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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."]}

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 II Dorsal coloration in life (Fig. 6a) medium brown with few darker spots; flanks greenish yellow, becoming paler ventrally with some darker mottling; arms slightly lighter brown than dorsum, marbled with fine, irregular, dark pattern; hind limbs slightly lighter brown than dorsum with narrow transverse dark bars; posterior parts of thighs marbled; foot webbing greyish brown; throat and belly white; ventral side of legs pale beige; hind limbs ventrally patterned with indistinct brown spots; bright, white upper lip extending posterior to angle of mouth, below and posterior to tympanum. After four years in preservation (75% ethanol; Fig. 5), in general less saturated colors: brownish dorsal color darkened; greater contrast of marbled limb patterning; flanks lacking yellow coloration, now beige with small remnants of brownish speckles; white throat and lip more shiny. Variation. 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