changes towards tail tip, anterior part of tail axis and dorsal tail fin heavily covered with midsized dark dots; then dots get smaller but denser; overall posterior third of tail tip lighter but more densely dotted; anterior third of ventral fin translucent cream-white; venter lighter and with fewer dark patterns. Coloration in life. Light brown with many minute dark brown dots; snout brown or yellowish (Figs. 6, 14); dark line from snout tip to anterior corner of eye; funnel light brown to almost yellow; density of dark dots on tail axis decreasing towards tail tip; anterior third of tail axis with dark longitudinal line; fins very light, almost white, dorsal part with dark patterning; venter lighter; intestines shining through skin (Fig. 15). According to Amiet (1970) a greyish beige color with brown pattern on dorsal parts, the snout sometimes ochre, the lateral sacs are green in life (not differently colored in our specimens (Figs. 6, 14, 15). Tadpoles��� lateral line system of investigated Leptodactylodon species. As far as visible (depending on the preservation state of individuals) all examined Leptodactylodon tadpoles showed the following lateral lines (definitions sensu Lannoo 1987; compare also Fig. 11 in Amiet 1970 and Figs. in Cruz et al. 2013): Supraorbital, posteriororbital, dorsal, middle, ventral, longitudinal oral and angular lines present; angular line connected to posterior infraorbital line; second angular line running dorsal to spiracle, merging into middle line; middle line originating dorsal to lateral sacs, turns upward posteriorly and connects to dorsal line; supraorbital line s-shaped., Published as part of Mapouyat, Lissa, Hirschfeld, Mareike, R��del, Mark-Oliver, Liedtke, H. Christoph, Loader, Simon P., Gonwouo, L. Nono, Dahmen, Matthias, Doherty-Bone, Thomas M. & Barej, Michael F., 2014, The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae), pp. 29-53 in Zootaxa 3765 (1) on pages 43-45, DOI: 10.11646/zootaxa.3765.1.2, http://zenodo.org/record/285560, {"references":["Amiet, J. - L. (1970) Morphologie et developpement de la larve de Leptodactylodon ventrimarmoratus (Boulenger) (Amphibien Anoure). Annales de la Faculte des Sciences du Cameroun, 4, 53 - 71.","Amiet, J. - L. (1980) Revision du genre Leptodactylodon Andersson (Amphibia, Anura, Astylosterninae). Annales de la Faculte des Sciences de Yaounde, 27, 69 - 224.","Lannoo, M. J. (1987) Neuromast topography in anuran amphibians. Journal of Morphology, 191, 115 - 129. http: // dx. doi. org / 10.1002 / jmor. 1051910203","Cruz, D., Dawson, A. L. & Blackburn, D. C. (2013) Description of the tadpoles of two Cameroonian frogs, Leptodactylodon axillaris and L. perreti (Anura: Arthroleptidae). African Journal of Herpetology, 62, 28 - 39. http: // dx. doi. org / 10.1080 / 21564574.2013.777370"]}

Leptodactylodon boulengeri Nieden, 1910 We examined sixteen individuals collected near Fotabong and on Mount Manengouba: ZMB 78466 (one tadpole, Gosner stage 25, Manengouba village, 4 �� 58 ��� 23.76 ���N, 9 �� 52 ��� 31.8 ���E, 1290 m, 21 November 2010); ZMB 78469 ��� 71 (two tadpoles, Gosner stages 36 & 40, respectively; one tadpole Gosner stage not assessable; five tadpoles, Gosner stage 25; Fotabong, 5 �� 28 ��� 36.12 ���N, 9 �� 55 ��� 25.08 ���E, 1272 m, 3 November 2011); ZMB 78476 (seven tadpoles, Gosner stage 25���37, Fotabong, 5 �� 28 ��� 38.88 ���N 9 �� 55 ��� 24.48 ���E, 1267 m, 2 November 2011). All tadpoles of L. boulengeri were found on soaked sandy soil along small rivers or buried in sandy ground of these rivers in mid altitude (1267���1290 m), the sand being only covered by a very thin film of water. The description is based on four genotyped tadpoles (ZMB 78466, 78470, 78471, 78476) at Gosner stage 25. Body length/total length ratio, maximum tail length and height; as well as description of tail tip was based on nongenotyped specimens of the same developmental stage. Morphology. Long slender tadpole with narrow and long muscular tail; body elongate, elliptical in dorsal and depressed in lateral view (Fig. 5 a, b); body length 29.9 �� 2.0% (N= 10) of total length; body height 40.6 �� 7.5 % (N= 4) of body length; body width 52.2 �� 3.8 % (N= 4) of body length; maximum body width on level of spiracle insertion; snout nearly rounded to truncate in dorsal view; nostrils oval, situated laterally; nostrils equidistant from eye and snout tip; eye diameter 8.3 �� 2.7 % (N= 4) of body length; inter-orbital distance equals inter-nostril distance (inter-orbital distance of ZMB 78470 slightly exceeds inter-nostril distance); tail fins narrow; dorsal fin originating 0.3 ���1.0 mm (N= 3) anterior to tail base, narrow, reaching maximum depth after about first quarter, as high or only slightly deeper as ventral fin; ventral fin narrow, originates on level with tail base, almost parallel to tail axis; tail tip pointed; tail axis slender, edges almost parallel towards tail tip; body height 97.4���98.5 (N= 2) of total tail height; maximum height of tail axis 57.7 �� 2.7 % (N= 5) of total tail height; vent tube dextral; lateral sacs present, extending from spiracle to end of body, covering lower two thirds of flanks; short sinister spiracle, translucent, opening lateral, not visible in dorsal view, originating at app. mid-body; mouth opening frontal; labial tooth row formula 0/0; both jaw sheaths fully keratinized and of narrow almost rectangular shape; upper jaw distinctly serrated, serrations almost uniform, slightly larger serration towards edges; lower jaw flat U-shaped, distinctly serrated with a lateral pair of caniniform projections (fangs), median part with six to seven needle-like cusps (Fig. 5 d); one distinct serration abaxial to fangs; large semicircular posterior lip covered with 18 papillae; papillae symmetrically arranged along body axis; most papillae arranged in two semicircular rows, one near the lower jaw sheath, the second closer to posterior margin of lower lip (Fig. 5 c); distinct short skin fold on lower lip, just posterior to lower jaw sheath; oral disc width 35.5���48.5 % (N= 3) of body length; mouth width 23.1���31.8 % (N= 3) of oral disc width. The largest tadpole at Gosner stage 25 (ZMB 78471) measured 24.4 mm total length (body length: 7.7 mm; tail length: 16.7 mm). The most advanced/developed individual (ZMB 78476, Gosner stage 37) measured 52.1 mm (body length. 14.1 mm; tail length: 38.0 mm). The smallest juveniles reported by Amiet (1980) measured 16���19 mm SVL. Coloration in preservation. Pale brown; dorsal parts of body and tail with irregular dark speckling; lower part of body and tail paler; anterior part of tail with longitudinal black line (not visible in life); only few dark spots on venter and on the last third of ventral tail fin; dorsal fin with dark spots, fins otherwise translucent cream-white. Coloration in life (Fig. 6). Pale brown with dark brown speckles, some of them lighter brown; funnel dark, almost black; tail axis with whitish-blue spots; ventral fin with yellow spots; tail tip almost white; venter with light reddish coloration; some individuals with cream-white snout (like some L. bicolor and L. ventrimarmoratus)., Published as part of Mapouyat, Lissa, Hirschfeld, Mareike, R��del, Mark-Oliver, Liedtke, H. Christoph, Loader, Simon P., Gonwouo, L. Nono, Dahmen, Matthias, Doherty-Bone, Thomas M. & Barej, Michael F., 2014, The tadpoles of nine Cameroonian Leptodactylodon species (Amphibia, Anura, Arthroleptidae), pp. 29-53 in Zootaxa 3765 (1) on pages 34-36, DOI: 10.11646/zootaxa.3765.1.2, http://zenodo.org/record/285560, {"references":["Amiet, J. - L. (1980) Revision du genre Leptodactylodon Andersson (Amphibia, Anura, Astylosterninae). Annales de la Faculte des Sciences de Yaounde, 27, 69 - 224."]}

Ernst, Raffael, Nienguesso, Alvaro Bruno Toto, Lautenschläger, Thea, Barej, Michael F., Scmitz, Andreas, Hölting, Monique (2014): Relicts of a forested past: Southernmost distribution of the hairy frog genus Trichobatrachus Boulenger, 1900 (Anura: Arthroleptidae) in the Serra do Pingano region of Angola with comments on its taxonomic status. Zootaxa 3779 (2): 297-300, DOI: http://dx.doi.org/10.11646/zootaxa.3779.2.10

Phrynobatrachus rainerguentheri sp. nov. Figs 1���2 Holotype. 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��, Paratypes. 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. Additional material. 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. Diagnosis. The genetic results show that the new species belongs to the genus Phrynobatrachus. 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 Arthroleptis); lack of a median dorsal skin raphe (as in Arthroleptis); lack of parallel dorsal ridges (as in Ptychadena); neither finger nor toe tips heart shaped (as in Petropedetes) or expanded to discs. 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. Description of holotype (measures in mm). Typical small Phrynobatrachus, female with oval, compact body shape; snout-vent length: 18.8; short snout, rounded in dorsal and lateral view; canthus rostralis 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=IIColouration in life: 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. Colouration in formalin and ethanol after more than eight years: pattern slightly fainted, black head turned dark brown, reddish back turned to light brown. Variation (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. 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). Genetics. Phrynobatrachus rainerguentheri sp. nov. differs between 6.59-22.94 % in the investigated part of the 16 S rRNA gene from 33 West African and Central African species of the genus Phrynobatrachus (Table 2). The species with the lowest genetic divergence to P. rainerguentheri sp. nov. were P. ghanensis (mean difference 7.06 %), P. phyllophilus (7.06 %), P. guineensis (7.30 %) and P. al le n i (8.55 %). The intraspecific genetic variation of nine P. rainerguentheri sp. nov., one from the type locality (ZMB 77437, GenBank JQ 730857) and eight from Warri (ZMB 77737 -77744, GenBank JQ 954860 - JQ 954867) was 0-0.27 %. Habitat. 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 �� 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. Species GenBank # voucher p-distance min-max mean ��sd Other anuran species at the type locality where: Silurana tropicalis, Amietophrynus maculatus, A. regularis, Aubria subsigillata, Ptychadena cf. aequiplicata, Phrynobatrachus latifrons, P. francisci, Arthroleptis sp., Leptopelis sp., Afrixalus dorsalis, Hyperolius concolor and H. fusciventris (Onadeko & R��del 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 Phrynobatrachus spp., are clearly indicating a heavily altered state of this forest habitat. 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. Distribution. Phrynobatrachus rainerguentheri sp. nov. is known from the type locality and one further locality app. 250 km south-east from the type locality (Fig. 4). Etymology. The species is dedicated to Dr. sc. Rainer G��nther, from 1985-2006 curator of herpetology at the Museum f��r Naturkunde Berlin, in recognition for his outstanding contributions to herpetology., Published as part of R��del, 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 on pages 56-62, DOI: 10.5281/zenodo.281218, {"references":["Onadeko, A. B. & Rodel, M. - O. (2009) Anuran surveys in south-western Nigeria. Salamandra, 45, 1 - 14."]}

Astylosternus laticephalus sp. nov. R��del, Hillers, Leach��, Kouam��, Ofori-Boateng, Diaz & Sandberger Figs. 9���12 Holotype. ZMB 75449 (field and tissue #: ATE 10, female, 58.6 mm), Ghana, Atewa Forest Reserve, 6.23375 / - 0.56557, 14 April 2007, forest around stream next to road, coll. C. Ofori-Boateng, A. Hillers & G. Segniagbeto. Paratypes. Ivory Coast: MTD 48026 (field and tissue #: Ba04.24, female, 62.2 mm), ZMB 75454 -75458 (Ba04.26, subadult, 45.5 mm; Ba04.25, female, 62.5 mm; Ba04.21, female, 64.4 mm; Ba04.22, subadult, 43.3 mm; Ba04.23, female, 58.4 mm;), Banco National Park, 5.41667 / -4.10500, rainy season 2004, swampy forest, coll. N.E. Assemian, B. Toh�� & G. Kouam��; MNHN 1993.6073 (male, 53.8 mm), 1999.7333 (female, 54.8 mm), 1999.7334 (male, 48.7 mm), 1999.7335 (female, 59.8 mm), 1999.7736 (female, 52.6 mm), 1999.7337 (female, 51.8 mm), Banco National Park, Abidjan; Ghana: ZMB 75459 (AF 4, juvenile, 25.6 mm), ZMB 75459 -75460 (AF 3 juvenile; 26.7 mm; AF 2, tadpole), Afao Hills Forest Reserve, 6.25461 / - 2.29492, riparian forest around small stream in valley, coll. C. Ofori-Boateng & A. Hillers; ZMB 75450 -75453 (COB 90, juvenile, 34.7 mm; COB 318, subadult, 40.8 mm; COB 119, juvenile, 38.6 mm; COB 123, juvenile, 31.7 mm), Western Province, Ankasa Conservation Area, pristine forest along streams in dry season, coll. C. Ofori-Boateng; MVZ 244910 (female, 67 mm), Ankasa Conservation Area, 5.28173 / - 2.64022, 26 June 2004, wet evergreen forest area, trail adjacent to the bamboo cathedral, coll. A.D. Leach�� & R. Diaz; MVZ 244909 (juvenile, 35 mm), 28 June 2004, approximately 1 km farther from the bamboo cathedral, other data as MVZ 244909. Additional material. ZMB 77461 (field# AF 03), tadpole, Ghana, Afao Hills Forest Reserve, 6.25461 / - 2.29492, riparian forest around small stream in valley, coll. C. Ofori-Boateng & A. Hillers; ATE 7 (tissue only), juvenile, 25.8 mm; ATE 8 (tissue only), juvenile, 28.6 mm; ATE 27 (tissue only), male, 45.6 mm, Ghana, Atewa Forest Reserve, 6.23375 / - 0.56557, 15 April 2007, swampy area next to small stream, in valley in forest, coll. C. Ofori-Boateng, A. Hillers & G. Segniagbeto; COB 2202 (collection Ofori-Boateng, Kumasi, Ghana), Ghana, Atewa Forest Reserve, coll. C. Ofori-Boateng; three adult specimens without number (amphibian reference collection at University Abobo-Adjam��, Ivory Coast), male (51.0 mm), females (60.0, 67.0 mm), Ivory Coast, Banco National Park, 5.41667 / -4.10500, 5 May 2004, forest close to Banco River, coll. N.E. Assemian, N.G. Kouam�� & B. Toh��. Diagnosis. Astylosternus laticephalus sp. nov. has the typical body shape of frogs of the genus Astylosternus, however, with exceptionally broad heads, i.e. broader than in A. occidentalis; males of the new species without spines on throat and belly (always present in A. occidentalis males); males without nuptial black skin layer in pectoral region (present in male A. occidentalis from western Guinea); back brownish to brownish red, always with distinct red dots (red dots only rarely present in A. occidentalis); bicoloured iris, grey with a reddish upper third (A. occidentalis always with uniform grey iris). A. laticephalus sp. nov. differs from A. occidentalis by a mean of 3.2 % (range 2.9-3.7 %) in the investigated part of the 16 S rRNA gene. Genetic divergence to the morphologically most similar Central African species, A. diadematus, was 11.9 %. Description of the holotype [measures in mm]. The holotype is an adult female with a snout-vent length of 58.6; head width 22.7; head as long as broad; interorbital distance narrower than length of upper eyelid; snout rounded in dorsal view, obtuse in lateral view, longer than eye diameter; eyes large (7.9) and protuberant directed anterolaterally; interorbital distance 7.2; pupil vertically elliptical; eye diameter larger than tympanum diameter, 5.3; tympanum vertically elliptical; supratympanal fold from posterior vertical midpoint of orbit to meet with dorsal border of tympanic annulus, where descent is sharp and terminates at half the vertical diameter of the tympanic annulus dorsally to the axilla; nares closer to snout-tip (2.7) than to eye (5.5), nostrils oriented posterolaterally; internarial distance 4.4; canthus rostralis rounded and straight; loreal region slightly concave and gradually sloping laterally; single, small, tooth-like process at lower jaw symphysis, with corresponding socket in between premaxillae; upper premaxillae and maxillae with numerous teeth; vomerine teeth in form of two hemispherical odontophores, perpendicular to body axis, almost being in contact to each other medially, each with row of teeth like tips (5 left, 6 right); distance from odontophores to elliptical choanae slightly larger than maximum length of odontophores; choanae almost equal in size to odontophores; tongue broadly heart shaped, deeply notched anteriorly, densely beset with small papillae, extends over entire length of lower jaw; posterior 2 / 3 of tongue free. Forelimbs robust, fingers slender and long; prepollex absent; first finger 10.3, almost double the length of second finger (5.1); relative finger lengths I>III>II>IV; finger tips broadened without forming discs; subarticular tubercles large, subconical, protruding distally; thenar and palmar tubercles large and elliptical; supernumerary tubercles absent; number of subarticular tubercles on digits I-IV: 1, 1, 2, 2; no digital webbing; hind limbs short and robust; femur length 28.3, almost equal to tibia 28.4; tibia-fibula with longest toe 38.4; relative toe lengths IV>III>V>II>I; shortest toe (4.2) almost double the length of inner metatarsal tubercle (2.3); inner metatarsal tubercle large, elliptical; toe tips broadened without forming discs; number of subconical subarticular tubercles on toes I-V: 1, 1, 2, 3, 2; supernumerary tubercles absent; traces of webbing between toe bases; no skin fringes along toes. Tip of toe IV on left foot missing. Skin on head, back, belly and throat predominantly smooth; small whitish spines scattered on back, dorsal parts of extremities and eyelids; white warts posterior to angle of mouth, ventral from tympanum; posterior region of thighs areolate; irregularly spread narrow low tubercles along flanks; no tubercles or warts on forearms or interocular region; head with white scratches (scars). Base colour of dorsum in preservation is a light brown to reddish brown; back with irregularly arranged and shaped light beige spots; upper eyelids greyish; head laterally with two darker bars, the posterior one below eye; upper and lower lips white; supratympanic fold anteriorly rimmed thin black; warts on flanks beige; forelimbs dorsally light brown, slightly mottled beige, first two fingers almost uniform white; hind limb colour as forelimbs, dark bordered cross bars indicated in thighs (4) and lower legs (5-6); throat, belly, ventral parts of fore- and hind limbs uniform white or creamish; only ventral part of feet reddish brown, metatarsal tubercle and subarticular tubercles creamish white. Variation. Females seem to be larger (up to 67.0 mm; MVZ 244910) than males (53.8 mm), although the small sample size of the latter (N = 2) does not allow for generalization. The tympanum shape varies between vertically elliptical (most vouchers) to round; the supratympanic fold is sometimes thin to absent; sometimes tympanic membrane prominent within faint tympanic annulus. Other measures and indices identical between the two sexes; values are summarized in Tabs. 1 and 2. Dorsal and lateral skin texture can consist of many irregularly spaced small roundish warts, not arranged in rows, sometimes fused to small ridge like structures (Fig. 11). Males have paired subgular vocal sacs, visible in preserved frogs as slightly loose, granular black skin near the angles of the mouth; males with large brown nuptial pads on the external side of the thumb. Canthus rostralis in males more prominent bulging and rounded; males with more massive forearms than females. In contrast to other western West African Astylosternus, A. laticephalus sp. nov. males in breeding condition, lack spines on throat and belly. The dorsal base colouration in life consists of a light grey, brown, reddish brown or almost black; all specimens exhibit a more or less dense pattern of smaller to larger, irregularly shaped red spots on back; smaller black or brown spots may be present, in particular in juveniles; iris greyish, upper third reddish to orange; the anterior part of the eyelids bright yellow to red blotches, connected by a very conspicuous interorbital stripe of the same colour; snout tip and area posterior to interorbital stripe, including posterior part of eyelid, darker than rest of back, this pattern sometimes consisting of a symmetrical pair of triangular shaped black spots (Fig. 11); frenal area dark as snout tip; loreal area distinctly lighter coloured than snout tip; light infraorbital batch; supratympanal ridge usually bordered by thin black line; fore and hind limbs with or without distinct black cross bars; lateral colour gradually fainting from back colour to whitish venter, with irregularly scattered black points; ventral coloration (throat and abdomen) creamish-white; ventral surface of thighs and arms creamish-yellow with reddish brown mottling around lateral margins of limbs. In preservative all patterns and coloration almost as in life, but faded; the red dorsal spots remain as light beige spots. Tadpole [measures in mm; description based on ZMB 75460]. Exotrophic, lentic tadpole; Gosner stage 39; total length 62.1; body broad almost as long as wide, slightly elliptical in dorsal, slightly depressed in lateral view (possibly a preservation artefact); body length 23.8; body width 24.2; sides of body slightly concave, snout in dorsal view broadly rounded, a bit more pointed in lateral view; large lateral sacs originating posterior to eyes run along flanks; small eyes, positioned dorsolaterally, pupil round; nares small, positioned dorsolaterally, closer to snout trip than to eyes; oral apparatus in anteroventral position; dorsal lip wide and smooth, with large anterior gap between marginal papillae; lateral papillae ventral lip with large, uni- or biserial marginal papillae; upper jaw sheath massif, broad U-shaped, strongly serrated margin, most central ���tooth��� most prominent; lower jaw sheath massif, V-shaped, margin strongly serrated; labial tooth-row formula 1: 1 + 1 / 3; all keratodont rows on skin sheaths; labial keratodonts unidenticulate, connected by a hyaline skin; vent tube dextral; spiracle sinistral (almost invisible); very long tail axis (> 2.5 times body length); tail axis height largely exceeding height of dorsal and ventral fin, both fins very narrow; dorsal fin originates slightly posterior to tail body junction; dorsal fin almost parallel to tail axis up to rounded tip; no pores (neuromast canals) visible (possibly due to poor preservation or advanced developmental stage); body more or less beige, mottled with dark brown, tail fin brown to almost black in last third of tail. Newly metamorphosed froglets measured 25.6���28.7 mm (N = 2). Natural history. Not much is known about the biology of the new species. Astylosternus laticephalus sp. nov. lives in lowland rainforest habitats (Fig. 13), mostly close to small or mid sized streams (R��del et al. 2005; Assemian et al. 2006). In Banco National Park, Ivory Coast we recorded the new species in the leaf litter of swampy areas in closed canopy forest, open forest and in natural forest gaps, both in close vicinity to the River Banco and in larger distance (about 1 km) from the river (Assemian et al. 2006). A total of 12 specimens were only observed during the rainy season, 11 of them during the night. In the Ankasa Conservation Area we found the new species in wet evergreen forest on a trail adjacent to the Bamboo Cathedral (Fig. 13 b). Distribution. So far A. laticephalus sp. nov. is known from one forest in eastern Ivory Coast, the Banco National Park (Assemian et al. 2006), and various sites in Ghana: Afao Hills Forest Reserve, Draw River Forest Reserve and Ankasa Conservation Area in south-western Ghana (R��del et al. 2005; this paper), and Atewa Range in southern-central Ghana (Fig. 8). The latter site was chosen as type locality of the new taxon to underline the outstanding importance of this mountainous region for the Ghanaian biodiversity. Conservation status. The distribution of the species from eastern Ivory Coast to southern-central Ghana, including various protected areas (i.e. Banco National Park, Afao Hills Forest Reserve, Ankasa Conservation Area), seems to imply a IUCN RedList classification of ���Least Concern���. However, rainforests in south-eastern Ivory Coast are under intense logging pressure and highly fragmented. Forests in south-western Ghana are better protected but likewise fragmented and the Atewa Range is currently under threat because of small scale mining activities and further plans to mine for gold, diamonds and bauxite on an industrial scale. We thus suggest classifying the new species as ���Near Threatened���. Etymology. The name derives from the Latin words latus for wide and the Greek K������� for head, referring to the conspicuous wide head of the new species., Published as part of R��del, Mark-Oliver, Barej, Michael F., Hillers, Annika, Leach��, Adam D., Kouam��, N'Goran G, Ofori-Boateng, Caleb, Assemian, Emmanuel, Toh��, Blayda, Penner, Johannes, Hirschfeld, Mareike, Doumbia, Joseph, Gonwouo, Legrand Nono, Nopper, Joachim, Brede, Christian, Diaz, Raul, Fujita, Matthew K., Gil, Marlon & H, Gabriel, 2012, The genus Astylosternus in the Upper Guinea rainforests, West Africa, with the description of a new species (Amphibia: Anura: Arthroleptidae), pp. 1-29 in Zootaxa 3245 on pages 16-22, DOI: 10.5281/zenodo.280506, {"references":["Rodel, M. - O., Gil, M., Agyei, A. C., Leache, A. D., Diaz, R. E., Fujita, M. K. & Ernst, R. (2005) The amphibians of the forested parts of south-western Ghana. Salamandra, 41, 107 - 127.","Assemian, N. E., Kouame, N. G., Tohe, B., Gourene, G. & Rodel, M. - O. (2006) The anurans of the Banco National Park, Cote d'Ivoire, a threatened West African rainforest. Salamandra, 42, 41 - 51."]}

Rödel, Mark-Oliver, Barej, Michael F., Hillers, Annika, Leaché, Adam D., Kouamé, N'Goran G, Ofori-Boateng, Caleb, Assemian, Emmanuel, Tohé, Blayda, Penner, Johannes, Hirschfeld, Mareike, Doumbia, Joseph, Gonwouo, Legrand Nono, Nopper, Joachim, Brede, Christian, Diaz, Raul, Fujita, Matthew K., Gil, Marlon, H, Gabriel (2012): The genus Astylosternus in the Upper Guinea rainforests, West Africa, with the description of a new species (Amphibia: Anura: Arthroleptidae). Zootaxa 3245: 1-29, DOI: 10.5281/zenodo.280506

Astylosternus laticephalus sp. nov. Rödel, Hillers, Leaché, Kouamé, Ofori-Boateng, Diaz & Sandberger Figs. 9–12 Holotype. ZMB 75449 (field and tissue #: ATE 10, female, 58.6 mm), Ghana, Atewa Forest Reserve, 6.23375 / - 0.56557, 14 April 2007, forest around stream next to road, coll. C. Ofori-Boateng, A. Hillers & G. Segniagbeto. Paratypes. Ivory Coast: MTD 48026 (field and tissue #: Ba04.24, female, 62.2 mm), ZMB 75454 -75458 (Ba04.26, subadult, 45.5 mm; Ba04.25, female, 62.5 mm; Ba04.21, female, 64.4 mm; Ba04.22, subadult, 43.3 mm; Ba04.23, female, 58.4 mm;), Banco National Park, 5.41667 / -4.10500, rainy season 2004, swampy forest, coll. N.E. Assemian, B. Tohé & G. Kouamé; MNHN 1993.6073 (male, 53.8 mm), 1999.7333 (female, 54.8 mm), 1999.7334 (male, 48.7 mm), 1999.7335 (female, 59.8 mm), 1999.7736 (female, 52.6 mm), 1999.7337 (female, 51.8 mm), Banco National Park, Abidjan; Ghana: ZMB 75459 (AF 4, juvenile, 25.6 mm), ZMB 75459 -75460 (AF 3 juvenile; 26.7 mm; AF 2, tadpole), Afao Hills Forest Reserve, 6.25461 / - 2.29492, riparian forest around small stream in valley, coll. C. Ofori-Boateng & A. Hillers; ZMB 75450 -75453 (COB 90, juvenile, 34.7 mm; COB 318, subadult, 40.8 mm; COB 119, juvenile, 38.6 mm; COB 123, juvenile, 31.7 mm), Western Province, Ankasa Conservation Area, pristine forest along streams in dry season, coll. C. Ofori-Boateng; MVZ 244910 (female, 67 mm), Ankasa Conservation Area, 5.28173 / - 2.64022, 26 June 2004, wet evergreen forest area, trail adjacent to the bamboo cathedral, coll. A.D. Leaché & R. Diaz; MVZ 244909 (juvenile, 35 mm), 28 June 2004, approximately 1 km farther from the bamboo cathedral, other data as MVZ 244909. Additional material. ZMB 77461 (field# AF 03), tadpole, Ghana, Afao Hills Forest Reserve, 6.25461 / - 2.29492, riparian forest around small stream in valley, coll. C. Ofori-Boateng & A. Hillers; ATE 7 (tissue only), juvenile, 25.8 mm; ATE 8 (tissue only), juvenile, 28.6 mm; ATE 27 (tissue only), male, 45.6 mm, Ghana, Atewa Forest Reserve, 6.23375 / - 0.56557, 15 April 2007, swampy area next to small stream, in valley in forest, coll. C. Ofori-Boateng, A. Hillers & G. Segniagbeto; COB 2202 (collection Ofori-Boateng, Kumasi, Ghana), Ghana, Atewa Forest Reserve, coll. C. Ofori-Boateng; three adult specimens without number (amphibian reference collection at University Abobo-Adjamé, Ivory Coast), male (51.0 mm), females (60.0, 67.0 mm), Ivory Coast, Banco National Park, 5.41667 / -4.10500, 5 May 2004, forest close to Banco River, coll. N.E. Assemian, N.G. Kouamé & B. Tohé. Diagnosis. Astylosternus laticephalus sp. nov. has the typical body shape of frogs of the genus Astylosternus, however, with exceptionally broad heads, i.e. broader than in A. occidentalis; males of the new species without spines on throat and belly (always present in A. occidentalis males); males without nuptial black skin layer in pectoral region (present in male A. occidentalis from western Guinea); back brownish to brownish red, always with distinct red dots (red dots only rarely present in A. occidentalis); bicoloured iris, grey with a reddish upper third (A. occidentalis always with uniform grey iris). A. laticephalus sp. nov. differs from A. occidentalis by a mean of 3.2 % (range 2.9-3.7 %) in the investigated part of the 16 S rRNA gene. Genetic divergence to the morphologically most similar Central African species, A. diadematus, was 11.9 %. Description of the holotype [measures in mm]. The holotype is an adult female with a snout-vent length of 58.6; head width 22.7; head as long as broad; interorbital distance narrower than length of upper eyelid; snout rounded in dorsal view, obtuse in lateral view, longer than eye diameter; eyes large (7.9) and protuberant directed anterolaterally; interorbital distance 7.2; pupil vertically elliptical; eye diameter larger than tympanum diameter, 5.3; tympanum vertically elliptical; supratympanal fold from posterior vertical midpoint of orbit to meet with dorsal border of tympanic annulus, where descent is sharp and terminates at half the vertical diameter of the tympanic annulus dorsally to the axilla; nares closer to snout-tip (2.7) than to eye (5.5), nostrils oriented posterolaterally; internarial distance 4.4; canthus rostralis rounded and straight; loreal region slightly concave and gradually sloping laterally; single, small, tooth-like process at lower jaw symphysis, with corresponding socket in between premaxillae; upper premaxillae and maxillae with numerous teeth; vomerine teeth in form of two hemispherical odontophores, perpendicular to body axis, almost being in contact to each other medially, each with row of teeth like tips (5 left, 6 right); distance from odontophores to elliptical choanae slightly larger than maximum length of odontophores; choanae almost equal in size to odontophores; tongue broadly heart shaped, deeply notched anteriorly, densely beset with small papillae, extends over entire length of lower jaw; posterior 2 / 3 of tongue free. Forelimbs robust, fingers slender and long; prepollex absent; first finger 10.3, almost double the length of second finger (5.1); relative finger lengths I>III>II>IV; finger tips broadened without forming discs; subarticular tubercles large, subconical, protruding distally; thenar and palmar tubercles large and elliptical; supernumerary tubercles absent; number of subarticular tubercles on digits I-IV: 1, 1, 2, 2; no digital webbing; hind limbs short and robust; femur length 28.3, almost equal to tibia 28.4; tibia-fibula with longest toe 38.4; relative toe lengths IV>III>V>II>I; shortest toe (4.2) almost double the length of inner metatarsal tubercle (2.3); inner metatarsal tubercle large, elliptical; toe tips broadened without forming discs; number of subconical subarticular tubercles on toes I-V: 1, 1, 2, 3, 2; supernumerary tubercles absent; traces of webbing between toe bases; no skin fringes along toes. Tip of toe IV on left foot missing. Skin on head, back, belly and throat predominantly smooth; small whitish spines scattered on back, dorsal parts of extremities and eyelids; white warts posterior to angle of mouth, ventral from tympanum; posterior region of thighs areolate; irregularly spread narrow low tubercles along flanks; no tubercles or warts on forearms or interocular region; head with white scratches (scars). Base colour of dorsum in preservation is a light brown to reddish brown; back with irregularly arranged and shaped light beige spots; upper eyelids greyish; head laterally with two darker bars, the posterior one below eye; upper and lower lips white; supratympanic fold anteriorly rimmed thin black; warts on flanks beige; forelimbs dorsally light brown, slightly mottled beige, first two fingers almost uniform white; hind limb colour as forelimbs, dark bordered cross bars indicated in thighs (4) and lower legs (5-6); throat, belly, ventral parts of fore- and hind limbs uniform white or creamish; only ventral part of feet reddish brown, metatarsal tubercle and subarticular tubercles creamish white. Variation. Females seem to be larger (up to 67.0 mm; MVZ 244910) than males (53.8 mm), although the small sample size of the latter (N = 2) does not allow for generalization. The tympanum shape varies between vertically elliptical (most vouchers) to round; the supratympanic fold is sometimes thin to absent; sometimes tympanic membrane prominent within faint tympanic annulus. Other measures and indices identical between the two sexes; values are summarized in Tabs. 1 and 2. Dorsal and lateral skin texture can consist of many irregularly spaced small roundish warts, not arranged in rows, sometimes fused to small ridge like structures (Fig. 11). Males have paired subgular vocal sacs, visible in preserved frogs as slightly loose, granular black skin near the angles of the mouth; males with large brown nuptial pads on the external side of the thumb. Canthus rostralis in males more prominent bulging and rounded; males with more massive forearms than females. In contrast to other western West African Astylosternus, A. laticephalus sp. nov. males in breeding condition, lack spines on throat and belly. The dorsal base colouration in life consists of a light grey, brown, reddish brown or almost black; all specimens exhibit a more or less dense pattern of smaller to larger, irregularly shaped red spots on back; smaller black or brown spots may be present, in particular in juveniles; iris greyish, upper third reddish to orange; the anterior part of the eyelids bright yellow to red blotches, connected by a very conspicuous interorbital stripe of the same colour; snout tip and area posterior to interorbital stripe, including posterior part of eyelid, darker than rest of back, this pattern sometimes consisting of a symmetrical pair of triangular shaped black spots (Fig. 11); frenal area dark as snout tip; loreal area distinctly lighter coloured than snout tip; light infraorbital batch; supratympanal ridge usually bordered by thin black line; fore and hind limbs with or without distinct black cross bars; lateral colour gradually fainting from back colour to whitish venter, with irregularly scattered black points; ventral coloration (throat and abdomen) creamish-white; ventral surface of thighs and arms creamish-yellow with reddish brown mottling around lateral margins of limbs. In preservative all patterns and coloration almost as in life, but faded; the red dorsal spots remain as light beige spots. Tadpole [measures in mm; description based on ZMB 75460]. Exotrophic, lentic tadpole; Gosner stage 39; total length 62.1; body broad almost as long as wide, slightly elliptical in dorsal, slightly depressed in lateral view (possibly a preservation artefact); body length 23.8; body width 24.2; sides of body slightly concave, snout in dorsal view broadly rounded, a bit more pointed in lateral view; large lateral sacs originating posterior to eyes run along flanks; small eyes, positioned dorsolaterally, pupil round; nares small, positioned dorsolaterally, closer to snout trip than to eyes; oral apparatus in anteroventral position; dorsal lip wide and smooth, with large anterior gap between marginal papillae; lateral papillae ventral lip with large, uni- or biserial marginal papillae; upper jaw sheath massif, broad U-shaped, strongly serrated margin, most central “tooth” most prominent; lower jaw sheath massif, V-shaped, margin strongly serrated; labial tooth-row formula 1: 1 + 1 / 3; all keratodont rows on skin sheaths; labial keratodonts unidenticulate, connected by a hyaline skin; vent tube dextral; spiracle sinistral (almost invisible); very long tail axis (> 2.5 times body length); tail axis height largely exceeding height of dorsal and ventral fin, both fins very narrow; dorsal fin originates slightly posterior to tail body junction; dorsal fin almost parallel to tail axis up to rounded tip; no pores (neuromast canals) visible (possibly due to poor preservation or advanced developmental stage); body more or less beige, mottled with dark brown, tail fin brown to almost black in last third of tail. Newly metamorphosed froglets measured 25.6–28.7 mm (N = 2). Natural history. Not much is known about the biology of the new species. Astylosternus laticephalus sp. nov. lives in lowland rainforest habitats (Fig. 13), mostly close to small or mid sized streams (Rödel et al. 2005; Assemian et al. 2006). In Banco National Park, Ivory Coast we recorded the new species in the leaf litter of swampy areas in closed canopy forest, open forest and in natural forest gaps, both in close vicinity to the River Banco and in larger distance (about 1 km) from the river (Assemian et al. 2006). A total of 12 specimens were only observed during the rainy season, 11 of them during the night. In the Ankasa Conservation Area we found the new species in wet evergreen forest on a trail adjacent to the Bamboo Cathedral (Fig. 13 b). Distribution. So far A. laticephalus sp. nov. is known from one forest in eastern Ivory Coast, the Banco National Park (Assemian et al. 2006), and various sites in Ghana: Afao Hills Forest Reserve, Draw River Forest Reserve and Ankasa Conservation Area in south-western Ghana (Rödel et al. 2005; this paper), and Atewa Range in southern-central Ghana (Fig. 8). The latter site was chosen as type locality of the new taxon to underline the outstanding importance of this mountainous region for the Ghanaian biodiversity. Conservation status. The distribution of the species from eastern Ivory Coast to southern-central Ghana, including various protected areas (i.e. Banco National Park, Afao Hills Forest Reserve, Ankasa Conservation Area), seems to imply a IUCN RedList classification of “Least Concern”. However, rainforests in south-eastern Ivory Coast are under intense logging pressure and highly fragmented. Forests in south-western Ghana are better protected but likewise fragmented and the Atewa Range is currently under threat because of small scale mining activities and further plans to mine for gold, diamonds and bauxite on an industrial scale. We thus suggest classifying the new species as “Near Threatened”. Etymology. The name derives from the Latin words latus for wide and the Greek Kεφάλɩ for head, referring to the conspicuous wide head of the new species.

Phrynobatrachus ruthbeateae sp. nov. Figs. 1–2 Holotype. ZMB 77457 (field # 0 801, adult male), Cameroon, near Dja Biosphere Reserve, N 03° 24.257 ’, E 013°08.314’, 659 m a.s.l., primary swampy evergreen rainforest, 30 April 2009, coll. M.-O. Rödel, T. Doherty-Bone, M. Talla Kouete, P. Janzen, K. Garrett, R. Browne. Paratypes. ZMB 77458 -77459 (field # 0 808, adult male; 0 817, adult female), 1 May 2009, other data as holotype; ZMB 77460 (field # 0 826, adult female), N 03° 23.139 ’, E 013°07.608’, 650 m a.s.l., 2 May 2009, other data as holotype. Additional material. LG 0 785, female, SVL: 13.3. mm, N 03° 28.052 ’, E 013°03.425’, 684 m a.s.l, small swamp and river near campsite, 28 April 2009, other data as holotype; LG 0 825, female, SVL: 14.6 mm, other data as ZMB 77460. Diagnosis. The 16 S rRNA sequence data indicates that the new species is a member of the genus Phrynobatrachus. Morphologically this is visible by the body shape; presence of a tarsal tubercle; presence of webbing; lack of a median dorsal skin raphe, lack of a black spot in the tympanal region; lack of parallel dorsal ridges; neither finger nor toe tips heart shaped. The new species is characterized by a combination of small size; compact body shape; short round snout; black lateral face mask; males with a black throat, the anterior part beset with large flat black spines; females with white throat and uniform black lower mandible; absence of eyelid cornicle; larger warts on upper eyelids; scapular ridges converging in straight line; rest of dorsal skin slightly granular to smooth; flanks with large white warts; white belly; groin area and lower parts of thighs yellow; absence of nuptial pads in males; distinct discs on finger and toe tips; well developed webbing and 16 S rRNA sequence. Description of holotype [measures in mm]. Typical, small adult male Phrynobatrachus with long oval, compact body shape; snout-vent length: 17.8; short snout, rounded in dorsal and lateral view; canthus rostralis rounded; loreal region straight; head-width directly behind the eyes: 6.3; medium sized eyes positioned laterally; eye-diameter: 2.8; distance eye-nostril: 1.5; nares small and round, positioned laterally; distance nostril-snout tip: 1.0; nostril closer to snout than to eye; internarial distance: 2.1; interorbital distance: 2.2; pupil round; supratympanal ridge ranging from posterior corner of eye to forearm bases; tympanum diameter: 1.0, round and indistinct, much smaller than diameter of eye; upper and lower maxillae and praemaxillae with minute teeth, hidden by lips; vomerine teeth absent; broad cordiform tongue, tip distinctly notched, median papilla near anterior attachment of tongue; small round choanae. Forelimbs slender; upper arm: 4.0; lower arm: 3.4; hand to tip of finger III: 4.6; hand with large carpal tubercles, inner oval, outer round; carpal gland absent; fingers with small roundish subarticular tubercles, no additional tubercles on hands; relative finger length: II=IVColouration in life (Fig. 1 a–b): Basic colour of dorsal surface of head, upper part of flank, back and dorsal surfaces of extremities being a beige brown; only dorsal part of feet brown-orange; black lateral face mask extending from snout tip, below canthus, eye and supratympanal ridge to forearm bases, covering tympanum; continuing below arms as a lateroventral black band to almost groin region; posterior part of interorbital space with a indistinct band being darker than remaining parts of back; anterior to this band a narrower lighter line in-between eyelids; black pupil surrounded by lighter red-golden ring; iris golden; dorsal part of thighs with five (right) and four (left) dark cross-bars; shanks with four (right) and three (left) dark cross-bars; posterior part of dorsal skin on thighs uniform dark brown; vent surrounded by broad black triangle, extending towards belly; throat black; pectoral region white with few minute black points; belly white, bordered by black ventrolateral band; ventral part of lower arm black; other ventral surfaces of forelimbs gray; groin area and lower surfaces of thighs yellowish to orange; lower part of thighs anterior laterally bordered by black line; ventral surfaces of feet dark brown. Colouration in ethanol after almost two years: pattern only slightly fainted. Variation (Fig. 1 c–h). Snout-vent length in males 17.2–17.8 mm and 13.3–19.7 mm in females (smallest one presumably not adult); body dimensions generally as in holotype (Table 1). Females with thinner fingers, i.e. finger I; discs on fingers more conspicuous than in males; relative finger length in paratypes: IGenetics. Phrynobatrachus ruthbeateae sp. nov. differed between 4.5 –23.0% in the sampled region of the 16 S rRNA gene from 34 West African and Central African species of the genus Phrynobatrachus (Table 2). The species with highest genetic similarity to P. ruthbeateae sp. nov. all occur in Central Africa. The uncorrected pairwise sequence divergence to the most similar species was: P. chukuchuku (4.9 %), P. w e r n e r i (5.1 %), P. steindachneri (5.2 %), P. schioetzi (5.6 %), P. batesii (5.9 %), P. cricogaster (5.5 %), P. danko (6.1 %), and P. manengoubensis (6.1 %), respectively. The intraspecific variation of P. ruthbeateae sp. nov. ranged from 0.0– 1.2 % (Table 2). continued next page TABLE 2. (continued) Distribution and conservation status. Phrynobatrachus ruthbeateae sp. nov. is so far only known from the type locality (Fig. 3). However, it is very likely that the new species is widespread in western Central African lowland rainforests, south of the Sanaga River at least. For instance Burger et al. (2004) list and illustrate a small Phrynobatrachus (P. sp. 2) from forests between 350–660 m a.s.l. on the Monts Doudou, Gabon. These frogs might be conspecific with the species described herein; the only visible difference being a large yellow blotch on upper lip and loreal region. The Gabon frogs were calling during day and sunset in muddy terrain near small streams. Given the presumed large range in huge tracks of still undisturbed rainforest, the new species probably is of Least Concern according to the IUCN RedList criteria (IUCN 2001). However, currently P. ruthbeateae sp. nov. must be classified as Data Deficient until this predicted range is confirmed. Natural history. All specimens of P. ruthbeateae sp. nov. were observed in the rainy season during daytime in swampy primary rainforest (Fig. 4). Four individuals were recorded under closed canopy at the edge of large palm swamps. One female was collected within a large swamp in an area with small trees, palms forming an open canopy. Males were calling during the day, sitting very close to small puddles, well concealed below leaves. Calling males were widely spaced from each other. Unfortunately the call could not be recorded, however, to the human ear it sounded almost identical to the call of P. phyllophilus, consisting of a fast series of brief metallic clicks (Rödel & Ernst 2002). The presence of parasitic, subcutaneous mites on the posterior part of the belly and on the thighs (Fig. 1 b, f) indicates that the species is indeed mostly living in swamps: this family of Endotrombiculidae occur on many African Phrynobatrachus species, but are most abundant on species dwelling in very wet or humid environments (Spieler & Linsenmair 1999; Wohltmann et al. 2007; MOR, unpubl. data). Other Phrynobatrachus species living in syntopy with the new species were P. africanus, calling from more central parts of the swamps, and P. cornutus, living in drier parts at the edge of the swamps. Annual rainfall at the type locality summed to app. 1700 mm with highest peaks of precipitation from March-July and September-November. Annual mean of daily minimum temperature was about 19 °C (rarely dropping to 16 °C, never reaching 21 °C); the annual mean of daily maximum temperature was about 26 °C (rarely dropping to 23 °C, never exceeding 31 °C). Etymology. The first author dedicates this species to his mother, Ruth-Beate Rödel. Without her continuous support his scientific career would have never been possible., Published as part of Rödel, Mark-Oliver, Doherty-Bone, Thomas, Kouete, Marcel Talla, Janzen, Peter, Garrett, Katherine, Browne, Robert, Gonwouo, Nono Legrand, Barej, Michael F. & Sandberger, Laura, 2012, A new small Phrynobatrachus (Amphibia: Anura: Phrynobatrachidae) from southern Cameroon, pp. 54-68 in Zootaxa 3431 on pages 55-60, DOI: 10.11646/zootaxa.3431.1.4, http://zenodo.org/record/282047, {"references":["Burger, M., Branch, W. R. & Channing, A. (2004) Amphibians and reptiles of Monts Doudou, Gabon: species turnover along an elevational gradient. pp. 145 - 186, In: Fisher, B. L. (Ed.) Monts Doudou, Gabon: a floral and faunal inventory with reference to elevational variation, California Academy of Sciences Memoir, 28, 295 pp.","IUCN (2001) Red List Categories and Criteria version 3.1. http: // www. iucnredlist. org (accessed at 28 December 2011).","Rodel, M. - O. & Ernst, R. (2002) A new Phrynobatrachus from the Upper Guinean rain forest, West Africa, including a description of a new reproductive mode for the genus. Journal of Herpetology 36, 561 - 571.","Spieler, M. & Linsenmair, K. E. (1999) The larval mite Endotrombicula pillersi (Acarina: Trombiculidae) as a species-specific parasite of a West African savannah frog (Phrynobatrachus francisci). American Midland Naturalist, 142, 152 - 161.","Wohltmann, A., du Preez, L., Rodel, M. - O., Kohler, J. & Vences, M. (2007) Endoparasitic mites of the genus Endotrombicula Ewing, 1931 (Acari: Prostigmata: Parasitengona: Trombiculidae) from African and Madagascan anurans, with description of a new species. Folia Parasitologica, 54, 225 - 235."]}

Detailed tadpole descriptions for toads of the genus Werneria Poche, 1903 have been only published for W. preussi (Matschie, 1893) (Mertens 1938) and W. tandyi (Amiet, 1972) (Rödel et al. 2004), the latter description assigned to this species with reservation. A few characters of W. bambutensis (Amiet, 1972) tadpoles were mentioned by Amiet (1972). So far the assignment of tadpoles to a particular Werneria species was based on geography (locality and altitude). Three Werneria species, W. mertensiana, W. tandyi and W. bambutensis (Fig. 1), co-occur on Mount Manengouba in south-western Cameroon. There they inhabit forests and river edges in different but slightly overlapping altitudes (Rödel et al. 2004). During recent field work (November / December 2010) on Mount Manengouba we collected Werneria tadpoles which are herein described.

Amietophrynus channingi sp. nov. Figs. 1, 2 c, 4 c, 10 a���d, 11 a���d, 12 a���c. Holotype. MTSN 9674 (male) Democratic Republic of Congo, West Mwana, Itombwe Massif, 0 28.13875 E, 0 3.83781 S, 1302 m a.s.l., coll. Joseph Matunguru. Paratypes. IRSNB-KBIN 1982 (female) Democratic Republic of Congo, Beni; IRSNB-KBIN 1983 (subadult) Democratic Republic of Congo, Mutsora; IRSNB-KBIN 1984 (female) Democratic Republic of Congo, Bukavu; IRSNB-KBIN 1985 (juvenile) Democratic Republic of Congo, Irangira; MHNG 2520.25 (juvenile) Democratic Republic of Congo, Nduye, Alamby; ZFMK 62573 (male) Democratic Republic of Congo, Irangi; ZFMK 63895 - 6 (2 juveniles) Democratic Republic of Congo, Irangi. Additional material. IRSNB-KBIN 634 (male) Democratic Republic of Congo, Lulenga; IRSNB-KBIN 2216 (female) Democratic Republic of Congo, Zobia; IRSNB-KBIN 2217 (female) Democratic Republic of Congo, Zobia; IRSNB-KBIN 11363 (female) Democratic Republic of Congo, Walikale; IRSNB-KBIN 11364 (female + 2 juveniles) same data as IRSNB-KBIN 1985; MNHN 1933.027 (female) Congo Belge (= Dem. Rep. Congo), locality unknown; ZFMK 34376 (juvenile) Democratic Republic of Congo, Poko; ZFMK 63894 (male) same data as ZFMK 63895. Remark: The MNHN received a collection from ��� Congo Belge ��� in 1933 from Guy Babault (Bull. Mus. Natl. Hist. Nat. p. 18), today Democratic Republic of Congo, without further details on the collection localities. According to publications dealing with various taxa donated to the MNHN by G. Babault (Pellegrin 1933, 1935; Babault 1935; Berlioz 1935; Germain 1936) the locality in ��� Congo Belge ��� can be assigned to the Kivu area in eastern Democratic Republic of the Congo. Consequently, we assign MNHN 1933.027 to this new taxon. Diagnosis. Genetically the taxon belongs to the African toad genus Amietophrynus (Frost et al. 2006). As part of the Amietophrynus superciliaris -species complex, the taxon differs from all other members of the genus by a smooth dorsal skin in adult specimens, straight loreal region and large size. Medium to large sized toad with smooth dorsal skin in adults, granular in juveniles; body shape compact; tympanum distinct, drop-shaped, smaller than eye-diameter; parotids prominent, broad, width of parotid glands hardly reducing backwards, posterior tips of parotid glands rounded; eyelid slightly triangular in dorsal view, eyelid in lateral view forming a intimated triangular process; anterior and posterior part of back with different pattern; pair of dark spots present on posterior back region; extremities slender. Amietophrynus channingi sp. nov. is most similar to A. s. superciliaris as both species exhibit a triangular eyelid process in adults, a structure lacking in A. s. chevalieri (Figs. 2 a���c) and share the symmetric pair of abdominal spots (Figs. 3 c, e���g; 11 b, c). The new species differs from A. s. superciliaris by the following characters: shape of eyelid being more pointed and elongated in A. s. superciliaris; the eyelid width / eye diameter ratio shows no significant difference (Kruskal-Wallis-test; p> 0.05, N A.s.s.= 28, 0.24���0.39, x= 0.32 �� 0.04; N A.c.= 10, 0.23���0.38, x= 0.34 �� 0.04; H= 12.28); shape of parotid glands, much diminishing in width posteriorly and possessing a pointed tip in A. s. superciliaris and being more bulged and rounded at the posterior tip in A. channingi sp. nov.; colouration of the back, which is darker in the posterior region of the back and possesses a darker mid-dorsal line in A. channingi sp. nov., whereas the dorsum is more or less uniformly coloured in A. s. superciliaris (Figs. 3 c, e���g; 11 a���c); colouration of flanks, being reddish in A. s. superciliaris and brownish in the new species (Figs. 2 b���c). Moreover, juveniles without a fully developed eyelid process can be differentiated by the following colour patterns: interorbital pattern in juvenile A. channingi sp. nov. a depressed dark, V-like figure, covering the eyelids, while juvenile A. s. superciliaris usually possess two clearly separated interorbital spots, which may even be broken into numerous smaller spots (Figs. 4 b-c). The new species can be distinguished from A. s. chevalieri by the following characters: eyelid process being absent in A. s. chevalieri (present in A. channingi sp. nov.; Figs. 2 a, c); the eyelid width / eye diameter ratio is significantly higher in adults of the new species (Kruskal-Wallis-test; p A. s. chevalieri; A. channingi sp. nov. possess a symmetric pair of abdominal spots (lacking in A. s. chevalieri; Figs. 9 a���f, h; 11 a���c); lateral colouration being reddish-purple in A. s. chevalieri and brownish in A. channingi sp. nov. (Figs. 2 a, c). Description of the holotype. The holotype is an adult male in breeding condition; dorsal skin smooth, lateral and ventral skin rough (Figs. 10 b, c); compact and sturdy body shape; SUL 111.8 mm; head without bony ridges; canthus rostralis distinct, angular; loreal region slightly concave; snout very short, almost straight in lateral view; head width 44.1 mm; eye-diameter 12.2 mm; interorbital-distance 9.5 mm; nares closer to snout than to eye; eyelid with weak triangular process (Fig. 11 a), measuring 4.9 mm from lower eyelid border to tip of eyelid process; parotid length 33.5 mm; parotid width 11.8 mm; parotids bi-coloured, upper part pale brown like dorsum, lower part dark grey; gland openings present only on upper part, reaching colouration border; posterior to parotids a glandular dorsolateral fold, extending to groin area; fingers and toes simple; subarticular tubercles simple; relative length of fingers: III> I> II ��� IV; femur length 42.6 mm; tibia length 42.0 mm; foot length incl. longest toe 43.6 mm; inner metatarsal tubercle oval and prominent; outer metatarsal tubercle rounded, indistinct; manual webbing absent (Fig. 10 c), relative toe length: IV> III> V> II> I; pedal webbing rudimentary (Fig. 10 d): 1 (0.5), 2 (1 - 0.5), 3 (2 - 1.25), 4 (2.75-3), 5 (1.25); tarsal fold absent; secondary sexual characters: large dark nuptial pads on fingers I and II; miniscule spines scattered on lateral side of head and flanks below parotid glands, minuscule spines denser behind mouth gap; further minuscule spines cover anterior and dorsal side of upper arms; colouration in preservation: back pale brown; interorbital space with darker V-shaped pattern, the tips of the V pointing on the eyelid processes; dark vertebral line running backwards (Figs. 10 b; 11 b); posterior part of back with dark triangular figure and a symmetric pair of dark spots; skin of dark spots different in texture to remaining of back skin, blunt and ���fleecy���; dorsal and lateral colouration distinctly different, separated by thin white line; flanks coloured dark brown, this pattern ranging from snout-tip backwards and dissolving in the inguinal area, here replaced by dark brown spots (three on left, two on the right side); dark spots on anterior part of upper hind limb and on posterior side of lower hind limb; venter uniform pale. Variation. Medium to large sized toad with compact body shape; females being larger than males (SUL in males: 106.7���111.8 mm, in females: 100.3 ���143.0 mm; SUL of specimens from Congo according to Noble (1924: males: 98���112 mm, females: 133���149 mm); mean head width similar in both sexes, in males 38 % of SUL, in females 39 %; snout in lateral view straight almost truncate; distance eye-snout larger than eye diameter; eyelid in adult specimens more or less triangular shaped; tympanum more or less distinct, positioned in a concave area of the cheeks; tympanum vertically prolonged and drop shaped, smaller in horizontal diameter than eye-diameter; dorsal skin smooth in adults, granular in juveniles; parotid glands prominent, bulged, width posteriorly slightly reducing, usually rounded at posterior tip (parotid length / parotid width ratio in males: 2.82, in females: 2.73); glandular dorsolateral fold behind the parotids may be present; parotids bicoloured, lower parts like flanks, upper parts like back; border of the two parts forms a sharp ridge; parotid gland openings on upper part only; fingers and toes simple; subarticular tubercles simple; relative length of fingers: III> I> II ��� IV (mean length of finger I / length of finger III ratio in males: 89 %, in females: 94 %); manual webbing absent; extremities slender in comparison to body shape; mean tibia length 40 % of SUL in males, in females 41 %; inner metatarsal tubercle present, oval and prominent; outer metatarsal tubercle rounded, sometimes indistinct; relative toe length: IV> III> V> II> I; webbing rudimentary: 1 (0.5), 2 (1 - 0.5), 3 (2 - 1) or 3 (2 - 1.5), 4 (2.75 - 2.75) or 4 (2.75-3), 5 (1.25); males in breeding condition with hypertrophied forearms and swellings on fingers I and II. Colouration. Dorsum dirty yellowish-browinish, extending from the tip of the snout, along the upper side of head and parotids backwards to the posterior part of the back (Fig. 11); dark V-shaped pattern on head covering the eyelids, usually a vertebral line running from this pattern backwards, forming a dark triangular pattern on the posterior part of the back (Figs. 11 b, c); back with symmetric pair of dark spots; loreal area, lower parts of parotids and flanks brownish; upper parts of lateral colouration darker than ventral part, partly almost black, posterior most part of flanks orange-brown; forearms light purple-greyish above, usually bordered by white line, and dark below; inguinal region with small black spots; upper hind limbs and feet with black transversal bars (Figs. 11 c, d); forearms with distinct white line between pale dorsal and dark ventral colouration; throat and venter pale yellow; juveniles coloured like adults but may possess additional pairs of spots on the back and the interorbital V-pattern, covering large parts of the eyelids; this pattern is more distinct in juveniles than in adults; juveniles with additional transversal bars on tibiofibula, this pattern disappearing in larger specimens. Orts (1970) figured two specimens after completed metamorphosis: the first uniform brown-blackish coloured and the second with a ���typical toad pattern��� consisting of symmetric pairs of spots (as in A. regularis, A. camerunensis, etc.). According to Orts (1970) the typical interorbital V-pattern develops within the first two weeks. Colouration in preservation. Preserved specimens still exhibit the separation of dark and light colour as in life; dark V-pattern between eyes, vertebral line and darker triangular posterior region of back may be hardly or not visible; flanks turn greyish or yellowish, and the brownish colour may turn almost black; brownish flank colouration sometimes recognizable when the lateral skin got folded and remains of former colouration has been covered between folds; pair of dark spots on the posterior part of the back sometimes bleached out and hardly recognizable. Tadpole description. The tadpole of this new taxon has been briefly described by Orts (1970). According to Orts��� (1970) description and figures: body oval in dorsal view in young stages, becoming more elongated in later stages (Figs. 12 a���c); body slightly flattened; mascular tail slender, converging caudally; dorsal and ventral fin present, tail (fins included) of similar height as body; tail fin rounded at tip of tail; tail 1.33 to 1.75 as long as body; eyes dorsally positioned, moving into more lateral position during growth, distance snout-tip to eye approx. 20���25 % of body length; spiraculum positioned at midbody; mouth ventrally positioned, surrounded by robust lips; colouration at 10 mm total length: grey above and cr��me-white, translucent below; tadpoles of 10.5 mm length (4 mm body length, Gosner stage 25; Fig. 12 a) with keratodont formula: 1: 1 + 1 // 1 + 1: 2; development of posterior pats at length of 13 mm (body length 5 mm); keratodont formula: 1: 1 + 1 // 3 (tadpole length> 13 mm); same keratodont formula in later stages; metamorphosis finished at body length (measured from tip of snout to anus) of 9 to 10 mm; for more details see Orts (1970). Natural history. Although the taxonomic status of this species has so far been overlooked, much more about the natural history of this taxon has been published compared to the other taxa of the Amietophrynus superciliaris - species complex. A. channingi sp. nov. is active in the rainy season, and can then be observed almost everywhere in the forest (Noble 1924). The species is nocturnal at least in captivity, where also cannibalistic behaviour was recorded (Orts 1970). When disturbed, the toad may be feigning death by ���lowering the head to the ground, and dropping the prominent upper eyelid��� (Noble 1924). The diet comprises various invertebrate taxa (Noble 1924). The species seems to feed preferably on snails around the Irangi research station (DRC) as local research assistants regularly found the species in great numbers in pits dug by road workers to raise snails for food consumption. According to Noble (1924) oviposition may occur during several months, breeding thus potentially may take place throughout most of the year. Eggs are very numerous in comparison to other forest toads, 1200���1500 in Amietophrynus tuberosus (referred to as A. polycercus in Noble 1924) and up to three times that number in A. channingi sp. nov., and egg size ranges from 1.6 ���2.0 mm (envelopes removed). According to own measures, egg size ranges from 1.40���1.95 mm (N= 114, mean= 1.69, SD= 0.11). Orts (1970) recorded tadpoles of 10 mm in Irangi in Democratic Republic of Congo in mid-June, at the end of the rainy season. He observed more than 3000 tadpoles in a puddle of only of 40 x 25 cm and 7 cm depth, covered with vegetation, where tadpoles moved on the ground and among leaf litter. Other tadpoles inhabited a series of puddles of a diameter of approximately one meter in mid- April. These breeding sites were connected to small rivers, which may transform into large currents after rain and then wash the tadpoles away (Orts 1970). 1 2 3 4 5 6 7 8 9 A. brauni (N= 3) 0 �� 0 GenBank A. poweri (N= 2) 0.0428 - 0.0446 0.0018 GenBank 0.0439 �� 0.0007 A. rangeri (N= 2) 0.0611 - 0.0720 0.0648 - 0.0700 0.0018 GenBank 0.0668 �� 0.0050 0.0674 �� 0.0022 A. channingi (N= 2) 0.0805 - 0.0849 0.0648 - 0.0673 0.0648 - 0.0685 0.0000 D.R. Congo 0.0831 �� 0.0020 0.0687 �� 0.0017 0.0666 �� 0.0021 A. s. chevalieri (N= 4) 0.0772 - 0.0814 0.0669 - 0.0682 0.0557 - 0.0612 0.0218 - 0.0219 0 �� 0 Guinea, Liberia 0.0793 �� 0.0022 0.0676 �� 0.0007 0.0584 �� 0.0029 0.0218 �� 0.0000 A. s. superciliaris (N= 2) 0.0732 - 0.0779 0.0673 - 0.0719 0.0574 - 0.0650 0.0274 - 0.0286 0.0085 - 0.0101 0.0017 Cameroon, Nigeria 0.0754 �� 0.0018 0.0696 �� 0.0020 0.0612 �� 0.0041 0.0280 �� 0.0007 0.0093 �� 0.0008 A. camerunensis (N= 2) 0.0734 - 0.0780 0.0687 - 0.0730 0.0634 - 0.0725 0.0776 - 0.0809 0.0759 - 0.0789 0.0787 - 0.0823 0.0035 GenBank 0.0751 �� 0.0019 0.0709 �� 0.0023 0.0679 �� 0.0043 0.0792 �� 0.0018 0.0774 �� 0.0016 0.0802 �� 0.0016 A. gutturalis (N= 2) 0.0807 - 0.0864 0.0828 - 0.0861 0.0756 - 0.0840 0.0757 - 0.0774 0.0825 - 0.0840 0.0797 - 0.0839 0.0502 - 0.0540 0.0018 GenBank 0.0839 �� 0.0021 0.0843 �� 0.0015 0.0791 �� 0.0036 0.0765 �� 0.0009 0.0833 �� 0.0008 0.0818 �� 0.0019 0.0521 �� 0.0016 A. tuberosus (N= 1) 0.1049 - 0.1067 0.1138 - 0.1153 0.0873 - 0.0939 0.0927 - 0.0930 0.0946 - 0.0962 0.0894 - 0.0932 0.0902 - 0.0926 0.0928 �� 0.0000 --- GenBank 0.1055 �� 0.0010 0.1146 �� 0.0010 0.0906 �� 0.0047 0.0928 �� 0.0002 0.0954 �� 0.0012 0.0913 �� 0.0027 0.0914 �� 0.0017 Distribution. Amietophrynus channingi sp. nov. is known from various localities in the eastern Democratic Republic of Congo, East Lower Guinea (Boulenger 1919; Wheeler 1922; Noble 1924; Loveridge 1936; Orts 1970). One specimen (IRSNB-KBIN 10385) is labelled as originating from ���Mutsora, Rwanda ���, but, given coordinates reveal that the locality is situated in the Democratic Republic of Congo. Due to the vicinity of localities of A. channingi sp. nov. close to the eastern border of the Democratic Republic of Congo (Fig. 1), the distribution of the toad may cover westernmost parts of Uganda, Rwanda and Burundi in suitable habitats. However, recent herpetological works failed to detect the species in the Albertine Rift (Drewes & Vindum 1994; Vonesh 2001). Ethnozoology. According to Noble (1924) natives fear the whitish secretion, as they might go blind. Etymology. The specific epithet channingi is a patronym. We name the species in honour of Prof. Dr. Alan Channing from the University of the Western Cape (South Africa) in recognition of his invaluable contributions to our knowledge of African amphibians., Published as part of Barej, Michael F., Schmitz, Andreas, Menegon, Michele, Hillers, Annika, Hinkel, Harald, B��hme, Wolfgang & R��del, Mark-Oliver, 2011, Dusted off ��� the African Amietophrynus superciliaris - species complex of giant toads, pp. 1-32 in Zootaxa 2772 on pages 18-24, DOI: 10.5281/zenodo.276862, {"references":["Pellegrin, J. (1933) Poissons de la region du Kivu adresses par M. Guy Babault. Bulletin de la Societe Zoologique de France, 58, 169 - 175.","Pellegrin, J. (1935) Description de deux poissons nouveaux de la region du Kivu appartenant au genre Varicorhinus. Revue de zoologie et de botanique africaines, 26, 401 - 403.","Babault, G. (1935) Note sur les Boocercus du Kivu. Bulletin du Museum National d'Histoire Naturelle, 2 e Serie, 7, 224 - 225.","Berlioz, J. (1935) Etude d'une collection d'oiseaux du Congo Belge. Bulletin du Museum National d'Histoire Naturelle, 2 e Serie, 7, 159 - 164.","Germain, L. (1936) Contributions a la faune malacologique de l'Afrique Equatoriale. Bulletin du Museum National d'Histoire Naturelle, 2 e Serie, 8, 146 - 157.","Frost, D., Grant, T., Faivovich, J., Bain, R. H., Haas, A., Haddad, C. F. B., de Sa, R. O., Channing, A., Wilkinson, M., Donnellan, S., Raxworthy, C. J., Campbell, J. A., Blotto, B. L., Moler, P., Drewes, R. C., Nussbaum, R. A., Lynch, J. D., Green, D. M. & Wheeler, W. C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1 - 291.","Orts, S. G. (1970) Description et ecologie des formes larvaires de Bufo superciliaris Blgr (Amphibia, Bufonidae). Revue de zoologie et de botanique africaines, 81, 207 - 219.","Boulenger, G. A. (1919) Batraciens et reptiles recueillis par le Dr. C. Christy au Congo Belge dans les districts de Stanleyville, Haut-Uele et Ituri en 1912 - 1914. Revue Zoologique Africaine (avec supplement botanique), 7, 1 - 29.","Wheeler, W. M. (1922) Chpt. II. The ants collection by the American Museum Congo Expedition. In: Wheeler, W. M. (Ed.) Ants of the American museum Congo Expedition. A contribution to the myrmecology of Africa. Bulletin of the American Museum of Natural History, 45, 39 - 269.","Loveridge, A. (1936) African reptiles and amphibians in Field Museum of Natural History. Zoological Series of Field Museum of Natural History, 22, 1 - 111.","Drewes, R. C. & Vindum, J. V (1994) Amphibians of the Impenetrable Forest, southwest Uganda. Journal of African Zoology, 108, 55 - 70.","Vonesh, J. R. (2001) Natural history and biogeography of the amphibians and reptiles of Kibale National Park, Uganda. Contemporary Herpetology, 4, http: // www. contemporaryherpetology. org / ch / 2001 / CH _ 2001. htm."]}

Amietophrynus superciliaris superciliaris (Boulenger, 1888 ��� 1887 ���) Figs. 1, 2 b, 3 a���g, 4 b, 5, 6e���g, 7 b���c. Bufo superciliaris Boulenger, 1888 ��� 1887 ���, Proc. Zool. Soc. London 1887: 565 Bufo laevissimus Werner, 1897, Sitzungsber. Akad. Wiss. M��nchen 27: 212 Amietophrynus superciliaris ��� Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sa, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, & Wheeler, 2006, Bull. Am. Nat. Hist. 297: 363 Type locality. ���Rio del Rey, Cameroons ��� Material examined. MHNG 757.61 (juvenile) Cameroon, Efulen; MHNG 917.99 - 100 (2 juveniles) Cameroon, Sangmelima, Foulassi; MHNG 955.88 - 89 (female + juvenile) Cameroon, Sangmelima; MHNG 1018.74 - 75, MHNG 1018.78, MHNG 1018.88, MHNG 1018.91 (2 males + 3 juveniles) Cameroon, Sangmelima; MHNG 1018.77 (female) Cameroon, Ambam (Ntem); MHNG 1390.94 - 96 (male + 2 females) Cameroon, Okola, Yaound��; MHNG 1390.97 (juvenile) Cameroon, Kolmon Nga, Yaound��; MHNG 1467.9 (female) Gabon, exact locality unknown; MHNG 2094.10 (female) Cameroon, Ongot; MHNG 2207.55 - 57 (2 females + juvenile) Gabon, Ogou��e- Ivindo, Makokou; MHNG 2207.62 (female) Gabon, Loa Loa-Ivindo; MNHN 1946.157 (juvenile) Gabon, exact locality unknown; ZFMK 4578 (female) Cameroon, Kumba; ZFMK 8328 (female) Cameroon, Victoria; ZFMK 14905 - 6 (2 females) Cameroon, Mezam, Bafut; ZFMK 15990 (female) Cameroon, exact locality unknown; ZFMK 18682 (juvenile) Cameroon; ZFMK 57800 (female) Cameroon, Kumba; ZFMK 67268 (juvenile) Cameroon, Meked; ZFMK 73192 (juvenile) Gabon, Barrage de Kingu��l��; ZFMK 90569 (female) Cameroon, Rumpi Hills, Mofako Balue; ZMB 3907 (female) Cameroon, exact locality unknown; ZMB 19919 - 20, ZMB 19924, ZMB 71139 - 40 (formerly part of ZMB 19920) (4 females + juvenile) Cameroon, Bipindi; ZMB 19921 (female) Cameroon, exact locality unknown; ZMB 19922 - 23 (2 females) Cameroon, Kribi; ZMB 19971 - 73 (3 juveniles) Cameroon, Bipindi; ZMB 20074 (juvenile) Cameroon, exact locality unknown; ZMB 20076 (juvenile) Cameroon, exact locality unknown; ZMB 20694 (juvenile) Cameroon, south Cameroon; ZMB 20718 (male) Cameroon, south Cameroon; ZMB 28775, ZMB 74317 - 8 (formerly part of ZMB 28775) (female + 2 juveniles) Cameroon, Bipindihof; ZMB 32037 (juvenile) Cameroon, Bipindi; ZMB 74319 (juvenile) Nigeria, Cross River; ZMB 74522 (juvenile) Cameroon, Nfakwo; ZSM 148 / 1989 / 1-2 (syntypes of Bufo laevissimus) (2 juveniles) Cameroon, exact locality unknown. Problematic material. MNHN 1923.50 (juvenile) French Congo, locality unknown (today territories of Gabon, Central African Republic, Republic of the Congo); MNHN 1933.16 (adult) Equatorial Africa, locality unknown; MNHN 2002.0655 (adult) Central African Republic, La Mabok��, Mboye. As MNHN 2002.0655 was completely dried out, the taxon specific characters are irrecognizable. Diagnosis. Genetically the taxon belongs to the African toad genus Amietophrynus (Frost et al. 2006). As part of the Amietophrynus superciliaris -species complex, the taxon differs from all other members of the genus by a smooth dorsal skin in adult specimens, straight loreal region and large size. Large toad; dorsal skin smooth in adults, granular in juveniles; body shape very broad, ovoid; tympanum distinct, drop-shaped, smaller than eye diameter; parotids prominent, shape of parotids like slender drop, parotid glands distinctly converging caudally, posterior end of parotid glands pointed; eyelid triangular in dorsal view, enlarging from edge to middle of eyelid in lateral view, forming a triangular eyelid process (Fig. 2 b); flank colouration reddish-purple; posterior part of back coloured like anterior part of dorsum; pair of spots on abdominal region present; extremities slender; males with nuptial swelling on fingers I and II. A. s. superciliaris can be distinguished from A. s. chevalieri and the new species A. channingi sp. nov. by colouration and body shape. For characters distinguishing between A. s. superciliaris and the two other taxa see below. Description. Large sized Amietophrynus species, with very robust body shape; females distinctly larger than males (maximum snout-urostyle-length ��� SUL ��� in males: 116.0��� 128.4 mm, in females: 108.1���153.4 mm); mean head width similar in both sexes, in males 38 % of SUL, in females 39 %; snout in lateral view short, slightly rounded; head without bony ridges; canthus rostralis distinct, angular; loreal region straight, slightly concave; distance eye-snout similar to eye diameter; nares closer to snout tip than to eye; eyelid in adult specimens of prominent triangular shape from dorsal and in lateral view, possessing a distinct prolongation in the middle; eyelid process in juveniles less distinct and almost absent in very small specimens; tympanum more or less distinct, positioned at a concave part of the cheeks; tympanum vertically prolonged, horizontal diameter smaller than eye diameter; dorsal skin smooth in adults, warty in juveniles; parotid glands very prominent, shaped like a slender drop, converging backwards, usually with a pointed tip (parotid length / parotid width ratio in males: 3.17, in females: 3.23); in some specimens posterior to parotids a glandular bulge extending to the groins as a dorsolateral fold; parotids distinctly bi-coloured: like flanks on lateral lower part and like back on dorsal surface; gland openings only on dorsal part of parotids (rarely extending onto the borderline of the colour transition); border between the two colours sharp; fingers and toes simple, not enlarged at the end; subarticular tubercles simple; relative length of fingers: III> I> II ��� IV (mean length of finger I / length of finger III ratio in males: 88 %, in females: 94 %); manual webbing absent; femora short and slender; mean tibia length in both sexes 38 % of SUL; inner and outer metatarsal tubercle present; relative toe length: IV> III> V> II> I; webbing rudimentary: 1 (0.5), 2 (1 - 0.5), 3 (2 - 1) or 3 (2 - 1.25), 4 (2.75 - 2.75), 5 (1) or 5 (1.25); females growing larger than males; males with nuptial pads on fingers I and II. Boulenger (1888) mentions the presence of a tarsal fold in his description of the species, while Werner (1897) states that the tarsal fold is absent in Bufo laevissimus (current synonym of A. s. superciliaris, see below); a character also used within his bufonid key. A distinct tarsal fold was absent in all examined specimens, only an indistinct bulging of the skin has been observed in some specimens. In dehydrated specimens, this somehow concave skin might be mistaken for a tarsal fold. Juveniles possess a granular skin till they reach a size of about 40 mm (Amiet & Perret 1969). The largest specimen still showing minuscule warts on the dorsum measured 46 mm (MHNG 917.99), in few subadult and predominantly smooth specimens, single tiny bulges were recognizable on the eyelids. The eyelid projection can be very small and almost absent in juveniles of A. s. superciliaris, forming only a very indistinct ���swelling��� on the eyelid. Colouration. Dorsum a pale yellowish colour or marbled orange yellow (Figs. 3 b, c, e���g), extending from the tip of the snout, along the upper side of the parotids backwards to the abdominal region; a pair of dark spots on posterior third of back (Figs. 3 a, c, e���g); additional smaller spots on anterior part of back or between eyes in few adult specimens (Figs. 3 c, e); intense red lateral colouration from tip of snout, continuing beneath the canthus rostralis and along entire flanks; sometimes parts of flank colouration darker, red-purple; ventral colouration ranging from light to intense reddish colour (Fig. 3 d); extremities dark purple; inguinal region rarely with small spots; upper hind limbs and feet with white transversal bars; anterior extremities dark purple ventrally; gular region coloured like venter; juveniles coloured like adults but with additional transversal bars on tibiofibula, usually less distinct in adults (Figs. 3 a���c, e); juveniles with more spots in addition to the abdominal pair on anterior part of back and in the interorbital region (Figs. 3 a; 4; 5), these dark spots of different structure than surrounding skin, ���drier and softer���. Interorbital markings of juvenile specimens usually consist of a pair of small blotches (sometimes dissolved into several smaller spots), caudally converging towards each other and barely reaching the eyelids (Figs. 4 b; 5). Exceptionally these blotches are fused (Fig. 4 b), a pattern also reported by Andersson (1905). Boulenger (1887) describes a ���very fine lighter vertebral line��� in some juveniles. According to Amiet & Perret (1969) juveniles possess a brown-black gular colouration and breast, while adults show an overall pale belly. Perret & Mertens (1957) comment on the absence of the pair of dark spots on the posterior part of the back in two specimens. Werner (1898) mentions a female from Cameroon (referred to as Bufo laevissimus) with pale red-brown upper surface and chocolate brown flanks. Colouration in preservation. Preserved specimens may loose their reddish colouration, which turns brownish. The typical reddish colouration was only observable where it remained covered between skin folds; in other specimens it completely disappeared. Nonetheless, in many specimens a clear differentiation between the lateral and dorsal colouration is recognizable. Very old specimens may be bleached out completely, lacking any colour pattern; e.g. even the pair of abdominal spots then may only be recognizable by a closer examination of the skin structure. Natural history. Amietophrynus s. superciliaris inhabits the forest floor in primary forest vegetation along small rivers (Fr��tey & Blanc 2001; Lasso et al. 2002; Figs. 6 e���g). Akani et al. (2004) report the finding of the species in the rainy season in a pristine swamp site. But the species also inhabits secondary growth with dense vegetation or even plantations (Amiet 1976 a, 1986; R��del et al. 2004). Occasionally the species can be seen in more open places searching for food (Sanderson 1936 a). Similarly, local farmers in western Cameroon reported to the senior author on occasional findings of the species in plantations. During the late rainy season (September 2010) M. Hirschfeld and M.- O. R��del (pers. obs.) observed an adult male (SUL 124 mm) in a near pristine part of the Ebo forest, Cameroon, over a period of four days. During this time the toad only moved about 25 m, the largest daily distance being about 15 m. During two consecutive nights the male climbed onto the top of two different bare rocks of 50���60 cm height, respectively and seemed to stay there motionless for the night. The other days it was observed sitting under a smaller tree, turning its body only once into another direction. According to Amiet (1976 a) breeding males possess slightly hypertrophied anterior extremities and black callosities on fingers I and II. A. s. superciliaris males lack a vocal sac (Perret & Amiet 1971) and the species is assumed to utter no advertisement call (Perret & Mertens 1957; Tandy & Keith 1972). A published sonogram of a A. s. superciliaris vocalization is not regarded to show an advertisement call (Amiet 1976 a). This vocalization, a dull buzzing noise, lasted for almost 1��� 1.5 s, comprising a long series of pulses (Amiet 1976 a, b). Amiet only observed movements of the flanks during sound emission, but no movements of the gular region (Amiet 1976 a). Hence, Amiet doubts that this noise represents an advertisement call, as it can be heard on short distances only. However, it may function in short distance recognition (Amiet 1976 a, 1989). Clutches are deposited and tadpoles may develop in slowly running streams (Amiet 1976 a, 1986, 1989; Gossmann et al. 2002; Figs. 6 e���f). Egg size ranges from 1.40���1.95 mm (N= 102, mean= 1.67, SD= 0.09). Amiet (1976 a) assumes that reproduction takes place in the dry season (January ��� March) as most specimens have been found during that time and only at that time males show their nuptial dress. Juveniles have been found in January in Nigeria (M.- O. R��del & A.B. Onadeko, pers. obs; Figs. 3 a; 6 e) and Cameroon (Schmitz 1998). Amiet & Perret (1969) report on juveniles (Prosicuophora basoglui, Nyctotheroides ptychadenae) found in adult A. s. superciliaris. Distribution. Amietophrynus s. superciliaris inhabits the western Lower Guinean rain forest (Fig. 1). At present the taxon is known from localities in Nigeria (Boulenger in Perret & Mertens 1957; Schi��tz 1963; Akani et al. 2004; this paper), Cameroon (e.g. M��ller 1910; Barbour 1911; Parker 1936; Sanderson 1936 b; Mertens 1940; Perret 1966; Herrmann et al. 2005, this paper), Gabon (Fr��tey & Dewynter 1998; Fr��tey & Blanc 2001; L��tters et al. 2001; Blanc & Fr��tey 2004; Pauwels & R��del 2007), and Equatorial Guinea (Nieden 1910; Lasso et al. 2002). According to Fr��tey & Blanc (2000) the species is also known from the Republic of Congo and the Central African Republic, but the authors do not provide localities. Joger (1990) lists A. s. superciliaris (MHNP 1968.331) for the Central African Republic (Oubangi-Chari = without specification), but the respective collection number is associated with a Ptychadena trinodis (mentioned with this number in the very publication). The only specimen of A. s. superciliaris with a similar collection number (MHNP 1968.359) refers to a specimen from ���Afrique���. However, a specimen in very bad condition from the Central African Republic is deposited in the MNHN collection (MNHN 2002.0655; see remark above). The placement of MNHN 2002.0655 in the distribution map (Fig. 1) has been matched according to publications referring to the same locality (Joger 1990; Rasmussen et al. 2000). Hence, we tentatively assign A. s. superciliaris to the herpetofauna of the western part of the Central African Republic, but suggest verifying the taxonomic status of this population as soon as new material becomes available. The wide distribution of A. s. superciliaris in western lowland rain forests of Central Africa (Fig. 1) makes it likely that this toad is indeed living in the south-western Central African Republic and possibly likewise in western Congo. Taxonomy. Amietophrynus superciliaris was described based on a series of juvenile specimens from ���Rio del Rey Cameroons ��� (Boulenger 1887; Fig. 5). The description of Bufo laevissimus was based on juveniles and one adult specimen from Cameroon (Werner 1897), and in a later publication Werner (1898) describes a second adult toad. Werner (1897) distinguishes the two species on the size of the parotid glands and the lack of a dorsolateral fold in B. laevissimus. According to Frost (2010) B. laevissimus was synonymised with B. superciliaris by Andersson (1905). Indeed, Andersson (1905) discusses the consistency in morphometrics between juveniles of B. laevissimus and those of B. superciliaris and remarks that distinguishing characters presented by Werner (1897) are based on the presence of an adult specimen in his material only. He concluded that the differences in the descriptions may have been the result of allometric growth (Andersson 1905). Earlier, Boulenger (1900) remarks that ���adult specimens [of A. s. superciliaris] have been redescribed by Werner under the name of B. laevissimus, from Cameroons ���. Boulenger (1900) reports on a toad of about 120 mm and already lists B. laevissimus as a synonym of B. superciliaris. Andersson (1905) later provides a more detailed comparison and confirmed the conspecificity of the two species. He states that ���therefore it can be that both Boulenger���s and Werner���s descriptions correspond with the case in this same species, that of the former with the young, the latter with the older ones���. He also cites Boulenger���s (1900) publication. Consequently, the synonymy of Bufo laevissimus Werner, 1897 with Amietophrynus s. superciliaris (Boulenger, 1887), as defined in this work, is based on Boulenger (1900). The syntypes of Bufo laevissimus Werner, 1897 are deposited in the ZSM collection. Two juvenile syntypes (ZSM 148 / 1989 / 1-2) are still present in the ZSM collection (and have been examined herein), while the only adult female syntype (ZSM 1113 /0) is lost (Glaw & Franzen 2006; M. Franzen 18.01. 2010 in litt.). Our examination of the two available syntypes revealed no characters distinguishing them from other juvenile A. s. superciliaris. The available specimens also correspond in the shape of the eyelid (Figs. 7 b, c) and we regard these specimens as conspecific with A. s. superciliaris. Amietophrynus s. superciliaris is morphologically most similar to A. channingi sp. nov., the second taxon from Central Africa. Amietophrynus s. superciliaris differs in colouration and morphology from A. s. chevalieri and A. channingi sp. nov. (see respective paragraphs and the key). Ethnozoology. Perret & Mertens (1957) report that indigenous people fear the toad because of its venom. According to Herrmann et al. (2005) A. s. superciliaris plays an important role in traditional medicine for people in the Mt. Nlonako area, but they also report on the people���s fear that the toad transmits leprosy. This is similar to the belief in villages close to the Ebo forest, Cameroon, where people think that the saliva of the toad may damage human skin (M. Hirschfeld & M.- O. R��del, unpubl. data). Lawson (1993) describes a local belief from western Cameroon that toad bones are used to treat poisoning and mentally deranged people. Traditional healers of the Bakossi people dry toads and pound them for native medicine (A. Schmitz, pers. obs.). In southern Cameroon Bakola-pygmees use the parotid secretion to envenom their arrowheads for hunting (Medjo, WWF-guide at Campo Ma���an NP, pers. comm.). Lawson (1993) reports the local belief in south-western Cameroon where people assume that these toads indicate the change of season by changing position into the opposite direction, while they otherwise rest for long times in the same place and position (see above). Local hunters in southern Cameroon reported to the senior author on large frogs sitting for months without movement in the forest., Published as part of Barej, Michael F., Schmitz, Andreas, Menegon, Michele, Hillers, Annika, Hinkel, Harald, B��hme, Wolfgang & R��del, Mark-Oliver, 2011, Dusted off ��� the African Amietophrynus superciliaris - species complex of giant toads, pp. 1-32 in Zootaxa 2772 on pages 6-9, DOI: 10.5281/zenodo.276862, {"references":["Boulenger, G. A. (1888) A list of the reptiles and batrachians collected by Mr. H. H. Johnston on the Rio del Rey, Cameroons District, W. Africa. Proceedings of the Royal Society, London, 1887, 564 - 565.","Werner, F. (1897) Ueber einige neue oder seltene Reptilien und Frosche der zoologischen Sammlung des Staates Munchen. Sitzungsberichte der mathematisch-physikalischen Classe der koniglich bayerischen Akademie der Wissenschaften zu Munchen, 27, 203 - 220.","Frost, D., Grant, T., Faivovich, J., Bain, R. H., Haas, A., Haddad, C. F. B., de Sa, R. O., Channing, A., Wilkinson, M., Donnellan, S., Raxworthy, C. J., Campbell, J. A., Blotto, B. L., Moler, P., Drewes, R. C., Nussbaum, R. A., Lynch, J. D., Green, D. M. & Wheeler, W. C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1 - 291.","Amiet, J. - L. & Perret, J. - L. (1969) Contributions a la faune de la region de Yaounde (Cameroun) II. - Amphibiens anoures. Annales de la Faculte des Sciences du Cameroun, 3, 117 - 137.","Perret, J. - L. & Mertens, R. (1957) Etude d'une collection herpetologique faite au Cameroun de 1952 a 1955. Bulletin de l'Institut Francais d'Afrique Noire. Serie A, 19, 548 - 601.","Werner, F. (1898) Ueber Reptilien und Batrachier aus Togoland, Kamerun und Tunis aus dem kgl. Museum fur Naturkunde in Berlin (Teil II.). Verhandlungen der kaiserlich - koniglichen zoologisch - botanischen Gesellschaft in Wien, 48, 191 - 230.","Fretey, T. & Blanc, C. P. (2001) Inventaire systematique des amphibiens anoures du centre du Gabon. Bulletin de la Societe Zoologique de France, 126, 375 - 390.","Lasso, C. A., Rial, A. I., Castroviejo, J. & de la Riva, I. (2002) Herpetofauna del Parque Nacionale de Monte Alen (Rio Muni, Guinea Equatorial). Graellsia, 58, 21 - 34.","Akani, G. C., Politano, E. & Luiselli, L. (2004) Amphibians recorded in forest swamp areas of the river Niger delta (southeastern Nigeria), and the effects of habitat alteration from oil industry development on species richness and diversity. Applied Herpetology, 2, 1 - 22.","Amiet, J. - L. (1976 a) Voix d'Amphibiens camerounais V. Bufonidae; genres Bufo, Werneria et Nectophryne. 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Amietophrynus superciliaris chevalieri (Mocquard, 1908) Figs. 1, 2 a, 4 a, 6 a���d, 7 a, 8 a���d, 9 a���h. Bufo chevalieri Mocquard, 1908, Bull. Mus. Natl. Hist. Nat. Paris 14: 262 Type locality. ��� Ivory Coast ���. Material examined. MNHN 1908.0033 (holotype, juvenile) Ivory Coast, locality unknown; MNHN 1979.6148 - 50 (male + 2 females) Guinea, Nimba; MNHN 1980.1277 (female) Ivory Coast, locality unknown; MNHN 1989.4045 (female) Liberia, Mt. Tocadeh; MNHN 9026 (female) Guinea, locality unknown; ZFMK 31801 (male) Ghana, locality unknown; ZFMK 56292 (juvenile) Guinea, S��r��dou, For��t de Ziama; ZFMK 60724 (juvenile) Guinea, S��r��dou, For��t de Ziama; ZFMK 66282 (juvenile) Guinea, S��r��dou, For��t de Ziama; ZMB 74320 (juvenile) Guinea, D��r�� Classified Forest; ZMB 74321 (juvenile) Guinea, Di��ck�� Classified Forest; ZMB 74322 (juvenile) Guinea, Mont B��ro Classified Forest. Holotype. The holotype of Amietophrynus s. chevalieri (MNHN 1908.0033) is a juvenile (SUL: 35.8 mm) in poor condition. The skin is bleached and thinned, eyes and eyelids sunken, colouration not recognizable (neither on flanks nor on dorsum any visible pattern) and a long cut on the right flank opened the guts (Figs. 8 a, b). Nonetheless, the shape of the eyelid is recognizable. The eyelids are rounded in dorsal view and not thickened medially, thus not forming a triangular prolongation (Fig. 7 a). Skin on hand and feet are shrunken, hence, webbing between toes hardly visible (Figs. 8 c, d). Measures of characters which are accessible: head width 13.6 mm; parotid length 8.8 mm and 9.6 mm; length of first finger 4.0 mm; length of third finger 4.9 mm; femur length 12.8 mm; tibia length 12.9 mm; length of shortest toe 1.6 mm, length of inner metatarsal tubercle 0.9 mm. Diagnosis. Genetically the taxon belongs to the African toad genus Amietophrynus (Frost et al. 2006). As part of the Amietophrynus superciliaris -species complex, the taxon differs from all other members of the genus by a smooth dorsal skin in adult specimens, straight loreal region and large size. Large toad; dorsal skin smooth in adults, granular in juveniles; body shape very broad, ovoid; tympanum distinct, drop-shaped, smaller than eye diameter; parotids prominent, shape of parotids like broad drop, width of parotid glands slightly reduced posteriorly, posterior part of parotid glands rounded; eyelid rounded in dorsal view, slender in lateral view, triangular prolongation absent; posterior part of back with dark colouration, thus different from anterior part; pair of spots on posterior part of back absent; extremities slender; males with nuptial swellings on fingers I and II. A. s. chevalieri can be distinguished from the nominal subspecies by the following characters: shape of the eyelid (with the exception of few juveniles) being rounded and of equal width along the eye in A. s. chevalieri and of triangular shape and prolonged in the middle in A. s. superciliaris (Figs. 2 a, b); the eyelid width / eye diameter ratio is significantly higher in the nominal subspecies (Kruskal-Wallis-test; p A. s. chevalieri than in A. s. superciliaris; parotid glands being usually rounded at the posterior tip in A. s. chevalieri and pointed in A. s. superciliaris; posterior part of the back in A. s. chevalieri coloured darker (a triangular shaped figure, pointing cranially) than the anterior part of the dorsum, in A. s. superciliaris back of uniform colouration (generally pale yellowish); in addition, a pair of abdominal spots present in A. s. superciliaris (exceptionally lacking one or even both) and absent in A. s. chevalieri. For characters distinguishing A. s. chevalieri and A. channingi sp. nov. see below. Description. Large Amietophrynus with very broad body shape; females being larger than males (SUL in males: 116.4���137.6 mm, in females: 130.6 ���156.0 mm); mean head width similar in both sexes, in males 41 % of SUL, in females 40 %; snout in lateral view straight or marginally rounded; head without bony ridges; canthus rostralis distinct, angular; loreal region slightly concave; distance eye-snout similar to eye diameter; nares closer to snout tip than eye; eyelid in adult specimens rounded in dorsal view; eyelid of almost identical width along the eye in lateral view, without a process in the middle (only MNHN 9026 possesses a very slight angular shaped eyelid; see also Fig. 9 d); tympanum more or less distinct, positioned at the concave cheek region; tympanum vertically prolonged and drop shaped, its horizontal diameter smaller than eye-diameter; dorsal skin smooth in adults, warty in juveniles; parotid glands very prominent, drop or bean shaped, width slightly diminishing backwards, usually with rounded posterior tip (mean parotid length / parotid width ratio in males: 2.86, in females: 3.00); parotid pattern bicoloured, ventral part coloured like flanks, dorsal part like back, both colours separated by sharp angle; parotid gland openings on upper part, sometimes reaching parotid angle; fingers and toes simple, not enlarged at the end; subarticular tubercles simple; relative length of fingers: III> I> II ��� IV (mean length of finger I / length of finger III ratio in both sexes 96 %); manual webbing absent; extremities slender in comparison to body; mean tibia length 39 % of SUL in males, 37 % in females; inner and outer metatarsal tubercle present; relative toe length: IV> III> V> II> I; webbing rudimentary: 1 (0.5) or 1 (0), 2 (1 - 0.5) or 2 (1 -0), 3 (2 - 1), 4 (2.75 - 2.75) or 4 (3 - 3), 5 (1); tarsal fold absent; males in breeding condition with hypertrophied forearms and nuptial swellings on fingers I and II; juveniles with smaller and fewer warts on back than Central African specimens (R��del et al. 2004). Colouration. Dorsum pattern yellowish, extending from the tip of the snout, along the upper side of the parotids backwards to the vent region; posteriormost back region with darker marking, brownish or purple-blackish (Fig. 9); lateral colouration ranging from tip of snout, below the canthus rostralis and along the flanks to the groins: dark purple, purple-brownish or intense red colour; ventral colouration reddish; extremities dark purple; inguinal region with small spots; gular region coloured like venter; anterior extremities dark purple below; juveniles orange or light brown on dorsum; flanks in juveniles generally reddish or purple, sometimes greyish (Figs. 9 a���c, f); juveniles often with distinct white transversal bars on upper hind limbs and feet (Figs. 9 c, h); juvenile interorbital marking a single V-shaped spot (Fig. 4 a). Colouration in preservation. Preserved specimens may loose their colouration, turning brownish. However, a clear differentiation between the lateral and dorsal colouration is usually still recognizable. The darker posterior part of the back is recognizable in only few preserved specimens, as the whole back turns dark. Natural history. Amietophrynus s. chevalieri inhabits primary forests (Guib�� & Lamotte 1958; B��hme 1994; R��del et al. 2004). In contrast to A. s. superciliaris male A. s. chevalieri are known to expel loud calls during the breeding season (I. Herbinger in R��del et al. 2004). Unfortunately no recordings are available. R��del et al. (2004) report on juveniles and subadults along large rivers in primary forests in November / December in Guinea (dry season). B��hme (1994) finds freshly metamorphosed and still smooth skinned juveniles in October, at the end of the rainy season in Guinea and Hillers & R��del (2007) mention juveniles in December in Liberia. Distribution. Amietophrynus s. chevalieri (formerly usually referred to as A. superciliaris) is known from forests in Upper Guinea. At present the taxon is known from Liberia (Hillers & R��del 2007), Sierra Leone (this paper; Figs. 1; 9g); Guinea (Guib�� & Lamotte 1958; B��hme et al. 1994; R��del & Bangoura 2004 a, b; R��del et al. 2004), Ivory Coast (Mocquard 1908; Ernst et al. 2006) and Ghana (Schi��tz 1964; Hughes 1988; R��del & Agyei 2003). Bailo et al. (2009) list A. s. chevalieri (as A. superciliaris) for the Badiar National Park in northern Guinea. However, this area seems not suitable for the toad and the occurrence of the species is thus very unlikely. Other species names mentioned in this paper indicate further misidentifications. Hence, the locality has not been considered in the overall distribution (Fig. 1). According to Segniagbeto et al. (2006) the species should be deleted from the list of possible species in Togo. Amietophrynus s. chevalieri is hence a strict endemic of the Upper Guinean forest region. Taxonomy. Bufo chevalieri has been described by Mocquard (1908) from the Ivory Coast. According to Frost (2010) A. chevalieri has been synonymized with A. superciliaris by Tandy & Keith (1972) without comment. However, already Guib�� & Lamotte (1958) report that their examination of the juvenile type of A. chevalieri revealed that it corresponds to a juvenile specimen of A. superciliaris. The type of A. s. chevalieri differs from typical A. superciliaris only by the lack of the palpebral process, which however is often lacking in juveniles of the A. superciliaris -species complex (Guib�� & Lamotte 1958; this paper). The synonymy of A. chevalieri with A. superciliaris has been accepted in later publications (e.g. Perret 1966; Tandy & Keith 1972). However, the two taxa are distinctly different in morphology (see Diagnosis section above). Ethnozoology. According to B��hme (1994) locals believe the toad to be dangerous. This fear also exists in south-eastern Sierra Leone, where some people believe that this toad is evil and can kill people or at least cause disease to people who are close when it opens its mouth and exhales its evil breath. According to a widespread belief of non-related Upper Guinean ethnic groups the toad gives birth to the rainbow. These tales are known from Malink�� people in Guinea, from Oubi and Guer�� people in western Ivory Coast (R��del & Bangoura 2004 b) and from different ethnic groups in Liberia and Sierra Leone. In the latter two countries the toad is also called ���rainbow toad���., Published as part of Barej, Michael F., Schmitz, Andreas, Menegon, Michele, Hillers, Annika, Hinkel, Harald, B��hme, Wolfgang & R��del, Mark-Oliver, 2011, Dusted off ��� the African Amietophrynus superciliaris - species complex of giant toads, pp. 1-32 in Zootaxa 2772 on pages 9-18, DOI: 10.5281/zenodo.276862, {"references":["Mocquard, M. F. (1908) Description de quelques reptiles et d'un batracien nouveaux de la collection du museum. Bulletin du Museum national d'Histoire naturelle, 14, 259 - 262.","Frost, D., Grant, T., Faivovich, J., Bain, R. H., Haas, A., Haddad, C. F. B., de Sa, R. O., Channing, A., Wilkinson, M., Donnellan, S., Raxworthy, C. J., Campbell, J. A., Blotto, B. L., Moler, P., Drewes, R. C., Nussbaum, R. A., Lynch, J. D., Green, D. M. & Wheeler, W. C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1 - 291.","Guibe, J. & Lamotte, M. (1958) La reserve naturelle integrale du Mont Nimba. XII. Batraciens (sauf Arthroleptis, Phrynobatrachus et Hyperolius). Memoires de l'Institut Francais d'Afrique Noire, 53, 241 - 273.","Bohme, W. (1994) Frosche und Skinke aus dem Regenwaldgebiet Sudost-Guineas, Westafrika I. Einleitung; Pipidae, Arthroleptidae, Bufonidae. herpetofauna, 16 (92), 11 - 19.","Hillers, A. & Rodel, M. - O. (2007) The amphibians of three national forests in Liberia, West Africa. Salamandra, 43, 1 - 10.","Rodel, M. - O. & Bangoura, M. A. (2004 a) A conservation assessment of amphibians in the Foret Cassee du Pic de Fon, Simandou Range, southeastern Republic of Guinea, with description of a new Amnirana species (Amphibia Anura Ranidae). Tropical Zoology, 17, 201 - 232.","Rodel, M. - O. & Bangoura, M. (2004 b) Rapid survey of amphibians and reptiles in the Foret Classee du Pic de Fon, Guinea. In: McCullough, J. (Ed.): A Rapid Biological Assessment of the Foret Classee du Pic de Fon, Simandou Range, South-eastern Republic of Guinea / Une Evaluation Biologique Rapide de la Foret Classee du Pic de Fon, Chaine du Simandou, Guinee. RAP Bulletin of Biological Assessment 35, Conservation International, Washington D. C., 51 - 60 + appendices 3 - 5, 232 - 237.","Ernst, R., Linsenmair, K. E. & Rodel, M. - O. (2006) Diversity erosion beyond the species level: Dramatic loss of functional diversity after selective logging in two tropical amphibian communities. Biological Conservation, 133, 143 - 155.","Schiotz, A. (1964) A preliminary list of amphibians collected in Ghana. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening I Kobenhavn, 127, 1 - 17.","Hughes, B. (1988) Herpetology in Ghana (West Africa). British Herpetological Society Bulletin, 25, 29 - 38.","Rodel, M. - O. & Agyei, A. C. (2003) Amphibians of the Togo-Volta highlands, eastern Ghana. Salamandra, 39, 207 - 234.","Bailo, D., Alphonse, N. & Gu, Y. (2009) An inventory of biodiversity in the Badiar National Park, Guinea Conakry: Implication for conservation. Research Journal of Biological Sciences, 4, 948 - 951.","Frost, D. (2010) Amphibian Species of the World: an Online Reference. Version 5.4 (8 April, 2010). Electronic Database accessible at http: // research. amnh. org / vz / herpetology / amphibia / American Museum of Natural History, New York, USA.","Tandy, M. & Keith, R. (1972) Evidence from osteology. In: Blair, W. F. (Ed.) Evolution in the genus Bufo. University of Texas Press, Dallas, 119 - 170.","Perret, J. - L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbucher (Abteilung fur Systematik, Okologie und Geographie der Tiere), 8, 289 - 464."]}

Barej, Michael F., Schmitz, Andreas, Menegon, Michele, Hillers, Annika, Hinkel, Harald, Böhme, Wolfgang, Rödel, Mark-Oliver (2011): Dusted off—the African Amietophrynus superciliaris - species complex of giant toads. Zootaxa 2772: 1-32, DOI: 10.5281/zenodo.276862

Amietophrynus channingi sp. nov. Figs. 1, 2 c, 4 c, 10 a–d, 11 a–d, 12 a–c. Holotype. MTSN 9674 (male) Democratic Republic of Congo, West Mwana, Itombwe Massif, 0 28.13875 E, 0 3.83781 S, 1302 m a.s.l., coll. Joseph Matunguru. Paratypes. IRSNB-KBIN 1982 (female) Democratic Republic of Congo, Beni; IRSNB-KBIN 1983 (subadult) Democratic Republic of Congo, Mutsora; IRSNB-KBIN 1984 (female) Democratic Republic of Congo, Bukavu; IRSNB-KBIN 1985 (juvenile) Democratic Republic of Congo, Irangira; MHNG 2520.25 (juvenile) Democratic Republic of Congo, Nduye, Alamby; ZFMK 62573 (male) Democratic Republic of Congo, Irangi; ZFMK 63895 - 6 (2 juveniles) Democratic Republic of Congo, Irangi. Additional material. IRSNB-KBIN 634 (male) Democratic Republic of Congo, Lulenga; IRSNB-KBIN 2216 (female) Democratic Republic of Congo, Zobia; IRSNB-KBIN 2217 (female) Democratic Republic of Congo, Zobia; IRSNB-KBIN 11363 (female) Democratic Republic of Congo, Walikale; IRSNB-KBIN 11364 (female + 2 juveniles) same data as IRSNB-KBIN 1985; MNHN 1933.027 (female) Congo Belge (= Dem. Rep. Congo), locality unknown; ZFMK 34376 (juvenile) Democratic Republic of Congo, Poko; ZFMK 63894 (male) same data as ZFMK 63895. Remark: The MNHN received a collection from “ Congo Belge ” in 1933 from Guy Babault (Bull. Mus. Natl. Hist. Nat. p. 18), today Democratic Republic of Congo, without further details on the collection localities. According to publications dealing with various taxa donated to the MNHN by G. Babault (Pellegrin 1933, 1935; Babault 1935; Berlioz 1935; Germain 1936) the locality in “ Congo Belge ” can be assigned to the Kivu area in eastern Democratic Republic of the Congo. Consequently, we assign MNHN 1933.027 to this new taxon. Diagnosis. Genetically the taxon belongs to the African toad genus Amietophrynus (Frost et al. 2006). As part of the Amietophrynus superciliaris -species complex, the taxon differs from all other members of the genus by a smooth dorsal skin in adult specimens, straight loreal region and large size. Medium to large sized toad with smooth dorsal skin in adults, granular in juveniles; body shape compact; tympanum distinct, drop-shaped, smaller than eye-diameter; parotids prominent, broad, width of parotid glands hardly reducing backwards, posterior tips of parotid glands rounded; eyelid slightly triangular in dorsal view, eyelid in lateral view forming a intimated triangular process; anterior and posterior part of back with different pattern; pair of dark spots present on posterior back region; extremities slender. Amietophrynus channingi sp. nov. is most similar to A. s. superciliaris as both species exhibit a triangular eyelid process in adults, a structure lacking in A. s. chevalieri (Figs. 2 a–c) and share the symmetric pair of abdominal spots (Figs. 3 c, e–g; 11 b, c). The new species differs from A. s. superciliaris by the following characters: shape of eyelid being more pointed and elongated in A. s. superciliaris; the eyelid width / eye diameter ratio shows no significant difference (Kruskal-Wallis-test; p> 0.05, N A.s.s.= 28, 0.24–0.39, x= 0.32 ± 0.04; N A.c.= 10, 0.23–0.38, x= 0.34 ± 0.04; H= 12.28); shape of parotid glands, much diminishing in width posteriorly and possessing a pointed tip in A. s. superciliaris and being more bulged and rounded at the posterior tip in A. channingi sp. nov.; colouration of the back, which is darker in the posterior region of the back and possesses a darker mid-dorsal line in A. channingi sp. nov., whereas the dorsum is more or less uniformly coloured in A. s. superciliaris (Figs. 3 c, e–g; 11 a–c); colouration of flanks, being reddish in A. s. superciliaris and brownish in the new species (Figs. 2 b–c). Moreover, juveniles without a fully developed eyelid process can be differentiated by the following colour patterns: interorbital pattern in juvenile A. channingi sp. nov. a depressed dark, V-like figure, covering the eyelids, while juvenile A. s. superciliaris usually possess two clearly separated interorbital spots, which may even be broken into numerous smaller spots (Figs. 4 b-c). The new species can be distinguished from A. s. chevalieri by the following characters: eyelid process being absent in A. s. chevalieri (present in A. channingi sp. nov.; Figs. 2 a, c); the eyelid width / eye diameter ratio is significantly higher in adults of the new species (Kruskal-Wallis-test; p A. s. chevalieri; A. channingi sp. nov. possess a symmetric pair of abdominal spots (lacking in A. s. chevalieri; Figs. 9 a–f, h; 11 a–c); lateral colouration being reddish-purple in A. s. chevalieri and brownish in A. channingi sp. nov. (Figs. 2 a, c). Description of the holotype. The holotype is an adult male in breeding condition; dorsal skin smooth, lateral and ventral skin rough (Figs. 10 b, c); compact and sturdy body shape; SUL 111.8 mm; head without bony ridges; canthus rostralis distinct, angular; loreal region slightly concave; snout very short, almost straight in lateral view; head width 44.1 mm; eye-diameter 12.2 mm; interorbital-distance 9.5 mm; nares closer to snout than to eye; eyelid with weak triangular process (Fig. 11 a), measuring 4.9 mm from lower eyelid border to tip of eyelid process; parotid length 33.5 mm; parotid width 11.8 mm; parotids bi-coloured, upper part pale brown like dorsum, lower part dark grey; gland openings present only on upper part, reaching colouration border; posterior to parotids a glandular dorsolateral fold, extending to groin area; fingers and toes simple; subarticular tubercles simple; relative length of fingers: III> I> II ≥ IV; femur length 42.6 mm; tibia length 42.0 mm; foot length incl. longest toe 43.6 mm; inner metatarsal tubercle oval and prominent; outer metatarsal tubercle rounded, indistinct; manual webbing absent (Fig. 10 c), relative toe length: IV> III> V> II> I; pedal webbing rudimentary (Fig. 10 d): 1 (0.5), 2 (1 - 0.5), 3 (2 - 1.25), 4 (2.75-3), 5 (1.25); tarsal fold absent; secondary sexual characters: large dark nuptial pads on fingers I and II; miniscule spines scattered on lateral side of head and flanks below parotid glands, minuscule spines denser behind mouth gap; further minuscule spines cover anterior and dorsal side of upper arms; colouration in preservation: back pale brown; interorbital space with darker V-shaped pattern, the tips of the V pointing on the eyelid processes; dark vertebral line running backwards (Figs. 10 b; 11 b); posterior part of back with dark triangular figure and a symmetric pair of dark spots; skin of dark spots different in texture to remaining of back skin, blunt and “fleecy”; dorsal and lateral colouration distinctly different, separated by thin white line; flanks coloured dark brown, this pattern ranging from snout-tip backwards and dissolving in the inguinal area, here replaced by dark brown spots (three on left, two on the right side); dark spots on anterior part of upper hind limb and on posterior side of lower hind limb; venter uniform pale. Variation. Medium to large sized toad with compact body shape; females being larger than males (SUL in males: 106.7–111.8 mm, in females: 100.3 –143.0 mm; SUL of specimens from Congo according to Noble (1924: males: 98–112 mm, females: 133–149 mm); mean head width similar in both sexes, in males 38 % of SUL, in females 39 %; snout in lateral view straight almost truncate; distance eye-snout larger than eye diameter; eyelid in adult specimens more or less triangular shaped; tympanum more or less distinct, positioned in a concave area of the cheeks; tympanum vertically prolonged and drop shaped, smaller in horizontal diameter than eye-diameter; dorsal skin smooth in adults, granular in juveniles; parotid glands prominent, bulged, width posteriorly slightly reducing, usually rounded at posterior tip (parotid length / parotid width ratio in males: 2.82, in females: 2.73); glandular dorsolateral fold behind the parotids may be present; parotids bicoloured, lower parts like flanks, upper parts like back; border of the two parts forms a sharp ridge; parotid gland openings on upper part only; fingers and toes simple; subarticular tubercles simple; relative length of fingers: III> I> II ≥ IV (mean length of finger I / length of finger III ratio in males: 89 %, in females: 94 %); manual webbing absent; extremities slender in comparison to body shape; mean tibia length 40 % of SUL in males, in females 41 %; inner metatarsal tubercle present, oval and prominent; outer metatarsal tubercle rounded, sometimes indistinct; relative toe length: IV> III> V> II> I; webbing rudimentary: 1 (0.5), 2 (1 - 0.5), 3 (2 - 1) or 3 (2 - 1.5), 4 (2.75 - 2.75) or 4 (2.75-3), 5 (1.25); males in breeding condition with hypertrophied forearms and swellings on fingers I and II. Colouration. Dorsum dirty yellowish-browinish, extending from the tip of the snout, along the upper side of head and parotids backwards to the posterior part of the back (Fig. 11); dark V-shaped pattern on head covering the eyelids, usually a vertebral line running from this pattern backwards, forming a dark triangular pattern on the posterior part of the back (Figs. 11 b, c); back with symmetric pair of dark spots; loreal area, lower parts of parotids and flanks brownish; upper parts of lateral colouration darker than ventral part, partly almost black, posterior most part of flanks orange-brown; forearms light purple-greyish above, usually bordered by white line, and dark below; inguinal region with small black spots; upper hind limbs and feet with black transversal bars (Figs. 11 c, d); forearms with distinct white line between pale dorsal and dark ventral colouration; throat and venter pale yellow; juveniles coloured like adults but may possess additional pairs of spots on the back and the interorbital V-pattern, covering large parts of the eyelids; this pattern is more distinct in juveniles than in adults; juveniles with additional transversal bars on tibiofibula, this pattern disappearing in larger specimens. Orts (1970) figured two specimens after completed metamorphosis: the first uniform brown-blackish coloured and the second with a “typical toad pattern” consisting of symmetric pairs of spots (as in A. regularis, A. camerunensis, etc.). According to Orts (1970) the typical interorbital V-pattern develops within the first two weeks. Colouration in preservation. Preserved specimens still exhibit the separation of dark and light colour as in life; dark V-pattern between eyes, vertebral line and darker triangular posterior region of back may be hardly or not visible; flanks turn greyish or yellowish, and the brownish colour may turn almost black; brownish flank colouration sometimes recognizable when the lateral skin got folded and remains of former colouration has been covered between folds; pair of dark spots on the posterior part of the back sometimes bleached out and hardly recognizable. Tadpole description. The tadpole of this new taxon has been briefly described by Orts (1970). According to Orts’ (1970) description and figures: body oval in dorsal view in young stages, becoming more elongated in later stages (Figs. 12 a–c); body slightly flattened; mascular tail slender, converging caudally; dorsal and ventral fin present, tail (fins included) of similar height as body; tail fin rounded at tip of tail; tail 1.33 to 1.75 as long as body; eyes dorsally positioned, moving into more lateral position during growth, distance snout-tip to eye approx. 20–25 % of body length; spiraculum positioned at midbody; mouth ventrally positioned, surrounded by robust lips; colouration at 10 mm total length: grey above and crème-white, translucent below; tadpoles of 10.5 mm length (4 mm body length, Gosner stage 25; Fig. 12 a) with keratodont formula: 1: 1 + 1 // 1 + 1: 2; development of posterior pats at length of 13 mm (body length 5 mm); keratodont formula: 1: 1 + 1 // 3 (tadpole length> 13 mm); same keratodont formula in later stages; metamorphosis finished at body length (measured from tip of snout to anus) of 9 to 10 mm; for more details see Orts (1970). Natural history. Although the taxonomic status of this species has so far been overlooked, much more about the natural history of this taxon has been published compared to the other taxa of the Amietophrynus superciliaris - species complex. A. channingi sp. nov. is active in the rainy season, and can then be observed almost everywhere in the forest (Noble 1924). The species is nocturnal at least in captivity, where also cannibalistic behaviour was recorded (Orts 1970). When disturbed, the toad may be feigning death by “lowering the head to the ground, and dropping the prominent upper eyelid” (Noble 1924). The diet comprises various invertebrate taxa (Noble 1924). The species seems to feed preferably on snails around the Irangi research station (DRC) as local research assistants regularly found the species in great numbers in pits dug by road workers to raise snails for food consumption. According to Noble (1924) oviposition may occur during several months, breeding thus potentially may take place throughout most of the year. Eggs are very numerous in comparison to other forest toads, 1200–1500 in Amietophrynus tuberosus (referred to as A. polycercus in Noble 1924) and up to three times that number in A. channingi sp. nov., and egg size ranges from 1.6 –2.0 mm (envelopes removed). According to own measures, egg size ranges from 1.40–1.95 mm (N= 114, mean= 1.69, SD= 0.11). Orts (1970) recorded tadpoles of 10 mm in Irangi in Democratic Republic of Congo in mid-June, at the end of the rainy season. He observed more than 3000 tadpoles in a puddle of only of 40 x 25 cm and 7 cm depth, covered with vegetation, where tadpoles moved on the ground and among leaf litter. Other tadpoles inhabited a series of puddles of a diameter of approximately one meter in mid- April. These breeding sites were connected to small rivers, which may transform into large currents after rain and then wash the tadpoles away (Orts 1970). 1 2 3 4 5 6 7 8 9 A. brauni (N= 3) 0 ± 0 GenBank A. poweri (N= 2) 0.0428 - 0.0446 0.0018 GenBank 0.0439 ± 0.0007 A. rangeri (N= 2) 0.0611 - 0.0720 0.0648 - 0.0700 0.0018 GenBank 0.0668 ± 0.0050 0.0674 ± 0.0022 A. channingi (N= 2) 0.0805 - 0.0849 0.0648 - 0.0673 0.0648 - 0.0685 0.0000 D.R. Congo 0.0831 ± 0.0020 0.0687 ± 0.0017 0.0666 ± 0.0021 A. s. chevalieri (N= 4) 0.0772 - 0.0814 0.0669 - 0.0682 0.0557 - 0.0612 0.0218 - 0.0219 0 ± 0 Guinea, Liberia 0.0793 ± 0.0022 0.0676 ± 0.0007 0.0584 ± 0.0029 0.0218 ± 0.0000 A. s. superciliaris (N= 2) 0.0732 - 0.0779 0.0673 - 0.0719 0.0574 - 0.0650 0.0274 - 0.0286 0.0085 - 0.0101 0.0017 Cameroon, Nigeria 0.0754 ± 0.0018 0.0696 ± 0.0020 0.0612 ± 0.0041 0.0280 ± 0.0007 0.0093 ± 0.0008 A. camerunensis (N= 2) 0.0734 - 0.0780 0.0687 - 0.0730 0.0634 - 0.0725 0.0776 - 0.0809 0.0759 - 0.0789 0.0787 - 0.0823 0.0035 GenBank 0.0751 ± 0.0019 0.0709 ± 0.0023 0.0679 ± 0.0043 0.0792 ± 0.0018 0.0774 ± 0.0016 0.0802 ± 0.0016 A. gutturalis (N= 2) 0.0807 - 0.0864 0.0828 - 0.0861 0.0756 - 0.0840 0.0757 - 0.0774 0.0825 - 0.0840 0.0797 - 0.0839 0.0502 - 0.0540 0.0018 GenBank 0.0839 ± 0.0021 0.0843 ± 0.0015 0.0791 ± 0.0036 0.0765 ± 0.0009 0.0833 ± 0.0008 0.0818 ± 0.0019 0.0521 ± 0.0016 A. tuberosus (N= 1) 0.1049 - 0.1067 0.1138 - 0.1153 0.0873 - 0.0939 0.0927 - 0.0930 0.0946 - 0.0962 0.0894 - 0.0932 0.0902 - 0.0926 0.0928 ± 0.0000 --- GenBank 0.1055 ± 0.0010 0.1146 ± 0.0010 0.0906 ± 0.0047 0.0928 ± 0.0002 0.0954 ± 0.0012 0.0913 ± 0.0027 0.0914 ± 0.0017 Distribution. Amietophrynus channingi sp. nov. is known from various localities in the eastern Democratic Republic of Congo, East Lower Guinea (Boulenger 1919; Wheeler 1922; Noble 1924; Loveridge 1936; Orts 1970). One specimen (IRSNB-KBIN 10385) is labelled as originating from “Mutsora, Rwanda ”, but, given coordinates reveal that the locality is situated in the Democratic Republic of Congo. Due to the vicinity of localities of A. channingi sp. nov. close to the eastern border of the Democratic Republic of Congo (Fig. 1), the distribution of the toad may cover westernmost parts of Uganda, Rwanda and Burundi in suitable habitats. However, recent herpetological works failed to detect the species in the Albertine Rift (Drewes & Vindum 1994; Vonesh 2001). Ethnozoology. According to Noble (1924) natives fear the whitish secretion, as they might go blind. Etymology. The specific epithet channingi is a patronym. We name the species in honour of Prof. Dr. Alan Channing from the University of the Western Cape (South Africa) in recognition of his invaluable contributions to our knowledge of African amphibians.

Key to the taxa of the Amietophrynus superciliaris- species complex (see also Fig. 13) 1 eyelid process present, eyelid triangular in dorsal and lateral view (Figs. 2 b, c); pair of symmetric spots present on posterior part of back (Figs. 3; 11); posterior part of back coloured like anterior one or darker; palmar tubercles prominent; occurring in Lower Guinea (= Central Africa; Fig. 1)................................................................... 2 1 ’ eyelid process absent, eyelid rounded in dorsal and lateral view (Fig. 2 a); pair of symmetric spots absent on posterior part of back (Fig. 9); posterior part of back darker coloured like anterior one (Fig. 9); palmar tubercles extremely prominent; occurring in Upper Guinea (= West Africa; Fig. 1)........................................... A. superciliaris chevalieri 2 body shape very broad in dorsal view; eyelid process distinctly pointed (Fig. 2 b); parotid glands shaped like slender drop, posterior tip pointed; back colour more or less uniform; flanks reddish-purple (Fig. 2 b); juveniles usually with two distinct interorbital spots, rarely these spots may be fused (Fig. 4 b); occurring in West Lower Guinea (= western Central Africa; Fig. 1).................................................................................. A. superciliaris superciliaris 2 ’ body shape less broad in dorsal view; eyelid process slightly pointed (Fig. 2 c); parotid glands bulged, posterior end rounded; back with lighter anterior and darker posterior pattern; flanks brownish (Fig. 11); juveniles with a distinct single depressed Vlike, dark, interorbital figure covering large parts of the eyelids (Fig. 4 c); occurring in East Lower Guinea (= eastern Central Africa; Fig. 1)....................................................................... A. channingi sp. nov.

Key to the taxa of the Amietophrynus superciliaris- species complex (see also Fig. 13) 1 eyelid process present, eyelid triangular in dorsal and lateral view (Figs. 2 b, c); pair of symmetric spots present on posterior part of back (Figs. 3; 11); posterior part of back coloured like anterior one or darker; palmar tubercles prominent; occurring in Lower Guinea (= Central Africa; Fig. 1)................................................................... 2 1 ��� eyelid process absent, eyelid rounded in dorsal and lateral view (Fig. 2 a); pair of symmetric spots absent on posterior part of back (Fig. 9); posterior part of back darker coloured like anterior one (Fig. 9); palmar tubercles extremely prominent; occurring in Upper Guinea (= West Africa; Fig. 1)........................................... A. superciliaris chevalieri 2 body shape very broad in dorsal view; eyelid process distinctly pointed (Fig. 2 b); parotid glands shaped like slender drop, posterior tip pointed; back colour more or less uniform; flanks reddish-purple (Fig. 2 b); juveniles usually with two distinct interorbital spots, rarely these spots may be fused (Fig. 4 b); occurring in West Lower Guinea (= western Central Africa; Fig. 1).................................................................................. A. superciliaris superciliaris 2 ��� body shape less broad in dorsal view; eyelid process slightly pointed (Fig. 2 c); parotid glands bulged, posterior end rounded; back with lighter anterior and darker posterior pattern; flanks brownish (Fig. 11); juveniles with a distinct single depressed Vlike, dark, interorbital figure covering large parts of the eyelids (Fig. 4 c); occurring in East Lower Guinea (= eastern Central Africa; Fig. 1)....................................................................... A. channingi sp. nov., Published as part of Barej, Michael F., Schmitz, Andreas, Menegon, Michele, Hillers, Annika, Hinkel, Harald, B��hme, Wolfgang & R��del, Mark-Oliver, 2011, Dusted off ��� the African Amietophrynus superciliaris - species complex of giant toads, pp. 1-32 in Zootaxa 2772 on page 24, DOI: 10.5281/zenodo.276862

Petropedetes 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 juliawurstnerae = 10, N perreti = 15, p P. juliawurstnerae sp. nov. (tympanum diameter / eye diameter in males: N juliawurstnerae = 12, N perreti = 13, p 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 juliawurstnerae = 10, N perreti = 15, p 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 juliawurstnerae = 10, N cameronensis = 17, p 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 juliawurstnerae = 10, N cameronensis = 17, p 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., R��del, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., B��hme, 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.193288

Key to adult Central African Petropedetes species 1 toes fully webbed.......................................................................................................................................................... 2 1 ’ toes half-webbed or rudimentary-webbed.................................................................................................................... 3 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) ...... P. perreti 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)................................................................ P. palmipes 3 toes half-webbed........................................................................................................................................................... 4 3 ’ toes rudimentary-webbed.............................................................................................................................................. 5 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) .............................................................................................. P. juliawurstnerae sp. nov. 4 ’ tympanum indistinct; males without tympanic papillae or carpal spike (Fig. 8 b); femoral gland ovoid (Fig. 9 b), larger in males ....................................................................................................................................... P. cameronensis 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................ ..................................................................................................................................................................... P. johnstoni 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 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: P. vulpiae sp. nov. 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 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).... P. parkeri 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) ....... P. euskircheni sp. nov.

Key to adult Central African Petropedetes species 1 toes fully webbed.......................................................................................................................................................... 2 1 ��� toes half-webbed or rudimentary-webbed.................................................................................................................... 3 2 tympanum distinct; in males �� of eye diameter or larger, males with tympanic papillae (Fig. 8 a) and carpal spike in breeding season, tympanum in females up to �� of eye diameter; femoral glands large (larger in males) ...... P. perreti 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)................................................................ P. palmipes 3 toes half-webbed........................................................................................................................................................... 4 3 ��� toes rudimentary-webbed.............................................................................................................................................. 5 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) .............................................................................................. P. juliawurstnerae sp. nov. 4 ��� tympanum indistinct; males without tympanic papillae or carpal spike (Fig. 8 b); femoral gland ovoid (Fig. 9 b), larger in males ....................................................................................................................................... P. cameronensis 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................ ..................................................................................................................................................................... P. johnstoni 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 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: P. vulpiae sp. nov. 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 7 femoral gland small but distinct (Fig. 2 c), in males about 22 % of femur length; tympanum in males �� of eye diameter or larger than eye, occurs in lower altitudes of south-western Cameroon and eastern Nigeria (Fig. 4).... P. parkeri 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) ....... P. euskircheni sp. nov., Published as part of Barej, Michael F., R��del, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., B��hme, 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 41, DOI: 10.5281/zenodo.193288

Petropedetes cameronensis Reichenow, 1874 Figs. 3 b, 4, 8b, 9 b, 10 c, 12, 13 Platymantis cameronensis ��� Peters, 1875, Montasber. Preuss. Akad. Wiss. Berlin, 1875: 211 Hylambates cameronensis ��� Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 137 Hylambates (Platymantis) cameronensis ��� Knauer, 1883, Naturgesch. Lurche: 111 Petropedetes cameronensis ��� Boulenger, 1900, Proc. Zool. Soc. London, 1900: 439 Petropedetes obscurus Ahl, 1924, Zool. Anz., 58: 81-82 Material examined. 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, Mahol��, 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. B��hme; ZFMK 14980 (female), Cameroon, Fako, Buea, 20.- 22.III. 1974, coll. W. B��hme & W. Hartung; ZFMK 15427 (male), Cameroon, Batoki, 28.II. 1954, coll. M. Eisentraut; ZFMK 61343 (female), Cameroon, Ekundu-Kundu, 21.- 27.II. l 989, 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 et al.; 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 Petropedetes obscurus 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. R��del; ZMB 73727 - 72728 (2 females), Nigeria, Mbe Mountains, VI/ VII. 2005, coll. A. Onadeko. Problematic material: ZSM 159 / 1988 / 1-2 (2 specimens, sex unknown), Cameroon, Victoria, II. 1978, coll. F. Sauer. Diagnosis. Small sized Petropedetes 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 Petropedetes 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). Description. Small sized Petropedetes 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; canthus rostralis 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). Coloration. 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). Tadpole. According to Boulenger���s (1906) figure of the mouth of an advanced P. c a m e ro n e n s i s 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. Schi��tz (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 (Schi��tz 1966). 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 Schi��tz (1966) additionally describes the call of adult specimens of P. cameronensis. Natural history. Petropedetes cameronensis occurs mainly in the lowland, but is also known from localities up to 1400 m, where it was found together with P. p e r re t i (Amiet 1973). On Bioko the species is known from mid-altitude, but may also occur in the coastal zone (Boulenger 1906; Mertens 1965). Schi��tz (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 P. cameronensis has been described as a whistle (Schi��tz 1966) or a trill that drowns the noise of torrents (Amiet 1989). Eggs of P. cameronensis are deposited on tree trunks or leaves close to humid rocks, where the saxicolous (living on rocks) tadpoles develop (Amiet 1989). Herrmann et al. (2005) observe males guarding clutches from February to May on Mt. Nlonako. Schi��tz (1966) assumes that the muscular tail without or only narrow fin allows the tadpole to jump on stones. Distribution. 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. Schi��tz (1963) calls the type locality ���Bimba, Cameroons ���, which is probably a misspelling. This mistake has been further developed as ���Bima, Cameroons ��� by Frost (2009). P. c a m e ro n e n s i s is also known from moderate altitudes (Lawson 1993), even reaching heights above 1100 m on Mt. Nlonako (Herrmann et al. 2005). The species is known from areas in eastern Nigeria (Schi��tz 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. P. cameronensis has also been recorded from Bioko (Boulenger 1906; Mertens 1965). Genetics. The genetic comparison for the uncorrected p-distances of the mitochondrial 16 S rRNAfragment (Table 4) gave the following results for Petropedetes cameronensis: interspecific comparison between P. cameronensis and all other Central African Petropedetes taxa ranged between 5.96 %- 9.25 %, while the intraspecific variation within P. cameronensis was much lower at 0.00%- 0.50 % (N= 4). Systematic remarks. Ahl (1924) describes Petropedetes obscurus from Tscharra, Dana River and distinguishes this half-webbed species from P. cameronensis by the size of its tympanum, which is only one third of the eye diameter in P. obscurus and half the diameter in P. cameronensis. Perret (1984) who studies the validity of P. obscurus, 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 P. obscurus in the synonymy of P. cameronensis. Our examination of the type specimen of P. obscurus confirmed Perret���s (1984) decision. Hence, an affiliation of the two specimens to the East African Arthroleptides, 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. Herrmann (1989) publishes a list of amphibian types in the Zoological Museum, Greifswald (ZIMG) and lists among others, Petropedetes cameronensis. 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). 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 P. cameronensis 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 P. johnstoni, 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 P. cameronensis, collected by Reichenow (1874) himself is stored in the ZMB. Petropedetes cameronensis is half-webbed and possesses an indistinct tympanum lacking a tympanic papilla in males. Hence, P. cameronensis is easy to distinguish from the rudimentary-webbed species with a tympanic papilla P. euskircheni sp. nov., P. johnstoni, P. p a r k e r i and P. vulpiae sp. nov. The species could potentially be mistaken with P. palmipes due to its indistinct tympanum, but differs from P. palmipes by its smaller size, less developed webbing (toes fully webbed in P. p a l m i p e s) and lacking a carpal spike (Amiet 1973). Although having a similar webbing, P. c a m e ro n e n s i s can be unambiguously distinguished from P. juliawurstnerae sp. nov. alike (see the following species description)., Published as part of Barej, Michael F., R��del, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., B��hme, 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 30-35, DOI: 10.5281/zenodo.193288, {"references":["Reichenow, A. (1874) Eine Sammlung Lurche und Kriechthiere von Westafrika. Archiv fur Naturgeschichte, 1874, 287 - 298.","Boulenger, G. A. (1900) A list of the batrachians and reptiles of the Gaboon (French Congo), with descriptions of new genera and species. Proceedings of the Zoological Society of London, 1900, 433 - 456.","Ahl, E. (1924) Zur Kenntnis der Froschfauna Afrikas. Zoologischer Anzeiger, 58, 81 - 82.","Schiotz, A. (1966) On a collection of Amphibia from Nigeria. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening I Kobenhavn, 129, 43 - 48.","Boulenger, G. A. (1906) Report on the batrachians collected by the late L. Fea in West Africa. Annali del Museo Civico di Storia Naturale di Genova, 42, 157 - 172 + 2 plates.","Amiet, J. - L. (1973) Caracteres diagnostiques de Petropedetes perreti, nov. sp. et notes sur les autres especes camerounaises du genre (Amphibiens Anoures). Bulletin de l'Institut Francais d'Afrique Noire, Serie A, 35, 462 - 474.","Mertens, R. (1965) Die Amphibien von Fernando Poo. Bonner Zoologische Beitrage, 16, 14 - 29.","Schiotz, A. (1963) The amphibians of Nigeria. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening I Kobenhavn, 125, 1 - 92.","Amiet, J. - L. (1975) Ecologie et distribution des Amphibiens Anoures de la region de Nkongsamba. Annales de la Faculte des Sciences de Yaounde, 20, 33 - 107.","Amiet, J. - L. (1989) Quelques aspects de la biologie des Amphibiens Anoures du Cameroun. L'annee biologique, 28, 73 - 136.","Herrmann, H. - W., Bohme, W., Herrmann, P. A., Plath, M., Schmitz, A. & Solbach, M. (2005) African biodiversity hotspots: The amphibians of Mt. Nlonako, Cameroon. Salamandra, 41, 61 - 81.","Perret, J. - L. (1984) Identification de Petropedetes obscurus Ahl, 1924 (Amphibia, Phrynobatrachinae), conserves au Museum de Berlin. Bulletin de la Societe Neuchateloise des Sciences Naturelles, 107, 165 - 170.","Frost, D. (2009) Amphibian Species of the World: an Online Reference. Version 5.2 (12 February, 2009). The American Museum of Natural History. Available from http: // research. amnh. org / herpetology / amphibia / index. php (assessed 18 February 2009).","Lawson, D. P. (1993) The reptiles and amphibians of the Korup National Park Project, Cameroon. Herpetological Natural History, 1, 27 - 90.","Perret, J. - L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbucher (Abteilung fur Systematik, Okologie und Geographie der Tiere), 8, 289 - 464.","Klemens, M. W. (1998) The male nuptial characteristics of Arthroleptides martiensseni Nieden, an endemic torrent frog from Tanzania's Eastern Arc Mountains. Herpetological Journal, 8, 35 - 40.","Herrmann, H. - J. (1989) Die Amphibientypen des Zoologischen Museums Greifswald. Veroffentlichungen des Naturhistorischen Museums Schloss Bertholdsburg, Schleusingen, 4, 13 - 14."]}

Petropedetes vulpiae Barej, R��del, Gonwouo, Pauwels, B��hme 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 Yaound��) (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. johnstoni, Published as part of Barej, Michael F., R��del, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., B��hme, 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.193288

Petropedetes euskircheni Barej, R��del, Gonwouo, Pauwels, B��hme and Schmitz, 2010 Assignments before official naming of species: Euskirchen et al. (1999): referred to as P. parkeri Herrmann et al. (2000): referred to as P. parkeri Plath et al. (2004): referred to as P. parkeri Herrmann et al. (2005): referred to as P. parkeri Petropedetes perreti Amiet, 1973 Assignments before official naming of species, problematic assignment: Perret & Mertens (1957): as P. palmipes Lawson (1993): P. p e r re t i from Rumpi Hills, locality unsure, Published as part of Barej, Michael F., R��del, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., B��hme, 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.193288

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, Kingu��l��, 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 Kingu��l��, app. 165 m a.s.l., 25.- 27.I. 2000, coll. S. L��tters & V. Gossmann; ZMB 73891 Gabon (male), Province du Woleu-Ntem, Tchimb��l��, 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. Schi��tz (1966) describes similarities in the morphology of advanced larval stages of P. palmipes and P. cameronensis, possessing identical teeth formulae. Schi��tz (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 (L��tters et al. 2001; Pauwels & R��del 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., R��del, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., B��hme, 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.193288, {"references":["Boulenger, G. A. (1905) Descriptions of new West-African frogs of the genera Petropedetes and Bulua. The Annals and Magazine of Natural History, 15, 281 - 283.","Lamotte, M., Perret, J. - L. & Dzieduszycka, S. (1959) Contribution a l'etude des Batraciens de l'Ouest africain. - IX. Les formes larvaires de Petropedetes palmipes, Conraua goliath et Acanthixalus spinosus. Bulletin de l'Institut Francais d'Afrique Noire, Serie A, 21, 762 - 776.","Gossmann, V., Lotters, S., Obame, F. & Bohme, W. (2002) Zur Herpetofauna Gabuns - Teil II: Kommentierte Artenliste der gefundenen Reptilien, Bemerkungen zur Artenvielfalt. Herpetofauna, 24, 19 - 33.","Schiotz, A. (1966) On a collection of Amphibia from Nigeria. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening I Kobenhavn, 129, 43 - 48.","Amiet, J. - L. (1989) Quelques aspects de la biologie des Amphibiens Anoures du Cameroun. L'annee biologique, 28, 73 - 136.","Perret, J. - L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbucher (Abteilung fur Systematik, Okologie und Geographie der Tiere), 8, 289 - 464.","de la Riva, I. (1994) Anfibios anuros del Parque Nacional de Monte Alen, Rio Muni, Guinea Equatorial. Revista Espanola de Herpetologia, 8, 123 - 139.","Lasso, C. A., Rial, A. I., Castroviejo, J. & de la Riva, I. (2002) Herpetofauna del Parque Nacionale de Monte Alen (Rio Muni, Guinea Equatorial). Graellsia, 58, 21 - 34.","Lotters, S., Gossmann, V., Obame, F. & Bohme, W. (2001) Zur Herpetofauna Gabuns Teil I: Einleitung, Untersuchungsgebiet und Methodik, kommentierte Artenliste der gefundenen Froschlurche. Herpetofauna, 23, 19 - 34.","Pauwels, O. S. G. & Rodel, M. - O. (2007) Amphibians and National Parks in Gabon, western Central Africa. Herpetozoa, 19, 135 - 148.","Perret, J. - L. & Mertens, R. (1957) Etude d'une collection herpetologique faite au Cameroun de 1952 a 1955. Bulletin de l'Institut Francais d'Afrique Noire, Serie A, 19, 548 - 601."]}

Petropedetes perreti Amiet, 1973 Figs. 3 c, 4, 8a, 9 a, 10 a, 11 a, b Petropedetes perreti Amiet, 1973, Bull. I.F.A.N. s��r. A., 2: 463 Material examined. 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 Medji��, 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. R��del & 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. R��del & 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. R��del & 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. R��del. Diagnosis. Medium sized Petropedetes 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). Description. Medium sized Petropedetes 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; canthus rostralis 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). Coloration. 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. Tadpole. 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). Natural history. Petropedetes perreti is a forest species (Herrmann et al. 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 et al. (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). We observed P. p e r re t 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. Distribution. Petropedetes perreti is endemic to Cameroon. The species is found on the southern slopes of the Bamil��k�� Plateau, Mt. Manengouba and Mt. Nlonako (Fig. 4), areas which are characterized by a very high precipitation through the rainy season (Amiet 1973; Herrmann et al. 2005). Genetics. The genetic comparison for the uncorrected p-distances of the mitochondrial 16 S rRNAfragment (Table 4) gave the following results for Petropedetes perreti: interspecific comparison between P. perreti and all other Central African Petropedetes taxa ranged between 2.84 %- 10.16 %, while there was no intraspecific variation between the included sequences of the P. p e r re t i vouchers (0.00%; N= 6). The lowest interspecific difference has been detected between P. perreti and P. juliawurstnerae sp. nov. ranging between 2.84 %- 3.59 %, but species can be clearly differentiated on the morphological basis alone (see species description of P. juliawurstnerae sp. nov.). Systematic remarks. Petropedetes perreti is one of to the two species of the genus with fully developed webbing. In contrast, P. euskircheni sp. nov., P. johnstoni, P. parkeri and P. vulpiae sp. nov. show a mere rudiment of web. It can be easily distinguished from P. p a l m i p e s (fully webbed) and P. c a m e ro n e n s i s (halfwebbed) by its bigger and distinct tympanum and the tympanic papilla in males (see Fig. 8 a, 8 b, 8 d). The most similar species is P. juliawurstnerae sp. nov. (see respective paragraph for distinguishing characters). Lawson (1993) notes the occurrence of P. p e r re t i from the Rumpi Hills in western Cameroon. However, the taxonomic status of these frogs needs confirmation. Etymology. 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., Published as part of Barej, Michael F., R��del, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., B��hme, 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 28-30, DOI: 10.5281/zenodo.193288, {"references":["Amiet, J. - L. (1973) Caracteres diagnostiques de Petropedetes perreti, nov. sp. et notes sur les autres especes camerounaises du genre (Amphibiens Anoures). Bulletin de l'Institut Francais d'Afrique Noire, Serie A, 35, 462 - 474.","Herrmann, H. - W., Bohme, W., Herrmann, P. A., Plath, M., Schmitz, A. & Solbach, M. (2005) African biodiversity hotspots: The amphibians of Mt. Nlonako, Cameroon. Salamandra, 41, 61 - 81.","Lamotte, M., Perret, J. - L. & Dzieduszycka, S. (1959) Contribution a l'etude des Batraciens de l'Ouest africain. - IX. Les formes larvaires de Petropedetes palmipes, Conraua goliath et Acanthixalus spinosus. Bulletin de l'Institut Francais d'Afrique Noire, Serie A, 21, 762 - 776.","Amiet, J. - L. (1975) Ecologie et distribution des Amphibiens Anoures de la region de Nkongsamba. Annales de la Faculte des Sciences de Yaounde, 20, 33 - 107.","Amiet, J. - L. (1983) Une espece meconnue de Petropedetes du Cameroun: Petropedetes parkeri n. sp. (Amphibia Anura: Ranidae, Phrynobatrachinae). Revue Suisse de Zoologie, 90, 457 - 468.","Amiet, J. - L. (1989) Quelques aspects de la biologie des Amphibiens Anoures du Cameroun. L'annee biologique, 28, 73 - 136.","Lawson, D. P. (1993) The reptiles and amphibians of the Korup National Park Project, Cameroon. Herpetological Natural History, 1, 27 - 90."]}

Petropedetes parkeri Amiet, 1983 Figs. 1 c, 2 c, 4, 7a, 7 b, 7 f Material examined. 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. Problematic material (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, Tchimb��l��, 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, Yaound��, 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 Tchimb��l��, coll. V. Gossmann & S. L��tters; ZMB 73892 (female), same data as IRSNB-KBIN 14.941. Diagnosis. Large sized Petropedetes; 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. Description. Large sized Petropedetes 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; canthus rostralis 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). Coloration. 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. Natural history. After the first work on the ecology of the species by Sanderson (1936, referred to as P. johnstoni), additional data have been gathered by Amiet. However, some observations by Amiet (e.g. 1975, 1983, 1986) are based on populations of ��� P. parkeri ��� from Mt. Kala, the Bamil��k�� 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 P. johnstoni). 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. The advertisement call of P. p a r k e r i has been published by Narins et al. (2001). Narins et al. (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 P. parkeri is finished before the end of the rains or at the beginning of the dry season (Parker 1936). Petropedetes parkeri 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 P. aff. parkeri from Gabon (see below, Fig. 7 f) have been collected during the day in a gallery forest. P. aff. parkeri is called ���barking frog��� by locals in western Gabon (V. Gossmann pers. comm.). Distribution. We can only assign populations from western Cameroon and eastern Nigeria to P. parkeri with certainty (Fig. 4). At these locations species inhabit altitudes below 1000 m. Genetics. The genetic comparison for the uncorrected p-distances of the mitochondrial 16 S rRNAfragment (Table 4) gave the following results for Petropedetes parkeri: interspecific comparison between P. parkeri and all other Central African Petropedetes taxa ranged between 5.94 %- 12.39 %, while the intraspecific variation within P. p a r k e r i was much lower at 0.00%- 0.22 % (N= 4). Systematic remarks. Petropedetes parkeri has long been confused with ��� P. newtonii ��� (described herein as P. v u l p i a e sp. nov.) and P. johnstoni (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. P. parkeri and P. euskircheni sp. nov. 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. P. parkeri is only rudimentary-webbed and therefore easily distinguishable from more fully webbed species (P. palmipes and P. p e r re t i, both fully webbed; P. cameronensis and P. juliawurstnerae sp. nov., both half-webbed). Moreover, the presence of a distinct tympanum and a tympanal papilla seperates P. parkeri from P. cameronensis and P. palmipes, which lack these characters. The size of the femoral gland distinguishes P. parkeri from P. vulpiae sp. nov. and P. johnstoni, which possess much larger ones. Due to a similar appearance, P. p a r k e r i has been confused with a cryptic species in the past. Distinguishing characters between P. p a r k e r i and P. euskircheni sp. nov. are presented in the description of the new species (see below). As holotype, Amiet (1983) has chosen a P. parkeri 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, L��na (Yaound�� 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, Yaound�� 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 P. parkeri may overlap with the altitudinal range of P. euskircheni sp. nov. (type locality at about 900 m; see below). Without genetic comparisons, so far the P. parkeri -like populations in Central Cameroon region around Yaound�� and the northern populations of the Manengouba Mountains cannot be assigned with certainty, neither to P. parkeri nor to P. euskircheni sp. nov. Due to the finding of P. euskircheni sp. nov. 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. Fr��tey & Blanc 2001; L��tters et al. 2001) and Equatorial Guinea (de la Riva 1994; Lasso et al. 2002) to P. aff. parkeri (Fig. 4). Etymology. 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., Published as part of Barej, Michael F., R��del, Mark-Oliver, Gonwouo, Legrand Nono, Pauwels, Olivier S. G., B��hme, 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 17-22, DOI: 10.5281/zenodo.193288, {"references":["Amiet, J. - L. (1983) Une espece meconnue de Petropedetes du Cameroun: Petropedetes parkeri n. sp. (Amphibia Anura: Ranidae, Phrynobatrachinae). Revue Suisse de Zoologie, 90, 457 - 468.","Sanderson, I. T. (1936) The amphibians of the Mamfe Division, Cameroon. - II. Ecology of the frogs. Proceedings of the Zoological Society of London, 165 - 208.","Amiet, J. - L. (1975) Ecologie et distribution des Amphibiens Anoures de la region de Nkongsamba. Annales de la Faculte des Sciences de Yaounde, 20, 33 - 107.","Amiet, J. - L. (1986) La batrachofaune sylvicole d'un secteur forestier du Cameroun: la region de Yaounde. Memoires du Museum National d'histoire Naturelle, 132, 29 - 42.","Parker, H. W. (1936) The amphibians of the Mamfe Division, Cameroons. - I. Zoogeography and systematics. Proceedings of the Zoological Society of London, 135 - 163.","Narins, P. M., Lewis, E. R., Purgue, A. P., Bishop, P. J., Minter, L. R. & Lawson, D. P. (2001) Functional consequences of a novel middle ear adaptation in the Central African frog Petropedetes parkeri (Ranidae). The Journal of Experimental Biology, 204, 1223 - 1232.","Shine, R. (1979) Sexual selection and sexual dimorphism in the amphibia. Copeia, 1979, 297 - 306.","Boulenger, G. A. (1900) A list of the batrachians and reptiles of the Gaboon (French Congo), with descriptions of new genera and species. Proceedings of the Zoological Society of London, 1900, 433 - 456.","Amiet, J. - L. (1978) Les amphibiens anoures de la region de Mamfe (Cameroun). Annales de la Faculte de Sciences du Cameroun, 25, 189 - 219.","Fretey, T. & Blanc, C. P. (2001) Inventaire systematique des amphibiens anoures du centre du Gabon. Bulletin de la Societe Zoologique de France, 126, 375 - 390.","Lotters, S., Gossmann, V., Obame, F. & Bohme, W. (2001) Zur Herpetofauna Gabuns Teil I: Einleitung, Untersuchungsgebiet und Methodik, kommentierte Artenliste der gefundenen Froschlurche. Herpetofauna, 23, 19 - 34.","de la Riva, I. (1994) Anfibios anuros del Parque Nacional de Monte Alen, Rio Muni, Guinea Equatorial. Revista Espanola de Herpetologia, 8, 123 - 139.","Lasso, C. A., Rial, A. I., Castroviejo, J. & de la Riva, I. (2002) Herpetofauna del Parque Nacionale de Monte Alen (Rio Muni, Guinea Equatorial). Graellsia, 58, 21 - 34."]}