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Amnirana parva sp. nov. Lesser White-Lipped Frog Figs. 5–7 Holotype. ZMB 88458 (field #: RG27, GenBank #: 16S: OQ400914), adult male, Liberia, Krahn-Bassa Proposed Protected Area, 5°39′02.1′′N, 8°39′05.0′′W, 28 March 2018, leg. M.-O. Rödel & J. Glos. Paratypes. Twelve males, ten females: ZMB 71273 (field #: MOR ANK 15, GenBank #: 16S: MG552248), adult female, Ghana, Ankasa National Park, 5°16′38.5212′′N, 2°38′41.8812″W, 5 August 2005, leg. A.C. Agyei & A. Hillers; ZMB 71274 (field #: MOR OWS 28, GenBank #: 16S: MG552262), adult female, Ghana, Owabi Wildlife Sanctuary, 6°44′50.5212″N, 1°42′11.6388″W, July 2005, leg. A.C. Agyei & A. Hillers; ZMB 71276 (field #: MOR LE 11, GenBank #: 16S: MG552287), adult female, Ghana, Leklebi-Agbesia, 6°56′ 1.3812″N, 0°29′1.5612″W, 19 July 2005, leg. A.C. Agyei & A. Hillers; ZMB 79187 (field #: GO14), adult male, Liberia, Gola National Forest, 7°27′10.6812″N, 10°41′31.3188″W, 28 November 2005, leg. A. Hillers; ZMB 79197 (field #: G84), adult female, Ghana, Volta Region, 22 August 2001, leg. M.-O. R ̂del; ZMB 79210 (field #: GOL 060, GenBank #: 16S: MG552310), adult male, Sierra Leone, Gola North Forest Reserve, 7°35′10.14″N, 11°1′30.9612″W, 15 September 2005, leg. A. Barrie & A. Hillers; ZMB 79253 (field #: GRE 010, GenBank #: 16S: MG552307), adult female, Liberia, Grebo National Forest, 5°24′6.48″N, 7°44′0.6612″W, 7 December 2005, leg. A. Hillers; ZMB 87105–06 (field #: P.LI 12 004–P.LI 12 005, GenBank #: 16S: MG552296 & MG552305), adult female and male, Liberia, Gba Community Managed Forest, 7°28′52.6584″N, 8°34′28.5564″W, 28 January 2012, leg. J. Penner; ZMB 87107 (field #: P.LI 12 077, GenBank #: 16S: MG552312), adult male, Liberia, Gba Community Managed Forest, 7°30′34.5636″N, 8° 41′50.7408″W, 3 February 2012, leg. J. Penner; ZMB 87108 (field #: P.LI 12 321, GenBank #: 16S: MG552315), adult male, Liberia, Sapo National Park, 5°29′42.8352″N, 8°21′55.1988″W, 5 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87109 (field #: P.LI 12 466, GenBank #: 16S: MG552323), adult male, Liberia, Mount Ghi South, 5°38′34.9944″N, 8°12′30.1428″W, 29 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87110–11 & 87116 (field #: P.LI 12 503–P.LI 12 505, GenBank #: 16S: MG552319 & MG552320), adult male, female and male, Liberia, Mount Ghi South, 5°39′11.124″N, 8°12′21.708″W, 3 October 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87112 (field #: P.LI 12 300, GenBank #: 16S: MG552322), adult male, Liberia, Sapo National Park, 5°31′20.8272″N, 8°20′46.2084″W, 14 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87113 (field #: P.LI 12 377, GenBank #: 16S: MG552317), adult male, Liberia, Sapo National Park, 5°30′18.5652″N, 8°20′14.9208″W, 18 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87114 (field #: P.LI 12 432, GenBank #: 16S: MG552316), adult female, Liberia, Mount Swa, 6°26′18.0636″N, 9°3′25.9416″W, 24 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 87115 (field #: P.LI 12 279, GenBank #: 16S: MG552314), adult female, Liberia, Sapo National Park, 5°31′34.8564″N, 8°20′53.1636″W, 12 September 2012, leg. J. Penner & N.L. Gonwouo; ZMB 88438 (field #: RG07), adult female, Liberia, Krahn-Bassa Proposed Protected Area, 06°02′34.8″N, 08°20′10.3″W, 25 March 2018, leg. M.-O. R ̂del & J. Glos; ZMB 88512 (field #: RG80), adult male, Liberia, Foya Proposed Protected Area, 08°00′21.3″N, 10°25′20.2″W, 5April 2018, leg. M.-O. Rödel & J. Glos; ZMB 88549 (field #: RG118, GenBank #: 16S: OQ400915), adult male, Liberia, Foya Proposed Protected Area, 08°03′18.6″N, 10°23′04.0″W, 8 April 2018, leg. M.-O. R ̂del & J. Glos. Additional material. Ten males, ten females; ZMB 90052 (field #: N035), adult female, Guinea, Keoulenta, 7°41′20.6”N, 8°21′19.0”W, 21 September 2009, leg. J. Doumbia, L. Sandberger, K. Camara & F. Gbêmou; ZMB 90053 (field #: N064), adult female, Guinea, Foromota, 7°42′58.3”N, 8°21′41.6”W, 3 November 2009, leg. J. Doumbia, L. Sandberger, K. Camara & F. Gbêmou; ZMB 90054 (field #: S4- Dian 91), adult male, Guinea, Diandian Missidé, 10°59′10.356”N, 13°47′44.556”W, 3 October 2016, leg. J. Doumbia & K. Camara; ZMB 90055 (field #: Ban _0362), adult female, Guinea, Banco, 8°31′6.312”N, 8°56′15.864”W, 17 May 2019, leg. J. Doumbia & K. Camara; ZMB 90056 (field #: G009), adult female, Liberia, Gangra, 7°33′47.556”N, 8°38′15.144”W, 14 January 2009, leg. J. Doumbia, K. Camara & A. T. Johnson; ZMB 90057 (field #: Wata 2_0201), adult female, Guinea, Wataférédou, 8°39′15.192”N, 8°51′11.196”W, 8 May 2019, leg. J. Doumbia & K. Camara; ZMB 90059 (field #: G003), subadult female, Liberia, Gangra, 7°32′40.992”N, 8°37′35.58”W, 12 January 2009, leg. J. Doumbia, K. Camara & A. T. Johnson; ZMB 90060 (field #: G009), adult male, Liberia, Gangra, 7°33′47.556”N, 8°38′15.144”W, 14 January 2009, leg. J. Doumbia, K. Camara & A. T. Johnson; ZMB 90062 (field #: Obay _0548), adult male, Liberia, Obayanmai, 8°8′59.82”N, 9°52′49.764”W, 7 April 2019, leg. J. Doumbia, K. Camara, F. Gbêmou & F. B. Beyan; ZMB 90063 (field #: Obay _0611), adult male, Liberia, Obayanmai, 8°8′56.04”N, 9°52′26.436”W, 9 April 2019, leg. J. Doumbia, K. Camara, F. Gbêmou & F.B. Beyan; ZMB 90159 (field #: PG.L.15/040), adult female, near Bouaflé, Ivory Coast, 7°0′15.1488”N, 5°34′37.0344”W, 7 May 2015, leg. J. Penner & N.L. Gonwouo; ZMB 90161 (field #: PG.L.13 033), subadult female, Liberia, Dugbe, 5°5′15”N, 8°36′29.88”W, 26 August 2013, leg. J. Penner & N.L. Gonwouo; ZMB 90162 (field #: PG.L.13 109), adult male, Liberia, Dugbe, 5°8′40.92”N, 8°29′57.12”W, 3 September 2013, leg. J. Penner & N.L. Gonwouo; ZMB 90163 (field #: LI10 014), adult male, Liberia, Tokadeh, 7°26′40.704”N, 8°39′27.972”W, 11 November 2010, leg. J. Penner; ZMB 90164 (field #: PG.L.13 054), adult female, Liberia, Dugbe, 5°4′20.28”N, 8°33′23.76”W, 28 August 2013, leg. J. Penner & N.L. Gonwouo; ZMB 90165 (field #: LI10 121), adult male, Liberia Tokadeh, 7°27′52.164”N, 8°39′53.928”W, 27 November 2010, leg. J. Penner; ZMB 90166 (field #: LI10 114), adult male, Liberia, Eastern Nimba Nature Reserve, 7°29′3.264”N, 8°34′36.264”W, 26 November 2010, leg. J. Penner; ZMB 90167 (field #: LI10 136), adult male, Liberia, Geipa, 7°28′59.196”N, 8°32′13.344”W, 3 December 2010, leg. J. Penner; ZMB 90168 (field #: PG.L.13 188, adult male, Liberia, Dugbe, 5°5′45”N, 8°29′32.64”W, 13 September 2013, leg. J. Penner & N.L. Gonwouo; ZMB 90169 (field #: LI10 120, adult female, Liberia, Eastern Nimba Nature Reserve, 7°29′3.264”N, 8°34′36.264”W, 28 November 2010, leg. J. Penner. Diagnosis. Amnirana parva sp. nov. differs from other West African Amnirana genetically, morphologically, and acoustically. The uncorrected pairwise-divergence in 16S ribosomal RNA between the new species and its closest congener, A. fonensis, is 5.8%. The new species can be distinguished from A. fonensis by several morphological traits in male specimens. SVL of male A. parva sp. nov. (41.9–50.4 mm) is smaller than in A. fonensis (48.7–65.3 mm). Area of humeral gland is smaller in A. parva sp. nov. (3.7–14.8 mm 2) than in A. fonensis (13.5–26.4 mm 2) as well. Males of A. occidentalis and A. galamensis reach larger SVL (67 and up to 80 mm, respectively; females of these species reach 96 and 86 mm, respectively), have differing color pattern and completely smooth dorsal skin (R̂del & Bangoura 2004; Channing & R̂del 2019). Most males of A. parva sp. nov. have dark spots on the dorsum. Breeding males of A. fonensis differ by a yellow dorsal coloration. The ranges of two Central African Amnirana species that are morphologically similar to A. parva sp. nov., A. albolabris (comprising various lineages, see Jongsma 2018) and A. asperrima (Perret, 1977), do not overlap with the new species. Nominotypical A. albolabris from Central Africa show a larger body size in males (Perret 1977 reports 44–57 mm SVL for A. albolabris from Cameroon). Amnirana asperrima can also be distinguished by both its larger adult body size (SVL of males: 46.7–60.1 mm) and the texture of its dorsal skin, which is densely covered with tiny spines. Other Central African Amnirana species show distinct morphological traits and cannot be confused with Amnirana parva sp. nov.: A. amnicola (Perret, 1977) adults are larger (SVL of males and females: 53–72 mm) and have a more pointed snout; and A. lepus (Andersson, 1903) adults are much larger (SVL: 70–100 mm), have finely granular skin, fully webbed toes, and much longer hind limbs. Last, the new species also differs from populations previously referred to A. longipes (Perret, 1960) that are much larger (SVL 77–87 mm) and have a more robust body and longer hind limbs. Although we follow Jongsma (2018) in recognizing A. longipes as a junior synonym of A. albolabris, we include it here in anticipation that this may be recognized as a valid taxon if one or several lineages of Central African A. albolabris are recognized as distinct species. Description of the holotype. Figs. 5–6; measurements in mm. Adult male with long and slender body, slightly dorsoventrally flattened; snout–vent length 42.9; snout acuminate with rounded tip in dorsal view, narrowly rounded in lateral view; head length 15.8, approximately 27% of SVL; head width 13.6; eye–snout distance 6.7; nostrils directed laterally; nostril closer to the snout (1.7) than to eye (4.7); eye diameter 6.1, slightly larger than tympanum diameter (5.2); interorbital distance 4.3; approximately one third of eye projecting above dorsal margin of head in lateral view; canthus rostralis distinct and sharply protruding; loreal region concave; anterocranial part of upper arm with large protruding gland (3.6 long, 2.0 wide, surface: 7.2 mm 2); long and slender fingers, tips broader than subarticular tubercles, forming discs, reaching 130–200% of finger width, inner metacarapal tubercle large and elongated, outer metacarpal tubercle smaller and more rounded; manual webbing absent; finger formula IIVariation. Measurements (in mm) of all individuals are summarized in Table 5. Overall, the male paratypes were similar to the holotype in external appearance and coloration. Male SVL ranged from 41.9–50.4 (mean ± sd: 45.9 ± 2.9, N= 13). The humeral gland area spanned between 3.7–14.8 mm 2 (9.4 ± 2.7 mm 2, N= 13). Female SVL ranged from 53.6–75.6 (63.9 ± 7.8, N= 10) thus markedly exceeding male SVL (Fig. 7). Seven of 13 showed slightly more extended (1-1), and a single male paratype showed slightly less extended webbing (2-2) of the fourth toe. In female paratypes seven of 10 individuals exhibited a webbing formula with slightly more extended webbing on toe IV(1-1). The dorsal skin was covered with small tubercles in all individuals, but females had fewer, smaller, and less spiny tubercles. In some individuals, the skin appeared almost smooth at first sight, but tiny granules were always present upon detailed inspection. The dorsolateral ridge was interrupted in the majority of specimens: females (six of 10) and males (nine of 13). Females showed a brighter brown dorsum and few to no dark spots (see Figs. 6b, 7b, 8e, g, 11c). On the dorsum of males, the ground color varied from chocolate-brown to dark brown and all males had at least some dark spots. The ventral coloration varied greatly among males. Among the paratypes, some had a plain white throat (e.g., ZMB 88549, 87112), similar to the holotype, whereas others exhibited dusty brown speckling covering almost the entire throat (e.g., ZMB 87113). The belly of all males were white to cream, and some individuals had irregular brown spots (e.g., ZMB 87110) that were densest near the legs, throat, or flanks, but were absent at the center of the venter. Females showed some color variation on the venter. In ZMB 87114, the belly and throat were white, whereas ZMB 79253 and ZMB 79197 had a cream-colored belly and a dusty brown speckled throat. Concerning life coloration (Figs. 8, 11), the dorsum exhibited various shades of brown, sometimes with darker spots (especially in males, most females without darker spots). The flanks in males were greenish yellow (Fig. 8f, h) to chocolate brown dorsally (Fig. 8b), usually turning lighter ventrally (but see Fig. 8a). The flanks in females were olive green, turning paler ventrally (Figs. 8c, e, g, 11c), and becoming almost cream-colored towards the belly (then often with some greenish or brownish mottling). The tympanum was brown in both sexes. Males had a narrow green line surrounding the eyes. In both sexes the iris was golden, sometimes with a reddish tinge, with the lower half of the iris being darker. The limbs and dorsum were a similar shade of brown in both sexes. The limbs sometimes were somewhat lighter, but with darker patterning. The number of conspicuous dark transverse bars on the hind limbs varied among individuals (e.g. compare Figs. 8b & 8h). The venter was beige or cream. Both sexes had a light cream or white upper lip, and rarely the anterior part of the lip was brownish (Fig. 8h). Juveniles showed a more contrasting pattern compared to adults. It consisted of a greenish brown dorsum with black spots and an almost black loreal region and flank. The ventral part of the flank and the belly were white with black dots. The juveniles had more conspicuous dark cross bands on the hind limbs and blackish dots on the forelegs; a bright white upper lip, and a bright red iris (uppermost part golden) (Fig. 11f). Acoustics. The advertisement call of Amnirana parva sp. nov. consists of a whiny-voiced croaking sound with ascending frequency modulation from 932 to 1092 Hz (mean ± sd: 1030 ± 86, N= 3). The sound consists of up to five harmonics of which the second-lowest is the dominant one in the first half, the lowest one in the second half (Fig. 4a). The call duration ranges from 0.26– 0.78 s (0.52 ± 0.26, N= 3). Dominant frequency at the beginning of the call ranges from 948–1034 Hz (1004 ± 49, N= 3). All call characteristics are summarized in Table 4. A call description by Schiøtz (1964c) from central Sierra Leone seems to refer to our new species. In that paper, Schiøtz describes three different sections of the call (such a clear distinction of sections was not detected by us). Call duration and complexity, however, seem to increase with the motivation of the frogs (MOR pers. obs.). Schiøtz (1964c) also mentions that these calls were different from those he recorded in Cameroon for A. albolabris, though he did not provide further details. Distribution. Confirmed A. parva sp. nov. records so far are known from western Sierra Leone to southerncentral Ghana, all within the belt of the (former) rainforest zone, bordering the southern humid Guinean savanna (Fig. 9). So far, no confirmed genetic records exist for Ivory Coast. However, morphologically we could assign frogs from the Taï National Park, the Azagny National Park, the Banco National Park, the Haute Dodo, Cavally and Tanoé forest reserves, as well as the Ivorian parts of Mount Nimba to the new species. Other formerly published records of A. albolabris (see literature cited below) from this country from habitats that include primary lowland evergreen and humid but semi-deciduous forest cannot be unambiguously assigned to either A parva sp. nov. or A. fonensis (see below). Life history. It is difficult in many cases to assign previous field observations and literature records to A. parva sp. nov. or A. fonensis. We summarize observations based on confirmed records (morphology and genetics of vouchers, and/or acoustics) as well as records, which originate from primary humid-evergreen rainforest. The latter decision was based on the results of the environmental niche modelling (see below). All observed and collected individuals of A. parva sp. nov. were encountered in closed primary or slightly degraded lowland rainforest, on the banks of or near to small to medium-sized streams (Fig. 10). In Banco National Park, southeastern Ivory Coast, the species was recorded in the forest, forest patches dominated by bamboos, and in man-made ponds of a fish-farm in a forest clearing (Fig. 11a, b). Although we often heard small choruses, Published as part of Griesbaum, Frederic, Jongsma, Gregory F. M., Penner, Johannes, Kouamé, N'Goran Germain, Doumbia, Joseph, Gonwouo, Nono L., Hillers, Annika, Glos, Julian, Blackburn, David C. & Rödel, Mark-Oliver, 2023, The smallest of its kind: Description of a new cryptic Amnirana species (Amphibia, Anura, Ranidae) from West African rainforests, pp. 301-339 in Zootaxa 5254 (3) on pages 309-319, DOI: 10.11646/zootaxa.5254.3.1, http://zenodo.org/record/7727536, {"references":["R ˆ del, M. - O., Bangoura, M. A. & B ˆ hme, W. (2004) The amphibians of south-eastern Republic of Guinea (Amphibia: Gymnophiona, Anura). Herpetozoa, 17, 99 - 118.","Jongsma, G. F., Barej, M. F., Barratt, C. D., Burger, M., Conradie, W., Ernst, R., Greenbaum, E., Hirschfeld, M., Leache, A. D., Penner, J., Portik, D. M., Zassi-Boulou, A. - G., R ˆ del, M. - O. & Blackburn, D. C. (2018) Diversity and biogeography of frogs in the genus Amnirana (Anura: Ranidae) across sub-Saharan Africa. Molecular Phylogenetics and Evolution, 120, 274 - 285. https: // doi. org / 10.1016 / j. ympev. 2017.12.006","Perret, J. - L. (1977) Les Hylarana (Amphibiens, Ranides) du Cameroun. Revue suisse de Zoologie, 84, 841 - 868. https: // doi. org / 10.5962 / bhl. part. 91429","Schiotz, A. (1964 c) The voices of some West African amphibians. Videnskabelige Meddelelser fra dansk Naturhistorik Forening, 127, 35 - 83.","Lamotte, M., Lauwarier, G. & Perret, J. - L. (1957) Contribution a l'etude des batraciens de l'Ouest Africain V. - Le developpement larvaires de Rana (Hylarana) albolabris. Bulletin de l'Institut fondamental d'Afrique Noire, Ser. A, 19, 1312 - 1327.","Guibe, J. & Lamotte, M. (1958) La reserve naturelle integrale du Mont Nimba. XII. Batraciens (sauf Arthroleptis, Phrynobatrachus et Hyperolius). Memoires de l'Institut Francais d'Afrique Noire, 53, 241 - 273.","Channing, A., R ˆ del, M. - O. & Channing J. (2012) Tadpoles of Africa. The biology and identification of all known tadpoles in sub-Saharan Africa. Edition Chimaira, Frankfurt am Main, Germany, 402 pp.","McIntyre, P. (1999) Hylarana albolabris, predation. Herpetological Review, 30, 223."]}

Redescription of A. fonensis (Rödel & Bangoura, 2004) Figs. 12–14 Holotype. SMNS 11788, adult male, Guinea, Simandou Range, northwest flank of Pic de Fon, 8°36′N, 8°51′W; 6 November 2002, leg. M.A. Bangoura. Paratypes. SMNS 11790, adult male, Guinea, Simandou Range, village situated to the northwest of Pic de Fon, 3 November 2002, leg. M.A. Bangoura; ZFMK 64481–64482, adult males, Guinea, Park National du Haut Niger, November 1996 – February 1997, leg. G. Nikolaus. Referenced specimens. Six males, one female; ZMB 79185 (field #: TI 60, GenBank #: 16S: MG552336), adult female, Sierra Leone, Tingi Hills, 8°52′2.1″N, 10°47′32.1612″W, 4 June 2007, leg. J. Johnny & A. Hillers; ZMB 79203 & 84733 (field #: SI 032–033, GenBank #: 16S: MG552334 & MG552340), two adult males, Guinea, Korombadou, 9°16′49.62″N, 9°6′ 52.0812″W, 5 October 2008, leg. J. Doumbia & A. Hillers; ZMB 79208 (field #: TI 020, GenBank #: 16S: MG552335), adult male, Sierra Leone, Tingi Hills, 8°53′25.44″N, 10°47′24.7812″W, 1 June 2007, leg. J. Johnny & A. Hillers; ZMB 90058 (field #: N044, 09-609), adult male, Guinea, N’Zérékore, 7°35′33.3″N, 8°28′5.4″W, 24 October 2009, leg. J. Doumbia, L. Sandberger, K. Camara & F. Gbêmou; ZMB 90158 (field #: LI10 088), adult male, Liberia, Tokadeh, 7°26′40.704″N, 8°39′27.972″W; 22 November 2010, leg. J. Penner & N.L. Gonwouo; ZMB 90160 (field #: PG.L. 15/041), adult male, Ivory Coast, 7°0′15.1488″N, 5°34′37.0344″W, 7 May 2015, leg. J. Penner & N.L. Gonwouo. For diagnostic differences to A. parva sp. nov., please see the diagnosis of the new species above. Description (based on the type series and seven additional specimens; measurements in mm, see Table 5). Medium-sized, slender frog, SVL of males range from 48.7 to 65.3 (mean ± sd: 57.2 ± 5.2, N= 10); a female measured 68.0; head slightly longer than broad, approximately one third of SVL; snout acuminate with rounded tip in dorsal view, narrowly rounded in lateral view; tympanum diameter around 85% of eye diameter, in two type specimens (SMNS 11788 & 11790) tympanum slightly larger than eye; inter-orbital distance slightly smaller than eye diameter; nostril closer to snout than to eye; canthus rostralis distinct and rounded; loreal region concave; dorsolateral ridge broad and protruding, only two male individuals show slight discontinuities in the posterior third of the ridge; males with large humeral glands on anterocraniad part of upper arms; area of humeral gland spans between 13.5–26.4 mm 2 (20.5 ± 4.86, N= 10); fingers long and thin, without webbing; tips rounded, forming small discs; finger formula: II– d), dark spots on dorsum and bands on hind limbs visible but paler; limbs and yellowish brown; flanks yellow, lighter towards cream belly; lip and throat bright yellow; dorsolateral ridge yellow to reddish brown; tympanum of breeding males with yellow center bordered by pale brown ring; tympanum of females dark or reddish brown (Fig. 13e, f). Acoustics. The advertisement call of male A. fonensis consists of a whiny-voiced croaking sound with an ascending frequency modulation of 548–723 Hz (mean ± sd: 633 ± 87, N= 3). The sound comprises up to five harmonics of which the lowest is the dominant one (see Fig. 4b) and lasts 0.55– 0.92 s (0.7 ± 0.19, N = 3). All call characteristics are summarized in Table 4. Distribution. Amnirana fonensis was described from the Pic de Fon, Simandou Range, and from the Haut Niger National Park, both located in Guinea. Our new records add further localities, including the first records from eastern Sierra Leone, the Nimba Mountains in northern Liberia, and central Ivory Coast. However, we cannot at this point clarify which previous West African records of Amnirana albolabris refer to either A. fonensis or A. parva sp. nov. (but see below). Life history. The observed and collected individuals were found near forest streams. These forests were less humid than in confirmed A. parva sp. nov. records and comprise semi-deciduous and degraded forest fragments (Fig. 14d) surrounded by savanna. Amnirana fonensis males differ from breeding A. parva sp. nov. males, by a bright yellow breeding coloration. Similar yellow breeding color has been reported from many, unrelated frogs across the tropics. In West Africa such color change has been observed in Phrynobatrachus alleni Parker, 1936 (R̂del 2003; Kanga 2021), Sclerophrys maculata (Hallowell, 1854) (Hillers & R̂del 2007) and S. togoensis (Ahl, 1924) (R̂del & Bangoura 2004). Amnirana fonensis breeds in water bodies of different sizes but seems to prefer stagnant or slow-flowing water. In a savanna-like habitat (Foma), males called from inundated patches, well concealed between grasses and reeds. In degraded gallery forests at Wataférédou, males called while floating on the water surface (Fig. 14a), well concealed between roots of trees and shrubs, or between low vegetation at the water edge. One to two males initiated calling and many other males then joined the chorus. Amnirana fonensis called at night as well as during the day. For instance, between 20:35 GMT and 23:00 GMT, we encountered many calling males (N> 100); the peak of calling activity was between 20:40 GMT and 21:30 GMT. Likewise between 14:45 and 16:00 GMT, we observed numerous calling males (N> 70); the peak of calling activity was at 14:57 GMT. In Foma, females attached eggs to grasses emerging from flowing water. After a few hours these eggs became submerged. In Wataférédou, the eggs were deposited between roots of trees and shrubs in the forest streams. The dark brown and yellow eggs were deposited in large clumps (Fig. 14b, c). We counted 809, 1074, and 2017 eggs in three clutches. The egg diameter ranged from 0.9‒2.4 mm (mean ± sd: 2.1 ± 0.42 mm mm; N= 100). We observed aquatic heteropterans (family Nepidae) preying on the eggs. It is likely that the tadpoles described by Lamotte (1957) from the Simandou range, can actually be assigned to A. fonensis. At Wataférédou, where A. fonensis was breeding, the forest stream was regularly polluted by people washing laundry. The edges of this gallery forest were degraded by subsistence farming, and cattle, and cleared for charcoal production and fuelwood., Published as part of Griesbaum, Frederic, Jongsma, Gregory F. M., Penner, Johannes, Kouamé, N'Goran Germain, Doumbia, Joseph, Gonwouo, Nono L., Hillers, Annika, Glos, Julian, Blackburn, David C. & Rödel, Mark-Oliver, 2023, The smallest of its kind: Description of a new cryptic Amnirana species (Amphibia, Anura, Ranidae) from West African rainforests, pp. 301-339 in Zootaxa 5254 (3) on pages 319-322, DOI: 10.11646/zootaxa.5254.3.1, http://zenodo.org/record/7727536, {"references":["Rodel, M. - O. & Bangoura, M. A. (2004) A conservation assessment of amphibians in the Foret Classee du Pic de Fon, Simandou Range, southeastern Republic of Guinea, with the description of a new Amnirana species (Amphibia Anura Ranidae). Tropical Zoology, 17, 201 - 232. https: // doi. org / 10.1080 / 03946975.2004.10531206","R ˆ del, M. - O. (2003) The amphibians of Mont Sangbe National Park, Ivory Coast. Salamandra, 39, 91 - 110.","Kanga, K. P., Kouame, N. G., Zogbasse, P., Gongomin, B. A. - I., Agoh, K. L., Kouame, A. M., Konan, J. C. B. Y. N., Adepo-Gourene, A. B., Gourene, G. & R ˆ del, M. - O. (2021) Amphibian diversity of a West African biodiversity hotspot: an assessment and commented checklist of the batrachofauna of the Ivorian part of the Nimba Mountains. Amphibian and Reptile Conservation, 15 (1) [General Section]: 71 - 107 (e 275).","R ˆ del, M. - O., Bangoura, M. A. & B ˆ hme, W. (2004) The amphibians of south-eastern Republic of Guinea (Amphibia: Gymnophiona, Anura). Herpetozoa, 17, 99 - 118.","Lamotte, M., Lauwarier, G. & Perret, J. - L. (1957) Contribution a l'etude des batraciens de l'Ouest Africain V. - Le developpement larvaires de Rana (Hylarana) albolabris. Bulletin de l'Institut fondamental d'Afrique Noire, Ser. A, 19, 1312 - 1327."]}

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

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

Phrynobatrachus afiabirago sp. nov. Figs 2–3 Holotype. UWBM 0 5925, field number: ADL 3999), a male from Atewa Range, Ghana, 6.24246°N, - 0.5571°E (date = WGS84), 769 m asl, 26 May 2011, coll. C. Ofori-Boateng & A.D. Leaché. Paratypes (15). UWBM 9042–9046; ZMB 86155– 86159 (males) and UWBM 9041, UWBM 5924, ZMB 86154 (females) collected from north-eastern (6°14'32.856"N, 0°33'25.56"W) and north-western part of the Atewa Range (6°7'40.7634"N, 0°37'44.04"W) respectively, April 2016, coll. C. Ofori-Boateng & A.D. Leaché; ZMB 86160 (male), Atewa range, Atiwiredu, 06°12’22.7”N, 0°34’39.2’’W, 817 m asl, 7 June 2006, coll. N.G. Kouamé & C. Ofori-Boateng; ZMB 73708 (male) and ZMB 73709 (female), Jukwa forest, south-western Ghana, 5°14’44.13’’N, 1°22’42.31’’W, coll. C. Ofori-Boateng & A. Hillers. Diagnosis. The molecular results place the new species within Phrynobatrachus. Morphologically this genus is distinguished from the similar looking and often syntopic Arthroleptis and juvenile Ptychadena by the presence of a tarsal tubercle (absent in Arthroleptis and Ptychadena); presence or absence of webbing (always absent in Arthroleptis); lack of a black spot in the tympanal region (present in Arthroleptis); lack of a median dorsal skin raphe (present in Arthroleptis); and the lack of parallel dorsal ridges (present in Ptychadena). The new species is a small frog ( The new species differs genetically from other West African Phrynobatrachus by 5–19% in the investigated part of the 16S gene (see below). Superficially the new species resembles P. latifrons, but can be told apart by the more pronounced pedal webbing and the yellow throat of breeding males in the latter species. Genetically and morphologically closest to the new species are three other West African species: P. gutturosus, P. fraterculus and P. maculiventris. P. afiabirago sp. nov. males of the new species share with P. maculiventris a throat, that is either completely covered with blackish spots or completely dusted in black (Figs. 2, 3, 4F); in P. gutturosus the black coloration of the throat in males is typically restricted to the anterior throat and does not extend onto the chest (Fig. 4B; Chabanaud 1921; Rödel 2000). The throat of P. fraterculus males is dirty yellowish grey (Fig. 4D). The belly of P. afiabirago sp. nov. always shows some dark patterning, ranging from very few small black dots to being densely beset with large dark blotches (Fig. 3). The belly of adult P. gutturosus is usually white or only exhibiting very few minute black dots (Fig. 4B); in P. fraterculus the belly is greyish white with few larger and smaller black dots along the edges and in the pectoral region (Fig. 4D); the belly of P. maculiventris is densely covered with huge black blotches, separated by white lines (Fig. 4F). Furthermore P. fraterculus and P. maculiventris differ from the new species by having a smooth or slightly granular back skin (Figs. 4C, E), in contrast to the presence of many distinct warts in P. afiabirago sp. nov. (Fig. 2). The most distinctive feature, differentiating P. afiabirago sp. nov. from P. gutturosus, P. fraterculus and P. maculiventris, is the lack of femoral glands in males of the new species, present in males of the other three species. From other West African Phrynobatrachus species P. afiabirago sp. nov. differs morphologically (for genetic differences see below) by small size (SVL versus adults equal or larger than 25 mm in: P. intermedius, P. liberiensis, P. plicatus, P. natalensis); absence of a black lateral face mask (black face mask present in P. intermedius, P. plicatus); absence of a spiny eyelid tubercle (present in P. annulatus, P. calcaratus, P. pintoi, P. taiensis, P. villiersi); presence of distinct black lateral bands on flanks (shared with most P. latifrons and some other, but smaller species); throat of males with black markings, but not entirely black (males with yellow throat: P. alleni, P. latifrons; uniform black or grey: P. liberiensis, P. plicatus, P. calcaratus, P. pintoi, P. villiersi, P. guineensis, P. phyllophilus, P. ghanensis, P. francisci, P. natalensis, P. tokba; white or greyish: P. annulatus; clear with darker stippling and without obvious vocal sac: P. brongersmai); males with disc-like vocal sac skin extending to anterior chest (only shared with P. gutturosus, P. fraterculus, P. maculiventris, P. pintoi); most individuals with larger black spots on white belly (shared with P. taiensis, P. ghanensis, P. pintoi, P. annulatus— at least some spots round with white centre, P. villiersi— black blotches on clear blue belly; some other Phrynobatrachus may have few small black spots); rough warty back skin (back skin smooth in P. fraterculus, P. maculiventris, P. hieroglyphicus; sometimes smooth in breeding P. latifrons, P. tokba); males without femoral glands (femoral glands present in males of P. gutturosus, P. fraterculus, P. maculiventris, P. taiensis, P. phyllophilus); no longer or Xshaped scapular or dorsal ridges (scapular ridges long and converging towards mid-body, almost X-shaped: P. alleni, P. plicatus); distinct but rudimentary webbing (webbing more developed in P. alleni, P. latifrons, P. francisci, P. natalensis, P. intermedius, P. liberiensis, P. plicatus, P. rainerguentheri). Holotype description (UWBM 5925, all measures in mm; Fig. 2 A –C). Typical, small, adult male Phrynobatrachus with short, slender-oval body shape; snout-vent length: 18.4 mm; short snout, rounded in dorsal and rounded to slightly protruding in lateral view; canthus rostralis rounded; loreal region slightly concave; headwidth directly behind the eyes: 7.1; eye-diameter: 2.0; distance eye-nostril: 1.7; distance eye-snout tip: 2.2; nostril closer to snout than to eye; tympanum present but indistinct, tympanum diameter: ~ 0.9, nearly half the size of eye diameter; femur: 8.2, shorter than tibio-fibulare: 10.9; foot including longest toe: 14.8; hand with large oval palmar and thenar tubercles; fingers with small roundish subarticular tubercles, no additional tubercles on hands; finger tips very slightly expanded without forming discs; relative finger length: III>I Dorsal skin slightly warty, two converging—in almost x-shape—pairs of scapular warts, followed by three pairs of smaller oval to roundish black warts along posterior back; pair of very small warts on posterior part of snout, edges of back, flanks and upper surfaces of hind limbs with smaller round warts on granular skin; eyelids faintly granular without a spiny tubercle; ventral skin smooth. Colour in life. Overall coloration of dorsum light to dark bronze brown with numerous black spots scattered particularly along edges of dorsal warts; light vertebral line extending from dark inter-orbital band to almost vent; anterior canthal region black; a conspicuous broad black band extends from behind eye across tympanum region along flanks to groin; narrow area between this band and back light brown; narrow area from tympanum to groin and between the lateral band and belly light brown; dorsal parts of hind limbs brownish with one broad and 2–3 narrow black cross-bars on either thighs and lower legs; anterior legs light brown with very faint black markings; brown upper lip with black bars, two white spots below anterior and posterior edge of eye; lower mandible almost completely black with few minute white spots; posterior part of thighs with light longitudinal lines; throat with black blotches on dark grey background; pectoral region grey; belly white with larger and smaller black spots and blotches, posterior part of belly slightly lighter than anterior one; groin area, posterior-most part of belly and lower part of thighs yellowish orange with few black spots; antero-ventral part of thighs with black longitudinal line. Throat covered by disc like vocal sac skin, slightly extending to pectoral region; throat skin slightly granular without spines; dorsal part of first finger with light nuptial pad; femoral glands absent. Colour in preservation: Pattern as in life but faded; black may turn into darker brown; yellowish-orange colour turns into beige or white. Paratype variation (measures in mm, summarized in Tab. 1; see Figs. 2D, 3). Body shape and dimensions and colouration, mostly as in holotype; male snout-vent length ranges from 17.2–20.3, females from 21.8–23.4 mm; canthal region with or without black line; back colour varies from bronze brown to greyish brown, median yellowish to orange vertebral line always present, broadest at mid of back, edges often indistinctly delimitated; two to three black, broad cross bars on thighs and lower legs; anterior part of thighs often with clear longitudinal band; narrow to broader yellowish line turning down from dorsal hind leg insertion to middle of posterior part of thighs, from there as longitudinal line with dark borders to knee, rarely interrupted; male throats range from entirely black to densely covered with black blotches on dark grey ground, tip sometimes with clear spot, sometimes longitudinal folds parallel to lower mandible; female throats smooth, from almost dark brown to black within irregular white median line and few white spots to very light with several dark blotches; dark lower lips of females partly interrupted by clearer spots; pectoral region, belly and lower side of hind legs from densely best with larger or smaller dark brown to black botches to almost white with few dark spots; some individuals with a fine light longitudinal line on posterior part of thighs and from mid lower leg to heel; females with slightly more expanded finger tips, compared to males; webbing may vary slightly, web between toe 4 and 5 often extending along 1.5 to 2 phalanges. In ZMB 73708 (male) and ZMB 73709 (female) from Jukwa, in life the yellowish-orange colouration of groin and ventral thighs was replaced by lemon green; posterior surfaces of thighs with lemon-green longitudinal bands; ventral spots on belly restricted to edges leaving 90% of belly clear white; in preservation colouration much faded, specimens partly shrunken due to preservation. Genetics. The 16S alignment contained 538 bp and 111 variable sites (75 parsimony-informative). There is 100% bootstrap support for the monophyly of P. afiabirago sp. nov. (Fig. 5). Phrynobatrachus afiabirago sp. nov. together with two undescribed taxa, P. aff. gutturosus (bootstrap = 84% and 83%), and P. maculiventris (bootstrap = 100%) formed a highly supported clade (bootstrap = 96%). The genetic diversity within P. afiabirago sp. nov. samples was low (average pairwise distance = 0.0016; F81 substitution model). The alignment for P. afiabirago sp. nov. comprised 533 bp with 3 variable sites (1 parsimony-informative). We found four unique haplotypes among the nine samples, but there was no geographic structure in the distribution of these haplotypes. For example, ZMB 73708 from Jukwa in western Ghana shared the same haplotype as UWBM 9043 from the Atewa Range in southeastern Ghana. We finally compared the 16S sequence of the new species to all 20 West African Phrynobatrachus species for which genetic data have been published (Table 2, average pairwise distance: mean + sd = 13.03 + 3.65%, range = 5.00–18.63%). Concerning described taxa, P. maculiventris (5.00% difference) and P. fraterculus (6.45%) were the genetically most similar taxa. Habitat and natural history. Phrynobatrachus afiabirago sp. nov. always occurred around large forest ponds and forest swamps (Fig. 6). The frogs were usually collected from around the stilt-like root system of trees along the bank of swamps, including palms. The altitudinal range of the species extended from 300–800 m asl in the Atewa Range, but they occurred as low as 135–250 m in the south-western Jukwa forest. The Atewa Range, type locality, is located in south-eastern Ghana, west of Lake Volta (Eastern Region, see Fig. 1). It is one of the highest upland forests in Ghana, characterized by an interconnecting series of hills that runs from north to south (Swaine & Hall 1977). This unique area comprises mountain forest like vegetation, not occurring anywhere else in West Africa (McCullough et al. 2007). Distribution. Phrynobatrachus afiabirago sp. nov. is so far known from only two localities in the Ghanaian forest zone; the Atewa Range in south-eastern Ghana and the small (0.5 ha) Jukwa forest, situated near the Kakum National Park in south-western Ghana (Fig. 1). We assume that it will occur in other forests of the area as well. Etymology. The specific epithet honours Afia Birago, the mother of the first author for overcoming all odds as a widow, woman and a single parent to provide the highest level of education to all her eight children. Her tenacity and love for nature is a source of inspiration to the first author’s professional career. The specific epithet is treated as a noun in apposition. We suggest Afia Birago’s Puddle Frog as common name. Phrynobatrachus alleni Parker, 1936 ZMB 73747 / GU457535 0.15141 Phrynobatrachus annulatus Perret, 1966 ZMB 73746 / GU457538 0.10581 Phrynobatrachus calcaratus (Peters, 1863) MVZ 245140 / EU075282 0.10177 Phrynobatrachus dispar (Peters, 1870) CAS 219386 / EU075278 0.08685 Phrynobatrachus francisci Boulenger, 1912 MVZ 249545 / GU457548 0.14781 Phrynobatrachus fraterculus (Chabanaud, 1921) ZMB 73761 / GU457551 0.06405 Phrynobatrachus ghanensis Schiøtz, 1964 ZMB 70721 / GU457553 0.16203 Phrynobatrachus guineensis Guibé & Lamotte, 1962 MOR T13 / GU457555 0.15837 Phrynobatrachus gutturosus (Chabanaud, 1921) MOR C11 / GU457556 0.11696 Phrynobatrachus intermedius Rödel, Boateng, Penner & Hillers, 2009 ZMB 71539 / GU457557 0.14615 Phrynobatrachus latifrons Ahl, 1924 ZMB 73753 / GU457559 0.18633 Phrynobatrachus liberiensis Barbour & Loveridge, 1927 ZMB 73768 / GU457562 0.15001 Phrynobatrachus maculiventris Guibé & Lamotte, 1958 ZMB 71592 / GU457563 0.05002 Phrynobatrachus natalensis (Smith, 1849) ZMB 73717 / GU457566 0.16584 Phrynobatrachus phyllophilus Rödel & Ernst, 2002 ZMB 73763 / GU457570 0.15328 Phrynobatrachus pintoi Hillers, Zimkus & Rödel, 2008 ZMB 76883 / JN813916 0.09848 Phrynobatrachus plicatus (Günther, 1858) ZMB 73742 / GU457575 0.15683 Phrynobatrachus rainerguentheri Rödel, Onadeko, Barej & Sandberger, 2012 ZMB 77742 / JQ954865 0.15515 Phrynobatrachus tokba (Chabanaud, 1921) ZMB 73774 / GU457585 0.14155 Phrynobatrachus villiersi Guibé, 1959 ZMB 73740 / GU457589 0.10777

Arthroleptis formosus sp. nov. Figs 1–3 Holotype. ZMB 75480 (field #: KD073, GenBank #: JN 408725; female, 18.6 mm), western Guinea, Préfecture Télimélé, N 10.8549167, W - 13.7417222, 286 m a.s.l., 9 October 2010, degraded gallery forest, small creak present, surrounding areas with corn, peanuts and rice fields, coll. N.G. Kouamé & J. Doumbia. Paratypes. ZMB 75481 (field #: KD038, GenBank #: JN 408726; male, 19.1 mm), ZMB 75482 (field#: KD049, GenBank #: JN 408727; juvenile, 15.7 mm), N 10.8083611, W - 13.8375833, 182 m a.s.l., 2 October 2010, tree and grass savanna, other data as holotype; ZMB 75483 (field #: KD048, male, 18.7 mm), N 10.8131667, W - 13.8350278, 359 m a.s.l., 3 October 2010, degraded gallery forest, river with waterfalls and rapids, other data as holotype; ZMB 75484 (field #: KD074, GenBank #: JN 408728; female, 20.8 mm), ZMB 75485 (field #: KD075, male, 18.3 mm), data as holotype; ZMB 75486 (field #: KD 132, GenBank #: JN 408730; juvenile, 13.2 mm), ZMB 75487 (field #: KD 133, GenBank #: JN 408729; juvenile, 11.6 mm), N 11.0883333, W - 13.6670278, 330 m a.s.l., 19 October 2010, tree and grass savanna, other data as holotype. Diagnosis. The presence of a fine vertebral skin raphe, lack of webbing and tarsal tubercles (Rödel 2000; Zimkus & Blackburn 2008), as well as molecular phylogenetic relationships (not shown), characterize the new species as a member of the genus Arthroleptis. From other Arthroleptis species it differs by small size and a unique colour pattern. From the most similar congener, A. aureoli, the species differs genetically by 4.5–5.1 % in the investigated part of the 16 S rRNA gene, smaller size, more enlarged toe and finger tips, the lack of hypertrophied third fingers and digital spines in males, and ventral colouration. The combination of small size and its unique colouration distinguishes it from all other African Arthroleptis. Genetically the new species differs from other African Arthroleptis by a minimum of 16.0%. Description of the holotype (measures in mm). An adult female Arthroleptis with a slender almost parallel body shape; snout-vent length: 18.6; short snout, roundish in dorsal and pointed in lateral view; canthus rostralis indistinct and rounded; loreal region straight; head-width directly behind the eyes: 5.6; eye large, approximately equal to distance eye-nostril, eye-diameter: 1.9; pupil round; distance eye-nostril: 1.8; distance nostril-snout tip: 1.0, nostril much closer to snout than to eye; tympanum small but distinct, tympanum-diameter: 1.2, smaller than diameter of eye; tongue slender, about twice as long as wide, slightly diverging towards tip, distinctly notched, no lingual papillae; no vomerine teeth; long and slender extremities; femur: 8.6, distinctly shorter than tibio-fibula: 10.1; foot including toe IV: 13.9; hand with small, ovoid palmar and thenar tubercles; fingers with round to ovoid subarticular tubercles, additional tubercles on bases of fingers I–III (Fig. 2 a); fingers without spines; relative finger length: I Basic dorsal colour pattern: red with large black spots and patches; a black spot in the centre of the snout; a black irregularly bordered interorbital line; followed posteriorly by a black U open towards snout tip; from scapular region towards urostyle a black long, goblet shaped blotch; large black inguinal blotches; further smaller roundish black spots, in particular along dorsolateral areas of dorsum; two transversal black bars on thighs and two to three black bars on lower legs; side of head, temporal region and region dorsal from forearm bases black; upper arms reddish; lower arms with very broad transversal black bars; hands and feet blackish with small white spots; many very small yellow-whitish spots within the reddish coloured dorsal and lateral surfaces; upper and lower mandible almost uniform black with a few very small white dots; iris dark golden with many small black points; throat, chest and belly, a swell as ventral parts of upper arms and lower legs dark grey; throat and chest with small white spots. Coloration in preservation: red colours turn into light brown; the yellowish small points turn white; large black spots and blotches persist; ventral colour persist with the exception of dark grey ventral parts of legs, which turn lighter than in life. Variation. The paratypes very closely resemble the holotype in most characters. Adult males may be slightly smaller (18.3–19.1 mm; N= 3) than adult females (18.6–20.8 mm, N= 2), have narrower heads and shorter tibiofibulae (Table 1). However, the sample sizes are too small to test the significance of this. Whereas both adult females had slightly lighter bellies and throats with comparatively many white spots (Fig. 1 e), males have dark brown almost blackish ventral surfaces with no or only very few white spots (Fig. 1 f). Females may have more and lighter reddish parts on dorsal surfaces than males (Fig. 1 a, b, d). In males the flanks were almost uniform black and the many small yellowish-white dorsal points of the females have a more bluish-greenish tinge; especially the light spots on the flanks seem to be mostly blue. From the pictures of live animals, males have slightly more granular skin on the dorsum than females. This is not visible in preserved specimens. Measurements and indices of measurements of adult specimens of the new species are summarized in Tab. 1. Genetics. When aligning the sequences of the new species to the data provided by Blackburn (2008 b), and further sequences generated for this study (Table 2) A. formosus sp. nov. clustered together with other species, currently placed within Arthroleptis (tree not shown). Based on the 402 investigated base pairs of the 16 S rRNA gene A. formosus sp. nov. was genetically most similar to A. aureoli. From this species the new species differed by a mean 4.7 % sequence divergence (Table 2). The genetic distance to other African Arthroleptis ranged from 16.0– 23.4 % (Table 2). continued. ZMB # sex SVL/HW SVL/FL SVL/TL SVL/FTL FL/TL LSF/ LTF 75480 f 3.3 2.2 1.8 1.3 0.9 0.6 75484 f 3.3 2.4 1.9 1.5 0.8 0.6 75481 m 3.4 2.4 2.1 1.4 0.9 0.6 75483 m 3.6 2.3 2.1 1.5 0.9 0.6 75485 m 3.5 2.3 2.2 1.5 1.0 0.6 nov. (for GenBank accession numbers see type series) and the sequences of 20 other African arthrolpetid species. Given are mean and range of sequence divergence. species GenBank accession numbers distance mean range Arthroleptis adelphus FJ 151092 (Cameroon) 0.181 0.176–0.184 A. aureoli EF 640989, EF 640990, FJ 151117 (Sierra Leone), JN 408731 0.047 0.045–0.051 (Guinea) A. poecilonotus FJ 151119 (Sierra Leone), FJ 151111 (Ghana), HM 238192 0.196 0.190–0.202 (Nigeria) Natural history. Between 2 nd to 19 th October 2010, which is the end of the rainy season, we found nine individuals (ZMB 75480 -75487, 1 specimen NGK) of the new species at four sites in the Telimélé region. All sites (see type series for exact geographic data) consisted of lowland habitats (180–360 m a.s.l.) which were strongly altered by human activities, i.e. timber extraction and agriculture (Fig. 3). Two sites were gallery forests with small rivers; the two others comprised humid grass and tree savanna. Five individuals were collected in grasses on rocky ground, one was recorded on a large rock in a disturbed gallery forest and three individuals were observed on dead wood in a gallery forest which was highly impacted by logging activities (type locality; Fig. 3 c & d). All savanna habitats were close to forest, indicating that A. formosus sp. nov. may migrate between the two habitat types. Similar to other frogs in West Africa (Arthroleptis spp., Phrynobatrachus gutturosus, P. tokba, Rödel 2000, 2003, own unpubl. data), this species may live in humid savanna during the rainy season and in nearby forests during the dry season. All individuals were encountered during twilight or at night, while being active as well while being hidden. The female holotype had a large, but completely empty ovarium. The largest female (ZMB 75484) carried only very small, unripe egg. It thus seems that, during the period of our survey, the reproductive period was already finished. This is in line with our failure to detect any calling males of the new species. ZMB 75483, however, had very well-developed testis and fat bodies. Distribution. Arthroleptis formosus sp. nov. is known from a total of four sites (Fig. 4) in the Télimélé region in western Guinea. Its occurrence in further forests and forested savannas within this region is likely. So far it is not known from any protected area. Etymology. The species name is a Latin adjective meaning “beautiful” and refers to the colour pattern of the new species. As English name we propose “Beautiful Squeaker”.