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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."]}

[EN] The identity of the dominant root-associated microbial symbionts in a forest determines the ability of trees to access limiting nutrients from atmospheric or soil pools, sequester carbon and withstand the effects of climate change. Characterizing the global distribution of these symbioses and identifying the factors that control this distribution are thus integral to understanding the present and future functioning of forest ecosystems. Here we generate a spatially explicit global map of the symbiotic status of forests, using a database of over 1.1 million forest inventory plots that collectively contain over 28,000 tree species. Our analyses indicate that climate variables—in particular, climatically controlled variation in the rate of decomposition—are the primary drivers of the global distribution of major symbioses. We estimate that ectomycorrhizal trees, which represent only 2% of all plant species, constitute approximately 60% of tree stems on Earth. Ectomycorrhizal symbiosis dominates forests in which seasonally cold and dry climates inhibit decomposition, and is the predominant form of symbiosis at high latitudes and elevation. By contrast, arbuscular mycorrhizal trees dominate in aseasonal, warm tropical forests, and occur with ectomycorrhizal trees in temperate biomes in which seasonally warm-and-wet climates enhance decomposition. Continental transitions between forests dominated by ectomycorrhizal or arbuscular mycorrhizal trees occur relatively abruptly along climate-driven decomposition gradients; these transitions are probably caused by positive feedback effects between plants and microorganisms. Symbiotic nitrogen fixers—which are insensitive to climatic controls on decomposition (compared with mycorrhizal fungi)—are most abundant in arid biomes with alkaline soils and high maximum temperatures. The climatically driven global symbiosis gradient that we document provides a spatially explicit quantitative understanding of microbial symbioses at the global scale, and demonstrates the critical role of microbial mutualisms in shaping the distribution of plant species., This work was made possible by the Global Forest Biodiversity Database, which represents the work of over 200 independent investigators and their public and private funding agencies (see Supplementary Acknowledgements), Supplementary information is available for this paper at https://doi.org/10.1038/s41586-019-1128-0.

In this Letter, the middle initial of author G. J. Nabuurs was omitted, and he should have been associated with an additional affiliation: ‘Forest Ecology and Forest Management Group, Wageningen University and Research, Wageningen, The Netherlands’ (now added as affiliation 182). In addition, the following two statements have been added to the Supplementary Acknowledgements. (1): ‘We would particularly like to thank The French NFI for the work of the many field teams and engineers, who have made extraordinary efforts to make forest inventory data publicly available.’ (1): ‘Sergio de Miguel benefited from a Serra- Húnter Fellowship provided by the Generalitat of Catalonia.’ Finally, the second sentence of the Methods section should have cited the French NFI, which provided a national forestry database used in our analysis, to read as follows: ‘The GFBi database consists of individual-based data that we compiled from all the regional and national GFBi forest-inventory datasets, including the French NFI (IGN—French National Forest Inventory, raw data, annual campaigns 2005 and following, https://inventaire-forestier.ign.fr/spip.php?rubrique159, site accessed on 01 January 2015)’. All of these errors have been corrected online. © 2019, The Author(s), under exclusive licence to Springer Nature Limited.

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

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