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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

Cacosternum nanogularum sp. nov. Small-throated Dainty Frog. (Figs. 3 I, 4 I) Holotype. An adult male MHNG 2740.77 collected by M. Burger, 3 December 2005, on the road between Eshowe and Nkandla, KwaZulu-Natal Province, South Africa, 28 �� 51 ��� 04��� S, 31 �� 12 ��� 27 ��� E. The specimen was collected from a roadside puddle. Paratypes. Six males; MHNG 2740.75 from 41 km from Kranskop on road to Qudeni, KwaZulu-Natal Province, South Africa, 28 �� 43 ��� 41 ��� S, 30 �� 56 ��� 35 ��� E; NMB A 7518 ��� 20, MHNG 2740.76 from 13 km from Kranskop along road to Qudeni, KwaZulu-Natal Province, South Africa, 28 �� 54 ��� 14 ��� S, 30 �� 51 ��� 15 ��� E; MHNG 2740.78 collected on the road between Eshowe and Nkandla, KwaZulu-Natal Province, South Africa, 28 �� 51 ��� 04��� S, 31 �� 12 ��� 27 ��� E; a female, MHNG 2740.74, from Dukuduku, KwaZulu-Natal Province, South Africa, 28 �� 21 ' 09" S, 32 �� 17 ' 53 " E. Diagnosis. The uncorrected p distances for 16 S from the other 14 species ranges from 3.8���6.9 %, while the within-clade p distances range from 0���0.4 (n= 7). The sample has five likely tyr haplotypes of which none are shared. This species lacks a distinct supratympanic fold, which is present in all the other species. It can be distinguished from C. capense by the absence of large lateral and posterior glands on the dorsum; from C. karooicum which has the nostrils at least three nostril diameters back from the snout margin; from C. leleupi which has a small amount of webbing; from C. namaquense which has a diffuse grey vocal sac anteriorly (dark in C. nanogularum sp. nov.) (Fig. 4). It has three supernumerary tubercles on the palm, less than the five of C. aggestum sp. nov. and C. leleupi. The inner metatarsal tubercle is equal to the proximal subarticular tubercle of the first toe, differing from the wider inner metatarsal tubercle of C. aggestum sp. nov., C. boettgeri and C. parvum. The rictal gland is indistinct or absent, which distinguishes it from the prominent glands of C. aggestum sp. nov., C. boettgeri, C. australis sp. nov., C. striatum, C. nanum, C. parvum, C. platys, C. kinangopensis and C. plimptoni. The advertisement call is a series of chirps, which distinguishes it from the regular single or multiple clicks of C. boettgeri, C. kinangopensis, C. leleupi, C. plimptoni and the brief chirp of C. nanum. It is unlike the 'bouncing marble' calls that consist of a number of pulses that speed up, as C. aggestum sp. nov., C. australis sp. nov. and C. platys, or the long complex call of C. rhythmum sp. nov. The call is not a creak as in C. capense, C. karooicum, C. namaquense, C. parvum, or C. striatum. Description of the holotype. The holotype is an adult male, SUL 19.7. Body widest at mid-belly, with a narrow head (HW/SUL 0.33). The head is bluntly rounded from above and in profile. Head length measured from the angle of the jaw is about one third of body length (HL/SUL 0.31). Canthus rostralis rounded, straight from eye to nostril, loreal region sloped outwards ventrally; nostrils small, rounded, directed laterally, and posteriorly. The nostrils are placed nearly midway between the snout than the eye (EN/SL 0.52). Internostril distance is greater than distance between eye and nostril (NN/EN 1.31). Eyes directed anterolaterally, the eyes protrude, and are visible from below, relatively small (ED/HW 0.35; ED/SUL 0.12), less than snout (ED/SL 0.74). Distance between anterior corners of eyes subequal to internostril distance (NN/EE 0.53). The angle of the jaw is posterior to a line drawn vertically from the back of the eye. The tympanum is not visible. Jaws without dentition; choanae small, round, located at anterior margins of roof of mouth; vomer processes and teeth absent; tongue long, narrow, slightly bifurcated distally. No median lingual papilla present. The dorsal surfaces of the head, trunk and limbs are smooth, with glands and skin folds present but indistinct; the rictal gland is smooth, continuing posteriorly as a series of bulges to the arm insertion. The supratympanic fold forms a forms a groove running over the arm to the leg insertion. Lines of broken skin glands form an hourglass pattern dorsally, narrowest at the scapular region. The underside is smooth, with a slightly pigmented vocal sac with lateral folds. The fore limb is slender, hand small (HAN/SUL 0.30), finger tips bluntly rounded without discs. Relative finger lengths IColour in life. The background colour is beige, overlain with a dark brown interorbital bar with a sharp tip pointing posteriorly, and a black band running from the nostril, through the eye to the arm insertion. The upper lip is white with black speckles, continuing as a white rictal gland. Colour in preservative. The patterns are visible as dark browns on a paler background. Paratype variation. The male paratypes have SUL 20.4���22.5, with the females 23.2 ���25.0. They are very similar to the holotype in having discontinuous bumpy rictal glands, large subarticular tubercles on the hand, and small conical subarticular tubercles at the base of the fourth toe. The belly spots are similar, with the female throats spotted but less pigmented than the males, while the body proportions show little variation. Advertisement call. The general call structure consists of a series (10 to> 20) of regularly repeated pulsed notes (chirps), with a note rate of 0.4��� 0.6 s - 1. The mean note duration is 89 ms, with the mean number of pulses per call of 9. The mean pulse rate is 89 s - 1. It has a mean emphasised frequency of 3.02 kHz. The illustrated call (Fig. 10) has a duration of 0.1 s, with 9 pulses at a pulse rate of 80 s - 1. Eggs and tadpoles. Unknown. Distribution. Molecular samples come from Nkandla and the Lebombo Mountains, with additional specimens known from Qudeni and Dukuduku (Fig. 9). The full extent of its range is unknown. The frogs were found in flooded temporary pools with emergent vegetation. Etymology. The specific name is derived from the Latin nanus (small), and gula (throat). It is an adjective. The name refers to the very small vocal sac that is produced during calling. Cacosternum poyntoni Lambiris, 1988 Genetics. The uncorrected p distances between this species and the other 14 range from 2.8 ���6.0%, while the within-clade p distances range from 0.0��� 0.4 (n= 20). The sample has 16 likely tyr haplotypes, of which one is shared between a single C. nanum and one C. rhythmum sp. nov. Advertisement call. Based on an analysis of 447 calls of 41 specimens, a calling bout usually consists of a long (15 to> 30) series of pulsed chirps that are rapidly repeated, about 1.2 s - 1 (range 0.6 ��� 2.0 s - 1). The mean call (chirp) duration is 52.6 ms (range 19.5���99.7 ms). The mean number of pulses per call is 11 (range 5���18), with a mean pulse rate of 200 s - 1 (range 95���288 s - 1). The mean emphasised frequency is 3.6 kHz (range 3.3���4.1 kHz) A spectrogram of a typical call is shown in Fig. 10. It has a duration of 0.04 s, consisting of 11 pulses at a pulse rate of 250 s - 1. Morphology. Females reach 18 mm SUL. The dorsum lacks protruding glands. Viewed from above, the nostrils are close to the anterior margin of the snout. The supratympanic fold continues as a straight line from eyelid to arm insertion. The rictal gland is interrupted, tapering to the arm insertion. There are four supernumerary tubercles on the palm. The inner metatarsal tubercle is low, with a width equal to the tip of the first toe. Breeding males possess an inconspicuous nuptial pad that extends to the last phalanx. The belly spots are the size of the lens of the eye, dense grey, often running together (Fig. 3). Distribution. This species is known from the south and east of South Africa, occurring in 276 quarter degree squares (Animal Demography Unit 2012). Molecular samples come from Baviaanskloof, Cedarville, Goukamma, Hogsback, Kubusi Forest Reserve, Maclear and Montague Pass in South Africa (Fig. 7). It breeds in a variety of water bodies (Scott 2004 b), usually those with emergent vegetation. Note. We regard Cacosternum poyntoni as a junior synonym of C. nanum, following the reasons presented by Minter (2004), Scott (2004 b) and Scott & Minter (2004 a, b). These include the lack of material apart from the type, after nearly 60 years, from a well-known locality, Pietermaritzburg; the consensus amongst local herpetologists that this is not a valid taxon; and the skeletal similarity to C. nanum from an unpublished report. Cacosternum parvum Poynton, 1963 Mountain Dainty Frog. (Figs. 3 K, 4 K) Genetics. The within-clade variation in uncorrected p distances for the 16 S fragment is 0.2 %, with the differences to the other 14 species being 3.9���7.6 %. The sample has two tyr haplotypes, of which none are shared. Advertisement call. The call is a brief chirp. The following description is based on an analysis of 172 calls of 17 specimens. A motivated calling bout usually consists of a long (10 to> 20) series of pulsed chirps that are repeated at a rate of about 0.6 s - 1 (range 0.4��� 0.7 s - 1). The mean call (chirp) duration is 60.2 ms, (range 27.6 ���109.0 ms). The mean number of pulses per call is 6 (range 3���9) with a mean pulse rate of 86.8 s - 1 (range 59.5���125 s - 1). The mean emphasised frequency is 3.99 (range 3.55���4.42 kHz). The illustrated call (Fig. 10) has a duration of 0.06 s, with seven almost equally spaced pulses, with a pulse rate of 100 s - 1. Morphology. Females do not exceed 16.5 mm SUL in our sample. Breeding males have dark vocal sacs with a paler margin. The dorsum is smooth, and, viewed from above the nostrils are positioned on the anterior edge of the snout. The supratympanic fold continues posteriorly to form a rounded saddle. The rictal gland is indistinct, interrupted. There are no supernumerary tubercles on the palm. There is no webbing, with the notch between the third and fourth toes reaching the proximal subarticular tubercle of the third toe. The outer metatarsal is absent or indistinct, while the inner metatarsal tubercle is low, twice the width of the tip of the first toe. The nuptial pad in breeding males is slightly pigmented but indistinct. The belly markings consist of groupings of brown speckles that form blotches. Distribution. Molecular samples come from Mariepskop (Fig. 9). The species appears to be a highland endemic in eastern South Africa (Scott & Minter 2004 b), where it breeds in high altitude grassland pools (Scott 2004 e). Note. The GenBank sequence DQ 022353 with the voucher identified as Cacosternum nanum parvum (Scott 2005) is a C. nanum sequence. Cacosternum platys Rose, 1950 Flat Dainty Frog. (Figs. 3 L, 4 L) Genetics. The within-clade variation in uncorrected p distance for the 16 S fragment varies from 0.0��� 0.2 %, but between C. platys and the other 14 species 1.8���6.6 %. The sample has nine likely tyr haplotypes, of which one is shared with a specimen of C. australis sp. nov. See the Discussion. Advertisement call. A typical advertisement call (Fig. 11) consists of a long series of pulses that increase in tempo. The duration of the illustrated call is 0.55 s, with 34 pulses. The initial and final pulse rates are 20 s - 1 and 80 - 1. Morphology. This is a small species, with females less than 14 mm in our sample. The vocal sac in breeding males is pale grey (Fig. 4). The dorsum is smooth. Viewed from above, the nostrils are situated on the anterior margin of the snout. The supratympanic fold continues posteriorly in a straight line from the eyelid to the arm insertion. The rictal gland is interrupted. There are two supernumerary tubercles on the palm. There is no webbing. The inner metatarsal tubercle is low, rounded, equal in width to the tip of the first toe. The nuptial pad is a dark swollen mass at the base of the first finger, tapering to the level of the proximal tubercle. The belly pattern consists of very small spots consisting of groups of speckles. Distribution. Molecular evidence shows the species is present on the Cape Peninsula (Noordhoek) and in Cape Town (Kenilworth) (Fig. 7). It is a fynbos species, breeding in low altitude temporary pools. Notes. The GenBank sequence DQ 283258 with the voucher identified as Cacosternum platys (Frost et al. 2006) is Microbatrachella capensis. The GenBank sequence DQ 022352 with the voucher identified as Cacosternum boettgeri (Scott 2005) is C. platys. Genetics. This species varies from the other 14 species for the 16 S fragment, by uncorrected p values of 2.5���5.9 %. A single specimen had only one likely tyr haplotype, that was not shared with any other species. Advertisement call. A series of 29 calls of four individuals from the Serengeti National Park, Tanzania showed a mean duration of 0.55 s (range 0.37���0.77) and the mean number of notes (pulse trains) 6.2 (range 4���8) (Channing et al. 2005). A typical advertisement call from Nairobi (Fig. 11) is a harsh clicking, where each series of notes consists of double pulses, apart from the initial note of three pulses. The call duration is 0.5 s, with six notes at a rate of 10 s - 1. Morphology. The female holotype has a SUL 19.7 (Channing et al. 2005). Breeding males have yellow vocal sacs without a pale margin (Fig. 4). The dorsum is smooth, and, viewed from above the nostrils are positioned on the anterior edge of the snout. The supratympanic fold is inconspicuous. The rictal gland is indistinct. There are 5 supernumerary tubercles on the palm. There is no webbing, with the notch between the third and fourth toes reaching the proximal subarticular tubercle of the third toe. The outer metatarsal is absent or indistinct, while the inner metatarsal tubercle is conical and protruding, twice the width of the tip of the first toe. The nuptial pad in breeding males is slightly pigmented but massive, reaching the proximal subarticular tubercle. The belly markings consist of groupings of brown speckles that form blotches with spots equal to the size of the eye lens. Distribution. The species appears to be present from northern Tanzania to the highlands of central Kenya (Channing et al. 2005, Channing & Schmitz 2009) (Fig. 6) and may be more widely distributed. It breeds in temporary and permanent pools with little emergent vegetation. Cacosternum rhythmum sp. nov. Rhythmic Dainty Frog. (Figs. 3 N, 4 N) Holotype. An adult male NMB A 7521, collected by M. Burger, 18 December 2001 at Nottingham Road (29 �� 21 ��� 21 ��� S, 29 �� 59 ��� 36 ��� E), KwaZulu-Natal Province, South Africa. The habitat is flat open terrain with shallow pools and flooded grassy patches. Paratypes. Eight males and one female, NMB A 7522���7530, from Nottingham Road, Nottingham Ford, and 28 km S of Harrismith. Diagnosis. This species differs from the other 14 species for the 16 S fragment, by uncorrected p values of 1.1��� 6.2 %. The within-clade variation ranges from 0.0��� 0.2 %. The small difference of 1.1 % with C. boettgeri will be considered in the Discussion. The sample had eight likely tyr haplotypes, of which two were shared between one C. nanum and one C. rhythmum sp. nov. The vocal sac of breeding males is white (Fig. 5), which distinguishes it from species where the sac is black, grey or yellowish-beige (C. capense, C. karooicum, C. leleupi, C. namaquense, C. nanogularum sp. nov., C. aggestum sp. nov., C. boettgeri, C. australis sp nov., C. nanum, C. parvum, C. platys, and C. kinangopensis. It is distinguished from C. capense by the absence of protruding dorsolateral glands and a pair just above the vent. Viewed from above, the nostrils are close to the anterior margin of the snout, distinguishing it from C. karooicum, which has the nostrils about three nostril diameters back from the anterior edge of the snout. The supratympanic fold continues posteriorly as a weak saddle, distinguishing it from C. nanogularum sp. nov., which has no saddle. The rictal gland is smooth, tapering to the arm insertion. This distinguishes it from those species where the rictal gland is interrupted (C. capense, C. karooicum, C. striatum, C. nanum, C. parvum and C.platys). There are no supernumerary tubercles on the palm, distinguishing it from species with 2���5 tubercles (C. capense, C. karooicum, C. leleupi, C. namaquense, C. nanogularum sp. nov., C. aggestum sp. nov., C. boettgeri, C. nanum and C. platys). It has no webbing, which distinguishes it from C. leleupi, which has a trace of webbing between the third and fourth toes. The inner metatarsal tubercle is small, conical, equal in width to the proximal subarticular tubercle of the first toe. This distinguishes it from species where the width of the inner metatarsal tubercle is equal to the width of the tip of the first toe (C. capense, C. karooicum, C. namaquense, C. nanogularum sp. nov., C. australis sp. nov., C. striatum, C. nanum, C. plimptoni and C. platys), and from those where the inner metatarsal tubercle is twice the width of the tip of the first toe (C. leleupi, C. aggestum sp. nov., and C. parvum. The belly is pale, with a few very small brown blotches. This distinguishes it from species where the spots are larger than the eye lens, sometimes running together to form blotches (C. capense, C. namaquense, C. nanogularum sp. nov., C. aggestum C. aggestum sp. nov.,) and from C. australis sp. nov. which has irregular yellow, Published as part of Channing, Alan, Schmitz, Andreas, Burger, Marius & Kielgast, Jos, 2013, A molecular phylogeny of African Dainty Frogs, with the description of four new species (Anura: Pyxicephalidae: Cacosternum), pp. 518-550 in Zootaxa 3701 (5) on pages 533-539, DOI: 10.11646/zootaxa.3701.5.2, http://zenodo.org/record/217052

Cacosternum striatum FitzSimons, 1947 Striped Dainty Frog. (Figs. 3 O, 4 O) Genetics. The uncorrected p within-clade variation in 16 S ranges from 0.0��� 0.2 % (n= 3). This species differs from the other 14 species by 2.5���7.3 %. The sample has four likely tyr haplotypes, of which one is shared with an individual C. boettgeri. Advertisement call. The advertisement call is a brief high-pitched chirp. A typical call (Fig. 11) shows that the pulses are arranged in doublets in the middle, with the initial and final few pulses being produced at a faster rate than the middle. The duration of this call is 0.33 s, with the 21 pulses starting at 100 s - 1, slowing to 60 s - 1, before speeding up slightly at the end of the call. Morphology. These are small frogs, with females reaching 13.7 mm SUL in our sample (n= 7). The vocal pouch of breeding males is pale (Fig. 4). The dorsum is smooth, lacking protruding glands. Viewed from above, the nostrils are within one nostril diameter of the anterior snout margin. The supratympanic fold continues posteriorly to form a saddle. The upper lip continues posteriorly as an indistinct interrupted rictal gland. The palm lacks supernumerary tubercles. The inner metatarsal tubercle is low, inconspicuous, as wide as the tip of the first toe. The nuptial pad of breeding males is low, inconspicuous, extending to the level of the proximal subarticular tubercle. The belly is silvery white, with some small black speckles. Distribution. The species is widespread in KwaZulu-Natal, South Africa, where it has been recorded from 13 quarter-degree cells (Animal Demography Unit, University of Cape Town). Our sequenced specimens come from Mpur Forest (Fig. 9). It breeds in shallow temporary pools that may be overgrown., Published as part of Channing, Alan, Schmitz, Andreas, Burger, Marius & Kielgast, Jos, 2013, A molecular phylogeny of African Dainty Frogs, with the description of four new species (Anura: Pyxicephalidae: Cacosternum), pp. 518-550 in Zootaxa 3701 (5) on page 540, DOI: 10.11646/zootaxa.3701.5.2, http://zenodo.org/record/217052

Channing, Alan, Schmitz, Andreas, Burger, Marius, Kielgast, Jos (2013): A molecular phylogeny of African Dainty Frogs, with the description of four new species (Anura: Pyxicephalidae: Cacosternum). Zootaxa 3701 (5): 518-550, DOI: http://dx.doi.org/10.11646/zootaxa.3701.5.2

Cacosternum namaquense Werner, 1910 Namaqua Dainty Frog. (Figs. 3 H, 4 H) Genetics. The within-clade uncorrected p distance for 16 S is 0.2 %, with the distances to the other 14 species ranging from 3.0��� 7.2 %. The sample has two likely tyr haplotypes, of which neither are shared. Advertisement call. The call is a high-pitched creak. The illustrated example (Fig. 8) has a duration of 0.3 s, with 18 pulses at a pulse rate of 56 s - 1. Morphology. Females reach 20 mm SUL in our sample. Breeding males have a vocal sac that is diffuse grey anteriorly, with the belly pattern of spots posteriorly. Viewed from above, the nostrils are within one nostril diameter of the snout margin. The supratympanic fold continues posteriorly to form a glandular saddle. The rictal gland is prominent, continuous with the upper lip. There are two supernumerary tubercles on the palm. There is no webbing, although the notch between the third and fourth toes passes the proximal subarticular tubercle of the third toe. The inner metatarsal tubercle is conical, equal to the width of the tip of the first toe. The nuptial pad of breeding males extends to the level of the proximal subarticular tubercle of the first finger. The belly spots are larger than the lens of the eye, many running together to form lines (Fig. 4). Distribution. Molecular samples are from Arakoep (Fig. 7). This species is morphologically distinct, unlikely to be confused with similar species, and is widespread in Namaqualand, known from 41 quarter-degree squares (Animal Demography Unit 2012). The species is known from arid rocky areas, where it breeds in temporary pools (Scott 2004 d)., Published as part of Channing, Alan, Schmitz, Andreas, Burger, Marius & Kielgast, Jos, 2013, A molecular phylogeny of African Dainty Frogs, with the description of four new species (Anura: Pyxicephalidae: Cacosternum), pp. 518-550 in Zootaxa 3701 (5) on pages 532-533, DOI: 10.11646/zootaxa.3701.5.2, http://zenodo.org/record/217052

Hyperolius veithi sp. nov. (Figs. 2 A, 4 A) Holotype. ZFMK 89607 (field number A 519), adult male, collected by J. Kielgast at a flooded area in the middle of primary forest away from rivers and streams in Salonga National Park (02.88 S, 20.41 E, ca. 415 m above sea level), Province of Bandundu, Equateur Kasaï Oriental and Occidental, Democratic Republic of Congo, 24 January 2008. Paratypes. Fifty-eight adult specimens of both sexes (ZFMK 89608 89645, ZMUC R 771393 R 771412) from the same locality; data as the holotype but collected 24 26 January 2008. Diagnosis. A sexually monochromatic species of Hyperolius based on the following characters: pupil horizontal; tips of toes and fingers broadened; last phalanx of fingers out of alignment; fingers not arranged in opposing pairs; gular flap in males (Schiøtz 1999; Channing & Howell 2006). It is defined by: (1) SVL males 25.7 ± 1.7 mm (22.7 28.4 mm, N = 32), females 33.9 ± 1.3 mm (31.7 36.5 mm, N = 27); (2) TIBL /SVL in males 0.54 ± 0.02 (0.52 0.61, N = 32) and in females 0.51 ± 0.02 (0.48 0.55, N = 27), HW/SVL in males 0.33 ± 0.02 (0.3 0.37, N = 32) and in females 0.33 ± 0.01 (0.29 0.35, N = 27); (3) dorsal surface smooth to coarse with few scattered pointed warts, most prominent and tubercular below eye; (4) snout shape dorsally and laterally rounded, nares visible from above; (5) E N/EYE in males 0.78 ± 0.09 (0.53 0.95, N = 32) and in females 0.85 ± 0.09 (0.71 0.99, N = 27), canthus rostralis convex from tip of snout to nostril and straight or slightly concave from nostril to eye; (6) tympanum covered by thick skin but with annulus tympanicus still visible; (7) FOOT / TIBL in males 0.67 ± 0.04 (0.57 0.75, N = 32) and in females 0.75 ± 0.05 (0.67 0.86, N = 27); (8) foot webbing formula 1 (1), 2 i (1) 2 e(½), 3 i (1 1 ½) 3 e(½ 1), 4 i (1 2) 4 e(1 1 ½), 5 (1 ½); (9) no sexual dichromatism; both sexes dorsally tan, olive brown to dark brown, usually lighter towards the sides and dorsally with more or less intense, diffuse dark markings; there is always a bright yellow dorsolateral line from tip of snout (lines from both sides of the body meet here) to sacral region which is commonly bordered by dark brown; it is broader posterior to eye and often contains tiny dark brown spots which may continue posterior into groin and onto hind limbs; always a cream to yellow spot on the heel; ventral sides are white or cream; a red marking on the inner femur is lacking; the gular flap of males is yellow; the iris is bronze to dark brown; (10) for DNA barcodes access GenBank (http://www.ncbi.nlm.nih.gov): GU 443988 (male holotype, ca. 530 bp), GU 443975 (female paratype, ZFMK 89629, ca. 480 bp). By being brownish and having a light dorsolateral lines, H. veithi males and females (Fig. 2 A) are similar in colour and pattern to numerous Hyperolius species (see Schiøtz 1999), including PhJ H. cinnamomeoventris (Fig. 2 C D) and its suggested synonyms H. fimbriolatus Buchholz & Peters, 1876 and H. ituriensis Laurent, 1943 (Lötters et al. 2001, Frost 2009). Hyperolius schoutedeni from the Congo basin deserves special attention, as this reed frog, like H. veithi, is also sexually monochromatic. The new species can be distinguished from any other Hyperolius species which are brownish and have a light dorsolateral lines (including H. cinnamomeoventris and H. schoutedeni) as follows: (i) dorsolateral lines relatively short and broad ending in the sacral region (versus in the groin in other species); (ii) dorsolateral lines in life are bright yellow (versus whitish to tan in other species); (iii) dorsolateral lines often contain dark spots (absent in most other species); (iv) presence of a light spot on the heel, cream to yellow in life (absent in other species); (v) absence of a in life red marking on the inner femur (present at least in H. cinnamomeoventris and its suggested synonyms and occasionally in H. schoutedeni; Schiøtz 2006). In addition to (i) to (v), in H. schoutedeni, a mid-dorsal line is common (absent in H. veithi) and the new species is larger than H. schoutedeni and develops less foot webbing. Five individuals (three males, two females from N'Sele and Garamba, DRC; Schiøtz 2006) and the female holotype of H. schoutedeni (Fig. 4 B) have maximum SVL in males 22.9 (minimum SVL in H. veithi 22.7 mm) and in females 25.2 mm (minimum SVL in H. veithi 31.7 mm); holotype foot webbing formula is 1 (1), 2 i (1) 2 e(½), 3 i (1 ½) 3 e(1), 4 i (2) 4 e(1 ½), 5 (1) (for H. veithi see above) (note that measurements and foot webbing data for H. schoutedeni differ slightly from those given by Schiøtz 2006). Several additional Hyperolius species have been described from DRC (Schiøtz 1999; Frost 2009) but remain little known as they are based on one to a few specimens only and poor original descriptions. Of these, H. atrigularis Laurent, 1941, H. kibarae Laurent, 1957 and H. polli are similar to H. veithi in colour and pattern in preservative, but none has continuous bright dorsolateral lines (based on holotype examinations and Laurent 1941, 1957). In terms of uncorrected p-distances of the 16 S mitochondrial rRNA, H. veithi is little differentiated genetically from some, but not all populations assigned to H. cinnamomeoventris (Table 2). We reject conspecifity, however, as H. veithi syntopically occurs with H. cinnamomeoventris (uncorrected p-distance 2.8 4.4 %, but see comment to Table 2) and amplexus was observed 27 times between H. veithi individuals but never between H. veithi and H. cinnamomeoventris. Additional arguments include presence of monochromatism in the new species versus dichromatism in H. cinnamomeoventris (Fig. 2 A versus Fig. 2 B D) and H. veithi having white eggs (Fig. 2 A) versus the half-pigmented eggs in H. cinnamomeoventris (Lötters et al. 2004). The genetically related H. molleri and H. thomensis (Table 2) are dorsally brownish to greenish, lack dorsolateral lines (present in the new species) and possess a more coarse skin texture (Fig. 2 A versus Fig. 2 E F). Description. The type series contains 32 males and 27 females. In external morphology, mature sexes differ in size (Table 3) and the presence of gular flap and vocal sac in males only. Body slender with sacrum width about one fifth of SVL; head short and broad with distance from tip of snout to posterior corner of eye about one sixth of SVL; HW about one third of SVL; snout tip in dorsal and lateral views rounded; nostril dorsolateral, protruding and visible from above; choanae rounded; maxillary teeth present, vomerine teeth lacking; tongue about as long as wide, free for half of its length and anteriorly bifurcated; canthus rostralis convex from nostril to tip of snout, straight and longer from nostril to anterior corner of eye; EYE> E N; loreal area barely concave; tympanic membrane covered by thick skin but tympanic annulus visible, TYMP Hyperolius species (see Schiøtz 1999). Tibia long, about half of SVL, tibiotarsal articulation extending to loreal region when hind limb adpressed forward along body; FOOT For measurements and proportions see Table 3. In life (Fig. 2 A), both sexes are dorsally tan, olive brown to dark brown, usually lighter towards the sides and dorsally with more or less intense, diffuse dark markings. A yellow dorsolateral line always runs from the Hyperolius cinnamomeoventris and related reed frogs studied in this paper (Fig. 3; Table 1). For meaning of #, ## and E see Table 1. Distinct difference of the two H. veithi sp. nov. samples against the other samples is because the sequence of GU 443975 is ca. 480 bp only (versus ca. 530 bp in GU 443988). Sample, locality (GenBank accession number) 1 2 3 4 5 6 7 8 9 10 11 12 13 1 H. cinnamomeoventris E, Kenya: Kakamega - Forest (AY 323925) 2 H. cinnamomeoventris E, Uganda: Bundibugyo 1.6 - (GU 443994) 3 H. cinnamomeoventris E, Uganda: Semliki 2.2 1.2 - (GU 443993) 4 H. cinnamomeoventris E, Uganda: Kampala 0.2 1.5 2.2 - (GU 443996) 5 H. cinnamomeoventris E, Uganda: Bwindi 1.6 1.1 1.6 1.5 - (AY 603985) 6 H. cinnamomeoventris E, Uganda: Kibale 0.6 1.8 2.6 0.4 1.9 - (GU 443995) 7 H. cinnamomeoventris E, DRC: Salonga 3.4 3.0 3.4 3.4 4.0 3.8 - National Park (GU 443998) 8 H., cinnamomeoventris E, CAR: Dzanga- 4.4 4.0 4.2 4.3 3.6 4.7 2.1 - Sangha Reserve (GU 443997) 9 H. cinnamomeoventris #, Gabon: Lamabréné 8.4 8.5 8.1 8.3 8.6 8.7 7.5 7.6 - (FJ 594077) 10 H. cinnamomeoventris ##, Angola: Kalandula 3.8 3.8 3.4 3.8 4.2 4.2 4.2 3.8 5.6 - (GU 443990) 11 H. molleri, São Tomé (AY 603990) 6.0 5.7 5.6 6.1 5.1 6.5 6.2 6.2 6.6 4.7 - 12 H. thomensis, São Tomé (AY 603991) 7.0 6.4 6.2 6.8 5.9 5.8 6.7 6.8 7.2 4.9 1.4 - 13 H. veithi sp. nov., DRC: Salonga National 9.4 5.2 8.5 5.4 4.6 5.8 4.4 8.7 8.6 3.8 6.2 6.4 - Park (GU 443988) 14 H. veithi sp. nov., DRC: Salonga National 4.2 3.5 2.9 3.9 3.3 4.4 2.8 3.5 7.3 2.2 4.9 4.8 0.0 Park (GU 443975) tip of snout (lines from both sides of the body meet here) to the sacral region; this line is commonly bordered by dark brown colour and is broader posterior to eye where it often contains tiny dark brown spots. Spots may continue posterior to the dorsolateral lines into groin and onto hind limbs. The heel always has a cream to yellow spot. Ventral sides are white or cream; a red marking on inner femur is lacking; the gular flap of males is yellow. The iris is bronze to dark brown. In preservative, colours fade as follows: tan and brown to greyish, yellow to cream; dark brown, cream and white remain. In characters with intraspecific variation, male holotype conditions are as follows: gular flap and vocal sac present; foot webbing formula 1 (1), 2 i (1) 2 e(½), 3 i (1) 3 e(½), 4 i /e(1), 5 (½); SVL 26.3 mm, TIBL 13.9 mm, HW 9.2 mm, E N 2.4 mm, EYE 3.7 mm, TYMP 1.4 mm, FOOT 10.6 mm. For individual aspects of colour in preservative and pattern see Figure 4 A; individual information on life colour was not recorded. For molecular genetic studies, some muscle tissue had been taken from the ventral side of the left femur. Distribution and natural history. The new species is only known from Salonga National Park in the central Congo basin, DRC (Fig. 1). The habitat is lowland rainforest (ca. 400 m above sea level) with seasonal rainfall (short dry period in February, long dry period in June August); annual precipitation is ca. 1,865 mm and monthly mean temperature ca. 25.0 °C (data obtained from WorldClim; Hijmans et al. 2005). Due to limited sampling efforts in the Congo basin and an expected lack of distribution barriers to Hyperolius veithi, we expect this species to show a geographically wider range than currently known. Twenty-seven amplectant pairs and five single males were found at the end of the rainy season (January) at night on vegetation, ca. 1 m above water in a flooded but apparently permanent pond deep within primary forest. Frogs were not heard calling and some pairs were observed laying small clutches of whitish eggs on vegetation above water (Fig. 2 A). It is expected that tadpoles (unknown) develop in water. Syntopic Hyperolius species included H. cinnamomeoventris, H. cf. platyceps and another undescribed species, not closely related to H. cinnamomeoventris and H. veithi. Conservation status. Due to the limited knowledge on both the geographic range encompassed by this species and population trends, we consider it Data Deficient when applying IUCN Red List categories and criteria (IUCN 2008). Due to the apparent lack of physical borders to dispersal and the observation that numerous other amphibian species in the Congo basin show relatively large distributions, we expected that Hyperolius veithi occupies a larger geographic range than currently known in the central Congo basin (see above). If so, it may perhaps later well rank under Least Concern. Also, we do not consider chytridiomycosis, an emerging infectious disease causing amphibian decline in other regions of the planet, a potential threat to H. veithi. The central Congo basin shows limited suitability for the emergence of this fungal pathogen (Rödder et al. 2009) and elsewhere in tropical Africa, Hyperolius species well survive despite high prevalence and individual parasite load (Kielgast et al. 2010). Etymology. The specific name is a patronym for Michael Veith, acknowledging his support of amphibian research in tropical Africa. Paraphyly of Cinnamon-belly reed frogs. As shown in Figure 3, Hyperolius cinnamomeoventris is paraphyletic when H. molleri, H. thomensis and H. veithi are considered distinct species. In addition to H. veithi (see above), we concur with Drewes & Wilkinson (2004) that specific distinctness of the sexually monochromatic H. molleri and H. thomensis is warranted. Although both these reed frogs may show similar colour and pattern (Fig. 2 E F) to PhF H. cinnamomeoventris (Fig. 2 C) (which is more applicable to H. molleri), they possess a more coarse skin texture, lack (in life) a red marking on the inner femur (present in H. cinnamomeoventris) and have parts of the toe and fingers tips bright orange (not different to dorsal or ventral colours in H. cinnamomeoventris). Hyperolius thomensis is distinguished further by being larger in body size and having a black, white and orange marbled venter in life (cream in H. cinnamomeoventris), while that of H. molleri is white or red in life and thus can be similar to that of H. cinnamomeoventris. When comparing uncorrected p-distances of H. molleri and H. thomensis with other reed frogs discussed in this paper, divergence is ≥ 4.7 % and ≥ 4.9 %, respectively (Table 2). An operable threshold for the consideration of putative species (to be confirmed by an integrative approach) in the 16 S mitochondrial rRNA gene for anuran amphibians is at about 3 % (Fouquet et al. 2007, Vieites et al. 2009) but can even be lower between well accepted species (e.g. Zimkus & Schick 2010). Applying this threshold to reed frogs studied here, specific distinctness of both H. molleri and H. thomensis against H. cinnamomeoventris is supported. Likewise, divergence of H. veithi and H. cinnamomeoventris is ≥ 4.4 % (or ≥ 2.2 % in the shorter H. veithi sequence, but see comment to Table 2). In addition, specific distinctness is still supported by morphology (colour and pattern in life and monochromatism versus dichromatism) and by having white versus half-pigmented eggs (see above). As clades currently assigned to H. cinnamomeoventris are generally well supported, in a purely cladistic view, different names should be applied. Considering the available names in the synonymy of H. cinnamomeoventris (Frost 2009), the Gabon clade in Figure 3 could be referred to as H. olivaceus Buchholz & Peters, 1876 with H. fimbriolatus treated as a synonym (Lötters et al. 2001), since this genetic sample was obtained at Lambaréné which is the type locality of both these names (Table 1). Likewise, the Angola clade (Fig. 3) is based on samples from Kalandula which is the type locality of H. cinnamomeoventris and its synonym H. tristis Bocage, 1866 (Table 1). Therefore, this clade could be recognized as H. cinnamomeoventris sensu stricto. All samples from the eastern range of Cinnamon-belly reed frogs constitute a monophyletic group, here called Eastern Clade (Figs. 1, 3). Of the available names in the synonymy of H. cinnamomeoventris, two originate from the eastern portion of the Cinnamon-belly reed frog distribution, H. ituriensis Laurent, 1943 and H. wittei Laurent, 1947 (Frost 2009). Consequently, one of them could potentially be applicable to the Eastern Clade, apart from the fact that the type locality of H. wittei lays out of the suggested geographic range of Cinnamon-belly reed frogs, at all (Fig. 1). In spite of this a priori conclusive scenario, we momentarily refrain from taxonomic action for two reasons. First, delimitating and phenotypic diagnosing putative species remains a problem. Not only is the material available limited for comparisons (see Appendix) and call and other natural history data are sparse or lacking (Lötters et al. 2004) so that an integrative approach combining data from different fields to is hampered. It is also true that molecular data demonstrate considerable variation within putative species and likewise overlap between clades (Table 2). Second, with regard to tree-based taxonomy, it has to be considered that only a fragment of one mitochondrial gene and relatively few samples have been involved in our analysis. Although, the 16 S mitochondrial rRNA gene is useful for DNA barcoding and tree-based species recognition in anuran amphibians (Vences et al. 2005, Fouquet et al. 2007, Vieites et al. 2009), it has been shown that when analysing mitochondrial DNA sequences, paraphyly does not always uncover distinct taxa. There are various explanations for paraphyly or polyphyly within species when constructing mitochondrial gene trees, in particular introgression, hybrid speciation and (although less of a concern for mitochondrial than for nuclear loci) incomplete lineage sorting (Funk & Omland 2003). In these cases some alleles in a species may appear more close to those of another (related) species than to conspecifics. As a result, intraspecific variation is larger than variation at the species level and may lead to a mistaken consideration of variants for species when following the monophyly criterion (Funk & Omland 2003). Intensive sampling and analyzing phylogeographic data including related species are helpful for an appropriate inter