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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."]}

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

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

Toxicodryas blandingii (Hallowell, ���1844��� 1845) (Table 1, Figs. 5���8) Dipsas Blandingii: Hallowell (���1844��� 1845:170); type locality: ��� Liberia, West Africa. ��� Triglyphodon fuscum: Dum��ril, Bibron & Dum��ril (1854:1101); type locality: ��� Grand-Bassam, sur la C��te d���Ivoire (Guin��e)��� [Ivory Coast]. Dipsas fasciata: Fischer (1856:84); type locality: ��� Peki (West-Afrika)��� [Ghana]. Dipsas valida: Fischer (1856:87); type locality: ��� Edina (Grand Bassa County, West-Afrika)��� [Liberia]. Dipsas globiceps: Fischer (1856:89); type locality: ��� Edina (Grand Bassa County, Liberia, West-Afrika).��� Toxicodryas Blandingii: Hallowell (1857:60); comb. nov. Dipsas Fischeri: Jan in Dum��ril (1859:212); no type locality provided. Triglyphodon fuscum var. obscurum: Dum��ril (1861:211); type locality: ��� C��te d���Or ��� [Ghana]. Dipsas regalis: Jan (1871:3, Livraison 38, pl. vi, fig. 2) in Jan & Sordelli (1870 ���1881); type locality: ��� C��te d���Or ��� [Ghana]. Dipsas globiceps var. tumboensis: M��ller (1885:688); type locality: ��� Tumbo-Insel ��� [Guinea]. Boiga blandingi occidentalis: Stucki-Stirn (1979:377); inferred type locality: ���Besongabang��� Cameroon. Boiga blandingi subfulva: Stucki-Stirn (1979:381); type locality: none provided, but limited to Cameroon according to book title. Toxicodryas blandingii was originally described by Hallowell (1845:170) based on a single specimen collected by his friend, ���Dr. Blanding,��� in Liberia. The dorsum and venter of the specimen was noted to have a ���light yellow��� color with a series of blotches of ���leaden colour.��� This specimen reportedly possessed 2 preoculars, 2 postoculars, 272 ventrals, 131 subcaudals, body length (i.e., SVL) of 1.22 meters, and tail length of 0.39 meters. Hallowell (1854) provided additional details of the specimen���s teeth, noted it had 17 ���rows of scales,��� and corrected the tail length to 0.37 meters. Hughes & Barry (1969), Wallach et al. (2014) and Uetz et al. (2019) stated that the type was lost, which is consistent with Malnate (1971), who did not list a type specimen from the ANSP collection. Wallach et al. (2014) noted the type was a 1.67 m specimen, slightly longer than the total length of 1.61 m reported by Hallowell (1845) in the original description, but the longer measurement is likely a typographical error in reference to the latter citation (V. Wallach, pers. comm.). A query by EG to the Philadelphia Academy of Sciences in spring 2020 resulted in location of the type specimen (ANSP 10083, Fig. 6), and a redescription of this specimen is provided below. According to Loveridge (1957:269), the name Dipsas Fischeri was proposed by Jan (in Dum��ril 1859) to combine the minor color pattern variants Dipsas fasciata, D. valida, and D. globiceps named by Fischer (1856). Pel (1852:171) coined the name Dipsas regalis, and as translated by Savage & McDiarmid (2017:73), Pel stated, ���the third species of venomous snake, Naja atropos, belongs to cobras (spectacled snakes) and reaches a length of 6 to 7 feet [1.8���2.1 m]. Its color is entirely black... As this snake in general shows much similarity to a tree snake, Dipsas regalis, which equals it in color and size, but is not venomous.��� Perhaps because of this poor description, Boulenger (1896:78) attributed the latter name to Jan & Sordelli (1870 ���1881), who provided an illustration that served as an appropriate description. Jan listed the name in the Index des Planches for Livraison 38 as Dipsas cynodon Cuv. vari��t��? (D. regalis Schlegel), but according to Savage & McDiarmid (2017:73), the attribution to Hermann Schlegel is in error because he never used the name in any publication. M��ller (1885:687) seemed to suggest that Jan illustrated his specimen (collected by Dr. M��hly from ���Goldk��ste��� [i.e., Gold Coast or modern-day Ghana]) from Basel, but Hughes & Barry (1969:1020) listed a personal communication from M.S. Hoogmoed, who noted the type of D. regalis (specimen ���Leiden 958���) was collected by Pel in February 1844 from Accra, Ghana. Hallowell (1857:60) coined the genus Toxicodryas because he noticed that his specimen of T. blandingii had a ���single channelled posterior tooth on each side... and therefore...[it] cannot belong to the genus Triglophodon [sic] of Dum. and Bibron, which has three.��� Subsequent herpetological publications in the 19 th and early 20 th centuries seemingly ignored Hallowell���s new genus and continued to recognize the taxon in either the genus Dipsas (e.g., Mocquard 1896) or more commonly, Dipsadomorphus (e.g., Boulenger 1896, 1919; de Witte 1933). Schmidt (1923) transferred the taxon to the genus Boiga in his opus on Congolese snakes, recognizing B. (Toxicodryas) blandingii and B. (Toxicodryas) pulverulenta, an action that was followed by most subsequent authors for decades. Based on the placement of African Boiga in the ��� Dipsadidae: Lycodontinae ��� by Underwood (1967), Welch (1982) seems to have been the first to recognize the genus Toxicodryas for both species of the genus, an action followed by Meirte (1992) and observed by most herpetologists in the 21 st century (e.g., Uetz et al. 2020). Boulenger (1896) included all of the above West African, 19 th- century names in the synonymy of Dipsas (Toxicodryas) blandingii. Because the dubious subspecies described by Stucki-Stirn (1979) both occur in Cameroon, where one molecular sample (CAS 253611) from Allen et al. (in press) is recovered in a well-supported clade with West African samples (Fig. 2), we confirm the taxonomic nomenclature of Wallach et al. (2014) in treating these taxa as synonyms of T. blandingii. Marques et al. (2018) noted that northwestern Angolan records from ���Chinchoxo,��� ���Piri-Dembos,��� and ���Quirimbo��� by Peters (1877), Bocage (1895), Parker (1936), and Hellmich (1957a, b) are attributable to T. pulverulenta. However, the morphometric data provided by Hellmich (1957b) for Angolan snakes are inconsistent with the size and scale rows of the latter species, and herein, we consider his records to be attributable to T. vexator sp. nov. Diagnosis. Toxicodryas blandingii, as recognized herein, is restricted to West Africa and west-central Africa (west of the confluence of the Congo and Ubangi rivers), defined by the following combination of characters: maximum SVL> 1 meter (vs. maximum SVL T. pulverulenta and T. adamanteus sp. nov.); DSRN 23���25 (vs. 19���21 in T. pulverulenta and 18���23 in T. adamanteus sp. nov.); DSRM 21���25 (vs. 19���21 in T. pulverulenta and 18���21 in T. adamanteus sp. nov.); cloacal plate usually divided (vs. divided or undivided in T. vexator sp. nov., and always undivided in T. pulverulenta and T. adamanteus sp. nov.); adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters (vs. both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots in T. pulverulenta and T. adamanteus sp. nov.); hemipenis relatively short and massive (i.e., broad), proximal third covered with spines, distal two-thirds dimpled with a flattened apex (vs. relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, and with a domed apex in T. pulverulenta and T. adamanteus sp. nov.); venom toxicity LD 50 = 2.85���3.55 mg /kg in mice (vs. venom toxicity LD 50 = 4.88 mg /kg in mice for T. vexator sp. nov.). Redescription of the holotype. ANSP 10083 (Fig. 6) adult female in poor condition, 1330 mm SVL; head triangular and distinct from neck, 1.92% of SVL (25.5 mm); right loreal missing, left loreal partially obscured by supralabials due to cranial damage, upper side tapering superiorly; body triangular; tail moderately long (400 mm; 30.1% of SVL). Supralabials ���/9, ���/4 th, 5 th, and 6 th contacting orbit; infralabials 14/13, 1 st on each side in contact behind mental, 1 st ���4 th /1 st ���4 th contacting anterior chin shields and 4 th ���7 th /4 th ���7 th contacting posterior chin shields; 2 preoculars; 3 postoculars (on left, missing on right); temporals ���/2 + 2; 2 internasals; nasal divided (on left, missing on right); frontal slightly longer than wide, only left side undamaged; dorsal scale rows 23 one head length posterior to jaw rictus, 23 at midbody, 17 one head length anterior to vent, smooth and oblique, vertebral scales broad and apically flattened; ventrals 273 (standard), 271 (Dowling); cloacal plate undivided; paired subcaudals 132. Coloration (in preservative). After approximately 176 years in preservation, specimen is faded, with creamy tan background color in dorsal and ventral views. Brown markings on posterior edge of supralabials and dorsum of head. Irregular brown and dark brown blotches and saddles on dorsum from neck to tip of tail (Fig. 6). Variation. Morphometric variation of Toxicodryas blandingii is shown in Table 1. M��ller (1885:688) provided data for a snake from Ghana with 15 infralabials and noted that most of its scales have two ���Endgruben��� [terminal pits], which likely refer to apical pits. In his description of Dipsas globiceps var. tumboensis M��ller (1885) noted his specimen from Guinea (Fig. 7) had 147 subcaudals. In snakes from West Africa (without separating by sex), Angel (1933) noted a temporal formula of 2 + 2 or 2 + 3, 21���25 scales at midbody, 240���289 ventrals, 120���147 subcaudals, either an undivided or divided cloacal plate, and a maximum total size of 2290 mm, and nearly verbatim variation was listed by Villiers (1950a), Doucet (1963), Stucki-Stirn (1979), Chippaux (2006), and Trape & Man�� (2006). However, in snakes from Ghana, Swiecicki (1965:302) noted a maximum total length of 2450 mm for a ���black form��� individual, Gauduin (1970) listed the maximum total length as 2700 mm (600 mm tail) for Cameroon, Chirio & LeBreton (2007) provided a slightly larger total length of 2740 mm for Cameroon, and Luiselli et al. (1998a) noted a maximum size of 2800 mm, presumably for Nigeria. Villiers (1951) noted an unsexed individual from Benin with 115 subcaudals. Cansdale (1965) documented 21���25 scales at midbody, and Segniagbeto et al. (2011) documented snakes from Togo with 19���24 scales at midbody and 102���159 subcaudals. In our examined specimens, temporal formula includes the variation noted by Angel (1933), but is more extensive (2 +5, 3 + 4, 3 + 3, 1 + 5, 3 + 2, or 2 + 4), and either supralabials 3���5 or 4���6 contact the eye, which is consistent with the observations of Angel (1933), Villiers (1950b), and Chippaux (2006); the latter author also noted that sometimes only 2 supralabials contact the eye. Rasmussen (1997a) noted specimens with the 4th���5th, 5th���7 th, or 4th���7 th supralabials in contact with the eye, and in general, this species has sloping and smooth scales with apical pits, and the vertebral row is greatly enlarged. The holotype, one male and one female from Liberia (Loveridge 1941; Johnsen 1962), one male from Gabon (Pauwels et al. 2002b), one male from DRC (RBINS 10888), and a juvenile from Cameroon (Werner 1897) are unusual in having an undivided cloacal plate, because all other examined specimens have a divided cloacal plate, including the type specimens of Dipsas fasciata, D. valida, and D. globiceps (Fischer 1856). Rasmussen (1997a) remarked that his specimens have either a divided or undivided cloacal plate. In his book on West and Central African snakes, including countries west of the Congo River, Chippaux (2006:154) noted the anal [cloacal plate] is sometimes entire, but more often divided. Trape & Man�� (2006) stated that the cloacal plate is almost always divided. Segniagbeto et al. (2011) noted individuals with divided or undivided cloacal plates in Togo. Combined descriptions by Fischer (1856) of the teeth of Dipsas fasciata, D. valida, and D. globiceps (all now synonyms of T. blandingii) suggest the species has 9 maxillary teeth that increase in size posteriorly, followed by two fangs (three on the right side in one specimen), and 12 mandibular teeth, which decrease in size posteriorly. In snakes from Ghana, Leeson (1950) noted 10���11 maxillary teeth, becoming larger posteriorly, and two fangs followed by a shorter fang; fourteen large palatine and pterygoid teeth, and 15 mandibular teeth (anterior ones largest). Taylor & Weyer (1958:1217) described a Liberian specimen with 9���10 maxillary teeth (on different sides) that increase in size from the 1 st to 4 th tooth, and then become subequal; two fangs occur after this series of teeth, and after a short diastema, there is a third fang with only traces of a groove. A second Liberian specimen had 10 maxillary teeth followed by three fangs, the last of which had only ���a suggestion of a groove.��� Based on specimens ranging from Guinea to Congo, Rasmussen (1997a:98) noted 10 maxillary teeth followed by three enlarged, furrowed venomous teeth, with the 3 rd fang slightly smaller than the previous two. Fischer (1856) provided detailed descriptions of the color patterns of West African Dipsas fasciata, D. valida, and D. globiceps (all now synonyms of T. blandingii), which seem to suggest he examined a subadult male that had been kept in alcohol for a long (unspecified amount) time (D. fasciata), an adult female (D. valida), and a subadult that retained juvenile coloration (D. globiceps). In his description of Dipsas globiceps var. tumboensis M��ller (1885:689) noted his specimen from Guinea (Fig. 7) had a gray-reddish dorsum with 30 black transverse bands, usually containing milky white spots. The frontal, supraoculars, and occipital scales had large black spots, the labial scales and postoculars were edged with black, and the head shields had multiple milk-white speckles. The tail was bright red with dark, irregular transverse bands. Aspects of this coloration description are highly unusual for this species (e.g., bright red tail), and more typical of T. pulverulenta, but the black edging of the labial scales, number of preoculars (3), supralabials in contact with the eye (5 and 6), ventrals (269), and subcaudals (147) clearly indicate this taxon is a synonym of T. blandingii (Fig. 7, Tables 1���2). Mocquard (1887:80) described a recently collected, unsexed subadult (���la longueur du tronc��� [trunk length] 1.1 m) from Gabon as having a dorsal color of a general tint of Burgundy with slightly darker spots on the flanks that have a dirty white spot a little above their lower edge. Mertens (1938) described an adult male from Cameroon as solid black dorsally and ventrally, with the exception of the anterior third of the venter, which was white, but the ventral scales had dark gray edges. The labial scales were gray with vertical black borders. Villiers (1950b) described the color pattern of an unsexed individual from Ivory Coast as sooty black or brownish in places on the dorsum; underside iridescent dark gray posteriorly, becoming whitish anteriorly, with the posterior edge of the ventrals edged with gray; underside of head white, and labials whitish and edged in black. Another unsexed individual from Liberia was described as bluish black above, yellow below; supralabials yellow with black edges, and posterior part of venter and underside of tail black. A third unsexed individual from Liberia had identical coloration to the latter one, except for the presence of whitish bars on the neck. Leeson (1950) noted that snakes from Ghana have a dull green or gray dorsum. Monard (1951:162) described an unsexed individual from Cameroon as a beautiful light redbrown, ���barr����� [barred] with dark brown. Taylor & Weyer (1958:1217) described a Liberian brown-phase female with pale grayish green on the ventral side of the head and neck, merging into gray with ���a greenish cast��� 12.7 cm posteriorly, and at 40.6 cm behind the head, it transitioned into plain tan to the tip of the tail. Isemonger (1962:12) remarked that this species has a ���delicate bloom on the skin.��� Cansdale (1965:43) described a highly unusual color pattern for juveniles by noting that ���the young brown form is pink with irregular chocolate markings that break up its outline very effectively and make it difficult to pick out in a tree or shrub.��� Leston & Hughes (1968:753) described an unusual specimen from Ghana as ���pale grey with darker greyish-green transverse bands, the bands irregular but more or less diamond shaped on each side. The ventrals are also grey but more glossy.��� Groves (1973:107) described the coloration of hatchlings from a captive Liberian female as ���light grey background colour with pinkish undertones; black, roughly oval, lateral blotches narrowing as they approach the midline, where many of them fail to conjoin; top of head light grey; belly dark grey.��� Rasmussen (1997a:98) noted the scales of his specimens were dull and almost dusty, a sentiment also shared by Cansdale (1965). Adult males were solid black on the dorsum and yellow on the venter (becoming black posteriorly), whereas adult females were noted to be gray, brown or yellow-brown on the dorsum and yellow-brown on the venter, sometimes without transverse bands. Hughes (2000:8) noted juvenile and subadult (approximately 1 meter in total length or less) snakes had a dorsal coloration that was ���a distinctively bright and contrasting pattern of chocolate brown blotches.��� He noted that most male specimens lose this coloration as they age, becoming increasingly melanistic, and although exceptions are possible, this melanistic progression does not seem to occur in females. Chippaux (2006:154) noted there are two dorsal color morphs: (1) uniform black or dark blue with ���reflets velout��s��� [velvety reflections] or (2) gray with darker, poorly defined transverse spots. The venter was noted as dull yellow to charcoal gray, and juvenile coloration as light brown with darker transverse ring-shaped spots. Stucki-Stirn (1979), perhaps confused by the tw, Published as part of Greenbaum, Eli, Allen, Kaitlin E., Vaughan, Eugene R., Pauwels, Olivier S. G., Wallach, Van, Kusamba, Chifundera, Muninga, Wandege M., Aris- Tote, Mwenebatu M., Mali, Franck M. M., Badjedjea, Gabriel, Penner, Johannes, R��del, Mark-Oliver, Rivera, Jacqueline, Sterkhova, Viktoria, Johnson, Grant, Tapondjou, Walter P. & Brown, Rafe M., 2021, Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata Colubridae) with descriptions of two new cryptic species from Central Africa, pp. 1-44 in Zootaxa 4965 (1) on pages 7-17, DOI: 10.11646/zootaxa.4965.1.1, http://zenodo.org/record/4723024, {"references": ["Fischer, J. G. (1856) Neue Schlangen des Hamburgischen Naturhistorischen Museums. Abhandlungen aus dem Gebiete der Naturwissenschaften herausgegeben von Naturwissenschaftlichen Verein in Hamburg, 3, 79 - 116 + 3 pl.", "Hallowell, E. (1857) Notice of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of the Academy of Natural Sciences of Philadelphia, 9 (1857 - 1858), 48 - 72.", "Dumeril, A. H. A. (1859) Reptiles et poissons de l'Afrique occidentale. Etude precedee de considerations generales sur leur distribution geographique. Archives du Museum National d'Histoire Naturelle, 10, 137 - 240.", "Jan, G. & Sordelli, F. (1870 - 1881) Iconographie Generale des Ophidiens. Tome Troisieme (Livrais. 35 a 50), Contenant Quatre Vingt Seize Planches. Georges Jan and Ferdinand Sordelli, Milan, 96 pls.", "Muller, F. (1885) Vierter nachtrag zum Katalog der herpetologischen Sammlung der Basler Museums. Verhandlungen der Naturforschenden Gesellschaft in Basel (1887 - 1890), 7, 668 - 717. https: // doi. org / 10.5962 / bhl. part. 2388", "Stucki-Stirn, M. C. (1979) A Comparative Study of the Herpetological Fauna of the former West Comeroon [sic] / Africa: With a Classification and Synopsis of 95 Different Snakes and Description of Some New Sub-species. Snake Report 721. Herpeto- Verlag, Teuffenthal, Switzerland, 650 pp.", "Hallowell, E. (1854) Remarks on the geographical distribution of reptiles, with descriptions of several species supposed to be new, and corrections of former papers. Proceedings of the Academy of Natural Sciences of Philadelphia, 7 (1854 - 1855), 98 - 105.", "Hughes, B. & Barry, D. H. (1969) The snakes of Ghana: a checklist and key. Bulletin de l'Institut fondamental d'Afrique noire. Serie A, Sciences naturelles, 31 (3), 1004 - 1041.", "Wallach, V., Williams, K. L. & Boundy, J. (2014) Snakes of the World: A Catalogue of Living and Extinct Species. CRC Press, Boca Raton, Florida, xxvii + 1209 pp.", "Uetz, P., Cherikh, S., Shea, G., Ineich, I., Campbell, P. D., Doronin, I. V., Rosado, J., Wynn, A., Tighe, K. A., McDiarmid, R., Lee, J. L., Kohler, G., Ellis, R., Doughty, P., Raxworthy, C. J., Scheinberg, L., Resetar, A., Sabaj, M., Schneider, G., Franzen, M., Glaw, F., Bohme, W., Schweiger, S., Gemel, R., Couper, P., Amey, A., Dondorp, E., Ofer, G., Meiri, S. & Wallach, V. (2019) A global catalog of primary reptile type specimens. Zootaxa, 4695 (5), 438 - 450. https: // doi. org / 10.11646 / zootaxa. 4695.5.2", "Malnate, E. V. (1971) A catalog of primary types in the herpetological collections of the Academy of Natural Sciences, Philadelphia (ANSP). Proceedings of the Academy of Natural Sciences of Philadelphia, 123, 345 - 375.", "Loveridge, A. (1957) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya, Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, Harvard, 117 (2), 151 - 362 + i - xxxvi (index).", "Pel, H. S. (1852) Over de jagt aan de Goudkust, volgens eene tienjarige eigene ondervinding. Nederlandsch Tijdschrift voor Liefhebbers der Jagt und Visscherei, 1, 149 - 173.", "Savage, J. M. & McDiarmid, R. W. (2017) The Herpetological Contributions of Giorgio Jan (1791 - 1866): With an Introduction, Annotated Bibliography, Synopsis of Herpetological Taxa, and a Comprehensive Guide to the Iconographie generale des Ophidiens. Society for the Study of Amphibians and Reptiles, Villanova, Pennsylvania, viii + 926 pp.", "Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History). Vol. III. Containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae, and Viperidae. Trustees of the British Museum (Natural History), London, xiv + 727 pp.", "Mocquard, F. (1896) Notes sur quelques ophidiens de Conakry. Bulletin du Museum Paris, 20, 59 - 60.", "Boulenger, G. A. (1919) Batraciens et reptiles recueillis par le Dr C. Christy au Congo Belge dans le districts de Stanleyville, Haut-Uele et Ituri en 1912 - 1914. Revue Zoologique Africaine, 7, 1 - 29. https: // doi. org / 10.5962 / bhl. part. 15108", "de Witte, G. - F. (1933) Reptiles recoltes au Congo Belge par le Dr H. Schouteden et par M. G. - F. de Witte. Annales du Musee du Congo Belge, C. - Zoologie, 3 (1), 55 - 98.", "Schmidt, K. P. (1923) Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909 - 1915. Part II. - Snakes. Bulletin of the American Museum of Natural History, 49, 1 - 146, pls. I - XXII.", "Underwood, G. (1967) A Contribution to the Classification of Snakes. British Museum (Natural History), London, x + 179 pp.", "Welch, K. R. G. (1982) Herpetology of Africa: A Checklist and Bibliography of the Orders Amphisbaenia, Sauria and Serpentes. Robert E. Krieger Publishing Company, Malabar, Florida, 293 pp.", "Meirte, D. (1992) Cles de determination des serpents d'Afrique. Museum Royal d'Afrique Centrale, Tervuren Belgique Annual Series Octavo Science Zoologique, 267, 1 - 152.", "Uetz, P., Freed, P. & Hosek, J. (Eds.) (2020) The Reptile Database. Available from: http: // www. reptile-database. org (accessed 12 December 2020)", "Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola: Atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement II), 1 - 501.", "Peters, W. C. H. (1877) Ubersicht der Amphibien aus Chinchoxo (Westafrika), welche von der Afrikanischen Gesellschaft dem Berliner zoologischen Museum ubergeben sind. Monatsberichte der Koniglichen Preussischen Akademie der Wissenschaften zu Berlin, 1877, 611 - 621.", "Bocage, J. V. B. (1895) Herpetologie d'Angola et du Congo. Ministerio da Marinha e das Colonias, Lisbonne, XX + 203 pp., pls. I - XIX.", "Parker, H. W. (1936) Dr. Karl Jordan's expedition to South-West Africa and Angola: Herpetological collection. Novitates Zoologicae, 40, 115 - 146.", "Hellmich, W. (1957 a) Herpetologische Ergebnisse einer Forschungreise in Angola. Veroffentlichungen der Zoologischen Staatssammlung Munchen, 5, 1 - 92.", "Hellmich, W. (1957 b) Die Reptilienausbeute der hamburgischen Angola-Expedition. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 55, 39 - 80.", "Angel, F. (1933) Les Serpents d'Afrique Occidentale Francaise. Larose Editeur, Paris, 246 pp.", "Villiers, A. (1950 a) Initiations Africaines. II. Les Serpents de l'Ouest Africain. Institut Francais d'Afrique Noire, Dakar, Senegal, 148 pp.", "Doucet, J. (1963) Les serpents de la Republique de Cote d'Ivoire. Serpents venimeux. Acta Tropica, 20 (3 - 4), 297 - 340.", "Chippaux, J-P. (2006) Les Serpents d'Afrique Occidentale et Centrale. 3 rd Edition. IRD Editions, Paris, 311 pp.", "Trape, J. - F. & Mane, Y. (2006) Guide des Serpents d'Afrique Occidentale: Savane et Desert. IRD editions, Paris, 226 pp. https: // doi. org / 10.4000 / books. irdeditions. 37382", "Swiecicki, A. W. (1965) Snakes and snake bite in the western region, Ghana. Journal of Tropical Medicine and Hygiene, 68 (12), 300 - 304.", "Gauduin, P. (1970) Contributions a la faune de la region de Yaounde. Annales de la Faculte des Sciences du Cameroun, 4, 73 - 99.", "Chirio, L. & LeBreton, M. (2007) Atlas des Reptiles du Cameroun. Publications Scientifiques du MNHN, IRD Editions, Paris, 686 pp.", "Luiselli, L., Akani, G. C. & Barieenee, I. F. (1998 a) Observations on habitat, reproduction and feeding of Boiga blandingi (Colubridae) in south-eastern Nigeria. Amphibia-Reptilia, 19 (4), 430 - 436. https: // doi. org / 10.1163 / 156853898 X 00098", "Villiers, A. (1951) Mission A. Villiers au Togo et au Dahomey (1950). II. - Ophidiens. Etudes Dahomeennes, 5, 17 - 43.", "Cansdale, G. S. (1965) West African Snakes. Longmans, Green and Co. Ltd., London, vi + 74 pp.", "Segniagbeto, G. H., Trape, J. F., David, P., Ohler, A., Dubois, A. & Glitho, I. A. (2011) The snake fauna of Togo: Systematics, distribution and biogeography, with remarks on selected taxonomic problems. Zoosystema, 33 (3), 325 - 360. https: // doi. org / 10.5252 / z 2011 n 3 a 4", "Villiers, A. (1950 b) Catalogues. IV. La Collection de Serpents de l'I. F. A. N. Institut Francais d'Afrique Noire, Dakar, Senegal, 155 pp.", "Rasmussen, J. B. (1997 a) Afrikanske slanger (10). Boiga blandingii. Nordisk Herpetologisk Forening, 40 (3), 97 - 103.", "Loveridge, A. (1941) Report on the Smithsonian-Firestone expedition's collection of reptiles and amphibians from Liberia. Proceedings of the United States National Museum, 91 (3128), 113 - 140. https: // doi. org / 10.5479 / si. 00963801.91 - 3128.113", "Johnsen, P. (1962) Notes on African snakes, mainly from northern Rhodesia and Liberia. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn, 124, 115 - 130.", "Pauwels, O. S. G., Kamdem Toham, A. & Chimsunchart, C. (2002 b) Recherches sur l'herpetofaune des Monts de Cristal, Gabon. Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Biologie, 72, 59 - 66.", "Werner, F. (1897) Ueber Reptilien und Batrachier aus Togoland, Kamerun und Tunis aus dem kgl. Museum fur Naturkunde in Berlin. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien, 47, 395 - 408. https: // doi. org / 10.5962 / bhl. part. 11464", "Leeson, F. (1950) Identification of Snakes of the Gold Coast. The Crown Agents for the Colonies, London, x + 142 pp.", "Taylor, E. H. & Weyer, D. (1958) Report on a collection of amphibians and reptiles from Harbel, Republic of Liberia. The University of Kansas Science Bulletin, 38 (14), 1191 - 1229. https: // doi. org / 10.5962 / bhl. part. 10973", "Mocquard, F. (1887) Sur les ophidiens repportes du Congo par la Mission de Brazza. Bulletin de la Societe Philomathique de Paris, Serie 7, 11, 62 - 92.", "Mertens, R. (1938) Herpetologische Ergebnisse einer Reise nach Kamerun. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 442, 1 - 52.", "Monard, A. (1951) Resultats de la mission zoologique suisse au Cameroun. Reptiles. Batraciens. Memoires de l'Institut Francais d'Afrique Noire, Sciences Naturelles, (1), 123 - 170.", "Isemonger, R. M. (1962) Snakes of Africa: Southern, Central and East. Thomas Nelson and Sons (Africa) (Party) Ltd., Johannesburg, viii + 236 pp.", "Leston, D. & Hughes, B. (1968) The snakes of Tafo, a forest cocoa-farm locality in Ghana. Bulletin de l'Institut Francaise d'Afrique Noire, 30 A, 737 - 770.", "Groves, F. (1973) Reproduction and venom in Blanding's tree snake. International Zoo Yearbook, 13, 106 - 108. https: // doi. org / 10.1111 / j. 1748 - 1090.1973. tb 02114. x", "Hughes, B. (2000) Polymorphism, sex, size and other aspects of the African snake, Boiga blandingi (Hallowell, 1844). Herpetological Bulletin, 74, 8 - 12.", "Lawson, D. (1993) The reptiles and amphibians of the Korup National Park Project, Cameroon. Herpetological Natural History, 1 (2), 27 - 90.", "Greenbaum, E. & Carr, J. L. (2005) The herpetofauna of Upper Niger National Park, Guinea, West Africa. Scientific Papers, Natural History Museum, The University of Kansas, 37, 1 - 21. https: // doi. org / 10.5962 / bhl. title. 8470", "Pauwels, O. S. G., Pauly, A., Araldi, A., Ndonda Makemba, R., Fonteyn, D., Oslisly, R. & Whittaker, A. M. (2020) Miscellanea Herpetologica Gabonica XVI. Bulletin of the Chicago Herpetological Society, 55 (5), 93 - 99.", "Sternfeld, R. (1909) Die Fauna der deutschen Kolonien, Herausgegeben mit Unterstutzung des Reichs-Kolonialamtes vom Zoologischen Museum in Berlin. Reihe I. Kamerun. Heft I. Die Schlangen Kameruns. R. Friedlander & Sohn, Berlin, iv + 28 pp.", "Leache, A. D., Grummer, J. A., Miller, M., Krishnan, S., Fujita, M. K., Bohme, W., Schmitz, A., Lebreton, M., Ineich, I., Chirio, L., Ofori-Boateng, C., Eniang, E. A., Greenbaum, E., Rodel, M. - O. & Wagner, P. (2017) Bayesian inference of species diffusion in the West African Agama agama species group (Reptilia, Agamidae). Systematics and Biodiversity, 15 (3), 192 - 203. https: // doi. org / 10.1080 / 14772000.2016.1238018", "Cansdale, G. (1955) Reptiles of West Africa. Penguin Books, London, United Kingdom, 104 pp.", "Dekeyser, P. L. (1955) Les Mammiferes de l'Afrique Noire Francaise. 2 nd Edition. Institut Francais d'Afrique Noire, Initiations Africaines, Dakar, Senegal, 426 pp.", "Villiers, A. (1955) Note sur quelques Ixodidae et Gamasidae parasites des Vertebres rencontres en Afrique Occidentale francaise. Bulletin de l'Institut fondamental d'Afrique noire, Serie A, 17 (2), 444 - 454.", "Wickler, V. W. & Uhrig, D. (1969) Verhalten und okologische Nische der Gelbflugelfledermaus, Lavia frons (Geoffroy) (Chiroptera, Megadermatidae). Zeitschrift fur Tierpsychologie, 26 (6), 726 - 736. https: // doi. org / 10.1111 / j. 1439 - 0310.1969. tb 01972. x", "Happold, M. & Happold, D. (Eds.) (2013) Mammals of Africa. Vol. IV. Hedgehogs, Shrews and Bats. Bloomsbury, London, New Delhi, New York, and Sydney, 800 pp.", "Villiers, A. (1956) La collection de serpents de l'I. F. A. N. (Acquisitions 1954 - 1955). Bulletin de l'Institut fondamental d'Afrique noire, Serie A, 18 (3), 878 - 883.", "Wuster, W., Chirio, L., Trape, J-F., Ineich, I., Jackson, K., Greenbaum, E., Kusamba, C., Barron, C., Nagy, Z., Storey, R., Hall, C., Wuster, C., Barlow, A. & Broadley, D. G. (2018) Integration of nuclear and mitochondrial gene sequences and morphology reveal unexpected diversity in the forest cobra (Naja melanoleuca) species complex in Central and West Africa (Serpentes: Elapidae). Zootaxa, 4455 (1), 68 - 98. https: // doi. org / 10.11646 / zootaxa. 4455.1.3", "Pitman, C. R. S. (1958) Snake and lizard predators of birds. Bulletin of the British Ornithologists' Club, 78, 82 - 86.", "Woodward, S. F. (1960) A bat-eating snake. The Nigerian Field, 25, 172 - 174.", "Menzies, J. I. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 44.", "Cozens, B. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 44.", "Pitman, C. R. S. (1962) More snake and lizard predators of birds. Bulletin of the British Ornithologists' Club, 82, 45 - 55.", "Barry, D. H. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 42 - 43.", "Dunger, G. T. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 43.", "Greene, H. W. (1989) Ecological, evolutionary, and conservation implications of feeding biology in Old World cat snakes, Genus Boiga (Colubridae). Proceedings of the California Academy of Sciences, 46 (8), 193 - 207.", "Jones, T. (1961) Notes on bat-eating snakes. The Nigerian Field, 26, 69 - 70.", "Menzies, J. I. (1966) The snakes of Sierra Leone. Copeia, 1966 (2), 169 - 179. https: // doi. org / 10.2307 / 1441123", "Leston, D. (1970) Some snakes from the forest zone of Ghana. British Journal of Herpetology, 4 (6), 141 - 144.", "Greene, H. W. (1997) Snakes: The Evolution of Mystery in Nature. University of California Press, Berkeley, Los Angeles and London, xiii + 351 pp.", "Luiselli, L., Akani, G. C. & Capizzi, D. (1998 b) Food resource partitioning of a community of snakes in a swamp rainforest of south-eastern Nigeria. Journal of Zoology, London, 246 (2), 125 - 133. https: // doi. org / 10.1111 / j. 1469 - 7998.1998. tb 00141. x", "Bohme, W. (2000) Diversity of a snake community in a Guinean rain forest (Reptilia, Serpentes). In: Rheinwald, G. (Ed.), Isolated Vertebrate Communities in the Tropics. Proceedings of the 4 th International Symposium in Bonn. Bonner zoologische Monographien, 46, 69 - 78.", "Akani, G. C., Ebere, N., Luiselli, L. & Eniang, E. A. (2007) Community structure and ecology of snakes in fields of oil palm trees (Elaeis guineensis) in the Niger Delta, southern Nigeria. African Journal of Ecology, 46 (4), 500 - 506. https: // doi. org / 10.1111 / j. 1365 - 2028.2007.00885. x", "Pauwels, O. S. G., Carlino, P., Chirio, L., Daversa, D. R., Lips, J., Oslisly, R. & Testa, O. (2019 a) Amphibians and reptiles found in caves in Gabon, western Equatorial Africa. Cave and Karst Science, 46 (1), 3 - 12.", "Butler, J. A. & Reid, J. C. (1990) Records of snakes from Nigeria. The Nigerian Field, 55 (1), 19 - 40.", "Pauwels, O. S. G., Carlino, P., Chirio, L., Meunier, Q., Vivien Okouyi Okouyi, J., Orbell, C., Rousseaux, D. & Testa, O. (2017 c) Miscellanea Herpetologica Gabonica IX. Bulletin of the Chicago Herpetological Society, 52 (6), 97 - 102.", "Young, R. A. (1983) Feeding behavior of captive Boiga. Bulletin of the Chicago Herpetological Society, 18 (3 - 4), 89 - 91.", "Rodel, M. - O. & Glos, J. (2019) Herpetological surveys in two proposed protected areas in Liberia, West Africa. Zoosystematics and Evolution, 95 (1), 15 - 35. https: // doi. org / 10.3897 / zse. 95.31726", "Pauwels, O. S. G. & Vogel, G. (2011). Ein Fall von Thanatose (Totstellreflex) bei Boiga blandingii (Squamata, Serpentes, Colubridae) in Gabun. Elaphe, 4, 24 - 25.", "Pasqual, J. R. H. (1962) Correspondence: West African snakes. The Nigerian Field, 27, 189 - 190.", "Knoepffler, L. - P. (1966) Faune du Gabon (amphibiens et reptiles). I. Ophidiens de l'Ogooue-Ivindo et du Woleu N'tem. Biologia Gabonica, 2 (1), 3 - 23.", "Roux-Esteve, R. (1969) Les serpents de la region de Lamto (Cote-d'Ivoire). Annales de l'Universite d'Abidjan, Ecologie, 2 (1), 81 - 140.", "Perret, J. - L. (1961) Etudes herpetologiques africaines III. Bulletin de la Societe de Sciences Naturelles de Neuchatel, 84, 133 - 138.", "Hughes, B. (1983) African snake faunas. Bonner zoologische Beitrage, 34 (1 - 3), 311 - 356.", "Mertens, R. (1941) Zur Kenntnis der Herpetofauna von Fernando-Poo. 3. Beitrag zu den wissenschaftlichen Ergebnissen der Forschungreise H. Eidmann nach Spanisch-Guinea 1939 / 40. Zoologischer Anzeiger, 135, 275 - 281.", "Blackwell, K. (1967) A preliminary survey of the reptiles of the Ibadan area. British Journal of Herpetology, 3, 307 - 311.", "Butler, J. A. & Reid, J. C. (1986) Habitat preferences of snakes in the southern Cross River State, Nigeria. In: Rocek, Z. (Ed.), Studies in Herpetology. Charles University, Prague, pp. 483 - 488.", "Hughes, B. (1988) Herpetology in Ghana (West Africa). British Herpetological Society Bulletin, (25), 29 - 37.", "Rodel, M. - O., Kouadio, K. & Mahsberg, D. (1999) Die Schlangenfauna des Comoe-Nationalparks, Elfenbeinkuste: Erganzungen und Ausblick. Salamandra, 35 (3), 165 - 180.", "Leache, A. D. (2005) Results of a herpetological survey in Ghana and a new country record. Herpetological Review, 36 (1), 16 - 19.", "Auliya, M., Wagner, P. & Bohme, W. (2012) The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist. Bonn zoological Bulletin, 61 (2), 255 - 281.", "Akaffou, M. H., Chippaux, J. - P., Kouadio Allali, B., Coulibaly, Z. & Dosso, M. (2017) Peuplement ophidien des plantations d' Hevea brasiliensis d'Anguededou (sud-est de la Cote d'Ivoire). Bulletin de la Societe Herpetologique de France, 162, 31 - 38.", "Pauwels, O. S. G., Albert, J. - L., Arrowood, H., Mvele, C., Casanova, M., Dodane, J. - B., Morgan, J., Primault, L., Thepenier, L. & Fenner, J. N. (2017 a) Miscellanea Herpetologica Gabonica X. Bulletin of the Chicago Herpetological Society, 52 (8), 133 - 138.", "Leache, A. D., Oaks, J. R., Ofori-Boateng, C. & Fujita, M. K. (2020) Comparative phylogeography of West African amphibians and reptiles. Evolution, 74 (4), 716 - 724. https: // doi. org / 10.1111 / evo. 13941", "Taub, A. M. (1967) Comparative histological studies on Duvernoy's gland of colubrid snakes. Bulletin of the American Museum of Natural History, 138 (1), 1 - 50.", "Levinson, S. R., Evans, M. H. & Groves, F. (1976) A neurotoxic component of the venom from Blanding's tree snake (Boiga blandingi). Toxicon, 14 (4), 307 - 312. https: // doi. org / 10.1016 / 0041 - 0101 (76) 90027 - 1", "Weinstein, S. A. & Kardong, K. V. (1994) Properties

Toxicodryas pulverulenta (Fischer, 1856) (Table 2, Figs. 8, 11) Dipsas pulverulenta: Fischer (1856:11); type locality: ��� Edina, Grand Bassa County in Liberia (West-Afrika).��� Dipsadomorphus Boueti: Chabanaud (1917a:373); type locality: none provided, but subsequently clarified to ��� Porto-Novo ��� ��� Dahomey ��� [Porto-Novo, Benin] by Chabanaud (1917b:12). As noted by Cer��aco et al. (2018), the description of this species was based on two specimens (only one syntype remains today according to Hallermann 1998 and Uetz et al. 2019), one of which was depicted in plate 4 of livraison 38 by Jan & Sordelli (1871). In a footnote to the nomenclatural history of Boiga (Toxicodryas) pulverulenta, Hughes & Barry (1969:1020) wrote ���Dr. Ladiges [then curator at ZMH] tells us that the type is still in Hamburg and is from St. Thom��, not Edina, Grand Bassa County, Liberia as stated by Loveridge (1958:269).��� The latter citation is in error (Loveridge 1957). Cer��aco et al. (2018) deduced that the information from Ladiges was based on an old label associated with the presumed remaining type specimen. Because extensive fieldwork on the island of S��o Tom�� over the last century failed to document this species, Cer��aco et al. (2018) concluded that it does not occur there, and the original type locality from Fischer (1856) ���Edina, Grand Bassa County in Liberia (West-Afrika)��� is correct. Moreover, the former authors determined that the pholidosis from this presumed type (Cer��aco et al. 2018:fig. 1) is not consistent with the extensively detailed original description, and thus, it is not a type specimen (contra Uetz et al. 2019). Although we agree with this conclusion, an examination of this specimen (ZMH R04376) by J. Hallermann (pers. comm.) found some differences in the morphometric data provided by Cer��aco et al. (2018) (e.g., SVL = 635 mm, not 880 mm), and we used these updated data in our analyses of morphological data (Table 2). Chabanaud (1917a:375), convinced that the odd configuration of subcaudals (combination of single and double subcaudals near the vent) seen in one male and one female was unique to Colubridae, named the taxon Dipsadomorphus boueti from these specimens that were donated by ���Dr Bouet,��� who collected for the Paris Museum in Ivory Coast (1909) and Dahomey (1910���1913). Unfortunately no specific type locality was provided. Chabanaud (1917b) then clarified and corrected himself by stating that both types were males (the mistaken female was a juvenile), and they were collected from ���Porto-Novo��� in Dahomey, which is modern-day Porto-Novo, the capital city of Benin. Chabanaud (1917c) subsequently realized that his specimens of D. boueti represented T. pulverulenta individuals with abnormal subcaudals, and he synonymized the former taxon with the latter (as D. pulverulentus). We agree with this taxonomic placement, because the types of D. boueti have subcaudal counts (106 and 116) and TL/SVL ratios (0.25 and 0.28) that are most consistent with T. pulverulenta (Table 2). The Porto-Novo locality is on the eastern edge of the Dahomey Gap (sensu Demenou et al. 2016:fig. 1), a renowned biogeographic barrier to at least some forest-specialist species in West Africa. Because T. pulverulenta does not seem to be a strict forest specialist (see Habitat below), and the Porto-Novo locality occurs on the eastern edge of the Dahomey Gap, it is likely that the Gap is not a barrier to dispersal for this species. We therefore hypothesize that the Niger Delta is the biogeographic barrier separating T. pulverulenta from its cryptic sister taxon T. adamanteus sp. nov. Diagnosis. A species of Toxicodryas restricted to West Africa, west of the Niger Delta, defined by the following combination of characters: maximum SVL 1 meter in T. blandingii and T. vexator sp. nov.), DSRN 19���21 (vs. 23���25 in T. blandingii and 23���29 in T. vexator sp. nov.), DSRM 19���21 (vs. 21���25 in T. blandingii and T. vexator sp. nov.); cloacal plate undivided (vs. usually divided in T. blandingii, and divided or undivided in T. vexator sp. nov.); both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots (vs. adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters in T. blandingii and T. vexator sp. nov.); hemipenis relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, with a domed apex (vs. hemipenis relatively short and massive [i.e., broad], proximal third covered with spines, distal two-thirds dimpled with a flattened apex in T. blandingii and T. vexator sp. nov.). Variation. Morphometric variation of Toxicodryas pulverulenta is shown in Table 2. Chabanaud (1917a) first noted that some individuals (including the type of Dipsadomorphus boueti) can have a few single subcaudals posterior to the cloacal plate. Bogert (1940) provided morphometric data, but he did not distinguish between populations from Liberia (T. pulverulenta) and former French Cameroon (T. adamanteus sp. nov.), except that ���the Liberia specimens are distinctly more reddish brown than the Cameroon specimens.��� However, he listed the maximum total length of a male (1112 mm) and female (1050 mm) T. pulverulenta from Liberia. In snakes from West Africa (without noting sex), including Cameroon where T. adamanteus sp. nov. occurs, Angel (1933:146) noted temporal formula variation of 2 + 2 (rarely 2 + 3 or 3 + 2 or 1 +2), 236���276 ventrals, 96���132 subcaudals, and a maximum size of 1225 mm; nearly identical data were reported by Villiers (1950a), Doucet (1963), Stucki-Stirn (1979), and Chippaux (2006). Although all of our examined specimens and most literature records noted the 3 rd to 5 th supralabial in contact with the eye (e.g., Villiers 1950b), Chippaux (2006) documented individuals with the 4 th to 6 th supralabial in contact with the eye, and sometimes, only two scales in contact with the eye, but some of these specimens might be attributable to T. adamanteus sp. nov. Segniagbeto et al. (2011) reported snakes from Togo with 165���175 ventrals, which is undoubtedly erroneous. Fischer (1856:83) noted his type specimen had seven maxillary teeth that were oriented nearly backwards, becoming larger posteriorly. These were followed by two larger teeth (i.e., fangs) that were furrowed in a ���besonderen Hauttasche��� [special skin pocket]. There were 10���12 mandibular teeth, which were slightly curved towards the back of the mouth, and increased in size posteriorly. Bogert (1940:61) remarked ��� five specimens examined show variation from eleven to thirteen anterior subequal teeth followed after a very short diastema by two larger grooved fangs and a smaller fang, the total number of teeth being fourteen to eighteen.��� Because only two of his examined specimens originated from former French Cameroon (attributable to T. adamanteus sp. nov.), at least three of these specimens were from Liberia, which are attributable to T. pulverulenta. In snakes from Ghana, Leeson (1950) noted 11���13 maxillary teeth, becoming slightly larger posteriorly, followed by 2 fangs, and sometimes a 3 rd, smaller fang; 14���16 palatine and pterygoid teeth, and 15 mandibular teeth with the anteriormost ones largest. Johnsen (1962) reported that his specimen from Liberia had 11 maxillary teeth on one side, and six on the other. Based on a specimen from Liberia, Malnate (1972) noted the species lacks posterior hypapophyses. In his original description of the species, Fischer (1856:83) described the coloration in great detail as chocolate brown above, yellow below, finely dotted everywhere. On each side, near the back, there was a large number (60��� 70) of pale red spots that lacked dark edging and extended over 4���6 scales. Usually the spots on one side alternated with those on the other; sometimes they also were opposite to each other and in this case were connected to weak ���Querbinden��� [cross-ties] by bright red connecting strips that extended over the back. Usually there was a small black spot under each of these spots on the outermost tip of the corresponding ventral shields. The innumerable fine black points, with which the whole body was sown, were grouped on the abdominal shields on each side at the point where they bent over to rise sideways, to form a black spot, which in their succession looked like a black longitudinal band, whereby the narrow belly appeared delimited from the flanks. Head was brown and without black lines. Upper lip, lower lip, throat yellow, dotted with black. G��nther (1858:173) described the dorsum of two Nigerian specimens as ���brown with a strong cast of purple��� with elliptical transverse streaks, and at mid-body, these streaks had a small yellow spot in their center. In his description of Dipsadomorphus boueti, Chabanaud (1917a) noted the types were light brownish gray and dotted with brown, whereas the supralabials and venter were yellowish gray. Leeson (1950) described animals from Ghana as brownish red on the dorsum, with ���dull brown��� heads, and ���lightly coloured��� patches on the flanks that had a black edge on the lower margins. There were more narrow light gray bands on the anterior third of the body that gradually diminished to form light gray patches on the posterior two-thirds of the body. The ventral surface of the head was white or cream, whereas the venter of the body was pinkish with numerous brown spots. Brown lines ���commence���[ed] just before the middle of the body on each side of the ventrals, and continued along the edge of the ventrals to the tip of the tail. In his description of two specimens from Ivory Coast, Villiers (1950b) noted their color pattern was pale brownish gray, the dorsal scales dotted with black, the back with alternate dark brown diamond spots (some ocellated with white). Yellowish ventral surface with two black lateral lines. Doucet (1963) noted the dorsal coloration ranged from uniform reddish or yellowish or powdery brown, sometimes with dark bars, with ventral coloration ranging from yellowish to pinkish. Chippaux (2006) noted dorsal coloration as dark beige or reddish with occasional dark gray designs or crossbars; venter pinkish with two dark lateral lines. Based on photographs of an adult male from Guinea (Fig. 11B), the base of the tongue is orangish red, and the forked tip is silvery white with black edging. Our observations of live specimens suggest the dorsal scales have an almost satiny sheen, similar to the appearance of a spider web (MOR, pers. obs.). Hemipenis. Bogert (1940) did not note whether his hemipenis description was based on specimens from Liberia (T. pulverulenta) or former French Cameroon (T. adamanteus sp. nov.). Doucet (1963:301) described the hemipenis of a snake from Ivory Coast as very different from that of B. [Toxicodryas] blandingi. Quite elongated, thorny at the base and dimpled at the apex, it is not bifid. The spines on either side of the sperm groove are very long in the middle part and decrease in size towards the apex and the base. The apex is domed, not flattened. Diet. Villiers (1950a) noted both species of Toxicodryas in West Africa feed mainly on birds. Cansdale (1954) reported an individual from Ghana with two young mice in its stomach. Johnsen (1962:121) reported an adult male from Liberia with ���a small insect-eating bird��� in its stomach. In their paper on snakes of Ghana, Leston & Hughes (1968:754) noted the presence of undigested hair and a 10-cm long rodent tail in the stomachs of two specimens, and that ���this suggests the attraction of Cocoa is for its Rodent fauna.��� Another specimen contained the foot and tail of an Agama sp. lizard (sensu Leach�� et al. 2017). B��hme (2000) noted the diet of snakes from Guinea included small mammals (shrews and rodents), adult and nestling birds, and lizards with well-developed limbs. Greene (1989) suggested that there is an ontogenetic shift in diet from mostly lizards in young individuals to mostly birds and/or mammals as adults, and some individuals can ingest a third of their body weight if they eat more than one prey item in a nest or roost. Behavior. In a paper on Guinean snakes, B��hme (2000) classified the species as relatively common and nocturnal. Reproduction. An adult female found in Ghana in early September contained eggs ���about 8 mm maximum breadth and between 30 and 35 mm long��� attached to the left and right ovaries (Leston & Hughes 1968). Habitat. Menzies (1966) documented several specimens from forest in Sierra Leone, but one animal found at Rokupr seemed to occur in a habitat labeled as mangrove and coastal savanna. Leston & Hughes (1968:754) noted that this species was very common in cocoa tree pods in Ghana. Hughes & Barry (1969) listed the habitat as forest. Leston (1970:143) encountered several snakes in Ghana in trees (including one in a rotten cocoa pod), ���amidst old files on an office shelf,��� on a 1.5-meter high cocoa tree in a greenhouse, and in savanna. Hughes (1988) noted the species only from rainforest in Ghana. R��del & Mahsberg (2000) found snakes in Ivory Coast from thick undergrowth in a swamp bordered by fields and secondary forest, and swampy primary forest. Branch & R��del (2003) captured a juvenile in a funnel trap on the ground in closed-canopy forest in Ivory Coast. In Togo, Segniagbeto et al. (2011) noted the species is most common in forest, but some specimens were found outside of it, presumably in savanna. Geographic distribution and habitat. Based on molecular data from Allen et al. (in press) and patterns of our morphometric data (Table 2), we hypothesize that this species occurs west of the Niger Delta from Guinea to Nigeria. Venom. Venom composition and effects are unknown (Weinstein et al. 2011)., Published as part of Greenbaum, Eli, Allen, Kaitlin E., Vaughan, Eugene R., Pauwels, Olivier S. G., Wallach, Van, Kusamba, Chifundera, Muninga, Wandege M., Aris- Tote, Mwenebatu M., Mali, Franck M. M., Badjedjea, Gabriel, Penner, Johannes, R��del, Mark-Oliver, Rivera, Jacqueline, Sterkhova, Viktoria, Johnson, Grant, Tapondjou, Walter P. & Brown, Rafe M., 2021, Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata Colubridae) with descriptions of two new cryptic species from Central Africa, pp. 1-44 in Zootaxa 4965 (1) on pages 24-27, DOI: 10.11646/zootaxa.4965.1.1, http://zenodo.org/record/4723024, {"references":["Fischer, J. G. (1856) Neue Schlangen des Hamburgischen Naturhistorischen Museums. Abhandlungen aus dem Gebiete der Naturwissenschaften herausgegeben von Naturwissenschaftlichen Verein in Hamburg, 3, 79 - 116 + 3 pl.","Chabanaud, P. (1917 a) Enumeration des ophidiens non encore etudies de l'Afrique occidentale, appartenant aux collections du Museum avec la description des especes et des varietes nouvelles. Bulletin du Museum d'Histoire Naturelle, Paris, 22, 362 - 382. https: // doi. org / 10.5962 / bhl. part. 17132","Chabanaud, P. (1917 b) Note complementaire sur les ophidiens de l'Afrique occidentale, avec la description d'une espece nouvelle. Bulletin du Museum d'Histoire Naturelle, Paris, 23, 7 - 14.","Ceriaco, L. M. P., Marques, M. P. & Bauer, A. M. (2018) Miscellanea Herpetologica Sanctithomae, with a provisional checklist of the terrestrial herpetofauna of Sao Tome, Principe and Annobon islands. Zootaxa, 4387 (1), 091 - 108. https: // doi. org / 10.11646 / zootaxa. 4387.1.4","Hallermann, J. (1998) Annotated catalog of the type specimens of the herpetological collection in the Zoological Museum of the University of Hamburg. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 95, 197 - 223.","Uetz, P., Cherikh, S., Shea, G., Ineich, I., Campbell, P. D., Doronin, I. V., Rosado, J., Wynn, A., Tighe, K. A., McDiarmid, R., Lee, J. L., Kohler, G., Ellis, R., Doughty, P., Raxworthy, C. J., Scheinberg, L., Resetar, A., Sabaj, M., Schneider, G., Franzen, M., Glaw, F., Bohme, W., Schweiger, S., Gemel, R., Couper, P., Amey, A., Dondorp, E., Ofer, G., Meiri, S. & Wallach, V. (2019) A global catalog of primary reptile type specimens. Zootaxa, 4695 (5), 438 - 450. https: // doi. org / 10.11646 / zootaxa. 4695.5.2","Hughes, B. & Barry, D. H. (1969) The snakes of Ghana: a checklist and key. Bulletin de l'Institut fondamental d'Afrique noire. Serie A, Sciences naturelles, 31 (3), 1004 - 1041.","Loveridge, A. (1957) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya, Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, Harvard, 117 (2), 151 - 362 + i - xxxvi (index).","Chabanaud, P. (1917 c) Revision de quelques reptiles d'Afrique et description de trois especes nouvelles. Bulletin du Museum d'Histoire Naturelle, Paris, 23, 442 - 454.","Demenou, B. B., Pineiro, R. & Hardy, O. J. (2016) Origin and history of the Dahomey Gap separating West and Central African rain forests: Insights from the phylogeography of the legume tree Distemonanthus benthamianus. Journal of Biogeography, 43 (5), 1020 - 1031. https: // doi. org / 10.1111 / jbi. 12688","Bogert, C. M. (1940) Herpetological results of the Vernay Angola expedition. Bulletin of the American Museum of Natural History, 77, 1 - 107, pl. I.","Angel, F. (1933) Les Serpents d'Afrique Occidentale Francaise. Larose Editeur, Paris, 246 pp.","Villiers, A. (1950 a) Initiations Africaines. II. Les Serpents de l'Ouest Africain. Institut Francais d'Afrique Noire, Dakar, Senegal, 148 pp.","Doucet, J. (1963) Les serpents de la Republique de Cote d'Ivoire. Serpents venimeux. Acta Tropica, 20 (3 - 4), 297 - 340.","Stucki-Stirn, M. C. (1979) A Comparative Study of the Herpetological Fauna of the former West Comeroon [sic] / Africa: With a Classification and Synopsis of 95 Different Snakes and Description of Some New Sub-species. Snake Report 721. Herpeto- Verlag, Teuffenthal, Switzerland, 650 pp.","Chippaux, J-P. (2006) Les Serpents d'Afrique Occidentale et Centrale. 3 rd Edition. IRD Editions, Paris, 311 pp.","Villiers, A. (1950 b) Catalogues. IV. La Collection de Serpents de l'I. F. A. N. Institut Francais d'Afrique Noire, Dakar, Senegal, 155 pp.","Segniagbeto, G. H., Trape, J. F., David, P., Ohler, A., Dubois, A. & Glitho, I. A. (2011) The snake fauna of Togo: Systematics, distribution and biogeography, with remarks on selected taxonomic problems. Zoosystema, 33 (3), 325 - 360. https: // doi. org / 10.5252 / z 2011 n 3 a 4","Leeson, F. (1950) Identification of Snakes of the Gold Coast. The Crown Agents for the Colonies, London, x + 142 pp.","Johnsen, P. (1962) Notes on African snakes, mainly from northern Rhodesia and Liberia. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn, 124, 115 - 130.","Malnate, E. V. (1972) Observations on the vertebral hypapophyses and associated musculature in some snakes, with special reference to the Colubridae. Zoologische Mededelingen, 47 (18), 225 - 239.","Gunther, A. (1858) Catalogue of Colubrine Snakes in the Collection of the British Museum. Trustees of the British Museum, London, xvi + 281 pp. https: // doi. org / 10.5962 / bhl. title. 13272","Cansdale, G. (1954) Gold Coast snakes - a complete list. The Nigerian Field, 19, 118 - 132.","Leston, D. & Hughes, B. (1968) The snakes of Tafo, a forest cocoa-farm locality in Ghana. Bulletin de l'Institut Francaise d'Afrique Noire, 30 A, 737 - 770.","Leache, A. D., Grummer, J. A., Miller, M., Krishnan, S., Fujita, M. K., Bohme, W., Schmitz, A., Lebreton, M., Ineich, I., Chirio, L., Ofori-Boateng, C., Eniang, E. A., Greenbaum, E., Rodel, M. - O. & Wagner, P. (2017) Bayesian inference of species diffusion in the West African Agama agama species group (Reptilia, Agamidae). Systematics and Biodiversity, 15 (3), 192 - 203. https: // doi. org / 10.1080 / 14772000.2016.1238018","Bohme, W. (2000) Diversity of a snake community in a Guinean rain forest (Reptilia, Serpentes). In: Rheinwald, G. (Ed.), Isolated Vertebrate Communities in the Tropics. Proceedings of the 4 th International Symposium in Bonn. Bonner zoologische Monographien, 46, 69 - 78.","Greene, H. W. (1989) Ecological, evolutionary, and conservation implications of feeding biology in Old World cat snakes, Genus Boiga (Colubridae). Proceedings of the California Academy of Sciences, 46 (8), 193 - 207.","Menzies, J. I. (1966) The snakes of Sierra Leone. Copeia, 1966 (2), 169 - 179. https: // doi. org / 10.2307 / 1441123","Leston, D. (1970) Some snakes from the forest zone of Ghana. British Journal of Herpetology, 4 (6), 141 - 144.","Hughes, B. (1988) Herpetology in Ghana (West Africa). British Herpetological Society Bulletin, (25), 29 - 37.","Rodel, M. - O. & Mahsberg, D. (2000). Vorlaufige Liste der Schlangen des Tai-Nationalparks / Elfenbeinkuste und angrenzender Gebiete. Salamandra, 36 (1), 25 - 38.","Branch, W. R. & Rodel, M. - O. (2003) Herpetological survey of the Haute Dodo and Cavally forests, western Ivory Coast, Part II: Trapping results and reptiles. Salamandra, 39 (1), 21 - 38.","Weinstein, S. A., Warrell, D. A., White, J. & Keyler, D. E. (2011) \" Venomous \" Bites from Non-Venomous Snakes: A Critical Analysis of Risk and Management of \" Colubrid \" Snake Bites. Elsevier, Amsterdam, Boston, Heidelburg, London, New York, Oxford, Paris, San Diego, San Francisco, Singapore, Sydney and Tokyo, xxvii + 336 pp."]}

Toxicodryas adamanteus sp. nov. (Table 2, Figs. 8, 12–13) We hypothesize that this new, cryptic species occurs in western, central and eastern Africa, east of the Niger Delta, and it has been considered to be conspecific with Toxicodryas pulverulenta since it was first documented to occur in Angola by Peters (1877). The recognition of this new species is supported by evidence from molecular data (Fig. 2) and significant differences in subcaudal scale counts (see Results). Diagnosis. A species of Toxicodryas restricted to West, Central and East Africa, east of the Niger Delta, defined by the following combination of characters: maximum SVL 1 meter in T. blandingii and T. vexator sp. nov.), DSRN 18–23 (vs. 23–25 in T. blandingii and 23–29 in T. vexator sp. nov.), DSRM 18–21 (vs. 21–25 in T. blandingii and T. vexator sp. nov.); cloacal plate undivided (vs. usually divided in T. blandingii, and divided or undivided in T. vexator sp. nov.); both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots (vs. adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters in T. blandingii and T. vexator sp. nov.); hemipenis relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, with a domed apex (vs. hemipenis relatively short and massive [i.e., broad], proximal third covered with spines, distal twothirds dimpled with a flattened apex in T. blandingii and T. vexator sp. nov.). Holotype. UTEP 22204 (field no. ELI 2213; Figs. 8, 12–13), adult male from Npenda village, NE of Lake Tumba (00.7465° S, 18.2243° E, 311 m), Equateur Province, DRC, collected by local Twa people and brought to Eli Greenbaum, Chifundera Kusamba, Wandege M. Muninga, and Mwenebatu M. Aristote on 8 July 2013. Paratopotype. UTEP 22203 (field no. ELI 2212) adult female with same collection details as the holotype. Paratype. RBINS 2699 (formerly RBINS 9127) (field no. Paul Leloup #26), adult female from Région Tshabondo (2.690861° S, 27.341972° E), South Kivu Province, DRC, collected by Paul Leloup on 11 October 1958. Description of the holotype. Adult male, 812 mm SVL; head weakly triangular and moderately distinct from neck, 2.4% of SVL (19.3 mm); interocular distance 13.4 mm, pupil elliptical, eye diameter 5.2 mm; loreal nearrectangular, shorter (1.7 mm) than high (2.0 mm), slightly tapering superiorly and vertically; body semi-triangular; tail moderately long (27.7% of SVL). Supralabials 8/8, 3 rd –5 th /3 rd –5 th contacting orbit; infralabials 13/12, 1 st on each side in contact behind mental, 1 st –5 th /1 st –5 th contacting anterior chin shields and 5 th –7 th /5 th –7 th contacting posterior chin shields; 1/1 preocular; 2/2 postoculars; temporals 2 + 3 + 3/2 + 3 + 3; 2 internasals; nasal divided; frontal width and length equal (6.6 mm); dorsal scale rows 21 one head length posterior to jaw rictus, 19 at midbody, 15 one head length anterior to vent, smooth and oblique with apical pits visible on the neck, vertebral scales broad and apically flattened; ventrals 264 (standard), 262 (Dowling); cloacal plate undivided; paired subcaudals 117; both hemipenes everted. Coloration (in life) of the holotype. Silvery gray-brown over the entire length of the dorsum including the head, with similarly colored but darker diamonds occurring laterally over the full length of the neck, body, and tail. These diamonds are elongated vertically, have pale gray centers, and often have black spots at the top and bottom corners. Between each diamond on the body and neck is a more vaguely defined, pale gray diamond or stripe. The lateral and antero-dorsal sides of the head are speckled, as is the ventral side of the head, neck, body, and tail. The venter is otherwise white, with the speckles forming two pale stripes running down either side of the ventrals and subcaudals. The base of the tongue is orangish red, and the forked tip is silvery white with black edging. Coloration (in preservative) of the holotype. This specimen’s coloration has become slightly darker and browner than it was in life, with less pronounced patterning on the flanks, but otherwise the appearance is similar to the coloration in life. Variation. Morphometric variation of Toxicodryas adamanteus sp. nov. is shown in Table 2. We observed extensive temporal scale variation, including 1 + 1, 1 + 2, 2 + 2 (most commonly), 2 + 3, 2 + 5, 3 + 2, 3 + 3, and 4 + 3. Chabanaud (1917c) described a male from Gabon with supralabials 3–6 contacting the eye, and this specimen seems to be the size record at 1,225 mm total length (995 mm SVL, 230 mm tail length). Schmidt (1923) listed ventral counts of 251–269 in snakes from DRC, and noticed one individual that had a preocular fused to the supraocular on one side, and in two individuals, the loreal was fused to the lower portion of the preocular and thus contacted the eye. Laurent (1956) noticed that snakes from DRC (i.e., T. adamanteus sp. nov.) had more subcaudals than snakes from West Africa (i.e., T. pulverulenta). Skinner (1973) noted ventral scale counts range from 240–269, subcaudal scale counts range from 105–126 (identical to Pitman 1974), and the maximum size is about 2 meters, substantially larger than all other published records, and thus, highly doubtful. De Witte (1975) provided data for ventral scale counts of DRC snakes ranging from 235–242 (males) and 239–249 (females), and subcaudal counts of 112–120 (both sexes). Spawls & Branch (2020) listed the maximum size as “about 1.25 m ” but no specific locality or record was provided. Although most of our examined specimens and literature records noted the 3 rd to 5 th supralabial in contact with the eye, some individuals have the 4 th to 6 th supralabial in contact with the eye (e.g., Loveridge 1937) and one specimen from Gabon (CAS 258155) had only the 4 th and 5 th supralabial in contact with the eye. In snakes from Uganda, Pitman (1974) reported ventral scale counts of 240–269 and subcaudal counts of 105–126 (not distinguished by sex). In de Witte’s (1975) study of snakes from Virunga National Park in eastern DRC, he noted ventrals range from 235–242 in males and 239–249 in females; subcaudals ranged from 112–120 in both sexes. Rasmussen (1997b:106) described individuals with 3 or 4 postoculars, 7–9 supralabials, 9–13 infralabials, 236–278 ventrals, and 96–132 subcaudals (sometimes undivided). In general, he noted this species has sloping and smooth scales with apical pits, and the vertebral row is more or less enlarged. Bogert (1940:fig. 8) illustrated the maxillary teeth of a specimen (AMNH 50590) from former French Cameroon, noting “ five specimens examined show variation from eleven to thirteen anterior subequal teeth followed after a very short diastema by two larger grooved fangs and a smaller fang, the total number of teeth being fourteen to eighteen.” Because only two of his examined specimens originated from former French Cameroon, at least three of these specimens are from Liberia, which are attributable to T. pulverulenta. Schmidt (1923:103) quoted field notes of Herbert Lang for DRC snakes as “coloration, in life, reddish brown above, head darker brown. Irregular dark gray lateral bars, wider in the middle, extend from the vertebral line to the venter, tipped above and below with black. A cream-colored central spot in the broad portion of each lateral bar. Faint narrow grayish crossbars between the wider ones, disappearing posteriorly. The wider crossbars are usually alternate, sometimes confluent on the back. Venter pinkish gray, heavily dotted with brown which forms two lines at the inner edges of the ventral edges of the ventral angle. These lines are more distinct beneath the tail.” Hellmich (1957b) described several individuals from Angola with a reddish brown to reddish blue-gray dorsum, with varying degrees of contrasting rhombic patterns and spots. Pitman (1974) provided a similar description for specimens from Uganda, noting they range from pinkish-brown to reddish-brown. Hedges (1983:21) described Kenyan specimens as, “a rather pretty pattern of coffee coloured diamond shaped markings on a mushroom pink body.” Rasmussen (1997b) described coloration in this species (presumably based on examined specimens from Nigeria and Cameroon) as reddish brown on the dorsum of the body; dorsum of the head darker brown; irregular, dark gray, diamondshaped spots on the flanks that are widest at mid-body; spots edged in black on superior and inferior edges with a cream spot in the center that sometimes fuse together in the midline; faint grayish transverse bands between spots that fade and disappear posteriorly; venter pinkish-gray, with dense brown spots that form a line just inside and parallel to the keeled ventrals, most salient on the tail. Pauwels et al. (2019b) noted a specimen from Gabon with an orange tongue in life. Spawls & Branch (2020:239) noted the dorsal color can range from pinkish to brown, redbrown or pinky gray with “darker” cross-bars that enclose a “pale” spot. The dorsum and venter are “finely dusted” with brown or black specks, whereas the venter is pale pink with “dashed dark lines” on each side of the ventrals. Contrary to Pauwels et al. (2019b), they noted the tongue is pink with a white tip. Based on our photographs of DRC snakes (e.g., Fig. 12G), the tongue is orangish red, and the forked tip is silvery white with black edging, which is consistent with the coloration of the holotype in life (EG pers. obs.). An unsexed individual from Banalia, DRC (Fig. 12F) is unique in having a golden yellow mid-dorsal stripe. Diet. Pitman (1938) mentioned a juvenile from Uganda that had a small mouse in its stomach. In Kenya, Hedges (1983) noted the species eats small chameleons, geckos, and frogs. Butler & Reid (1990) described an adult female from Nigeria that contained the remains of an Agama sp. lizard (sensu Leaché et al. 2017). Spawls & Branch (2020) listed the diet as arboreal lizards and rodents. Parasites. Pitman (1938) noted a juvenile from Uganda with numerous ticks. Behavior. Pitman (1974:128) described the species as “arboreal, nocturnal... amiable disposition,” and suggested that it might rely on camouflage to avoid predation, a sentiment also suggested by Gans (1961). Chirio & LeBreton (2007) suggested the species is shy and rarely bites. Spawls & Branch (2020) noted that when threatened, it elevates the anterior part of its body, “flickers its red tongue,” hisses, and strikes. Reproduction. Schmidt (1923) described a DRC female collected in June 1914 with an unspecified number of eggs that measured 11 x 29 mm. Butler & Reid (1990) described an adult female from Nigeria (captured 21 January 1988) that contained two developing eggs (34–35 mm long, 11 mm wide). Branch (2005) noted the species lays 2– 5 eggs. Habitat. Parker (1936) noted this species is restricted to rain forest and “its outliers.” Perret (1961) listed the species from forest in Cameroon. De Witte (1962) listed the habitats for DRC snakes as equatorial forest and gallery forest, and this classification was repeated by Thys van den Audenaerde (1965). Blackwell (1967) listed the species from forest, thickets, and gardens in Nigeria. Pitman (1974:128) described this snake as a “forest species” in Uganda, but also noted one individual from a “fowl house,” and another from an epiphytic fern (Platycerium angolense) on a forest tree. Butler & Reid (1986) listed the species only from forest in Nigeria. Lawson (1993) documented the species from forest and “farmbush” in Cameroon. Rasmussen (1997b) described the species as nocturnal and restricted to forest. Chirio & LeBreton (2007) recorded the species from forests and savanna-forest mosaic in Cameroon. Spawls & Branch (2020) included forest, woodland, and forest-savanna mosaic from sea level to about 2,000 m elevation. Among the 19 snake species recorded within the garden of a villa in Yenzi ( 2.77261° S, 10.03403° E), Gamba, Nyanga Province, southwestern Gabon (forest-savanna mosaic), inhabited by one of the authors from 2004 to 2011 (OSGP, unpubl. data), T. adamanteus sp. nov. was among the most rarely encountered species. Geographic distribution and habitat. Based on molecular data from Allen et al. (in press) and patterns of our morphometric data (Table 2), we hypothesize that this species occurs east of the Niger Delta in Nigeria, Cameroon, Equatorial Guinea, Gabon, CAR, Republic of Congo, DRC, Angola, Uganda, South Sudan, and Kenya. Ullenbruch & Böhme (2017) recently listed the species from South Sudan. Venom. Based on a specimen from Cameroon, Taub (1967) described the histological morphology of the Duvernoy’s gland. Venom composition and effects are unknown (Weinstein et al. 2011). Etymology. The specific epithet adamanteus is a Latin adjective referring to the diamond-shaped marks on the flanks and dorsum of this species.

Toxicodryas vexator sp. nov. (Table 1, Figs. 1, 8���10) We hypothesize that this new, cryptic species occurs east of the confluence of the Congo and Ubangi rivers, and it has been considered to be conspecific with Toxicodryas blandingii since it was first documented to occur east of these rivers in DRC (Boulenger 1919). The recognition of this new species is supported by molecular data (Fig. 2), several significant differences in scale counts (see Results), and less toxic venom in comparison to its sister taxon T. blandingii (see Venom). Diagnosis. A species of Toxicodryas restricted to east-central and East Africa (east of the confluence of the Congo and Ubangi rivers), defined by the following combination of characters: maximum SVL> 1 meter (vs. maximum SVL T. pulverulenta and T. adamanteus sp. nov.); DSRN 23���29 (vs. 19���21 in T. pulverulenta and 18���23 in T. adamanteus sp. nov.); DSRM 21���25 (vs. 19���21 in T. pulverulenta and 18���21 in T. adamanteus sp. nov.); cloacal plate divided or undivided (vs. usually divided in T. blandingii, and always undivided in T. pulverulenta and T. adamanteus sp. nov.); adult males glossy or velvety black with a yellow venter, and adult females light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters (vs. both sexes brown to pink with darker cross-bars that often enclose a whitish spot, and the dorsum and venter sprinkled with fine dark brown or black spots in T. pulverulenta and T. adamanteus sp. nov.); hemipenis relatively short and massive (i.e., broad), proximal third covered with spines, distal two-thirds dimpled with a flattened apex (vs. relatively long with long spines mid-way along the shaft that decrease in size towards the apex and base, and with a domed apex in T. pulverulenta and T. adamanteus sp. nov.); venom toxicity LD 50 = 4.88 mg /kg in mice (vs. venom toxicity LD 50 = 2.85���3.55 mg /kg in mice for T. blandingii). Holotype. UTEP 22196 (field number MUSE 10341; Fig. 10), adult male collected in Mulisi, Nzovu Est, Kahuzi-Biega National Park (02.447291�� S, 28.2825378�� E, 1101 m), South Kivu Province, DRC, collected by Guillain M. Mitamba, Deo Kujirakwinja, Emmanuel Muhindo, Radar Nushili, Wandege M. Muninga, and Andrew J. Plumptre on 13 November 2015. Paratypes. UTEP 22195 (field number EBG 1362; Figs. 1B, 9C), adult female collected in the vicinity of Irangi (01.8780�� S, 28.4524�� E, 811 m), South Kivu Province, DRC, collected from a tree by a resident and brought to Maurice Luhumyo, Chifundera Kusamba, Mwenebatu M. Aristote, Wandege M. Muninga, and Eli Greenbaum on 30 August 2007; RBINS 2698 (formerly RBINS 8621) (field no. Leloup #27), adult male from Bunyakiri (2.075630�� S, 28.573194�� E, 1000 m), South Kivu Province, DRC, collected by Paul Leloup on 6 March 1958. Description of the holotype. Adult male, 1438 mm SVL; head strongly triangular and distinct from the neck, 2.1% of SVL (30.9 mm); interocular distance 21.7 mm, pupil elliptical, maximum horizontal eye diameter 7.5 mm; loreal near-rectangular, longer (3.0 mm) than high (2.4 mm), slightly tapering superiorly; body triangular; tail moderately long (32.1% of SVL). Supralabials 9/9, 4 th, 5 th, and 6 th /4 th, 5 th, and 6 th contacting orbit; infralabials 14/14, 1 st on each side in contact behind mental, 1 st ���4 th /1 st ���4 th contacting anterior chin shields, 5 th ���8 th /5 th ���8 th contacting posterior chin shields, and 1 cuneate between the 5 th and 6 th left infralabials; 2/2 preoculars; 2/2 postoculars; temporals 2 + 2/2 + 2; 2 internasals; nasal divided on both sides; frontal slightly wider (10.1 mm) than long (9.2 mm); dorsal scale rows 23 one head length posterior to jaw rictus, 21 at midbody, 15 one head length anterior to vent, smooth and oblique with apical pits visible on the neck, vertebral scales broad and apically flattened; ventrals 261 (standard), 260 (Dowling); cloacal plate undivided; paired subcaudals 136 (terminal spine missing); both hemipenes everted. Coloration (in life) of the holotype. Solid black over the entire length of the dorsum, and black ventrally except for the anterior half of the body, where the ventral scales are pale yellow with a black posterior edge that increases in thickness in correlation with increased distance from the head. The head is black dorsally and laterally except for yellow on the lower, anterior region of the 5 th ���9 th supralabials, and yellow ventrally except for thin black rims on the posterior edges of the infralabials. Coloration (in preservative) of the holotype. The specimen���s coloration in preservative is nearly the same as in life, although the ventral yellow coloration has dulled to a yellowish off-white; some scales also seem cloudy under close lighting, indicative of some stage of ecdysis in life. Variation. For years, the size record for this species was attributed to Goodman (1985), who recorded a total length of 2515 mm for an adult female collected in Uganda. However, Hedges (1983:20) measured a specimen from Kenya that was 2707 mm total length, and said it ���is reputed to exceed 3300 mm.��� Weinstein et al. (2011) noted the species may exceed 3.5 meters. Spawls & Branch (1995, 2020) listed the maximum size as ���about 2.8 m ��� (no specific location provided), but Weinstein (pers. comm.) estimated a captive specimen from Kakamega was approximately 3.0 meters total length. Schmidt (1923:104) remarked that two specimens from DRC had fused prefrontals, creating a single transverse scale. Five of his 20 DRC specimens had a divided cloacal plate, ���with indications of a groove in two others,��� and as a result, he stated that the condition of the cloacal plate is ���obviously useless��� as a diagnostic character. Laurent (1956) noted that, with the exception of DRC specimens from Mayombe (north of the Congo River in Kongo Central Province and thus attributable to T. blandingii as recognized herein) with consistently divided cloacal plates, 13 of 31 snakes from other areas of DRC had an undivided cloacal plate. Our examined specimens confirm this sentiment, because 17/ 92 specimens have a divided cloacal plate, and an additional five specimens have a partially divided cloacal plate. Hellmich (1957b) noted maximum sizes of Angolan males (1740 mm SVL; 523 mm TL) and females (1730 mm SVL; 505 mm TL), and aberrant individuals that had 11 supralabials and 9���14 infralabials. De Witte (1966) noted that the species has 17 scale rows at midbody, but this datum is so aberrant that it is likely an error (Pitman 1974; Table 1). Skinner (1973) provided ranges of scale counts (161���274 ventrals, 86���147 subcaudals) that are so aberrant that they are clearly erroneous (at least for lower counts), and noted a maximum length of about 3 meters. For Uganda snakes, Pitman (1974) listed ventral scale counts from 240���260 (males) and 240���259 (females). De Witte (1975) noted a single individual with 25 scale rows at midbody, and Broadley et al. (2003) also noted 25 scale rows as the maximum amount of variation. Based on two specimens from Lukolela, Belgian Congo (AMNH 45907) and Akenge, Belgian Congo (AMNH 12243), Bogert (1940:61) described their maxillary teeth as ���ten in number, followed after a short diastema by three enlarged, grooved fangs, the posterior one of which is smaller than the other two.��� Schmidt (1923:105) described two color phases of DRC specimens, including (1) a black phase with the anterior portion of the venter yellow, each ventral scale bordered with black on its posterior edge, the border increasing in width until the yellow color disappears on the posterior two-thirds of the venter; and (2) a brownish phase, ���with more or less distinct wide dark cross-bars, confluent anteriorly, alternate posteriorly on the vertebral line.��� Pitman (1938:211) described an unsexed individual from Uganda as ���darkish gunmetal with a purplish mottling towards the tail, belly pale yellow from head to tail but with slight brown markings on the posterior half increasing in occurrence towards and on the tail.��� A female from Uganda was described as rich chestnut dorsally and ventrally, with handsome chocolate blotches on the flanks, each one containing a small white spot. On the posterior part of the body, including most of the tail, these blotches were confluent with pale edging. The ventral aspect of the head and the first 50 ventrals were ���paly yellowish tinged green.��� The dorsum of the head was dark brown, and the posterior supralabials were dull greenish grey. The supralabials had black edging posteriorly, except for the last one. There were two elongated dark brown blotches behind the eye, which was iridescent hazel. Bogert (1940:61) provided data for a specimen (AMNH 45907; determined to be a subadult male based on photographs examined by EG) that had been described by James P. Chapin as ���olive-brown, with dark brown patches; below grayish brown.��� Hellmich (1957b) described the coloration of several individuals from Angola, including an adult male that had a blue-reddish-black dorsum with a venter that was pale yellow in the front half of the body, with an ever-widening blue-black border that eventually enclosed the entire ventral scales towards the posterior side. Another adult male had a dark black-brown dorsum, and light yellow venter on the first third of the body, but starting on the 5 th ventral, there was a dark gray-blue spot that increased in size posteriorly until it covered the entire venter towards the posterior end of the body. An adult female���s dorsum was described as clay yellow with mostly alternating dark transverse bars and a pale yellow venter. A second adult female had similar coloration to the latter specimen, but the transverse bands were only faintly visible on the posterior sixth of the body and tail. A juvenile dorsum was described as gray-brown with transverse black-brown spots that converge in the anterior part of the body, but then alternate posteriorly, with a light yellowish-white spot in the lower center of the spots. Laurent (1956:195) listed a section about possible sexual dimorphism for this species, but stated that nothing glaring appears in his data. Pitman (1974:125) described several individuals from Uganda that were similar to the descriptions above, and added that both color phases ���exhibit a handsome suede effect.��� He noted that although subadult males might retain the brown phase, black females are unknown, and the black and brown color phases have ���sexual significance.��� The dark blotches of juveniles were noted to vary from ���blackish, chocolate, reddishbrown or dark or bright chestnut,��� with interspaces ranging from light gray, pale brown or chestnut, sometimes with a pinkish tinge. Goodman (1985:56) described two adult females from Uganda as ���dingy olive-brown.��� Hughes (2000) noted that Laurent (1964) seemed to imply that his largest male specimen from Dundo, Angola was not black, but this is not clear, because the latter author only mentioned in passing that a smaller male from Andrada, Angola represented the black phase. Spawls & Branch (1995, 2020) remarked that the yellow ventral pigmentation might form a stripe in the middle of the venter. In the brown color morph, the skin between the scales is bluish gray, and especially visible when the snake inflates its body during a threat display. The eye can be yellowish or brown. The black phase snakes are usually male, whereas the brown phase ones are usually female. Based on photos of an adult male from Banalia, DRC (individual shown in Fig. 9A), the base of the tongue is bluish black, and the forked tip is silvery gray. Hemipenis. Bogert (1940:61) described the hemipenis of an individual, presumably from DRC, as ���not bifurcate, extending to the tenth caudal, the undivided sulcus deeply buried between two fleshy fold [sic]. The basal half is heavily armed with stout spines but distally the organ is calyculate, the edges of the calyces crenulated.��� Hemipenes of our examined specimens had a simple, subcylindrical shape, simple sulcus spermaticus, and spinose ornamentation with a rough apical structure, as also noted for T. blandingii. Because only two specimens with everted hemipenes were available for both T. blandingii and T. vexator sp. nov., any finer-scale differences cannot be attributed to species-level differentiation (Dowling & Savage 1960; Doucet 1963). Diet. Laurent (1964) noted a juvenile snake contained the remains of a bird. In snakes from Uganda, Pitman (1974) noted the diet included weaver finches, robber birds��� eggs, small rodents, bats, frogs, large agamids, and chameleons. Hedges (1983) described a Kenyan snake that regurgitated a Nectarinia famosa sunbird. Spawls & Branch (1995, 2020) described the diet as birds, bird eggs, arboreal lizards, frogs, arboreal rodents, and bats. Nagy et al. (2011) documented an adult female from DRC that predated a Short-palated Fruit Bat (Casinycteris argynnis) from a bat net in secondary forest. Another male specimen from DRC was captured with a nestling bushshrike (Laniarius sp.) in its stomach. Parasites. Loveridge (1937) noted a tick on the scales of a specimen (ANSP 20504) from Belgian Congo. Parasites of Ugandan snakes included internal nematodes (Kalicephalus sp.) and cestodes, ectoparasites (ticks, including Aponomma latum), and captive specimens had bacterial infections, including mouth rot (Pitman, 1974). Goodman (1985:56) documented dozens of ticks, ���probably Aponomma sp.,��� and linguatulid worms from the posterior saccular lung cavity of an adult female from Uganda. Behavior. Spawls & Branch (1995, 2020) described the snake as arboreal, sometimes climbing to heights of 30 meters in trees, but also descending to the ground to cross open spaces and roads. It is mostly nocturnal, sheltering in leaf clumps and tree hollows during the day. When threatened, it will open its mouth widely and expose the pink lining inside, inflate its body, flatten the head, lift the anterior part of its body off the ground and into C-shaped coils, and strike at perceived threats. It is adept at smelling sleeping birds in nests at night, and will make a slow, deliberate approach to attack them. Reproduction. A personal communication from ���Leakey��� (most likely Richard Leakey) to Pitman (1974) noted that a female from Kakamega (Kenya) laid 9 eggs that were approximately 20 x 40 mm on 9 January. Spawls & Branch (1995) reported an average of 7��� 14 eggs in this species (3���14 in Spawls & Branch 2020), presumably based mostly on East African data, which is substantially more than clutch sizes reported for T. blandingii by Luiselli et al. (1998a). Habitat. Laurent (1954) remarked that an adult female from Dundo, Angola was captured after it fell out of a mango tree. Laurent (1960) stated that his colleague M. Leloup often encountered the species in eastern DRC. De Witte (1962) listed the habitats for DRC snakes as equatorial forest and gallery forest, a classification that was repeated by Thys van den Audenaerde (1965). Pitman (1974) also limited the species to forest in Uganda, but he remarked that the species frequently enters houses (where it hunts for bats), and one individual was killed in a tree over 20 meters above the ground. Broadley & Cotterill (2004) stated the species inhabits forests and wooded savannas. Spawls & Branch (1995, 2020) noted the species from forest, woodland, forest-savanna mosaic, riverine woodland, and human habitations from sea level to about 2,200 m elevation. The holotype was found basking on a shrub (ca. 4 meters above ground) in a clearing created by a fallen tree in primary forest. The Irangi paratype was found in secondary forest near a road. Geographic distribution. Based on molecular data from Allen et al. (in press) and patterns of our morphometric data (Table 1), we hypothesize that this species occurs east of the Congo and Ubangi Rivers in CAR, DRC, Angola, Zambia, South Sudan, Uganda, Kenya, and Tanzania. Although Boulenger (1896) listed a specimen from ��� Zanzibar,��� the locality likely referred to mainland records (Parker et al. 1940; Loveridge 1957; Pitman 1974). Venom. Wakeman (1966) documented a 2.59 m (presumably total length) captive adult from Uganda that struck an adult rat, which died after 8.5 minutes. Pitman (1974:127) described the case of a young European who was bitten on the thumb three times in rapid succession by a small (305 mm) individual (erroneously reported as 350 mm by Goodman 1985); only ���initial smarting��� at the puncture sites occurred. Spawls (1979) described the bite of a captive 2.1 meter snake on a man, including three fang punctures on his thumb, which resulted in mild swelling of his hand. Anecdotal observations of a bite by a captive snake on a mouse resulted in death in under a minute. Hedges (1983:20) was bitten, ���and although it drew a considerable amount of blood, I felt no effects whatsoever.��� He also remarked that this species ���chews��� when biting, ostensibly to ensure the rear fangs penetrate the flesh to inject venom. Based on venom collected from a snake originating from Kenya, Weinstein & Smith (1993) estimated the LD 50 value as 4.88 mg /kg in mice. Weinstein & Smith (1993:88) also noted ���the differences in lethal potencies between the two samples of B. blandingi [compare with LD 50 = 2.85���3.55 mg /kg in T. blandingii from West Africa] demonstrate variation of secretion toxicity in this species,��� suggesting possible species-level differences in venom composition of these sister taxa (Weinstein & Kardong 1994). Immunological cross reactivity with elapid venom antisera was demonstrated, but Weinstein & Smith (1993) did not distinguish between samples originating from West Africa (i.e., T. blandingii) or East Africa (i.e., T. vexator sp. nov.). Branch (2005) suggested bites cause nausea and headaches, but fatalities are unknown. Dashevsky et al. (2018) included a Toxicodryas from Tanzania in their proteomics study, and they noted its venom is dominated by three-finger toxins. Scott Weinstein (pers. comm., October 2020) clarified that a snakebite mentioned by Weinstein & Smith (1993) and a second one that occurred one year after this publication were caused by snakes originating from Kakamega forest, Kenya. ���The bite mentioned in the paper was delivered by [an approximately] 3.0 m jet black male that was a favourite member of my collection for nearly 7 years. He typically performed the whole gamut of open mouthed posturing and mock strikes, but was also a very aggressive feeder, and although I tried to condition him to receiving food (freeze-thawed chicks and rodents) out of his enclosure, he still occasionally would ���pour��� ou, Published as part of Greenbaum, Eli, Allen, Kaitlin E., Vaughan, Eugene R., Pauwels, Olivier S. G., Wallach, Van, Kusamba, Chifundera, Muninga, Wandege M., Aris- Tote, Mwenebatu M., Mali, Franck M. M., Badjedjea, Gabriel, Penner, Johannes, R��del, Mark-Oliver, Rivera, Jacqueline, Sterkhova, Viktoria, Johnson, Grant, Tapondjou, Walter P. & Brown, Rafe M., 2021, Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata Colubridae) with descriptions of two new cryptic species from Central Africa, pp. 1-44 in Zootaxa 4965 (1) on pages 17-21, DOI: 10.11646/zootaxa.4965.1.1, http://zenodo.org/record/4723024, {"references":["Boulenger, G. A. (1919) Batraciens et reptiles recueillis par le Dr C. Christy au Congo Belge dans le districts de Stanleyville, Haut-Uele et Ituri en 1912 - 1914. Revue Zoologique Africaine, 7, 1 - 29. https: // doi. org / 10.5962 / bhl. part. 15108","Goodman, J. D. (1985) Two record size Blanding's tree snakes from Uganda. East African Natural History Society Bulletin, 1985, 56 - 57.","Hedges, N. G. (1983) Reptiles and Amphibians of East Africa. Kenya Literature Bureau, Nairobi, Kenya, xii + 139 pp.","Weinstein, S. A., Warrell, D. A., White, J. & Keyler, D. E. (2011) \" Venomous \" Bites from Non-Venomous Snakes: A Critical Analysis of Risk and Management of \" Colubrid \" Snake Bites. Elsevier, Amsterdam, Boston, Heidelburg, London, New York, Oxford, Paris, San Diego, San Francisco, Singapore, Sydney and Tokyo, xxvii + 336 pp.","Spawls, S. & Branch, B. (1995) The Dangerous Snakes of Africa: Natural History, Species Directory, Venoms and Snakebite. Ralph Curtis - Books, Sanibel Island, Florida, 192 pp.","Spawls, S. & Branch, B. (2020) The Dangerous Snakes of Africa. Bloomsbury, London, Oxford, New York, New Delhi and Sydney, 336 pp.","Schmidt, K. P. (1923) Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909 - 1915. Part II. - Snakes. Bulletin of the American Museum of Natural History, 49, 1 - 146, pls. I - XXII.","Laurent, R. F. (1956) Contribution a l'herpetologie de la region des Grands Lacs de l'Afrique centrale. I. Generalites - II. Cheloniens - III. Ophidiens. Annales du Musee Royal du Congo Belge, Serie in- 8 °, 48, 1 - 390, pls. I - XXXI.","Hellmich, W. (1957 b) Die Reptilienausbeute der hamburgischen Angola-Expedition. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 55, 39 - 80.","de Witte, G. - F. (1966) Exploration du Parc National de la Garamba. Reptiles. Institut du Parcs Nationaux du Congo, 48, 1 - 108, pls. I - V.","Pitman, C. R. S. (1974) A Guide to the Snakes of Uganda, Revised Edition. Wheldon & Wesley, Ltd., Codicote, xxii + 290 pp.","Skinner, H. A. (1973) Snakes and Us: An Introduction to East African Herpetology. East African Literature Bureau, Nairobi, Kampala and Dar es Salaam, xvi + 146 pp.","de Witte, G. - F. (1975) Serpents, Boidae, Colubridae, Elapidae et Viperidae. Fondacion pour Favoriser les Recherches Scientifiques en Afrique, Serie 2, 24, 5 - 121.","Broadley, D. G., Doria, C. T. & Wigge, J. (2003) Snakes of Zambia: An Atlas and Field Guide. Edition Chimaira, Frankfurt am Main, 280 pp.","Bogert, C. M. (1940) Herpetological results of the Vernay Angola expedition. Bulletin of the American Museum of Natural History, 77, 1 - 107, pl. I.","Pitman, C. R. S. (1938) A guide to the snakes of Uganda. Part XI. (Conclusion). The Uganda Journal, 5, 160 - 244, pls. XV - XVIII.","Hughes, B. (2000) Polymorphism, sex, size and other aspects of the African snake, Boiga blandingi (Hallowell, 1844). Herpetological Bulletin, 74, 8 - 12.","Laurent, R. F. (1964) Subsidos para o estudo da biologia na Lunda, Museo do Dundo. Reptiles et amphibiens de l'Angola (Troisieme contribution). Publicacoes Culturais da Companhia de Diamantes de Angola, 67, 17 - 165.","Dowling, H. G. & Savage, J. M. (1960) A guide to the snake hemipenis: A survey of basic structure and systematic characteristics. Zoologica, 45 (Part 1), 17 - 28, pls. I - III.","Doucet, J. (1963) Les serpents de la Republique de Cote d'Ivoire. Serpents venimeux. Acta Tropica, 20 (3 - 4), 297 - 340.","Nagy, Z. T., Chifundera Kusamba, Z., Gembu Tungaluna, G. - C., Lotana Lokasola, A., Kolby, J. & Kielgast, J. (2011) Foraging acrobatics of Toxicodryas blandingii in the Democratic Republic of the Congo. Herpetology Notes, 4, 91 - 92.","Loveridge, A. (1937) Zoological results of the George Vanderbilt African expedition of 1934. Part VII - Reptiles and amphibians. Proceedings of the Academy of Natural Sciences of Philadelphia, 89, 265 - 296.","Luiselli, L., Akani, G. C. & Barieenee, I. F. (1998 a) Observations on habitat, reproduction and feeding of Boiga blandingi (Colubridae) in south-eastern Nigeria. Amphibia-Reptilia, 19 (4), 430 - 436. https: // doi. org / 10.1163 / 156853898 X 00098","Laurent, R. F. (1954) Reptiles et batraciens de la region de Dundo (Angola). Deuxieme note. Publicacoes Culturais da Compan- hia de Diamantes de Angola, 23, 35 - 84.","Laurent, R. F. (1960) Notes complementaires sur les cheloniens et les ophidiens du Congo Oriental. Annales du Musee Royal du Congo Belge, Serie in- 8 °, 84, 1 - 86.","de Witte, G. - F. (1962) Genera des serpents du Congo et du Ruanda-Urundi. Annales du Musee Royal de l'Afrique Centrale, Sciences Zoologique, Serie in Octavo, 104, i - viii + 1 - 203.","Thys van den Audenaerde, D. F. E. (1965) Les serpents des environs de Leopoldville. Revue de zoologie et de botanique africaines, 72 (3 - 4), 366 - 388.","Broadley, D. G. & Cotterill, F. P. D. (2004) The reptiles of southeast Katanga, an overlooked ' hot spot. ' African Journal of Herpetology, 53 (1), 35 - 61. https: // doi. org / 10.1080 / 21564574.2004.9635497","Boulenger, G. A. (1896) Catalogue of the Snakes in the British Museum (Natural History). Vol. III. Containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae, and Viperidae. Trustees of the British Museum (Natural History), London, xiv + 727 pp.","Parker, H. W., Moreau, R. E. & Pakenham, R. H. W. (1940) XXV. - Notes on the herpetology of the East African Islands. Journal of Natural History, 5 (27), 309 - 314. https: // doi. org / 10.1080 / 00222934008527048","Loveridge, A. (1957) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya, Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, Harvard, 117 (2), 151 - 362 + i - xxxvi (index).","Wakeman, B. N. (1966) Uganda's poisonous snakes: Further observations on feeding habits. The Uganda Journal, 30, 101 - 103","Spawls, S. (1979) Sun, Sand & Snakes. William Morrow and Company, Inc., New York, New York, 254 pp.","Weinstein, S. A. & Smith, L. A. (1993) Chromatographic profiles and properties of Duvernoy's secretions from some boigine and dispholidine colubrids. Herpetologica, 49 (1), 78 - 94.","Weinstein, S. A. & Kardong, K. V. (1994) Properties of Duvernoy's secretions from opisthoglyphous and aglyphous colubrid snakes. Toxicon, 32 (10), 1161 - 1185. https: // doi. org / 10.1016 / 0041 - 0101 (94) 90347 - 6","Branch, W. R. (2005) A Photographic Guide: Snakes Other Reptiles and Amphibians of East Africa. Ralph Curtis Publishing, Inc., Sanibel Island, Florida, 132 pp.","Dashevsky, D., Debono, J., Rokyta, D., Nouwens, A., Josh, P. & Fry, B. G. (2018) Three-finger toxin diversification in the venoms of cat-eye snakes (Colubridae: Boiga). Journal of Molecular Evolution, 86, 531 - 545. https: // doi. org / 10.1007 / s 00239 - 018 - 9864 - 6"]}

Leptopelis viridis (G��nther, 1869) (Fig. 3D) COLLECTION MATERIAL. ��� Burkina Faso ��� 2♀ (SVL 38.7-41.7 mm); Kadiogo province, Pabr��; 12.51602��N, ��� 1.59691��W; 300 m a.s.l.; M. Lamotte leg.; MNHN-RA-1999.7060, 1999.7061. FIELD MATERIAL. ��� Burkina Faso ��� 10 ♂ (SVL 30.5-37.9 mm); Kadiogo province, Pabr�� dam; 12.51605��N, ��� 1.59689��W; 290 m a.s.l.; 12.VII.2017; H. J. Ayoro & A. Ohler leg.; bank of Pabr�� dam; hand capture ��� 3 ♂ (SVL 36.1-38.7 mm); Oubritenga province, Loumbila dam; 12.49829��N, ��� 1.40310��W; 275 m a.s.l.; 14.VII.2017; same collectors as for preceding; around a plantain field downstream from the Loumbila dam; hand capture ��� 5 ♂ (SVL 32.9-36.1 mm); Koubri, ���Ancien barrage���; 12.22883��N, ��� 1.34522��W; 290 m a.s.l.; 16.VII.2017; same collectors as for preceding; in a maize field on the bank of ���Ancien barrage��� ��� 1 ♀ (SVL 54.5 mm); Koubri, the inn���s court of Monastery; 12.21685��N, ��� 1.35041��W; 285 m a.s.l.; 17.VII.2017; same collectors as for preceding; under shrubs; hand capture ��� 1 ♂; Kadiogo province, town hall of Saaba; 12.37209��N, ��� 1.40374��W; 305 m a.s.l.; 30.VII.2018; H. J. Ayoro leg.; on shrubs near habitations; acoustic recording ��� 2 ♂ (SVL 33.9-36.5 mm); Arly National Park, Bouliabou pond; 11.57890��N, 1.46393��E; 170 m a.s.l.; 12. VI.2018; same collector as for preceding; on a shrub; hand capture ��� 6 ♂ (SVL 32.2- 33.8 mm); Arly National Park, Bongola swampy valley; 11.53255��N, 1.48244��E; 170 m a.s.l.; 13. VI.2018; same collector as for preceding; on a shrub; hand capture ��� 3 ♀ (SVL 39.3-42.3 mm); Arly National Park, a floodplain (500 m north of Bouliabou pond); 11.57852��N, 1.46000��E; 170 m a.s.l.; 16. VI.2018; same collector as for preceding; on the edge of a small water body; hand capture ��� 1 ♂ (SVL 33.8 mm), 2 ♀ (SVL 45.3-45.7 mm); W National Park, Kabougou River (500 m north-west of Kabougou forest post office); 11.97062��N, 2.01624��E; 280 m a.s.l.; 18. VI.2018; same collector as for preceding; on the edge of a pond; hand capture ��� 1 ♂ (SVL 34.4 mm), 1 ♀ (SVL 39.8 mm); W National Park, Nangouli swampy valley; 11.93663��N, 1.99461��E; 310 m a.s.l.; 21. VI.2018; same collector as for preceding; around a small pond; hand capture ��� 1 ♂; Deux Bal��s National Park, downstream from the Soumbou dam; 11.69831��N, ��� 2.79705��W; 260 m a.s.l.; 8.XI.2017; same collector as for preceding; hand capture ��� 2 ♂ (SVL 30. 8-31.2 mm), 1 ♀ (SVL 40.1 mm); Tuy province, Koti; 11.38783��N, ��� 3.06404��W; 300 m a.s.l.; 23.VIII.2018; same collector as for preceding; near a maize field; hand capture ��� 12 ♂ (SVL 31.3-36.4 mm), 2 ♀ (SVL 43.1-43.6 mm); Como��-L��raba Forest, Baboro swampy valley; 9.92523��N, ��� 4.57562��W; 240 m a.s.l.; 4.VIII.2018; H. J. Ayoro & A. Ohler leg.; on shrubs; hand capture ��� 3 ♂ (SVL 29.5-33.5 mm); Como�� province, bank of Folonzo village pond; 9.95775��N, ��� 4.67451��W; 240 m a.s.l.; 6.VIII.2018; same collectors as for preceding; on grasses, hand capture ��� 1 ♂ (SVL 35.6 mm), 1 ♀ (SVL 48.7 mm); Nasso, Kou Forest; 11.18812��N, ��� 4.44390��W; 320 m a.s.l.; 9.VIII.2018; same collectors as for preceding; near a small water body; hand capture ��� Ioba province, Bontioli reserve; 10.93685��N, ��� 3.13801��W; 260 m a.s.l.; 2011; J. Penner leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Passor�� province, Yako; 12.96740��N, ��� 2.26151��W; 330 m a.s.l.; 1961 (Schi��tz 1967; B��hme et al. 1996) ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 58 specimens examined. Size of this species 33- 35 mm for males and 42-48 mm for females, according to Schi��tz (1967; 1999) and R��del (2000). Size slightly larger in our voucher specimens. Leptopelis with smoother skin, longer extremities and larger discs on toes and fingers (Fig. 3C, D). Dorsal colouration brownish or reddish brown with darker spots forming a mark of the shape of a reversed ���V���. Venter whitish. juveniles colour mostly green. For detailed descriptions, see Lamotte & Xavier (1981), Schi��tz (1967, 1999), R��del (2000) and Amiet (2012). HABITAT. ��� The species was fossorial during the day and arboreal at night. Leptopelis viridis was found in herbaceous and wooded savannahs (Fig. 2C, E). Males were caught on trees and shrubs around ponds and rivers at heights varying between 0.5- 3 m. In contrast to males, females were collected on the ground of the same habitats., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 552-554, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","AMIET J. L. 2012. - Les rainettes du Cameroun. Edition J. L. Amiet et La Nef des Livres, Nyons & Saint Nazaire, France, 591 p."]}

Amnirana galamensis (Duméril & Bibron, 1841) COLLECTION MATERIAL. — Burkina Faso • 1 ♀ (SVL 62.3 mm); Sanemantenga province, Kossoghin; 13.29944°N, – 1.05095°W; 320 m a.s.l.; VII.1962; M. Lamotte leg.; MNHN-RA-1995.5766 • 1 ♂ (SVL 61.3 mm); Samba; 12.70144°N, – 2.40390°W; 290 m a.s.l.; 13.VII.1962; M. Lamotte leg.; MNHN-RA-2006.2754 • 1 ♀ (SVL 73.6 mm), 1♂ (SVL 69.3 mm); Sanemantenga province, Vourbira (current name Gouropila); 12.72553°N, – 1.75252°W; 310 m a.s.l.; 10.VII.1961; M. Lamotte leg.; MNHN-RA-2006.2759, 2006.2750 • 1 subadult (SVL 41.1 mm); Fada N’Gourma; 12.06051°N, 0.34910°E; 290 m a.s.l.; II.1983; H. Meyer leg.; ZFMK 38700. FIELD MATERIAL. — Burkina Faso • 6 ♂ (SVL 63.8-66.7 mm), 2♀ (SVL 70.0- 71.9 mm); Pabré dam; 12.51122°N, – 1.59225°W; 290 m a.s.l.; 10.VII.2017; H. J. Ayoro & A. Ohler leg.; around the irrigation ditches; hand capture • 1 ♂ (SVL 68.6 mm); Koubri, “Ancien barrage”; 12.22843°N, – 1.34766°W; 280 m a.s.l.; 16.VII.2017; same collectors as for preceding; near water body; hand capture • 1 ♂ (SVL 55.1 mm), 8 subadults (SVL 27.6-44.8 mm); Deux Balés National Park, around “Baignade des Eléphants”; 11.71809°N, – 2.83793°W; 270 m a.s.l.; 8.XI.2017; H. J. Ayoro leg.; under dry grasses near water body; hand capture • 3 subadults (SVL 37.4-46.1 mm); Tiona pond; 11.63241°N, – 2.85437°W; 265 m a.s.l.; 9.XI.2017; same collector as for preceding; hand capture • 2 ♀ (SVL 54.3-68.1 mm); Mouhoun River bank; 11.77879°N, – 2.91244°W; 245m a.s.l.; 10.XI.2017; same collector as for preceding; near water body; hand capture • 1 subadult (SVL 41.6 mm); the backwater of Koti; 11.38206°N, – 3.05143°W; 305 m a.s.l.; 23.X.2017; same collector as for preceding; in a small water well; hand capture • 1 ♂ (SVL 54.1 mm); Arly National Park, the central forest post office; 11.57532°N, 1.45443°E; 190 m a.s.l.; 16. VI.2018; same collector as for preceding; on a trail; hand capture • 2 ♂ (SVL 58.3-61.5 mm), 3 ♀ (SVL 64.7-73.5 mm); W National Park, Kabougou River; 11.96860°N, 2.01259°E; 280 m a.s.l.; 18. VI.2018; same collector as for preceding; in a paddy field; hand capture • 2 ♂ (SVL 57.9-65.3 mm), 1 ♀ (SVL 62.9 mm); the protected part of Kabougou River; 11.97060°N, 2.01627°E; 265 m a.s.l.; 19. VI.2018; same collector as for preceding; under grasses; hand capture • 2 ♂ (SVL 64.3-65.7 mm), 1 ♀ (SVL 64.1 mm); around PK-27 bouli; 11.88744°N, 2.11913°E; 265 m a.s.l.; 20. VI.2018; same collector as for preceding; near water body; hand capture • 2 ♂ (SVL 59.4- 60.1 mm); Nangouli swampy valley; 11.89793°N, 2.00091°E; 250 m a.s.l.; 21. VI.2018; same collector as for preceding; under grasses; hand capture • 4 ♂ (SVL 68.9 -. 3 mm), 1 ♀ (SVL 75.6 mm); the flooded zone of the Comoé River (300 m west of Comoé bridge); 9.93239°N, – 4.60655°W; 220 m a.s.l.; 5.VIII.2018; H. J. Ayoro & A. Ohler leg.; under shrubs near water body; hand capture • 3 juveniles (SVL 26.0-27.0 mm); the edge of Dori pond, Toroodi; 11.96092°N, 1.99709°W; 280 m a.s.l.; 23.VIII.2018; H. J. Ayoro leg.; near water body; hand capture • Djikologo, close to Bontioli reserve; 10.93684°N, – 3.13800°W; 260 m a.s.l.; J. Penner leg. • Dano; 11.16601°N, – 3.08691°W; 340 m a.s.l.; 2011; J. Penner leg.; field observation. LITERATURE REPORTS. — Burkina Faso • Bobo-Dioulasso, Dahora; coordinates not available; 14.VI.1992 (Böhme et al. 1996) • Gourma province, Fada N’gourma; 12.06051°N, 0.34913°E; 300 m a.s.l.; II.1983, I-III.2008 (Böhme et al. 1996; Mohneke et al. 2011) • Ganzourgou province, Mogtedo; 12.31844°N, – 0.83601°W; 280 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) • Zorgho; 12.2483°N, – 0.6278°W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011) • Houet province, Karankasso Vigué; 11.00982°N, – 3.90001°W; 300 m a.s.l.; 1985 (Böhme & Heath 2018). DESCRIPTION. — 51 specimens studied. Males 54.1-71.3 mm and females 54.3-75.6 mm. Body moderately wide and a wide head. Snout relatively pointed; skin smooth, but flat warts present on flanks and the posterior part of the back. Easily recognized by the colour pattern and distinct, broad white or yellow dorsolateral ridges (Rödel 2000). Colouration in alcohol dark brown with two white dorsolateral ridges and some grey spots on the limbs. Belly white. Males with a pair of lateral vocal sacs and large glands on the dorsal side of the upper arm. HABITAT. — We recorded A. galamensis in many habitats, i.e. agricultural zones, in water filled depressions of shrubby, grassy and wooded savannah. During the rainy season, we encountered the species around ponds and small water bodies in areas with sand and clay. subadult specimens were caught between cracks and small hollows at the edge of a permanent pond. The species occurs mostly along more permanent, larger waters in humid and dry savannah, including agricultural areas (Rödel 2000). In the dry season, this species can be found along river banks. The distribution area of the species, as currently understood, extends from extreme western Africa across to the horn of Africa and south into Angola, Mozambique, Zambia and Zimbabwe (Rödel & Bangoura 2004; Segniagbeto et al. 2007; Mohneke et al. 2011; Onadeko 2016; Böhme & Heath 2018). However, results by Jongsma et al. (2018) indicate that East African populations are not conspecific with those from West Africa., Published as part of Ayoro, Halamoussa Joëlle, Segniagbeto, Gabriel Hoinsoudé, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, Rödel, Mark-Oliver, Kabré, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 574-575, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","BOHME W. & HEATH J. 2018. - Amphibian and reptilian records from south-central Mali and western Burkina Faso. Bonn zoological Bulletin 67: 59 - 69.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90.","ONADEKO B. A. 2016. - Distribution, diversity and abundance of anuran species in three different vegetations habitats in southwestern Nigeria. Ethiopian Journal of environmental Studies and Management 9: 22 - 34. http: // dx. doi. org / 10.4314 / ejesm. v 9 i 1.3","JONGSMA G. F. M., BAREJ F. M., BARRATT D. C., BURGER M., CONRADIE W., ERNST R., GREENBAUM E., HIRSCHFELD M., LEACHE D. A., PENNER J., PORTIK M. D., ZASSI- BOULOU A. G., RODEL M. - O. & BLACKBURN C. D. 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"]}

Ptychadena trinodis (Boettger, 1881) COLLECTION MATERIAL. ��� Burkina Faso ��� 4♀ (SVL 52.5-55.2 mm); Ouagadougou; 12.38011��N, ��� 1.51100��W; 300 m a.s.l.; VIII.1966; P��re Terrible leg.; MNHN- RA-1995.3233, 1995.3234, 1995.3235, 1995.3236 ��� 5 ♀ (SVL 48.1-56.9 mm), 1 ♂ (SVL 51.8 mm); Loumana; 10.58001��N, ��� 5.35044��W; 330 m a.s.l.; 3.IX.1970; J.J. Mor��re leg.; MNHN-RA-1996.3931, 1996.3942, 1996.3943, 1996.3944, 1996.3945, 1996.3946 ��� 2 ♀ (SVL 54.1-57.2 mm), 1♂ (SVL 55.7 mm); a trail between Bobo-Dioulasso and Kofila; 11.33755��N, ��� 3.92852��W; 290 m a.s.l.; 7.XI.1970; J. J. Mor��re leg.; MNHN-RA-1996.3939, 1996.3940, 1996.3941 ��� 2 subadults (SVL 30.2-34.7 mm); Samba; 12.70144��N, ��� 2.40391��W; 290 m a.s.l.; 2.I.1963; M. Lamotte leg.; MNHN-RA-1993.3572, 1993.5679. FIELD MATERIAL. ��� Burkina Faso ��� 1 ♀ (SVL 55.5 mm); Deux Bal��s National Park, downstream from the Soumbou dam; 11.69872��N, ��� 2.79738��W; 260 m a.s.l.; 8.XI.2017; H. J. Ayoro leg.; under a stone near water body; hand capture ��� 1 ♂ (SVL 53.6 mm); Arly National Park; 11.57044��N, 1. 45211��E; 190 m a.s.l.; 12. VI.2018; same collector as for preceding; near a small water body on a trail; hand capture ��� 1 ♂; Koubri, downstream from Nabazana dam; 12.20319��N, ��� 1.35105��W; 280 m a.s.l.; 17.VII.2017; H. J. Ayoro & A. Ohler leg.; in a paddy field; acoustic report ��� 1 ♀ (SVL 52.1 mm); Como��-L��raba Forest, Kotokoi swampy valley; 9.82802��N, ��� 4.62727��W; 230 m a.s.l.; 3.VIII.2018; same collectors as for preceding; near a small water pool; hand capture ��� 1 subadult (SVL 42.4 mm); Baboro swampy valley; 9.92521��N, ��� 4.5739��W; 230 m a.s.l.; 4.VIII.2018; same collectors as for preceding; on a trail; hand capture ��� 1 ♀ (SVL 59.1 mm), 6 subadults (SVL 35.7-45.2 mm); Folonzo village swampy valley; 9.88803��N, ��� 4.61969��W; 230 m a.s.l; 6.VIII.2018; same collectors as for preceding; around a vegetable garden; hand capture ��� 18 subadults (SVL 25.0- 44.1 mm); the flooded zone of Yakouta dam; 14.06117��N, ��� 0.13181��W; 270 m a.s.l.; 26.VIII.2018; H. J. Ayoro leg.; under grasses; hand capture ��� 4 subadults (SVL 19.1-46.4 mm); the edge of Dori pond, Toroodi; 14.00607��N, ��� 0.02846��W; 280 m a.s.l.; 24-27.VIII.2018; same collector as for preceding; under grasses; hand capture ��� 4 subadults (SVL 31.1- 46.3 mm); the edge of N���Djomga pond; 14.07225��N, ��� 0.04862��W; 265 m a.s.l.; 25.VIII.2018; same collector as for preceding; under grasses near water body; hand capture ��� 3 ♀ (SVL 52.3-57.3 mm); the edge of Fafo dam; 11.33470��N, ��� 3.01947��W; 300 m a.s.l.; 10.XI.2018; same collector as for preceding; in paddy field; hand capture ��� 1 ♀ (SVL 59.8 mm); Djindjerma, close to Koti; 11.38211��N, ��� 3.05145��W; 310 m a.s.l.; 1.XI.2018; same collector as for preceding; around a drink water well; hand capture. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 54 specimens studied. Stout and robust frogs, with a moderately pointed snout, strong and long limbs. An inner and an external metatarsal tubercle present and very protruding as indicated by R��del (2000) and S��nchez-Vialas et al. (2017). Webbing formula: I 0 - 1 �� II 0 - 1 III 0 - 1 �� IV 1 �� - 0 V. Four pairs of continuous dorsal ridges extending along the entire back. In life and alcohol, dorsal colouration from brown to dark grey. Presence of a wide white, yellow in life, vertebral line in all specimens. Species usually with a more granular dorsal skin, in comparison to other members of this genus (R��del 2000). HABITAT. ��� Specimens were captured in grasses near water bodies in Como��-L��raba Forest and Deux Bal�� National Park; on a trail near a water body in Arly National Park; under humid gasses at Dori pond and Yakouta dam banks. Ptychadena trinodis was one of the most abundant Ptychadena species at savannah ponds during the rainy season (Nago et al. 2006). The species is then virtually restricted to small or tiny water bodies which often lack vegetation (Guib�� & Lamotte 1957; Lamotte 1969; Schi��tz 1967; Perret 1981; R��del 2000). This species is distributed from West Africa east to northern Democratic Republic of Congo (R��del 2000; Padial & De La Riva 2004; S��nchez-Vialas et al. 2017; B��hme & Heath 2018)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 572-573, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","GUIBE J. & LAMOTTE M. 1957. - Revision systematique des Ptychadena (Batraciens Anoures Ranides) d'Afrique Occidentale. Bulletin de l'Institut fondamental d'Afrique noire, (A) 19: 937 - 1003.","LAMOTTE M. 1969. - Le parc national du Niokolo-Koba, Fascicule III; XXX. Amphibiens (deuxieme note). Memoire de l'Institut fondamental de l'Afrique noire (A) 84: 420 - 426.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","PERRET J. L. 1981. - Le statut de Ptychadena schubotzi (Sternfeld, 1917) (Amphibia, Ranidae). Bulletin de la Societe Neuchateloise des Sciences naturelles 104: 53 - 57.","PADIAL J. M. & DE LA RIVA I. 2004. - Annotated checklist of the amphibians of Mauritania (West Africa). Revista Espanola de Herpetologia 18: 89 - 99.","BOHME W. & HEATH J. 2018. - Amphibian and reptilian records from south-central Mali and western Burkina Faso. Bonn zoological Bulletin 67: 59 - 69."]}

Phrynomantis microps Peters, 1875 (Fig. 5D) COLLECTION MATERIAL. ��� Burkina Faso ��� 4 ♂ (SVL 38.2- 38.6 mm); Kourweogo province, Laye; 12.53081��N, ��� 1.77155��W; 290 m a.s.l.; VIII.1968; M. Lamotte leg.; MNHN-RA-2000.5667, 2000.5660, 2000.5657, 2000.5659 ��� 1 ♂ (SVL 41.1 mm); Sanemantenga province, Kossoghin; 13.29944��N, ��� 1.05088��W; 320 m a.s.l.; VIII.1966; M. Lamotte leg.; MNHN-RA-1995.7090. FIELD MATERIAL. ��� Burkina Faso ��� 1 ♂ (SVL 38.2 mm); Arly National Park, the floodplain; 11.57852��N, 1.46004��E; 255 m a.s.l.; 16. VI.2018; H. J. Ayoro leg.; near a small water body; hand capture ��� 1 ♀ (SVL 42.5 mm); Koti; 11.38788��N, ��� 3.06401��W; 310 m a.s.l.; 15.IX.2018; same collector as for preceding; in a peanut field; hand capture ��� 1 ♂ (SVL 42.3 mm), 1 ♀ (SVL 51.4 mm); Folonzo village swampy valley; 9.95777��N, ��� 4.67455��W; 240 m a.s.l.; 6.VIII.2018; H. J. Ayoro & A. Ohler leg.; anthill and scorpion���s holes; hand capture ��� 2 ♂; on the edge of Pabr�� dam; 12.51597��N, ��� 1.59923��W; 290 m a.s.l.; 11-12.VII.2017; same collectors as for preceding; under the trees near the dam; acoustic report ��� 1 ♂; near the ��� Ancien barrage���; 12.22843��N, ��� 1.34766��W; 285 m a.s.l.; 16.VII.2017; same collectors as for preceding; acoustic report ��� 1 ♂; in the Koubri monaster reserve around the Monastery Hostel; 12.21687��N, ��� 1.35048��W; 280 m a.s.l.; 17.VII.2017; same collectors as for preceding; acoustic report ��� 1♂; W National Park, near PK-27 bouli; 11.88744��N, 2.11913��W; 265 m a.s.l.; 20. VI.2018; H. J. Ayoro leg.; acoustic report ��� 1 ♂; the Kabougou village swampy valley; 11.96075��N, 1.99744��E; 280 m a.s.l.; 22. VI.2018; same collector as for preceding; near paddy field; acoustic report ��� 3 ♂; Kou Forest; 11.18888��N, ��� 4.43944��W; 260 m a.s.l.; 7-9.VIII.2018; H. J. Ayoro & A. Ohler leg.; in the savannah part and at the edge of this forest; acoustic report ��� Ioba province, Djikologo (close to Bontioli reserve); 10.94253��N, ��� 3.15788��W; 260 m a.s.l.; J. Penner leg. ��� Dano, Gbagba; 11.18133��N, ��� 3.05159��W; 300 m a.s.l.; 2011; J. Penner leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Yako; 12.96744��N, ��� 2.26155��W; 330 m a.s.l.; 18.VIII.1961 (Schi��tz 1967) ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l.; 1983, I-III.2008 (B��hme et al. 1996; Mohneke et al. 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 9 specimens examined. Size within the range given by R��del (2000). Characterized by a unique body shape and a red back (Fig. 5D). Presence of a fine dark brown vertebral line on the back. In alcohol, the red colouration getting a greyish. HABITAT. ��� The species calls often below shrubs and from holes. In Como��-L��raba Forest, males were collected in shrubs on the edge of a large dark scorpion hole where they were calling. It was also found on the edge of water under a tree. A female was found on an anthill ready to withdraw into a hole. The species has been reported to often occur within ant nests and scorpion burrows (R��del & Braun 1999; R��del et al. 2013). It is known that P. microps colonizes various savannah habitats including dense tree savannah where it is found in flooded meadows, in ponds and swamps (Schi��tz 1967; R��del 2000; Hirschfeld & R��del 2011). Barej et al. (2017) reported its occurence in West and Central Africa., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 562, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","RODEL M. - O. & BRAUN U. 1999. - Associations between anurans and ants in a West African savanna (Anura: Microhylidae, Hyperoliidae and Hymenoptera: Formicidae). Biotropica 31: 178 - 183. https: // doi. org / 10.2307 / 2663971","RODEL M. - O., BREDE C., HIRSCHFELD M., SCHMITT T., FAVREAU P., STOCKLIN R., WUNDER C. & MEBS D. 2013. - Chemical camouflage - A frog's strategy to co-exist with aggressive ants. PLoS ONE 8: e 81950. https: // doi. org / 10.1371 / journal. pone. 0081950","HIRSCHFELD M. & RODEL M. - O. 2011. - Variable reproductive strategies of an African savanna frog, Phrynomantis microps (Amphibia, Anura, Microhylidae). Journal of tropical Ecology 27: 601 - 609. https: // doi. org / 10.1017 / S 0266467411000320","BAREJ M. F., SANDBERGER- LOUA L., DOUMBIA J., KOUAME N. G., RODEL M. - O. & PENNER J. 2017. - Microhylidae: Phrynomantis microps (Peters, 1875), West African rubber frog in Guinea. African Herp News 64: 37 - 39."]}

Arthroleptis poecilonotus Peters, 1863 (Fig. 3A, B) COLLECTION MATERIAL. ��� Burkina Faso ��� 10 subadults (SVL 12.1- 20.1 mm); Bobo-Dioulasso, near Nasso; 11.18803��N, ��� 4.29741��W; 420 m a.s.l.; VII.1970; M. Lamotte leg.; MNHN-RA-2008.1055, 2008.1056, 2008.1057, 2008.1058, 2008.1059, 2008.1060, 2008.1061, 2008.1062, 2008.1063, 2008.1064. FIELD MATERIAL. ��� Burkina Faso ��� 2 ♀ (SVL 27.6-28.1 mm), 5 ♂ (SVL 24.4-26.5 mm), 3 subadults (SVL 16.6-18.3 mm); Nasso, Kou Forest; 11.18888��N, ��� 4.43935��W; 320 m a.s.l.; 7-9.VIII. 2018; H. J. Ayoro & A. Ohler leg.; gallery forest; hand capture. DESCRIPTION. ��� 20 specimens examined. Small frogs with a blunt snout, short head and a concave tympanum; slender limbs, the inner metatarsal tubercle long and prominent. Adult males and females with protruding palmar and supernumerary tubercles on fingers. Tubercles absent on toes. Dorsum feebly granulated. In life, dorsum grey, brown or reddish-brown coloured. Sometimes an hourglass pattern present on back (Fig. 3A, B). Venter with whitish or brownish, smooth skin. As other Arthroleptis species (e.g. Blackburn 2009; Blackburn et al. 2010), presence of an elongated third finger, and transparent spines on second and third fingers in adult males. HABITAT. ��� The historical specimens and the recent samples of A. poecilonotus were from Kou Forest (Fig. 2E). At this site, males were seen at night on leaves of shrub and grass at heights varying from 0.2- 1 m. Females and subadults were seen hiding in leaflitter. The species may occur in other gallery forests along the major rivers (Mouhoun, Como�� and Pendjari rivers) in the western and southern part of Burkina Faso as well (Lamotte 1967b). It was found in a forest close to a waterfall near Pendjari National Park in Benin (Nago et al. 2006), in Togo (Segniagbeto et al. 2007), in Ivory Coast (Lamotte & Perret 1963; Lamotte 1967b; Barbault & Trefaut-Rodrigues 1979; R��del 2000; R��del 2003; R��del & Ernst 2003) and in Nigeria (Schi��tz 1964, 1967; Leach�� et al. 2006; Onadeko 2016). TAXONOMIC REMARK. ��� As stated in R��del & Bangoura (2004) and Blackburn (2010), it is currently not possible to assign these West African Arthroleptis taxa with certainty to one of the various available names, as intra-specific variability equals or surpasses interspecific differences. As actually understood, A. poecilonotus has an extended distribution range, including West, Central and parts of East Africa (Frost 2019) and probably recovers a group of species. Type locality is in West Africa, in south-eastern Ghana. Further studies, in particular including topotypical material, are needed for taxonomic decisions., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 552, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["BLACKBURN D. C. 2009. - Diversity and evolution of male secondary sexual characters in African squeakers and long-fingered frogs. Biological Journal of the Linnean Society 96: 553 - 573. https: // doi. org / 10.1111 / j. 1095 - 8312.2008.01138. x","BLACKBURN D. C., GVOZDIK V. & LEACHE A. D. 2010. - A new squeaker frog (Arthroleptidae: Arthroleptis) from the mountains of Cameroon and Nigeria. Herpetologica 66: 335 - 348. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 10 - 00015.1","LAMOTTE M. 1967 b. - Les batraciens de la region de Gbakobo (Cote d'Ivoire). Memoire de l'Institut fondamental de l'Afrique noire (A) 29: 218 - 294.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90.","LAMOTTE M. & PERRET J. L. 1963. - Contribution a l'etude des batraciens de l'Ouest Africain XV - Le developpement direct de l'espece Arthroleptis poecilonotus Peters. Bulletin de l'Institut fondamental d'Afrique noire (A) 25: 277 - 284.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","RODEL M. - O. & ERNST R. 2003. - The amphibians of Marahoue and Mont Peko National Parks, Ivory Coast. Herpetozoa 16: 23 - 39.","SCHIOTZ A. 1964. - A preliminary list of amphibians collected in Ghana. Videnskabelige Meddelelser fra dansk Naturhistorik Forening 127: 1 - 17.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","LEACHE A., RODEL M. - O., LINKEM C. W., DIAZ R. E., HILLERS A. & FUJITA M. K. 2006. - Biodiversity in a forest island: reptiles and amphibians of the West African Togo hills. Amphibian and Reptile Conservation 4: 22 - 45. https: // doi. org / 10.1514 / journal. arc. 0040018","ONADEKO B. A. 2016. - Distribution, diversity and abundance of anuran species in three different vegetations habitats in southwestern Nigeria. Ethiopian Journal of environmental Studies and Management 9: 22 - 34. http: // dx. doi. org / 10.4314 / ejesm. v 9 i 1.3","FROST D. R. 2019. - Amphibian Species of World: an online reference. Version 6.0. Available at htt: // research. amnh. org / herpetology / amphibia / index. html. Accessed on 15 February 2019 and 2 November 2020."]}

Kassina senegalensis (Dum��ril & Bibron, 1841) (Fig. 5B, C) COLLECTION MATERIAL. ��� Burkina Faso ��� 2 ♂ (SVL 31.5 - 34.0 mm); Kourweogo province, Laye; 12.53079��N, ��� 1.77155��W; 290 m a.s.l.; VIII.1966; M. Lamotte leg.; MNHN-RA-1998.2009, 1998.2010. FIELD MATERIAL. ��� Burkina Faso ��� 1 ♂ (SVL 31.3 mm); W National Park, on the edge of the PK-27 bouli; 11.88744��N, 2.11913��W; 265 m a.s.l.; 20. VI.2018; H. J. Ayoro leg.; under shrubs; hand capture ��� 1 ♂ (SVL 32.2 mm); Como��-L��raba Forest; 9.93239��N, ��� 4.60655��W; 220 m a.s.l.; 5.VIII.2018; H. J. Ayoro & A. Ohler leg.; under grasses; hand capture ��� 4♂ (SVL 34.9-39.4 mm), 3 subadults (SVL 20.6-22.4mm); Dori, Toroodi (near the road from Dori to Ouagadougou); 14.00385��N, ��� 0.02411��W; 270 m a.s.l.; 24.VIII.2018; H. J. Ayoro leg.; in small holes and in dried footprints of oxen near a water body; hand capture ��� 1 ♀ (SVL 32.0 mm), 2 subadults (SVL 24.3-27.6 mm); N���Djomga (on the road from Dori to Gorom-Gorom); 14.0722��N, ��� 0.0486��W; 265 m a.s.l.; 25.VIII.2018; same collector as for preceding; under a heap of dead grasses in a vegetable garden; hand capture ��� 1 subadult (SVL 22.5 mm); Yakouta; 14.06111��N, ��� 0.13185��W; 267 m a.s.l.; 26.VIII.2018; same collector as for preceding; under a shrub; hand capture ��� 1 ♂; Koubri, around ���Ancien barrage���; 12.22828��N, ��� 1.34587��W; 320 m a.s.l; 16.VII.2017; same collector as for preceding; in a bush; acoustic report ��� 2 ♂; Arly National Park, floodplain (500 m north of Bouliabou pond); 11.57852��N, 1.46004��E; 255 m a.s.l.; 16. VI.2018; same collector as for preceding; acoustic report ��� 1 ♂; Kou Forest; in the savannah part of this Forest; 11.18815��N, ��� 4.44397��W; 375 m a.s.l.; 9.VIII.2018; H. J. Ayoro & A. Ohler leg.; in a bush; acoustic report ��� Dano, Gbagba; 11.18133��N, ��� 3.05163��W; 300 m a.s.l.; 2011; J. Penner leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Yako; 12.96744��N, ��� 2.26155��W; 330 m a.s.l.; 18.VIII.1961 (Schi��tz 1967). DESCRIPTION. ��� 15 specimens examined. Males from Laye, Como��-L��raba forest and W National Park measuring 31.3- 34.0 mm SVL. Males of Dori and Yakouta atteining larger size, 34.9-39.4 mm SVL. These sizes within the known size range of the species (Lamotte & Xavier 1981; Schi��tz 1999; R��del 2000; Channing 2001; Channing & Howell 2006; Amiet 2012). Dorsal colouration of K. senegalensis a light-green, yellow or green olive, usually with three dark longitudinal strips, either thin continuous or broken into dark broad blotches (Fig. 5B, C). HABITAT. ��� Kassina senegalensis are fast moving among grasses and shrubs. Two vouchers were collected under bushy grasses from where they called and were difficult to localize. In Dori and Yakouta area, K. senegalensis was found on the ground under piles of pebbles near water, as well as in small holes or cattle footprints between grasses close to a pond. We heard other males calling from the base of thorny shrubs and piles of dry grass near a millet field. Kassina senegalensis was also found in Sahelian savannahs in Mauritania (Padial et al. 2013). It shelters under rocks or termite mounds during dry periods (Poynton & Broadley 1987; R��del 2000)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 562, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","CHANNING A. 2001. - Amphibians of Central and Southern Africa. Cornell University Press, New York, USA, 470 p.","CHANNING A. & HOWELL K. M. 2006. - Amphibians of East Africa. Cornell University Press, New York, USA, 432 p.","AMIET J. L. 2012. - Les rainettes du Cameroun. Edition J. L. Amiet et La Nef des Livres, Nyons & Saint Nazaire, France, 591 p.","PADIAL J. M., CROCHET P. A., GENIEZ P. & BRITO J. C. 2013. - Amphibian conservation in Mauritania. Basic and applied Herpetology 27: 11 - 22. http: // dx. doi. org / 10.11160 / bah. 13002","POYNTON J. C. & BROADLEY D. G. 1987. - Amphibia Zambesiaca 3. Rhacophoridae and Hyperoliidae. Annals of Natal Museum 28: 161 - 229."]}

Sclerophrys regularis (Reuss, 1833) COLLECTION MATERIAL. ��� Burkina Faso ��� 6♀ (SVL 57.8-72.2 mm); Burkina Faso; XII.1959; J. Arnoult leg.; MNHN-RA-1970.646, 1970.647, 1970.648, 1970.649, 1970.650, 1970.651 ��� 4 subadults (SVL 57.4-61 mm); Houet province, Bobo-Dioulasso; 11.18801��N, ��� 4.29743��W; 420 m a.s.l.; XI.1969; M. Lamotte leg.; MNHN-RA-1995.4358, 1995.4367, 1995.4367, 1995.4373 ��� 16 ♀ (SVL 62.1-73.2 mm); Bobo-Dioulasso, Koro; 11.14430��N, ��� 4.19881��W; 350 m a.s.l.; 30.X.1970; M. Lamotte leg.; MNHNRA-1995.4355, 1995.4356, 1995.4357, 1995.4358, 1995.4359, 1995.4360, 1995.4361, 1995.4362, 1995.4363, 1995.4364, 1995.4365, 1995.4366, 1995.4369, 1995.4370, 1995.4371, 1995.4372 ��� 1 ♀ (SVL 89.1 mm); Boulgou province, Garango; 11.79981��N, ��� 0.55054��W; 280 m a.s.l.); 18. VI.1962; M. Lamotte leg.; MNHN-RA-1979.336 ��� 4 subadults (SVL 42-55.1 mm); Burkina Faso; 1987; M. Lamotte leg.; MNHN-RA-1989.3863, 1989.3864, 1989.3865, 1989.3866 ��� 1♀ (SVL 66.2 mm), 1 subadult (SVL 50.5 mm); Como�� province, Ti��fora; 10.62765��N, ��� 4.55773��W; 290 m a.s.l.; 1987; Harald Schreiber leg.; ZFMK 93968-69 ��� 1 ♂ (SVL 64.5 mm), 6 ♀ (SVL 74.1-82.3 mm); Kadiogo province, Ouagadougou; 12.38011��N, ��� 1.51105��W; 300 m a.s.l.; V.1975; NA leg.; MHNG 1537.7 - 13. FIELD MATERIAL. ��� Burkina Faso ��� 6 ♂ (SVL 77.0-92.0 mm), 1♀ (SVL 95.7 mm); Pabr�� dam; 11-12.VII.2017; H. J. Ayoro & A. Ohler leg.; around an irrigation ditch; hand capture ��� 2 ♂ (SVL 62.5- 72.2 mm), Loumbila, south-west of Loumbila dam; 13.VII.2017; same collectors as for preceding; near a water body; hand capture ��� 1 ♀ (SVL 110.9 mm); Koubri, ���Ancien barrage���; 16.VII.2017; same collectors as for preceding; on a trail near a small water body; hand capture ��� 1 ♂ (SVL 64.4 mm), 2 ♀ (SVL 66.6-88.5 mm); Kadiogo province, Saaba; 13.X.2017; H. J. Ayoro leg.; under the dustbin; hand capture ��� 1 ♂ (SVL 64.6 mm); Deux Bal��s National Park, Soumbou dam; 8.XI.2017; same collector as for preceding; on a lateritic rock near a water body; hand capture ��� 2 ♀ (SVL 60.1-74.6 mm); Deux Bal��s National Park, a flooded zone of Mouhoun River; 10.XI.2017; same collector as for preceding; on the edge of small pond; hand capture ��� 5 ♀ (SVL 59.8-86.4 mm); Koti, backwater; 23.X.2017; same collector as for preceding; on the edge of water well; hand capture ��� 1 subadult (SVL 49.7 mm); Koti; 17.VIII.2018; same collector as for preceding; on a trail; hand capture ��� 1 ♀ (SVL 82.2 mm); Arly National Park, Bouliabou pond; 16. VI.2018; same collector as for preceding; near water body; hand capture ��� 2 ♀ (SVL 69.6 - 81.0 mm); Kou Forest; 8.VIII.2018; H. J. Ayoro & A. Ohler leg.; under shrubs; hand capture ��� 1♀ (SVL 91.2 mm); Dori, Toroodi swampy valley; 24.VIII.2018; H. J. Ayoro leg.; near the tarmac road; hand capture ��� 1 ♀ (SVL 68.9 mm); Yaga province, Sebba; 9.VII.2017; A. Antarou leg.; hand capture ��� 1 ♀ (SVL 75.1 mm); same locality; 21.IX.2018; same collector as for preceding; hand capture ��� 2 ♂; D��dougou, SONATUR city (the road from D��dougou to Toughan); 22.VIII.2017; H. J. Ayoro leg.; in a gutter; direct observation ��� 1♀ (SVL 80.7 mm); Sourou province, Toughan; 28. VI.2017; R. Demb��l�� leg.; hand capture. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 69 specimens examined. Male body size from 59.0- 90.9 mm while female SVL between 57.4-95.7 mm. Sclerophrys regularis, the largest toads in Burkina Faso with long and smooth parotid glands. For detailed descriptions see Lamotte & Xavier (1981) and R��del (2000). HABITAT. ��� This is a common species and it is found in numerous habitats, ranging from protected areas, to agricultural and irrigated land and even urban areas in Burkina Faso., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 555-556, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p."]}

Leptopelis bufonides Schi��tz, 1967 (Fig. 3C) FIELD MATERIAL. ��� Burkina Faso ��� 8 ♀ (SVL 40.2-44.9 mm), 10 ♂ (SVL 30.5-37.5 mm); W National Park; 11.93664��N, 1.9946��E; 310 m a.s.l.; 21.VI.2018; H. J. Ayoro leg.; edge of savannah pond; hand capture ��� 1 ♀ (SVL 41.2 mm), 1♂ (SVL 32.1 mm); Tuy province, Koti; 11.38785��N, ��� 3.06404��W; 295 m a.s.l.; 23.VIII.2018; H. J. Ayoro leg.as for preceding;near small pond in bush; hand capture. LITERATURE REPORTS. ��� Burkina Faso ��� Passor�� province, Samba; 12.70143��N, ��� 2.40389��W; 300 m a.s.l.; 18.VIII.1961 (Schi��tz 1967) ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke et al. 2011) ��� Ganzourgou province, Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke et al. 2011). DESCRIPTION. ��� 20 specimens examined. Leptopelis bufonides with small to moderate size and warty skin, stocky body shape, and short legs. In contrast to most other members of that genus, the fingers with small discs on their tips. In life colouration greyish brown with dark spots, including a dark interorbital spot and elongated spots or bands on the back (Fig. 3C). These markings making almost a ���m��� pattern. The venter light to dark grey, with a dark throat in both sexes. For detailed descriptions, see Schi��tz (1967), R��del (2000) and Amiet (2012). HABITAT. ��� This species was usually found in grassy and wooded savannahs (Fig. 2C, D). Specimens were collected from the ground around temporary ponds with grass banks and shrubs. It was also Evans from dry savannahs in Senegal (Schi��tz 1967), Mali (B��hme & Heath 2018), Ghana (Schi��tz 1967), Benin (Nago et al. 2006), Togo (Segniagbeto et al. 2007) and Cameroon (Amiet 2012)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 552, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","AMIET J. L. 2012. - Les rainettes du Cameroun. Edition J. L. Amiet et La Nef des Livres, Nyons & Saint Nazaire, France, 591 p.","BOHME W. & HEATH J. 2018. - Amphibian and reptilian records from south-central Mali and western Burkina Faso. Bonn zoological Bulletin 67: 59 - 69.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90."]}

Xenopus fischbergi Evans et al., 2015 COLLECTION MATERIAL. ��� Burkina Faso ��� 11 tadpoles; Banfora; 10.64800��N, ��� 4.77679��W; 280 m a.s.l.; XII.1959; M. Lamotte leg.; MNHN-RA-1993.4057, 1993.4058, 1993.4059, 1993.4060, 1993.4061, 1993.4062, 1993.4063, 1993.4064, 1993.4065, 1993.4066, 1993.4067 ��� 8♀ (SVL 39.5-45.3 mm); Boulgou province, Garango; 11.79982��N, ��� 0.55055��W; 280 m a.s.l.; 18. VI.1962; M. Lamontellerie leg.; MNHN-RA-1993.4376, 1993.4377, 1993.4378, 1993.4379, 1993.4380, 1993.4381, 1993.4382, 1993.4383 ��� 3 subadults (SVL 34-36.8 mm), 1 ♀ (SVL 45.1 mm); Bobo-Dioulasso, Pala; 11.14978��N, ��� 4.22275��W; 280 m a.s.l.; XII.1970; M. Lamotte leg.; MNHN-RA-1997.0886, 1997.0887, 1997.0888, 1997.0889. FIELD MATERIAL. ��� Burkina Faso ��� 3 ♀ (SVL 39.9-47.7 mm); Koti; 11.38788��N, ��� 3.06401��W; 310 m a.s.l.; 10.X.2017; H. J. Ayoro leg.; in a small water near a garbage dump; hand capture ��� 1 ♀ (SVL 40.7 mm); same locality; 27.IX.2018; same collector as for preceding; in a small pond in a bush; hand capture ��� 2 ♀ (SVL 45.1- 46.8 mm), 2 subadults (SVL 34.3-37.1 mm); Loumbila, downstream from the Loumbila dam; 12.491656��N, ��� 1.40244��W; 280 m a.s.l.; 14.VII.2017; H. J. Ayoro & A. Ohler leg.; in an irrigated well; hand capture ��� 4 ♀ (SVL 41.7-58.5 mm), 4 subadults (SVL 23.5-36.7 mm); Tuy province, Fafo; 11.33470��N, ��� 3.01947��W; 300 m a.s.l.; 5.XI.2018; H. J. Ayoro leg.; in the backwater of Fafo; hand capture ��� 2 individuals; Ouagadougou, University of Joseph Ki-Zerbo; 12.378350��N, ��� 1.50128��W; 295 m a.s.l.; 12.VIII.2018; same collector as for preceding; in the water of the channel near the university restaurant; visual observation. LITERATURE REPORTS. ��� Burkina Faso ��� Banfora; 10.64800��N, ��� 4.77679��W; 280 m a.s.l.; Yako; 12.96744��N, ��� 2.26155��W; 330 m a.s.l. (Loumont 1984) ��� Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 39 specimens examined. Adult and subadult specimens of Xenopus fischbergi with flat head and smooth body. Presence of a long subocular tentacle and three black claws on toes. In alcohol, the back and limbs greyish to brown with few darker brown spots. The belly whitish, some darker spots occasionally on the throat and on the thighs; whitish zones usually yellow in life. Previous West African reports of this species under the name Xenopus muelleri. HABITAT. ��� We collected X. fischbergi in the water of irrigation wells, in plantain fields in Loumbila and in a small water body in Koti. These habitats were surrounded by grasses and had aquatic plants. During the dry season, X. fischbergi was found at the banks of rivers in savannah habitats (Schi��tz 1967; B��hme et al. 1996). It is known from various other West African countries such as Benin, Ghana, Ivory Coast, Nigeria and Togo (R��del 2000; Nago et al. 2006; Segniagbeto et al. 2007; Onadeko 2016)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 565, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["EVANS B. J., CARTER T. F., GREENBAUM E., GVOZDIK V., KELLEY D. B., MCLAUGHLIN P. J., PAUWELS O. S. G., PORTIK D. M., STANLEY E. L., TINSLEY R. C., TOBIAS M. L. & BLACKBURN D. C. 2015. - Genetics, morphology, advertisement calls, and historical records distinguish six new polyploid species of African clawed frog (Xenopus, Pipidae) from West and Central Africa. PLoS ONE 10: e 0142823. https: // doi. org / 10.1371 / journal. pone. 0142823","LOUMONT C. 1984. - Current distribution of the genus Xenopus in Africa and future prospects. Revue Suisse de Zoologie 91: 725 - 746. https: // doi. org / 10.5962 / bhl. part. 81578","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90.","ONADEKO B. A. 2016. - Distribution, diversity and abundance of anuran species in three different vegetations habitats in southwestern Nigeria. Ethiopian Journal of environmental Studies and Management 9: 22 - 34. http: // dx. doi. org / 10.4314 / ejesm. v 9 i 1.3"]}

Hyperolius concolor concolor (Hallowell, 1844) (Fig. 4D) COLLECTION MATERIAL. ��� Burkina Faso ��� 2♂; (SVL 27.2-28.4 mm); Houet province, Sokourani (near Hippopotamus pond); 11.51183��N, ��� 4.11611��W; 320 m a.s.l.; 8.XI.1970; M. Lamotte leg.; MNHNRA-1999.7211, 1999.7212. FIELD MATERIAL. ��� Burkina Faso ��� 2♂ (SVL 27.8-29.5 mm); Como��-L��raba Forest, Sirigou�� swampy valley; 9.88809��N, ��� 4.61954��W; 230 m a.s.l.; 2.VIII.2018; H. J. Ayoro & A. Ohler leg.; on flooded grasses; hand capture ��� 5 ♂ (SVL 30.3-31.7 mm); Como��-L��raba Forest, Baboro swampy valley; 9.92522��N, ��� 4.57575��W; 240 m a.s.l.; 4.VIII.2018; same collectors as for preceding; on high, flooded grasses; hand capture ��� 1♂ (SVL 29.7 mm); Kou Forest, near the Guinguette water source; 11.18720��N, ��� 4.44185��W; 340 m a.s.l.; 7.VIII.2018; same collectors as for preceding; on leaves of palm trees; hand capture. DESCRIPTION. ��� 10 specimens examined. Only males collected, with SVL 27.8-31.5 mm. Life colouration yellowish to greenish (Fig. 4D). Usually during the day, males more brownish to yellow, often with some dark dorsal pattern, whereas females showing a green colouration. At night individuals of both sexes with a yellow colour (R��del 2000). In alcohol, specimens whitish with small black spots, visible only with a dissecting microscope. HABITAT. ��� Hyperolius concolor concolor was found on flooded grasses in Como��-L��raba Forest and on palm leaves at the source of the Guinguette River in Kou Forest. Specimens were usually sitting comparatively higher than H. nitidulus, but see R��del et al. (2006) for calling site position in various West African hyperoliids. The species is known to occur in areas of degraded forests and humid savannah; it extends into drier savannah zones along gallery forests (Schi��tz 1999; R��del 2000). It is distributed in West Africa from Sierra Leone to northern Benin (Schi��tz 1999; R��del 2000; Nago et al. 2006)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 560, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","RODEL M. - O., LAMPERT K. P. & LINSENMAIR K. E. 2006. - Reproductive biology of the West African savannah frog Hyperolius nasutus Gunther, 1864 (Amphibia: Anura: Hyperoliidae). Herpetozoa 19: 3 - 12.","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108."]}

Afrixalus weidholzi (Mertens, 1938) (Fig. 4C) FIELD MATERIAL. ��� Burkina Faso ��� 23 ♂ (SVL 17.4-21.2 mm); Como��-L��raba Forest; 9.82802��N, ��� 4.62727��W; 230 m a.s.l; 1-7.VIII.2018; H. J. Ayoro & A. Ohler leg.; on grasses in a swampy valley; hand capture ��� 1 ♂; W National Park, Kabougou River (in its protected part); 11.96865��N, 2.0125��E; 280 m a.s.l.; 19. VI.2018; H. J. Ayoro leg.; in a bush near water body; acoustic report ��� 1♂; Koubri, downstream from the Nabazana dam; 12.20322��N, ��� 1.35101��W; 280 m a.s.l.; 17.VII.2017; H. J. Ayoro & A. Ohler leg.; close to a paddy field; acoustic report. DESCRIPTION. ��� 23 specimens examined. Frogs with SVL from 17.4-21.2 mm in 23 males, smaller than A. vittiger. Specimens with a yellow or a very intense silvery white back (Fig. 4C), often with a fine brown vertebral line, at least in the posterior part of the back. Laterally, a less distinct dark band from the snout to the groin. On the shank, presence of dark brown longitudinal bands. HABITAT. ��� This species was usually collected on grass at a height not exceeding 0.5-1.5 m (Fig. 2D). During the day, resting animals can be found above water on plants (Lamotte & Xavier 1981; Schi��tz 1967, 1999). Males call at night, usually on high grasses, a few meters from the pond edges (R��del et al. 2006). Afrixalus weidholzi is a common species which has a wide range in West African drier savannah zones (R��del 2000; Padial et al. 2013) and occurs occasionally in the same habitats as A. vittiger., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 558-560, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O., LAMPERT K. P. & LINSENMAIR K. E. 2006. - Reproductive biology of the West African savannah frog Hyperolius nasutus Gunther, 1864 (Amphibia: Anura: Hyperoliidae). Herpetozoa 19: 3 - 12.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","PADIAL J. M., CROCHET P. A., GENIEZ P. & BRITO J. C. 2013. - Amphibian conservation in Mauritania. Basic and applied Herpetology 27: 11 - 22. http: // dx. doi. org / 10.11160 / bah. 13002"]}

Kassina fusca Schi��tz, 1967 (Fig. 5A) COLLECTION MATERIAL. ��� Burkina Faso ��� 1♂ (SVL 29.5 mm), 1♀ (SVL 37.3 mm); Kourweogo province, Laye; 12.53083��N, ��� 1.77150��W; 290 m a.s.l.; 6.VIII.1966; M. Lamotte leg.;MNHNRA-1998.2011, 1998.2302. FIELD MATERIAL. ��� Burkina Faso ��� 3 ♂ (SVL 32.2-32.4 mm), 1♀ (SVL 35.8 mm); Arly National Park, Bongola swampy valley; 11.53252��N, 1.48241��E; 170 m a.s.l.; 13. VI.2018; H. J. Ayoro leg.; at the base of shrubs; hand capture ��� 1♂ (SVL 32.0 mm), 1 ♀ (SVL 31.4 mm); Arly National Park, in the floodplain; 11.57852��N, 1.46004��E; 255 m a.s.l.; 16. VI.2018; same collector as for preceding; in a bare space; hand capture ��� 1 ♂; W National Park, around the PK-27 bouli; 11.88744��N, 2.11913��W; 265 m a.s.l.; 20. VI.2018; same collector as for preceding; in a bush near the water body; acoustic report ��� 6 ♂ (SVL 32.2-35.5 mm), 4 ♀ (SVL 32.8-36.5 mm); W National Park, the edge of a temporary pond (1 km south-west of the Kabougou forest post office); 11.93669��N, 1.99464��E; 310 m a.s.l.; 21. VI.2018; same collector as for preceding; near water body; hand capture ��� 1 ♂ (SVL 29.8 mm); Como��-L��raba Forest; 9.88803��N, 4.61969��W; 230 m a.s.l.; 5.VIII.2018; H. J. Ayoro & A. Ohler leg.; near a maize field (50 m of the forest); hand capture ��� 1 ♂ (SVL 32.4 mm); Folonzo village swampy valley; 9.95777��N, ��� 4.67455��W; 240 m a.s.l.; 6.VIII.2018; same collectors as for preceding; in a cavity of tree; hand capture ��� 4♂ (SVL 32.8 -35.0 mm), 3 ♀ (SVL 41.7-42.7 mm); Koti; 11.38779��N, ��� 3.06401��W; 310 m a.s.l.; 22.VIII.2018; H. J. Ayoro leg.; near a bean field; hand capture ��� 3 ♂; Kou Forest, on the bank of Kou River in the savannah part; 11.18815��N, ��� 4.44397��W; 375 m a.s.l.; 9.VIII.2018; H. J. Ayoro & A. Ohler leg.; in a bush on the river bank; acoustic report ��� 2 ♂; Koubri, around ���Ancien barrage���; 12.22828��N, ��� 1.34587��W; 320 m a.s.l.; 16.VII.2017; same collectors as for preceding; acoustic report ��� Ioba province, near Dissin; 10.94250��N, ��� 3.15778��W; 260 m a.s.l.; 2011; J. Penner leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l.; 1983, I-III.2008 (B��hme et al. 1996; Mohneke et al. 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011; Mohneke et al. 2011); ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 29 specimens examined. According to Schi��tz (1999) and R��del (2000), SVL of this species ranging from 29- 40.5 mm in both sexes. In the observed specimens, males measure 29.8-35.5 mm and females 31.4-42.7 mm SVL. The back of living specimens of a light green-olive colouration with scattered larger and smaller darker spots (Fig. 5A). HABITAT. ��� This species calls at night from holes at the base of shrubs. After rain these frogs may be found moving on the open ground, between grasses and bushes during the night. This species was not found in Dori and Yakouta in the Sahel region of Burkina Faso. It is a common West African savannah species, with a distribution ranging from southern Senegal to western Nigeria (B��hme et al. 1996; Schi��tz 1999; R��del 2000; Nago et al. 2006; Segniagbeto et al. 2007)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 561-562, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90."]}

Pyxicephalus maltzanii (Boulenger, 1882) (Fig. 6E, F) COLLECTION MATERIAL. ��� Burkina Faso ��� 2 adults; Pabr��; 12.51000��N, ��� 1.57953��W; 290 m a.s.l.; 5. V.1970; M. Lamotte leg.; MNHN-RA-1989.4058, 1989.4059 ��� 2 adults; Samba; 2.70144��N, ��� 2.40389��W; 290 m a.s.l.; M. Lamotte leg.; MNHN-RA-1989.4062, 1989.4063 ��� 2 adults; Garango; 11.79985��N, ��� 0.55053��W; 280 m a.s.l.; 18. VI.1962; M. Lamotte leg.; MNHN-RA-1995.1901, 1995.1902. FIELD MATERIAL. ��� Burkina Faso ��� 1 subadult (SVL 61.9 mm); W National Park, Nangouli swampy valley; 11.89793��N, 2.00091��E; 250 m a.s.l.; 21. VI.2018; H. J. Ayoro leg., under a shrub; hand capture ��� 3 ♀ (SVL 95.3-129.5 mm); Kabougou village; 11.96070��N, 1.99742��E; 280 m a.s.l.; 22. VI.2018; same collector as for preceding; in a paddy field; hand capture ��� 7 juveniles (SVL 29.1-38.1 mm); the edge of Dori pond, Toroodi; 14.00554��N, ��� 0.02405��W; 280 m a.s.l.; 24.VIII.2018; same collector as for preceding; under grasses near water body; hand capture ��� 1 ♀ (SVL 112.5 mm); the edge of N���Djomga village pond; 14.07225��N, ��� 0.04862��W; 265 m a.s.l.; 25.VIII.2018; same collector as for preceding; near water body; hand capture. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke et al. 2010b; Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 20 specimens studied. A plump, large and compact frog with a short and wide head (Fig. 6E, F). Its skin faintly warty and discontinuous ridges present on back. Measures for four females 95.3-129.5 mm SVL, thus larger than frogs from Benin (Nago et al. 2006) (female 90 mm and males 106-110 mm). Tympanum moderately large. Limbs relatively short and strong. A large inner metatarsal tubercle present. In life, dorsal colour greyish to green. Dori subadults and juveniles with a greenish colour and a white mediodorsal line. In W National Park, subadults with a dominantly greyish colour with large dark grey spots on the back (Fig. 6E, F). Adult females with olive green or light grey dorsal colouration. Belly white and throat yellow. REMARKS. ��� The taxonomy of West African Pyxicephalus Tschudi, 1838 needs revision (Nago et al. 2006; B��hme & Heath 2018). The West African specimens of this genus were in the past traditionally assigned to P. edulis Peters, 1854 which type locality is in Mozambique. The oldest available nomen for West African specimens is Rana maltzanii Boulenger, 1882, a replacement name for Maltzania bufonia Boettger, 1881. At the time of Boulenger (1882), Pyxicephalus Tschudi, 1838 was considered a subjective synonym of the genus Rana Linnaeus, 1758. Thus Boulenger (1882) considered Maltzania bufonia Boettger, 1881 as a secondary homonym of Rana bufonia Merrem, 1820 (valid as Elachistocleis surinamensis (Daudin, 1802)) (see Article 59.3 of the International Code of zoological Nomenclature for nomina replaced before 1961 for validity), and cannot be used as valid for the West African Pyxicephalus. The epithet Maltzania bufonia, as proposed by B��hme & Heath (2018), is permanently invalid and cannot be used for this taxon because the replacement name was���in use��� (e. g. cited as valid by Loveridge [1950] and Channing et al. [2012] listed in various synonymies of Pyxicephalus adspersus or Rana edulis; but the expression ���in use��� is quite imprecise). If we follow this taxonomic change, the nomen Rana maltzanii Boulenger, 1882 should be used as the valid name for Pyxicephalus maltzanii (Boulenger, 1882). Rana (Pyxicephalus) reiensis Monard, 1951 from Cameroon would be available if the central African populations proove to be taxonomically distinct. Thus here, we applied the name Pyxicephalus maltzanii to our voucher specimens, although a formal taxonomic revision including detailed morphological and molecular data is still pending. HABITAT. ��� This species was captured among grasses close to Dori and N���Djomga ponds. It was also found in Kabougou village near W National Park. Pyxicephalus maltzanii was recorded by gardeners in a banana plantation, vegetable gardens and paddy fields at D��dougou, Koti, Loumila and Koubri. Other authors indicate its presence in arid savannahs where it apparently prefers sand and clay substrates and it spends most of the year buried, appearing only to breed (Perret 1966; B��hme 1975). It is widely distributed in West Africa but rarely recorded; the species is almost everywhere eaten by the local population (R��del 2000; Nago et al. 2006; Mohneke et al. 2010b; Padial et al. 2013; B��hme & Heath 2018)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 573, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["BOULENGER 1882. - Catalogue of the Batrachia Salientia S. Ecaudata in the collection of the British Museum. Taylor & Francis, London, i-xvi + 1 - 503, pl. 1 - 30. https: // doi. org / 10.5962 / bhl. title. 8307","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","MOHNEKE M., ONADEKO A. B., HIRSCHFELD M. & RODEL M. - O. 2010 b. - Dried or fried: amphibians in local and regional food markets in West Africa. Traffic Bulletin 22: 117 - 128.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","BOHME W. & HEATH J. 2018. - Amphibian and reptilian records from south-central Mali and western Burkina Faso. Bonn zoological Bulletin 67: 59 - 69.","LOVERIDGE A. 1950. - History and habits of the East African Bullfrog. Journal of the East Africa Natural History Society 19: 253 - 255.","CHANNING A., RODEL M. - O. & CHANNING J. 2012. - Tadpoles of Africa - The biology and identification of all known tadpoles in sub-Saharan Africa. Frankfurt Contributions to Natural History, Edition Chimaira, Vol. 55, Frankfurt / Main, 402 p.","PERRET J. L. 1966. - Les amphibiens du Cameroun. Zoologische Jahrbucher fur Systematik 8: 289 - 464.","BOHME W. 1975. - Zur Herpetofaunistik Kameruns, mit Beschreibung eines neuen Scinciden. Bonner zoologische Beitrage 26: 1 - 48.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","PADIAL J. M., CROCHET P. A., GENIEZ P. & BRITO J. C. 2013. - Amphibian conservation in Mauritania. Basic and applied Herpetology 27: 11 - 22. http: // dx. doi. org / 10.11160 / bah. 13002"]}

Phrynobatrachus francisci Boulenger, 1912 (Fig. 5E) COLLECTION MATERIAL. ��� Burkina Faso ��� 1 ♂ (SVL 16.1 mm), 1♀ (SVL 22.8 mm), 1 subadult (SVL 14.6 mm); L��raba province, Loumana; 10.58001��N, ��� 5.35044��W; 330 m a.s.l.; XI.1970; J.-J. Mor��re & M. Lamotte leg.; MNHN-RA-1996.8753, 1996.8750, 1999.7505 ��� 2♂ (SVL 18-18.3 mm), 7♀ (SVL 20.2-23 mm); Samba; 12.70145��N, ��� 2.40388��W; 290 m a.s.l.; 13.VII.1962; M. Lamotte leg.; MNHN-RA-2007.0598, 2007.0599, 2007.0600, 2007.0601, 2007.0602, 2007.0603, 2007.0604, 2007.0605, 2007.0606 ��� 6 subadults (SVL 14-15.8 mm); Tapoa province, Arly; 11.57975��N, 1.47072��W; 175 m a.s.l.; 27.XII.1972; M. Lamotte leg.; MNHNRA-2001.1371, 2001.1372, 2001.1373, 2001.1374, 2001.13752001.1376 ��� 3♂ (SVL 18.1-20.3 mm), 1 ♀ (SVL 16.9 mm), 2 subadults (SVL 14.3-15.6 mm); Baz��ga province, at 40 km south of Ouagadougou; 12.38011��N, 1.51100��W; 350 m a.s.l.; XII.1965; T. J. Papenfuss leg.; MVZ 82791, MVZ 82794 -98. FIELD MATERIAL. ��� Burkina Faso ��� 5 ♂ (SVL 20.1-22.6 mm); Pabr�� dam; 12.51159��N, ��� 1.58833��W; 290 m a.s.l.; 10.VII.2017; H. J. Ayoro & A. Ohler leg.; in an irrigation ditch; hand capture ��� 2 ♂ (SVL 22.2-24.5 mm), 1 ♀ (SVL 23.4 mm), 5 subadults; Deux Bal��s National Park, close to Ouahabou bouli; 11.60765��N, ��� 3.06033��W; 260 m a.s.l.; 8.XI.2017; H. J. Ayoro leg.; around water body; hand capture ��� 3 ♂ (SVL 20.0- 22.5 mm), 2 subadults (SVL 15.6-17.5 mm); Deux Bal��s National Park, the edge of Pampara pond; 11.65613��N, ��� 3.0303234��W; 270 m a.s.l.; 8.XI.2017; same collector as for preceding; around water body; hand capture ��� 1 ♂ (SVL 18.1 mm), 4 subadults (SVL 15.0��� 16.2 mm); Deux Bal��s National Park, downstream of Soumbou dam; 11.69872��N, ��� 2.79738��W; 260 m a.s.l.; 8.XI.2017; same collector as for preceding; on dry grasses near water body; hand capture ��� 3 ♂ (SVL 22.3- 23.3 mm), 3 ♀ (SVL 22.1 -23.0 mm); Deux Bal��s National Park, the edge of Petit Bal�� dam; 11.59998��N, ��� 2.92701��W; 240 m a.s.l.; 9.XI.2017; same collector as for preceding; in the cavities of lateritic stones near water body; hand capture ��� 7♂ (SVL 18.5-23.5 mm), 5 ♀ (SVL 18.52 - 0.8 mm), 3 subadults (SVL 15.5-16.8 mm); Deux Bal��s National Park, the edge of Koh��na pond; 11.67665��N, ��� 2.83611��W; 275 m a.s.l.; 9.XI.2017; same collector as for preceding; on the mud around water body; hand capture ��� 1 ♀ (SVL 21.3 mm); Arly National Park, Bouliabou pond; 11.57892��N, 1.46388��E; 170 m a.s.l.; 11. VI.2018; same collector as for preceding; near water body; hand capture ��� 1♂ (SVL 22.2 mm), 1 ♀ (SVL 23.5 mm); the bank of Arly River; 11.55946��N, 1.45624��E; 170m a.s.l.; 14. VI.2018; same collector as for preceding; under dead wood near water body; hand capture ��� 3 ♂ (SVL 18.4���21.2 mm); Kabougou River; 11.96860��N, 2.01259��E; 280 m. a.s.l; 18. VI.2018; same collector as for preceding; in a paddy field; hand capture ��� 1 ♂ (SVL 22.9 mm), 3 ♀ (SVL 22.2-23.7 mm); near the protected part of Kabougou River; 11.97060��N, 2.01626��E; 265 m a.s.l.; 19. VI.2018; same collector as for preceding; on dead leaves; hand capture ��� 4♂ (SVL 18.6- 22.7 mm), 1 ♀ (SVL 22.9 mm); W National Park, on the edge of PK-27 bouli; 11.88744��N, 2.11913��W; 265 m a.s.l.; 20. VI.2018; same collector as for preceding; on the mud near water body; hand capture ��� 10 subadults; the edge of PK-27 bouli; 3.II.2017; same collector as for preceding; on the mud near water body; visual observation ��� 1 ♂ (SVL 21.5 mm), 1 ♀ (SLV 22.5 mm); Baboro swampy valley; 9.92509��N, ��� 4.57577��W; 240 m a.s.l.; 4.VIII.2018; H. J. Ayoro & A. Ohler leg.; near water body; hand capture ��� 1♀ (SVL 21.4 mm); the flood zone of Como�� River (300 m west of the Como�� bridge); 9.93239��N, ��� 4.60655��W; 220 m a.s.l.; 5.VIII.2018; same collectors as for preceding; near small water body; hand capture ��� 7 ♀ (SVL 18.4���23.5 mm); Folonzo village swampy valley; 9.95775��N, ��� 4.67449��W; 245 m a.s.l.; 6.VIII.2018; same collectors as for preceding; near a small water body; hand capture ��� 2 subadults (SVL 12.0��� 13.1 mm); N���Djomga village pond; 14.07225��N, ��� 0.04862��W; 270 m a.s.l.; 25.VIII.2018; H. J. Ayoro leg.; on the mud surrounding the pond; hand capture ��� 1 ♀ (SVL 21.2 mm); Dori, Toroodi; 14.00606��N, ��� 0.02846��W; 280 m a.s.l.; 27.VIII.2018; same collector as for preceding; under small grasses; hand capture ��� 2 ♂; D��dougou, SONATUR city (the way from D��dougou to Toughan); 12.48972��N, ��� 3.43611��W; 300 m a.s.l.; 22.VIII.2017; same collector as for preceding; under submerged grasses near small rubbish dump; visual observation and acoustic report ��� Ioba province, Dano; 11.16600��N, ��� 3.08689��W; 340 m a.s.l.; 2011; J. Penner leg.; field observation. LITERATURE REPORTS. ��� Burkina Faso ��� Yako; 12.96744��N, ��� 2.26155��W; 330 m a.s.l.; 18.VIII.1961 (Schi��tz 1967) ��� Como�� province, near Kafolo; coordinates not available; 11. VI.1992 (B��hme et al. 1996) ��� Houet province, Koro; 11.14433��N, ��� 4.19888��W; 350 m a.s.l.; 14. VI.1992 (B��hme et al. 1996) ��� Houet province, Daroha; coordinates not available; 14. VI.1992 (B��hme et al. 1996). DESCRIPTION. ��� 96 specimens examined. Our vouchers within the known size range given by R��del (2000). A small, warty frog with a short and rounded snout. Presence of one or two pairs of chevron shaped warts on the neck (Fig. 5E). Colour of P. francisci usually brown to grey. Males with a dark vocal sac. HABITAT. ��� Phrynobatrachus francisci is restricted to humid habitats and is abundant in all seasons close to water. During the dry season, we collected these frogs on the edge of ponds and rivers, under pebbles and rocks (Fig. 2A, B). It is also found near wells, fountains and cattle troughs. This species has a wide distribution in drier savannahs of West Africa (R��del 2000; Onadeko 2016)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 564, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","ONADEKO B. A. 2016. - Distribution, diversity and abundance of anuran species in three different vegetations habitats in southwestern Nigeria. Ethiopian Journal of environmental Studies and Management 9: 22 - 34. http: // dx. doi. org / 10.4314 / ejesm. v 9 i 1.3"]}

Ptychadena schillukorum (Werner, 1908) (Figs 6D, 8A) FIELD MATERIAL. ��� Burkina Faso ��� 2 ♂ (SVL 33.3-35.9 mm); Arly National Park, the floodplain; 11.57852��N, 1.46004��W; 255 m a.s.l.; 16.VI.2018; H. J. Ayoro leg.; around a small water body; hand capture. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 2 specimens studied. Ptychadena with a compact and less slender body than P. pumilio. Limbs comparatively short; webbing formula: I 1 - 1�� II 0 - 2 III 1 - 2 �� IV 2 �� - 0 V. Absence of supernumerary tubercles. The back with discontinuous, partly indistinct rows of elongate warts (Fig. 6D; Fig. 8A). A discontinuous white lateral line, extending from above the tympanum to the rear back. Dorsal colour greyish with some darker spots. COMMENTS. ��� Specimens identification based on the discontinuous dorsal ridges and their size (see Nago et al. 2006). However, in other parts of Africa, this name is applied to frogs which look completely different (larger and showing continuous dorsal ridges): see e.g. Channing (2001). Werner (1908) indicated in the original description a size ranging from 35-43 mm for the four females from Sudan. HABITAT. ��� The two males were captured on the edge of a small water body in the floodplain from Arly National Park. Ptychadena schillukorum inhabits dry savannahs and grasslands (Nago et al. 2006; S��nchez-Vialas et al. 2017) and it breeds in swampy valleys and savannah ponds during the rainy season (Nago et al. 2009). The species, as currently defined, is found across sub-Saharan savannahs of West and East Africa (Stewart 1967; Hughes 1988; Channing 2001; Mohneke et al. 2011; Sow et al. 2017)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 570, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["WERNER F. 1908 (1907). - Ergebnisse der mit Subvention aus der Erbschaft Treitl unternommenen zoologischen Forschungsreise Dr. Franz Werner's nach dem agyptischen Sudan und Nord- Uganda. XII. Die Reptilien und Amphibien. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch- Naturwissenschaftliche Classe 116: 1823 - 1926.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","CHANNING A. 2001. - Amphibians of Central and Southern Africa. Cornell University Press, New York, USA, 470 p.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2009. - The tadpole of Ptychadena schillukorum (Werner, 1908 \" 1907 \") (Amphibia: Anura: Ptychadenidae). Zootaxa 2115: 65 - 68. https: // doi. org / 10.11646 / zootaxa. 2115.1.6","STEWART M. M. 1967. - The amphibians of Malawi. State University Press, New York, USA, 163 p.","HUGHES B. 1988. - Herpetology in Ghana (West Africa). British herpetological Society Bulletin 25: 29 - 38.","SOW A. S., GONCALVES D. V., SOUSA F. V., MARTINEZ- FREIRIA F., SANTAREM F., VELO- ANTON G., DIENG H., CAMPOS J. C., DIAGNE S. K., BORATYNSKI Z. & BRITO J. C. 2017. - Atlas of the distribution of amphibians and reptiles in the Diawling National Park, Mauritania. Basic and applied Herpetology 31: 101 - 116. http: // dx. doi. org / 10.11160 / bah. 87"]}

Hoplobatrachus occipitalis (G��nther, 1858) COLLECTION MATERIAL. ��� Burkina Faso ��� 11 ♀ (SVL 62.3-106.7 mm), 6 ♂ (SVL 69.6-85.8 mm), 6 juveniles (SVL 21.7-47.1 mm); Passor�� province, Samba; 12.70142��N, ��� 2.40390��W; 300 m a.s.l.; 18. VI.1962; M. Lamotte leg.;MNHN-RA-1979.1192, 1979.1193, 1979.1194, 1979.1200, 1979.1203, 1979.1204, 1979.1205, 1979.1207, 1979.1208, 1979.1209, 1979.1210, 1979.1211, 1979.1212, 1979.1213, 1979.1214, 1979.1215, 1979.1216, 1979.1217, 1979.1218, 1979.1219, 1979.1220, 1979.1221, 1979.1222 ��� 2 ♀ (SVL 75.7-86.2 mm), 1 ♂ (SVL 82.6 mm); Ouagadougou; 12.38014��N, ��� 1.51104��W; 300 m a.s.l.; 1987; M. Lamotte leg.;MNHN-RA-1990.4366, 1990.4367, 1990.4368 ��� 2♀ (SVL 65.7-73.4 mm); Como�� province, Ti��fora; 10.62763��N, ��� 4.55772��W; 290 m a.s.l.; II.1983; NA leg.; ZFMK 93958, 93967 ��� 2 ♂ (SVL 56.5-68.9 mm); Como�� province, D��r��gou��; 10.75013��N, ��� 4.07932��W; 320 m a.s.l.; II.1982; NA leg.; ZFMK 38698-99. FIELD MATERIAL. ��� Burkina Faso ��� 1 ♀ (SVL 77.6 mm); Loumbila, south-west part of Loumbila dam; 12.49844��N, ��� 1.40313��W; 280 m a.s.l; 13.VII.2017; H. J. Ayoro & A. Ohler leg.; around a water body; hand capture ��� 1 ♀ (SVL 105.8 mm); downstream from the Loumbila dam; 12.49844��N, ��� 1.40313��W; 280 m a.s.l.; 14.VII.2017; same collectors as for preceding; around a irrigation well; hand capture ��� 1 ♀ (SVL 106.1 mm), 1 ♂ (SVL 89.5 mm), 1 subadult (SVL 63.8 mm), 7 juveniles (SVL 29.7-35.3 mm); Koubri, on the edge of the ���Ancien barrage���; 12.22883��N, ��� 1.34525��W; 290 m a.s.l.; 16.VII.2017; same collectors as for preceding; hand capture ��� 2♂ (SVL 85.1-95.3 mm), 1 ♀ (SVL 81.1-103.3 mm); Koubri, downstream of the Nabazana dam; 12.20324��N, ��� 1.35103��W; 280 m a.s.l.; 17.VII.2017; same collectors as for preceding; hand capture ��� 10 individuals; Pabr��, Pabr�� dam; 12.51684��N, ��� 1.59690��W; 290 m a.s.l.; 12.VII.2017; same collectors as for preceding; in the water of the irrigation ditch upstream of the dam; field direct observation ��� 1 subadult (SVL 47.1 mm); Deux Bal��s National Park, Ouahabou bouli; 11.60765��N, ��� 3.06033��W; 260 m a.s.l.; 8.XI.2017; H. J. Ayoro leg.; hand capture ��� 1 ♀ (SVL 67.1 mm); Deux Bal��s National Park, Tiona pond; 11.63242��N, ��� 2.85433��W; 270 m a.s.l.; 9.XI.2017; same collector as for preceding; in the pond; hand capture ��� 4 ♀ (SVL 66.3-80.4 mm); 3 subadults (SVL 48.0- 55.1 mm); Deux Bal��s National Park, on the flooded zone of Mouhoun River; 11.77886��N, ��� 2.91243��W; 240 m a.s.l.; 10.XI.2017; same collector as for preceding; on the edge of a pond, hand capture ��� 2 ♀ (SVL 83.6- 92.1 mm); W National Park, Kabougou River (500 m north-west of the Kabougou forest post office); 11.97060��N, 2.01622��E; 280 m a.s.l.; 3.II.2017; same collector as for preceding; in the water of a irrigated well; hand capture ��� 2♂ (SVL 79.9-84.6 mm); same locality; 18. VI.2018; same collector as for preceding; hand capture ��� 1 ♂ (SVL 100.5 mm); W National Park, a small pond (1 km southwest of the Kabougou forest post office); 11.93675��N, 1.99463��E; 310 m a.s.l.; 21. VI.2018; same collector as for preceding; hand capture ��� 4 individuals; W National Park, PK-27 bouli; 11.88741��N, 2.11913��E; 280 m a.s.l. 20. VI.2018; same collector as for preceding; on the edge of the bouli; field direct observation ��� 5 individuals; W National Park, Nangouli swampy valley; 11.89804��N, 2.00101��E; 250 m a.s.l.; 21. VI.2018; same collector as for preceding; on the small river bank; field direct observation ��� 1 ♀ (SVL 82.2 mm); Arly National Park, the Bouliabou pond; 11.57890��N, 1.46393��E; 170 m a.s.l.; 11. VI.2018; same collector as for preceding; on the edge of the pond; hand capture ��� 1 ♀ (SVL 84.2 mm), 1♂; Arly National Park, on a floodplain (500 m north of the Bouliabou pond); 11.57854��N, 1.46000��E; 260 m a.s.l.; 16. VI.2018; same collector as for preceding; hand capture, field direct observation ��� 1 ♂ (SVL 79.1 mm), 1 ♀ (SVL 75.0 mm); Arly National Park, on the bank of the Arly River; 11.57840��N, 1.46143��E; 170 m a.s.l.; 17. VI.2018; same collector as for preceding; near a small water body; hand capture ��� 2 subadults (SVL 36.0- 43.4 mm); Como��-L��raba Forest, a flooded zone of Como�� River (300 m west of the bridge of Como�� River); 9.93242��N, ��� 4.60654��W; 220 m a.s.l.; 5.VIII.2018; H. J. Ayoro & A. Ohler leg.; near a water body; hand capture ��� 3 individuals; Kou Forest; 11.18888��N, ��� 4.43944��W; 260 m a.s.l.; 7-9.VIII.2018; same collectors as for preceding; near the water source of the Guinguette; field direct observation ��� 4♀ (SVL 67.1-104.8 mm), 1 subadult (SVL 39 mm); Dori, on the edge of Dori pond near Wiindou bridge; 14.03665��N, ��� 0.04933��W; 260 m a.s.l.; 22.VIII.2018; H. J. Ayoro leg.; hand capture ��� 4 ♂ (SVL 75.0- 102.1 mm), 5 ♀ (SVL 93.7-111.3 mm); the Dori pond, Toorodi; 14.00385��N, ��� 0.02411��W; 270 m a.s.l.; 24.VIII.2018; same collector as for preceding; hand capture ��� 1 ♂ (SVL 95.3 mm); N���Djomga village pond; 14.07222��N, ��� 0.04865��W; 270 m a.s.l.; 25.VIII.2018; same collector as for preceding, on the edge of the pond; hand capture ��� 1♂ (SVL 92.8 mm); Yakouta, downstream from the Yakouta dam; 14.07454��N, ��� 0.14290��W; 280 m a.s.l.; 26.VIII.2018; same collector as for preceding; hand capture ��� 2♀ (SVL 65.1-95.2 mm); Koti; 20.X.2017; same collector as for preceding; in the stagnant water around a water fountain; hand capture ��� 6 individuals; D��dougou; 12.49204��N, ��� 3.43575��W; 300 m a.s.l.; 22.VIII.2017; same collector as for preceding; in a wetland of SONATUR habitations; field direct observation ��� Ioba province, Djikologo in Bontioli reserve; 10.93684��N, ��� 3.13804��W; 260 m a.s.l.; 2011; J. Penner leg.; field direct observation ��� Dano, Batiara; 11.21322��N, ��� 3.02685��W; 280 m a.s.l.; 21.X.2003; M.-O. R��del & T. Moritz leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Passor�� province, Yako: 12.96740��N, ��� 2.26151��W; 330 m a.s.l.; 1962 (Schi��tz 1967) ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (B��hme et al. 1996; Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011) ��� Zam; 12.32885��N, ��� 0.82631��W; 280 m a.s.l (Mohneke 2011; Mohneke et al. 2011) ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke et al. 2010b; Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� On 84 specimens examined, males��� SVL between 66.0- 105.2 mm (19 specimens) while females��� sizes between 69- 111.3 mm (38 specimens). Large and robust aquatic frogs. Eyes very prominent, in dorsal position. Head moderately broad with a rounded tympanum. Legs very strong and long, toes fully webbed. Dorsal skin with short warty and discontinuous folds, rather irregularly arranged. The dominant colouration greenish or grey olive on the back and white with some dark spots on the belly. juveniles showing a characteristic light yellow to green interorbital line. HABITAT. ��� Hoplobatrachus occipitalis is usually collected near or in water (R��del 2000). In the dry season it lives in rivers, pond banks and in irrigated wells. It is found in practically all freshwater habitats in wet season. Migrations were related in the dry season to the edge of rivers and the in wet season to surroundings of ponds (Spieler 1997). The species was observed on clayey and sandy soils. This frog is an important source of animal protein in the nutrition of local population from the south-eastern part of Burkina Faso (Mohneke et al. 2010b, 2011)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 556-557, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","MOHNEKE M., ONADEKO A. B., HIRSCHFELD M. & RODEL M. - O. 2010 b. - Dried or fried: amphibians in local and regional food markets in West Africa. Traffic Bulletin 22: 117 - 128.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","SPIELER M. 1997. - Radio-telemetrische Untersuchungen zur Laichplatzwahl eines westafrikanischen Raniden, in HENLE K. & VEITH M. (eds), Naturschutzrelevante Methoden der Feldherpetologie, Mertensiella 7: 203 - 220."]}

Amnirana albolabris (west) Jongsma et al. (2018) (Fig. 8B) COLLECTION MATERIAL. ��� Burkina Faso ��� 1 juvenile (SVL 16.1 mm); Houet province, in the forest of small-seminary at Nasso; 11.20953��N, ��� 4.43951��W; 340 m a.s.l.; 4.IV.1968; M. Lamotte leg.; MNHNRA-1999.7575. DESCRIPTION. ��� 1 specimen examined. Rather slender body with a triangular, depressed head and slightly rounded snout (Fig. 8B). Dorsal skin finely granular under the magnifying glass; two distinct dorsolateral folds present, from the posterior border of the eye to the sacrum. Dorsal colour dark brown, with few darker brown spots. REMARKS. ��� The genus Amnirana has been recently revised by Jongsma et al. (2018). Based on molecular data this study restricts the name Amnirana albolabris (Hallowell, 1856) to Central African populations, the populations from West Africa are clearly distinct and they are provisionally named Amnirana albolabris (west) (Jongsma et al. 2018). The type locality given for A. albolabris is ������West Africa������ by Hallowell (1856). However, Jongsma et al. (2018) searched the route of the collector Du Chaillu of the syntypes, and identified the area ���north of the Ogoou�� River in Gabon ��� as the type locality. Therefore, following Jongsma et al. (2018), we use for the West African populations the name A. albolabris (west) here. A formal description of this taxon is prepared by R��del and co-workers. HABITAT. ��� Amnirana albolabris (west) was collected in a gallery forest. We considered that place of collection could be Kou Forest which we visited in August 2018, what would be the northernmost site of its range. In other parts of its range the species inhabits pristine to heavily degraded rainforests and gallery forests in the savannah zones (R��del & Bangoura 2004; Kpan et al. 2014; Onadeko 2016)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 574, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["JONGSMA G. F. M., BAREJ F. M., BARRATT D. C., BURGER M., CONRADIE W., ERNST R., GREENBAUM E., HIRSCHFELD M., LEACHE D. A., PENNER J., PORTIK M. D., ZASSI- BOULOU A. G., RODEL M. - O. & BLACKBURN C. D. 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","HALLOWELL E. 1856. - Notes on Reptilia in the collection of the Museum of the Academy of Natural Sciences. Proceedings of the Academy of Natural Sciences of Philadelphia 8: 146 - 153.","KPAN T. F., ADEBA P. J., KOUAME N. G., KONE I., KOUASSI P. K. & RODEL M. - O. 2014. - The anuran fauna of a volunteer nature reserve: the Tanoe-Ehy swamp forests, south-eastern Ivory Coast, West Africa. Zoosystematics and Evolution 90: 261 - 270. https: // doi. org / 10.3897 / zse. 90.8796","ONADEKO B. A. 2016. - Distribution, diversity and abundance of anuran species in three different vegetations habitats in southwestern Nigeria. Ethiopian Journal of environmental Studies and Management 9: 22 - 34. http: // dx. doi. org / 10.4314 / ejesm. v 9 i 1.3"]}

Hyperolius lamottei Laurent, 1958 (Fig. 7B) COLLECTION MATERIAL. ��� Burkina Faso ��� 1♀ (SVL 21.6 mm); L��raba province, Katiar�� (north-east of Ferk��) close to Ivorian border; 9.91200��N, ��� 4.94776��W; 250 m a.s.l.; 10.XI.1970; M. Lamotte leg.; MNHN-RA-1997.9889. DESCRIPTION. ��� Female with a SVL of 21.6 mm. A small savannah frog with a SVL of 20-27 mm of both sexes (Lamotte & Xavier 1981; Schiotz 1999). Data based on additional specimens from various West African localities: males smaller than females; Ivory Coast 38♂ (19.5-23.1 mm), 32 ♀ (20.3-26.1 mm); Guinea 32 ♂ (18.0- 23.4 mm), 27 ♀ (20.0- 24.4 mm); Liberia 27 ♂ (19.5-23.8 mm), 37♀ (20.3-28.1 mm); Sierra Leone 54 ♂ (19.3-23.7 mm), 28 ♀ (21.8-27.3 mm). Slender frog, as Afrixalus vittiger. Back with 9 longitudinal bands: four white and five brown (Fig. 7B); see also figures inSchi��tz (1999), R��del & Ernst (2003), Adeba et al. (2010) and Channing et al. (2013). HABITAT. ��� Hyperolius lamottei was collected in Katiar��, north-east of Ferk�� region. This locality is situated in Cascades region in Burkina Faso, characterized by savannah and gallery forest areas with waterfalls, hills and cliffs. Based on the collection data of MNHN specimens from others localities of West Africa, H. lamottei inhabits wet savannah zones and seems to have a preference for montane regions such as the Nimba, Loma and P��ko mountains (Lamotte 1971; R��del & Ernst 2003; R��del et al. 2004), where it inhabits short grassland savannahs, often on granite ground (Schi��tz 1967; R��del & Ernst 2003). The Burkina Faso record is thus very unexpected and future field work should try to confirm its presence at the locality mentioned above and other similar sites., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 560, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O. & ERNST R. 2003. - The amphibians of Marahoue and Mont Peko National Parks, Ivory Coast. Herpetozoa 16: 23 - 39.","ADEBA P. J., KOUASSI P. & RODEL M. - O. 2010. - Anuran amphibians in a rapidly changing environment - revisiting Lamto, Cote d'Ivoire, 40 years after the first herpetofaunal investigations. African Journal of Herpetology 59: 1 - 16. https: // doi. org / 10.1080 / 04416651003595337","CHANNING A., HILLERS A., LOTTERS S., RODEL M. - O., SCHICK S., CONRADIE W., RODDER D., MERCURIO V., WAGNER P., DEHLING J. M., DU PREEZ L. H., KIELGAST J. & BURGER M. 2013. - Taxonomy of the super-cryptic Hyperolius nasutus group of long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species. Zootaxa 3620: 301 - 350. https: // doi. org / 10.11646 / zootaxa. 3620.3.1","LAMOTTE M. 1971. - Le Massif des Monts Loma (Sierra Leone), Fasciule I; XIX. Amphibiens. Memoire de l'Institut fondamental d'Afrique noire (A) 86: 397 - 407.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346."]}

Sclerophrys pentoni (Anderson, 1893) (Fig. 3F) COLLECTION MATERIAL. ��� Burkina Faso ��� 20♀ (SVL 56.6-66.2 mm); Bougouriba province, Di��bougou; 10.98411��N, ��� 3.25603��W; 300 m a.s.l.; 3.VI.1976; NA leg.; MHNG 1537.14 - 33. FIELD MATERIAL. ��� Burkina Faso ��� 5 ♂ (SVL 48.1-56.6 mm); Arly National Park, near the flood plain; 11.57853��N, 1.46001��E; 260 m a.s.l.; 16. VI.2018; H. J. Ayoro leg.; around a small water body on a trail; hand capture ��� 6 ♂ (SVL 58.2-64.8 mm), 1 ♀ (SVL 73.2 mm); W National Park, 50 m of the back of the Kabougou forest post office; 11.95738��N, 2.01381��E; 290 m a.s.l.; 20. VI.2018; same collector as for preceding; in a small water body on a trail; hand capture ��� 1 subadult (SVL 37.8 mm); Como��-L��raba Forest, 50 m of the forest; 9.93240��N, ��� 4.60655��W; 220 m a.s.l.; 5.VIII.2018; H. J. Ayoro & A. Ohler leg.; on the edge of a maize field; hand capture ��� 1 subadult (SVL 29.3 mm); Yakouta, downstream from Yakouta dam; 14.07451��N, ��� 0.14290��W; 280 m a.s.l.; 26.VIII.2018; H. J. Ayoro leg.; near a sand heap; hand capture ��� 6♂ (SVL 56- 761.8 mm), 1 ♀ (SVL 66.1 mm), 1 subadult (SVL 36.4 mm); Koti; 23.VIII.2018; same collector as for preceding; near a pond in bush; hand capture ��� 1 individual; Pabr��; ���Hotel Rep��re���; 12.51152��N, ��� 1.59116��W; 300 m a.s.l.; 11.VII.2017; H. J. Ayoro & A. Ohler leg.; in a swimming pool; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Passor�� province, Samba; 12.70143��N, ��� 2.40389��W; 300 m a.s.l.; 18.VIII.1961 (Schi��tz 1967) ��� Yako; 12.96740��N, ��� 2.26151��W; 330 m a.s.l.; (Schi��tz 1967; B��hme et al. 1996) ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 42 specimens examined. Toad with a broad and round head, a very short snout, and very well delimited, broad parotid glands, especially in females. Limbs relatively shorter than in S. maculata and S. regularis. Presence of projecting round granules in the dorsal part in all specimens. Dominant colour brownish to dark beige. Dorsal glands or granules reddish (Fig. 3F). The back of the twenty females from MHNG very dark brown or black in alcohol. The venter whitish with a light brown throat. HABITAT. ��� Sclerophrys pentoni was found in many savannah and Sahelian habitats in Burkina Faso (Fig. 2). We collected it in protected areas (Arly National Park, W National Park) and in disturbed zones (near a maize field in Como��-L��raba and near a farmbush in Koti). It was also found in neighbouring countries, namely Benin (Nago et al. 2006) and Togo (Segniagbeto et al. 2007). Beside, the species was reported from the Sahel and Sahelian savannah of western and eastern parts of Africa (Padial & De La Riva 2004; Padial et al. 2013; Portik & Papenfuss 2015; Sow et al. 2017)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 555, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90.","PADIAL J. M. & DE LA RIVA I. 2004. - Annotated checklist of the amphibians of Mauritania (West Africa). Revista Espanola de Herpetologia 18: 89 - 99.","PADIAL J. M., CROCHET P. A., GENIEZ P. & BRITO J. C. 2013. - Amphibian conservation in Mauritania. Basic and applied Herpetology 27: 11 - 22. http: // dx. doi. org / 10.11160 / bah. 13002","PORTIK D. M. & PAPENFUSS T. J. 2015. - Historical biogeography resolves the origins of endemic Arabian toad lineages (Anura: Bufonidae): Evidence for ancient vicariance and dispersal events with the Horn of Africa and South Asia. Evolutionary Biology 15: 1 - 19. https: // doi. org / 10.1186 / s 12862 - 015 - 0417 - y","SOW A. S., GONCALVES D. V., SOUSA F. V., MARTINEZ- FREIRIA F., SANTAREM F., VELO- ANTON G., DIENG H., CAMPOS J. C., DIAGNE S. K., BORATYNSKI Z. & BRITO J. C. 2017. - Atlas of the distribution of amphibians and reptiles in the Diawling National Park, Mauritania. Basic and applied Herpetology 31: 101 - 116. http: // dx. doi. org / 10.11160 / bah. 87"]}

Ptychadena tournieri (Guib�� & Lamotte, 1955) REMARK. ��� We have not yet found this species, neither during field work nor in the collections. So the following notes are based on literature only (Mohneke 2011). LITERATURE REPORTS. ��� Burkina Faso ��� Ganzourgou province, Kougri; 12.39422��N, ��� 1.06500��W; 270 m a.s.l. and 12.40554��N, ��� 1.04391��W; 270 m a.s.l. (Mohneke 2011) ��� Wayen Rapadama Peulh; 12.36783��N, ��� 1.02033��W; 290 m a.s.l. (Mohneke 2011) ��� Gourma province, Pokiamanga; 12.08286��N, 0.44111��E; 290 m a.s.l. (Mohneke 2011) ��� Kompienga province, Kompienga; 11.65222��N, 0.52365��E; 260 m a.s.l.; I-III.2008 (Mohneke 2011). DESCRIPTION. ��� Small and extremely slender species, with 33- 35 mm SVL for males and 41-43 mm for females (R��del 2000). Limbs relatively thick and long as in P. pumilio. Webbing formula: I 1 - 1 �� II 1 - 2 III 1 - 2 IV 1 �� - 1 �� V. Supernumerary tubercles present but poorly visible on the underside of toes II, III and IV. Three pairs of continuous longitudinal ridges on the back and a continuous white lateral ridge. For detailed descriptions see R��del (2000) and Lamotte & Ohler (2000). HABITAT. ��� Ptychadena tournieri inhabits open savannahs (Schi��tz 1967) and breeds in temporary ponds (Perret 1991; R��del 2000; Nago et al. 2006). It is common in West African savannahs and has been recorded from southern Senegal to northern Benin (Bourgat 1979; Lamotte 1967a; R��del 2000; Nago et al. 2006; Segniagbeto et al. 2007)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 572, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","LAMOTTE M. & OHLER A. 2000. - Revision des especes du groupe de Ptychadena stenocephala (Amphibia, Anura). Zoosystema 22: 569 - 583. http: // sciencepress. mnhn. fr / fr / periodiques / zoosystema / 22 / 3 / revision-des-especes-du-groupe-de-ptychadenastenocephala-amphibia-anura","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","PERRET J. L. 1991. - Description de Ptychadena ingeri n. sp. (Anura, Ranidae) du Zaire. Archives des Sciences de Geneve 44: 265 - 281.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","BOURGAT R. 1979. - Trematodes d'Amphibiens du Togo. Bulletin du Museum national d'Histoire naturelle 3: 597 - 624.","LAMOTTE M. 1967 a. - Le probleme des Ptychadena (Famille des Ranidae) du groupe mascareniensis dans l'Ouest Africain. Bulletin du Museum national d'Histoire naturelle, 2 eme serie 39: 647 - 656.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90."]}

Phrynobatrachus latifrons Ahl, 1924 (Fig. 5F) COLLECTION MATERIAL. ��� Burkina Faso ��� 1♂ (SVL 16.3 mm); Como�� province, D��r��gou��; 10.75011��N, ��� 4.07934��W; 320 m a.s.l.; II.1983; J. Heath leg.; ZFMK 93966. FIELD MATERIAL. ��� Burkina Faso ��� 1 ♂ (SVL 16.5 mm); Pabr�� dam; 12.51159��N, ��� 1.58833��W; 290 m a.s.l.; 10.VII.2017; H. J. Ayoro & A. Ohler leg.; in an irrigation ditch; hand capture ��� 6 ♂ (SVL 16.1-19.5 mm); Koubri, downstream from Nabazana dam; 12.20319��N, ��� 1.35105��W; 280 m a.s.l.; 17.VII.2017; same collectors as for preceding; under a heap of dry grasses near paddy field; hand capture ��� 2♂ (SVL 17.8-18.8 mm); Deux Bal��s National Park, the edge of Ouahabou bouli; 11.60765��N, ��� 3.06033��W; 260 m a.s.l.; 8.XI.2017; H. J. Ayoro leg.; around water body; hand capture ��� 2 ♂ (SVL 19.5-20.3 mm); Deux Bal��s National Park, the edge of Koh��na pond; 11.67665��N, ��� 2.83611��W; 275 m a.s.l.; 9.XI.2017; same collector as for preceding; on the mud around water body; hand capture ��� 1 subadult (SVL 13.4 mm); W National Park, the bank of M��krou River (close to the Beninese border, near the Point Triple Residence); 11.89991��N, 2.40265��E; 240 m a.s.l.; 4.II.2017; same collector as for preceding; under a stone near water body, hand capture ��� 6 ♂ (SVL 16.9-18.1 mm), 7♀ (SVL 19.0- 22.2 mm); Arly National Park, the edge of Bouliabou pond; 11.57892��N, 1.46388��E; 170 m a.s.l.; 11. VI.2018; same collector as for preceding; around water body; hand capture ��� 2 ♂ (SVL 17.9-22.1 mm), 1 ♀ (SVL 22.2 mm); the bank of Arly River, 500 m north-east of the central forest post office; 11.57845��N, 1.46146��E; 170 m a.s.l.; 12. VI.2018; same collector as for preceding; under dead wood; hand capture ��� 2 ♂ (SVL 17.6-17.8 mm), 3 ♀ (SVL 19.0-21.0 mm); the bank of Arly River; 1 km west of the central forest post office; 11.57048��N, 1.45218��E; 160 m a.s.l.; 15. VI.2018; same collector as for preceding; near water body; hand capture ��� 1 ♂ (SVL 18.8 mm), 2 ♀ (SVL 18.1-21.9 mm); the floodplain, 500 m north of Bouliabou pond; 11.57852��N, 1.46004��E; 255 m a.s.l.; 16. VI.2018; same collector as for preceding; near water body; hand capture ��� 1 ♀ (SVL 23.3 mm); Como��-L��raba Forest, Baboro swampy valley; 9.92509��N, ��� 4.57577��W; 240 m a.s.l.; 4.VIII.2018; H. J. Ayoro & A. Ohler leg.; in open space; hand capture ��� 1 ♂ (SVL 18.7 mm); the flooded area of Como�� River; 9.93239��N, ��� 4.60655��W; 220 m a.s.l.; 5.VIII.2018; same collectors as for preceding; on a shrub; hand capture. DESCRIPTION. ��� 39 specimens examined. Voucher specimens morphologically similar to frogs described by R��del (2000). Similar to P. francisci but the warts on the neck often less distinct in P. latifrons. Breeding males with a completely smooth skin (Fig. 5F), (R��del 2000). A yellow vocal sac in males (black in P. francisci) and a more pronounced black lateral bands distinguishing P. latifrons from P. francisci. HABITAT. ��� This species was found on mud under dead wood and rocks at the edge of ponds in the dry season (Fig. 2A, B). During the rainy season, the species always lives close to water and we collected them on submerged grasses, under shrubs on the edge of little water bodies. Phrynobatrachus latifrons lives in tree and open savannahs, farm bush and agricultural areas. It avoids closed primary rainforest, but may be found in degraded rainforest (R��del 2000; Kpan et al. 2014; Onadeko 2016; R��del & Glos 2019)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 564-565, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","KPAN T. F., ADEBA P. J., KOUAME N. G., KONE I., KOUASSI P. K. & RODEL M. - O. 2014. - The anuran fauna of a volunteer nature reserve: the Tanoe-Ehy swamp forests, south-eastern Ivory Coast, West Africa. Zoosystematics and Evolution 90: 261 - 270. https: // doi. org / 10.3897 / zse. 90.8796","ONADEKO B. A. 2016. - Distribution, diversity and abundance of anuran species in three different vegetations habitats in southwestern Nigeria. Ethiopian Journal of environmental Studies and Management 9: 22 - 34. http: // dx. doi. org / 10.4314 / ejesm. v 9 i 1.3","RODEL M. - O. & GLOS J. 2019. - Herpetological surveys in two proposed protected areas in Liberia, West Africa. Zoosystematics and Evolution 95: 15 - 35. https: // doi. org / 10.3897 / zse. 95.31726"]}

Kassina cassinoides (Boulenger, 1903) (Fig. 4F) COLLECTION MATERIAL. ��� Burkina Faso ��� 1 ♀ (SVL 53.4 mm), 1 ♂ (SVL 44.3 mm); Samba; 12.70145��N, ��� 2.40387��W; 290 m a.s.l.; 13.VII.1962; M. Lamotte leg.;MNHN-RA-1998.2307, 1998.2308 ��� 1 juvenile (SVL 15.6 mm); Kourweogo province, Laye; 12.53084��N, ��� 1.77155��W; 290 m a.s.l.; VIII.1966; M. Lamotte leg.; MNHNRA-1998.2309. FIELD MATERIAL. ��� Burkina Faso ��� 1♂ (SVL 44.9 mm); Pabr�� dam, south-west of this dam; 12.51159��N, ��� 1.58833��W; 290 m a.s.l.; 10.VII.2017; H. J. Ayoro & A. Ohler leg.; under a stone on the edge of the dam; hand capture ��� 1 ♂ (SVL 42.4 mm); Arly National Park, close to the Bouliabou pond; 11.57892��N, 1.4638��E; 170 m a.s.l.; 16. VI.2018; H. J. Ayoro leg.; near a small water body; hand capture ��� 1 ♂ (SVL 40.1 mm); W National Park, Koubougou River (500 m north-west of the Koubougou forest post office); 11.96860��N, 2.01259��E; 280 ma.s.l.; 18. VI.2018; same collector as for preceding; in a paddy field; hand capture ��� 1 ♂ (SVL 44.5 mm); Folonzo village swampy valley; 9.88803��N, ��� 4.61969��W; 230 m a.s.l.; 6.VIII.2018; H. J. Ayoro & A. Ohler leg.; on a shrub, hand capture ��� 2 ♂; Loumbila dam; 12.49837��N, ��� 1.40313��W; 285 m a.s.l.; 15.VII.2017; same collectors as for preceding; in a bush near the dam; acoustic report ��� 1 ♂; Koubri, ���Ancien barrage���; 12.22843��N, ��� 1.34766��W; 280 m a.s.l.; 16.VII.2017; same collectors as for preceding; on a tree near the dam; acoustic report ��� 1♂; Koubri, downstream of Nabazana dam; 12.20319��N, ��� 1.35105��W; 280 m a.s.l.; 17.VII.2017; same collectors as for preceding; in vegetable garden near the dam; acoustic report ��� 2 ♂; D��dougou; 12.48972��N, ��� 3.43611��W; 300 m a.s.l.; 22.VIII.2017; H. J. Ayoro leg.; under shrubs near the SONATUR city; acoustic report ��� 1 ♂; W National Park, around PK-27 bouli; 11.88744��N, 2.11913��W; 265 m a.s.l.; 20. VI.2018; same collector as for preceding; acoustic report ��� 1 ♂; Como��-L��raba Forest, Sirigou�� swampy valley; 9.91358��N, ��� 4.62686��W; 240 m a.s.l.; 2.VIII.2018; H. J. Ayoro & A. Ohler leg.; on a tree; acoustic report ��� 3 ♂; Kou Forest, on the bank of Kou River in the savannah part of this Forest; 11.18815��N, ��� 4.44397��W; 375 m a.s.l.; 9.VIII.2018; same collectors as for preceding; on the trees; acoustic report ��� Dano dam; 11.16464��N, ��� 3.08633��W; 340 m a.s.l.; 2011; J. Penner leg.; field direct observation. DESCRIPTION. ��� 7 specimens examined. Larger than the two other Kassina species from Burkina Faso. SVL of the five observed males 40.1-44.9 mm. The female MNHN-RA-1998.2307 with 53.4 mm SVL exceeding the known body size (Schi��tz 1999; R��del 2000; Amiet 2012). This species distinguished known from all other savannah frogs by a double, black vertebral line (Fig. 4F). HABITAT. ��� Kassina cassinoides and K. fusca Schi��tz, 1967 were calling at night from holes at the base of shrubs, at the beginning of rainy season. Three of our vouchers were collected on the ground near water (in about 10-50 m distance) in June and July. One was found on a tree where it was calling (in about 100 m from the water) in August. R��del (2000) also reported that single individuals may call from trees. It is most abundant in denser vegetated parts of dry savannah (R��del 2000; Nago et al. 2006). This species was not found in the Sahelian zone (Dori and Yakouta). Its distribution range covers a large part of the Sudano-Sahelian region in West Africa from Senegal to northeast Cameroon (Schi��tz 1999; B��hme 2005; Amiet 2012)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 561, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","AMIET J. L. 2012. - Les rainettes du Cameroun. Edition J. L. Amiet et La Nef des Livres, Nyons & Saint Nazaire, France, 591 p.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","BOHME W. 2005. - Presence of Kassina cassinoides (Boulenger, 1903) in Senegal. Herpetozoa 18: 177 - 178."]}

Hildebrandtia ornata (Peters, 1878) (Fig. 6A) COLLECTION MATERIAL. ��� Burkina Faso ��� 1♂ (SVL 58.7 mm), 2 ♀ (SVL 56.5 -57.0 mm), 1 juvenile (SVL 17.8 mm); Passor�� province, Samba; 12.70144��N, ��� 2.40391��W; 290 m a.s.l.; 13.VII.1962; M. Lamotte / P��re Terrible leg.; MNHN-RA-2006.2859, 2006.2860, 1995.5755, 2006.2858. FIELD MATERIAL. ��� Burkina Faso ��� 5 juveniles (SVL 19.0- 22.8 mm), 1 subadult (SVL 38.8 mm); W National Park, Kabougou River; 11.96860��N, 2.01259��E; 280 m. a.s.l; 18-20. VI.2018; H. J. Ayoro leg.; around paddy field; hand capture ��� 1 subadult (SVL 37.8 mm), 1 ♂; Folonzo village swampy valley; 9.95777��N, ��� 4.67455��W; 240 m a.s.l.; 6.VIII.2018; H. J. Ayoro & A. Ohler leg.; on the edge of the water body; hand capture and acoustic report ��� 5 ♀ (54.1- 61.9 mm), 18 subadults (SVL 33.0- 45.9 mm); Koti; 11.38788��N, ��� 3.06401��W; 310 m a.s.l.; 24.VIII.2018; H. J. Ayoro leg.; on the edge of a groundnut field; hand capture ��� 1 individual; Pabr��, ���Hotel le Rep��re���; 12.53261��N, ��� 1.58286��W; 310 m a.s.l.; 11.VII.2017; H. J. Ayoro & A. Ohler leg.; in a swimming pool; visual observation ��� 1 ♂; Arly National Park, on the floodplain; 11.57852��N, 1.46004��E; 255 m a.s.l.; 16. VI.2018; under grasses on the edge of small water body; visual and acoustic report. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l.; (Mohneke 2011; Mohneke et al. 2011) ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 34 specimens examined. Large frog with stocky body and short head, presence of a characteristic colour pattern with a green broad middorsal band, often split by a series of discontinous brown stripes, sides of dorsum dark brown with a greenish brown band on flanks from tympanic region to groin; y-shaped white stripes on the black throat (Fig. 6A). HABITAT. ��� Hildebrandtia ornata was collected on garbage dumps filled after rainfalls on Kabougou River banks (W National Park). We found it as well under deadwood on the edge of a groundnut field in Koti. At Pabr��, it was seen in a swimming pool in the hostel court. We saw a calling male between grasses in Arly National Park. The frog is a fossorial species which emerges only temporarily during the breeding season (Nago et al. 2006; Mohneke et al. 2010a). It occurs in dry and more humid savannahs and it is absent from forest pockets and very rocky terrain within its range (R��del 2000; B��hme & Heath 2018). It maintains in disturbed habitats., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 565-566, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","MOHNEKE M., ZONGO B. & RODEL M. - O. 2010 a. - Les amphibiens du Burkina Faso, in THIOMBIANO A. & KAMPMANN D. (eds), Biodiversity atlas of West Africa, volume 2, Ouagadougou and Frankfurt / Main: 298 - 302.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","BOHME W. & HEATH J. 2018. - Amphibian and reptilian records from south-central Mali and western Burkina Faso. Bonn zoological Bulletin 67: 59 - 69."]}

Hemisus marmoratus (Peters, 1854) (Fig. 4A) COLLECTION MATERIAL. ��� Burkina Faso ��� 3♂ (SVL 26.8-31.8 mm), 4 ♀ (SVL 33.3-36.4 mm); Passor�� province, Samba; 12.70144��N, ��� 2.40390��W; 290 m a.s.l.; 14.VII.1962; M. Lamotte leg.; MNHNRA-1979.156, 1979.159, 1979.160, 1979.162, 1979.152, 1979.154, 1979.155 ��� 29 tadpoles; Samba; I.1963; (12.701480��N, ��� 2.40537��W; 290 m a.s.l.; M. Lamotte leg.; MNHN-RA-2006.4833, 2006.4834, 2006.4835, 2006.4836, 2006.4837, 2006.4838, 2006.4839, 2006.4840, 2006.4841, 2006.4842, 2006.4843, 2006.4844, 2006.4845, 2006.4846, 2006.4847, 2006.4848, 2006.4849, 2006.4850, 2006.4851, 2006.4852, 2006.4853, 2006.4854, 2006.4855, 2006.4856, 2006.4857, 2006.4858, 2006.4859, 2006.4860, 2006.4861. FIELD MATERIAL. ��� Burkina Faso ��� 3 ♂ (SVL 29.2-30.6 mm); Como��-L��raba Forest, Baboro swampy valley; 9.92521��N, ��� 4.57574��W; 240 m a.s.l.; 4.VIII.2018; H. J. Ayoro & A. Ohler leg.; under grasses close to a small water body; hand capture ��� 1 ♀ (SVL 41.7 mm); Koubri, ���Ancien barrage���; 12.22882��N, ��� 1.34527��W; 285 m a.s.l.; 16.VII.2017; same collectors as for preceding; at edge of a paddy field; hand capture ��� 1 ♀ (SVL 41.3 mm); W National Park; 11.95738��N, 2.01381��E; 290 m a.s.l.; 20. VI.2018; H. J. Ayoro leg.; near a trail; hand capture ��� 1 ♀ (SVL 30.3 mm); Dori, 300 m from the N���Djomga village pond; 14.07222��N, ��� 0.04860��W; 270 m a.s.l. 25.VIII.2018; same collector as for preceding; at the border of a millet field; hand capture ��� 1 subadult (SVL 26.7 mm); Yakouta, downstream from the Yakouta dam; 14.07452��N, ��� 0.14290��W; 280 m a.s.l.; 26.VIII.2018; same collector as for preceding; near the asphalted road; hand capture ��� 1 subadult (SVL 26.2 mm); Koti; 11.38778��N, ��� 3.06400��W; 310 m a.s.l.; 15.IX.2018; same collector as for preceding; in a bean field; hand capture ��� 1 ♂; Arly National Park, floodplain; 11.57851��N, 1.46000��E; 260 m a.s.l.; 16. VI.2018; same collector as for preceding; near a small water body surrounded by grasses; acoustic report ��� 2 ♂; W National Park; the edge of a small pond (1 km south-west of the Kabougou forest post office); 11.93671��N, 1.99463��E; 310 m a.s.l.; 21. VI.2018; same collector as for preceding; acoustic report. LITERATURE REPORTS. ��� Burkina Faso ��� Passor�� province, Yako: 12.96740��N, ��� 2.26151��W; 330 m a.s.l.; 1962 (Schi��tz 1967) ��� Bobo-Dioulasso, halfway between Bobo-Dioulasso and Ouagadougou; coordinates not available; 31.VII.1992 (B��hme et al. 1996) ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Zam; 12.32885��N, ��� 0.82631��W; 280 m a.s.l; (Mohneke 2011; Mohneke et al. 2011); ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke et al. 2010b; Mohneke 2011; Mohneke et al. 2011) ��� Houet province, Karankasso vigu��; 11.00978��N, ��� 3.90001��W; 300 m a.s.l.; 2015 (B��hme & Heath 2018). DESCRIPTION. ��� 15 specimens examined. Small frogs with a triangular head and a very pointed and short snout. Morphologically sturdy with strong limbs, adapted for burrowing (Fig. 4A). Skin soft and smooth (Laurent 1972; R��del 2000). Dorsal colouration of living frogs yellowish with some dark brown spots on the back. In alcohol, yellow colour whitish dorsally and ventrally. HABITAT. ��� Hemisus marmoratus was difficult to see during our survey periods and is probably much more abundant in convenient habitat and at the right saison. We found this fossorial species in the vicinity of a paddy field on the ���Ancien barrage��� edge in Koubri, on a trail leading to a small water body in W National Park. We heard the male calling under flooded grasses around a pond (1 km southwest of the Kabougou forest post office) in W National Park. The species was also found in Baboro swampy valley in Como��-L��raba where males could be heard calling among grasses nearby a small water body in Folonzo village swampy valley on 6 August 2018. This was unusual as Hemisus usually breeds in the early rainy season or even before the rains set in (see e.g. R��del et al. 1995; Kaminsky et al. 1999). In a floodplain near Bouliabou pond, we heard males calling and recorded three males in Arly National Park. In the Sahelian part, we captured this species during rainy weather on sandy soils close to a millet field in N���Djomga and on the roadside in Yakouta. Hemisus marmoratus is mostly fossorial, in relatively moist, loose soils and is often captured during earthworks such as plowing fields (Lamotte & Xavier 1981; R��del 2000). The species is common and reported having a wide distribution range from West Africa south to Angola and South Africa (Laurent 1972; R��del 2000). However, several cryptic taxa are probably mixed under this name (M.-O. R��del, unpublished data)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 557-558, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","MOHNEKE M., ONADEKO A. B., HIRSCHFELD M. & RODEL M. - O. 2010 b. - Dried or fried: amphibians in local and regional food markets in West Africa. Traffic Bulletin 22: 117 - 128.","BOHME W. & HEATH J. 2018. - Amphibian and reptilian records from south-central Mali and western Burkina Faso. Bonn zoological Bulletin 67: 59 - 69.","LAURENT R. F. 1972. - Tentative revision of the genus Hemisus Gunther. Musee Royal d'Afrique Centrale, Sciences Zoologiques 194: 1 - 67.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","RODEL M. - O., SPIELER M., GRABOW K. & BOCKHELER C. 1995. - Hemisus marmoratus (Peters, 1854) (Anura: Hemisotidae), Fortpflanzungsstrategien eines Savannenfrosches. Bonner zoologische Beitrage 45: 191 - 207.","KAMINSKY S. K., LINSENMAIR K. E. & GRAFE T. U. 1999. - Reproductive timing, nest construction and tadpole guidance in the African pig-nosed frog, Hemisus marmoratus. Journal ofHerpetology 33: 119 - 123. https: // doi. org / 10.2307 / 1565550","LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816."]}

Sclerophrys mauritanica (Schlegel, 1841) (Fig. 7C) COLLECTION MATERIAL. ��� Burkina Faso ��� 1 ♀ (SVL 104 mm); Boulgou province, Garango; 11.79979��N, ��� 0.55053��W; 280 m a.s.l.; 18.VI.1962; M. Lamontellerie leg.; MNHN-RA-1989.4038. DESCRIPTION. ��� 1 specimen studied. Dorsum yellowish with large brown patches, bordered with black (Fig. 7A). Tympanum large and interorbital space concave. Parotoid glands kidney-shaped and parallel. Presence of double subarticular tubercles on the fourth toe. Characters corresponding to Salvador���s (1996) description of Sclerophrys mauritanica for northern Africa. HABITAT. ��� This toad is known from a wide variety of habitats, including rocky areas, grasslands, Mediterranean zones, oases, agricultural lands and urban areas (Salvador 1996; Ben Hassine & Nouira 2012). This species is reported from throughout northern Africa in an apparently continuous distribution between the Tunisian and Moroccan borders (Geniez et al. 2000; Ben Hassine & Escoriza 2017; Escoriza & Ben Hassine 2019). The presence of this species was reported by Lamotte & Xavier (1981) from Sub-Saharan areas like Burkina Faso but these authors did not mention any voucher specimens. The specimen of S. mauritanica in the MNHN is labelled as having been collected at Garango, in central-east of Burkina Faso, in VI.1962 by Michel Lamontellerie (1928-2013). The collector was an army doctor and stayed for over 20 years in this place. 109 specimens donated by Lamontellerie are present in the MNHN collections coming from Burkina Faso, Mali and France, but none from northern Africa. Nevertheless the collection locality of S. mauritanica in Burkina Faso should be regarded with some doubt, as it is more than 1000 km outside its southern range limit. Future research should carefully check toads from Burkina Faso, but also northern Mali and Niger for the potential presence of this species, south of the Sahara., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 575, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SALVADOR A. 1996. - Amphibians of northwest Africa. Smithsonian herpetological Information Service 109: 1 - 45. https: // doi. org / 10.5479 / si. 23317515.109.1","BEN HASSINE J. & NOUIRA S. 2012. - The amphibians of Tunisia: Biodiversity, distribution, status and majors threats. Froglog 101: 32 - 34.","GENIEZ P., MATEO J. A. & BONS J. 2000. - A checklist of the amphibians and reptiles of Western Sahara (Amphibia, Reptilia). Herpetozoa 13: 149 - 163.","BEN HASSINE J. & ESCORIZA D. 2017. - Amphibians of Algeria: new data on the occurrence and natural history. Herpetological Bulletin 142: 6 - 18.","ESCORIZA D. & BEN HASSINE J. 2019. - Amphibians of North Africa. Academic Press, London, United Kingdom, 337 p.","LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816."]}

Afrixalus vittiger (Peters, 1876) (Fig. 4B) COLLECTION MATERIAL. ��� Burkina Faso ��� 2♀ (SVL 21.6-27.4 mm), 1 subadult (SVL 18 mm); Passor�� province, Samba; 12.70144��N, ��� 2.40390��W; 290 m a.s.l.; VII.1962; M. Lamotte leg.; MNHNRA-1979.4741, 1979.4742, 1979.4743. FIELD MATERIAL. ��� Burkina Faso ��� 8 ♂ (SVL 24.5 -27.0 mm); Pabr�� dam, downstream of the dam; 12.51679��N, ��� 1.59693��W; 290 m a.s.l.; 11.VII.2017; H. J. Ayoro & A. Ohler leg.; on flooded grasses; hand capture ��� 2 ♂ (SVL 23.7-24.5 mm); same locality; 12.VII.2017; same collectors as for preceding; on grasses in irrigation ditch; hand capture ��� 3 ♂ (SVL 24.3-25.2 mm), 1 ♀ (SVL 26.7 mm); Loumbila dam; 12.49844��N, ��� 1.40311��W; 280 m a.s.l.; 14.VII.2017; same collectors as for preceding; in a plantain plantation downstream from the dam; hand capture ��� 8 ♂ (SVL 24.3-26.7 mm); same locality; 15.VII.2017; same collectors as for preceding; on small plants around vegetable gardens; hand capture ��� 4 ♂ (SVL 23.4- 25.2 mm); Koubri, ���Ancien barrage���; 12.20322��N, ��� 1.35100��W; 280 m a.s.l.; 16.VII.2017; same collectors as for preceding; in a maize field; hand capture ��� 2 ♂ (SVL 22.8-24.3 mm); Koubri, downstream from the Nabazana dam; 12.20322��N, ��� 1.35101��W; 280 m a.s.l.; 17.VII.2017; same collectors as for preceding; on the edge of a paddy field; hand capture ��� 2 ♀ (SVL 23.4-27.3 mm); Koubri, downstream from Arzoumbaongo dam; 12.22064��N, ��� 1.29590��W; 270 m a.s.l.; 18.VII.2017; same collectors as for preceding; on shrubs near water body; hand capture ��� 1 subadult (SVL 17.1 mm); Deux Bal��s National Park, downstream from the Soumbou dam; 11.69830��N, ��� 2.79711��W; 260 m a.s.l.; 8.XI.2017; H. J. Ayoro leg.; on dry grasses near the water body; hand capture ��� 3 subadults; Deux Bal��s National Park, edge of Koh��na pond; 11.67661��N, ��� 2.83614��W; 270 m a.s.l.; 9.XI.2017; same collector as for preceding; on the mud near water body; direct observation in field ��� 2 ♀ (SVL 21.0- 21.7 mm); Arly National Park, a floodplain; 11.57855��N, 1.46001��E; 260 m a.s.l.; 16-17. VI.2018; same collector as for preceding; on stones near the water body; hand capture ��� 4 ♂ (SVL 21.9-24.8 mm), 2 ♀ (SVL 21.3-23.1 mm); W National Park, Kabougou River (500 m north-west of the Kabougou forest post office); 11.97060��N, 2.01622��E; 280 m a.s.l.; 18. VI.2018; same collector as for preceding; on grasses; hand capture ��� 11 ♂ (SVL 20.9-23.6 mm), 4 ♀ (SVL 23.8-24.8 mm); the protected part of Kabougou River (500 m north-east of the Kabougou forest post office); 11.96865��N, 2.0125��E; 280 m a.s.l.; 19. VI.2018; same collector as for preceding; on grasses ��� 2 ♂ (SVL 21.3 -22.0 mm); the edge of PK-27 bouli; 11.88744��N, 2.11913��E; 265 m a.s.l.; 20. VI.2018; same collector as for preceding; on grasses; hand capture ��� 4♂ (SVL 22.7-24.1 mm), 1♀ (SVL 25.2 mm); Como��-L��raba Forest, Sirigou�� swampy valley; 9.88811��N, ��� 4.61950��W; 230 m a.s.l.; 2.VIII.2018; H. J. Ayoro & A. Ohler leg.; on shrubs and grasses; hand capture ��� 2 ♂ (SVL 22.7-23.5 mm), 1 ♀ (SVL 25.9 mm); Kotokoi swampy valley; 9.82802��N, ��� 4.62727��W; 230 m a.s.l.; 3.VIII.2018; same collectors as for preceding; on grasses; hand capture ��� Kou Forest, bank of Kou River; 11.18720��N, ��� 4.44185��W; 340 m a.s.l.; 8.VIII.2018; same collectors as for preceding; on shrubs around the river bank; acoustic report ��� Ioba province, Gbagba (near Dano); 11.18133��N, ��� 3.05165��W; 300 m a.s.l.; 2011; J. Penner leg.; direct observation in field. LITERATURE REPORTS. ��� Burkina Faso ��� Ouagadougou; 12.38009��N, ��� 1.51100��W; 300 m a.s.l.; 30.VII.1992 (B��hme et al. 1996) ��� Yako; 12.96740��N, ��� 2.26151��W; 330 m a.s.l.; 18.VIII.1961 (Schi��tz 1967). DESCRIPTION. ��� 71 specimens examined. Some of these specimens slightly larger than others. Male���s body size between 20.9-27.3 mm while female���s size 21.3-26.7 mm. Agreeing with Pickersgill (2007), indicating males body size from 21-26 mm and 22-28 mm for females. Skin asperities, typical for the genus, present on dorsum but also on belly, prominent on tarsus and feet in males. On back of all specimens, seven vertical distinct bands, four brown separated by three whitish (Fig. 4B). Ventral skin colour whitish. HABITAT. ��� We observed A. vittiger in floodplains and swampy valleys. The species is present in protected areas such as Arly National Park, W National Park, Como��-L��raba Forest and Deux Bal��s National Park. In Koubri, Pabr�� and Loumbila, the species was found in paddy fields, around ponds on grasses and plants surfacing water (Fig. 2C, D, F). It is a common savannah species which has a large distribution range in West Africa (R��del 2000; Leach�� et al. 2006; Nago et al. 2006; Segniagbeto et al. 2007). This species has often been confused with its congener A. fulvovittatus. The two species partly overlap in the forest-savannah ecotone of West Africa. However, Afrixalus fulvovittatus occurs in heavily degraded forest areas whereas A. vittiger mostly lives in savannah areas (Pickersgill 2007; R��del & Glos 2019)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 558, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","PICKERSGILL M. 2007. - A redefinition of Afrixalus fulvovittatus (Cope, 1860) and Afrixalus vittiger (Peters, 1876) (Amphibia, Anura, Hyeroliidae). African Journal of Herpetology 56: 23 - 37. https: // doi. org / 10.1080 / 21564574.2007.9635551","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","LEACHE A., RODEL M. - O., LINKEM C. W., DIAZ R. E., HILLERS A. & FUJITA M. K. 2006. - Biodiversity in a forest island: reptiles and amphibians of the West African Togo hills. Amphibian and Reptile Conservation 4: 22 - 45. https: // doi. org / 10.1514 / journal. arc. 0040018","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90.","RODEL M. - O. & GLOS J. 2019. - Herpetological surveys in two proposed protected areas in Liberia, West Africa. Zoosystematics and Evolution 95: 15 - 35. https: // doi. org / 10.3897 / zse. 95.31726"]}

Hyperolius nitidulus Peters, 1875 (Fig. 4E) COLLECTION MATERIAL. ��� Burkina Faso ��� 1 ♀ (SVL 24.1 mm), 2♂ (SVL 19-27.5 mm); Passor�� province, Samba; 12.70144��N, ��� 2.40390��W; 290 m a.s.l.; 18.VII.1961; NA leg.; MNHNRA-2000.3113, 2000.3114, 1979.3724 ��� 4 ♂ (SVL 18.9-24 mm); Pabr��; 12.51000��N, ��� 1.57953��W; 290 m a.s.l.; 1963; M. Lamotte leg.; MNHN-RA-2000.3115, 2000.3116, 2000.5609, 2000.5606. FIELD MATERIAL. ��� Burkina Faso ��� 4 ♂ (SVL 27.3 -29.0 mm); Pabr�� dam, downstream of the dam; 12.51685��N, ��� 1.59687��W; 290 m a.s.l.; 11.VII.2017; H. J. Ayoro & A. Ohler leg.; on grasses; hand capture ��� 1 ♂ (SVL 29.4 mm); Loumbila dam; 12.49844��N, ��� 1.40311��W; 280 m a.s.l.; 14.VII.2017; same collectors as for preceding; on aquatic plants near a plantain field; hand capture ��� 8 ♂ (SVL 29.3 -32.0 mm); Koubri, ���Ancien barrage���; 12.22878��N, ��� 1.34522��W; 290 m a.s.l.; 16.VII.2017; same collectors as for preceding; in a maize field near water body; hand capture ��� 1 ♂; Deux Bal��s National Park, downstream from Soumbou dam; 11.69833��N, ��� 2.79713��W; 260 m a.s.l.; 8.XI.2017; H.J. Ayoro leg.; on aquatic plants; acoustic report ��� 2♂ (SVL 23.2-24.9 mm); W National Park, near the Kabougou forest post office; 11.95738��N, 2.01381��E; 290 m a.s.l.; 20. VI.2018; same collector as for preceding; on shrubs; hand capture ��� 14 ♂ (SVL 25.7-28.4 mm), 1 ♀ (SVL 30.6 mm); Como��-L��raba Forest, Sirigou�� swampy valley; 9.88809��N, ��� 4.61954��W; 230 m a.s.l.; 2.VIII.2018; H. J. Ayoro & A. Ohler leg.; around a small pond; hand capture ��� 5 ♂ (SVL 26.8-28.4 mm); Como��-L��raba Forest, Baboro swampy valley; 9.92522��N, ��� 4.57575��W; 240 m a.s.l.; 5.VIII.2018; same collectors as for preceding; in an untilled field; hand capture ��� 4 ♂ (SVL 25.5 -28.0 mm); near the Como�� River bridge; 9.93239��N, ��� 4.60655��W; 220 m. a.s.l; 5.VIII.2018; same collectors as for preceding; on aquatic plants in small stagnant waters on the trail; hand capture ��� 1 ♂ (SVL 26.2 mm); Folonzo village swampy valley; 9.88803��N, ��� 4.61969��W; 230 m a.s.l.; 6.VIII.2018; same collectors as for preceding; near a vegetable garden; hand capture ��� 5 ♂; Kou Forest; 11.18888��N, ��� 4.43944��W; 260 m a.s.l.; 7-9.VIII.2018; same collectors as for preceding; on shrubs and grasses around a water body; acoustic report ��� Ioba province, Bontioli reserve; 10.93682��N, ��� 3.13804��W; 260 m a.s.l.; J. Penner leg. ��� Dano; 11.16681��N, ��� 3.08870��W; 330 m a.s.l. and 11.16464��N, ��� 3.08633��W; 340 m a.s.l.; 2011; J. Penner leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011) ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011). DESCRIPTION. ��� 49 specimens examined.Size of vouchers (SVL 23.2- 31.3 for males and 28.8-30.9 mm for females) corresponding the values of Lampert (2001). In life, specimens with a yellowish back framed by dark lateral bands with narrow yellow edges, extending from the eyes back to above the legs (Fig. 4E). Sometimes, back showing a brown and greyish pattern with very small dark spots. Dorsal coloration lighter and with less contrasting pattern at night. HABITAT. ��� Hyperolius nitidulus was found on grasses and bushes in floodplains, in swampy valleys and farmland such as maize and paddy fields, plantain plantations and vegetable gardens. It was also found on plants above water at nearby ponds and rivers. This species inhabits shrublands, grasslands and wetlands in the Guinean and Sudanian savannahs, from central Guinea to Cameroon (Lamotte 1967b; R��del 2000; R��del et al. 2010a)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 560-561, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["LAMPERT K. P. 2001. - Alternative life history strategies in the West African reed frog, Hyperolius nitidulus. Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Bayerischen Julius-Maximilians-Universitat Wurzburg. Germany, 168 p.","LAMOTTE M. 1967 b. - Les batraciens de la region de Gbakobo (Cote d'Ivoire). Memoire de l'Institut fondamental de l'Afrique noire (A) 29: 218 - 294.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","RODEL M. - O., SANDBERGER L., PENNER J., MANE Y. & HILLERS A. 2010 a. - The taxonomic status of Hyperolius spatzi Ahl, 1931 and Hyperolius nitidulus Peters, 1875 (Amphibia: Anura: Hyperoliidae). Bonn zoological Bulletin 57: 177 - 188."]}

Ptychadena pumilio (Boulenger, 1920) (Fig. 6B, C) COLLECTION MATERIAL. ��� Burkina Faso ��� 6 ♀ (SVL 25-27 mm), 1 ♂ (SVL 29.2 mm); Nasso; 11.20950��N, ��� 4.43952��W; 340 m a.s.l.; 6.XII.1968; M. Lamotte leg.; MNHN-RA-1999.7572, 1999.7573, 1999.7574, 1996.3900, 1996.3901, 1996.3902, 1996.3923 ��� 6 ♀ (SVL 25.3 ���34.0 mm); Bobo-Dioulasso, Pala; 11.14985��N, ��� 4.22671��W; 430 m a.s.l.; XI.1970; M. Lamotte leg.; MNHNRA-1996.3921, 1996.3922, 1996.3924, 1996.3925, 1996.3926, 1996.3927 ��� 3 ♀ (SVL 25-27.4 mm); Loumana; 10.58001��N, ��� 5.35043��W; 330 m a.s.l.; 30.X.1970; M. Lamotte leg.; MNHNRA-1996.7506, 1996.7507, 1996.7508 ��� 6 ♂ (SVL 27.1-29.5 mm), 7 ♀ (SVL 27.2-34.7 mm), 8 subadults (SVL 18.8-24.1 mm); Samba; 12.70144��N, ��� 2.40390��W; 290 m a.s.l.; VII.1962; M. Lamotte leg.; MNHN-RA-1996.3885, 1996.3888, 1996.3889, 1996.3894, 1996.3895, 1996.3896, 1996.3886, 1996.3887, 1996.3890, 1996.3892, 1996.3893, 1996.3878, 1996.3806, 1996.3876, 1996.3877, 1996.3879, 1996.3880, 1996.3881, 1996.3882, 1996.3883, 1996.3884 ��� 1 subadult (SVL 25.2 mm); Dano, Batiara; 11.21325��N, ��� 3.02682��W; 280 m a.s.l.; 21.X.2003; M.-O. R��del leg.; ZMB 84696. FIELD MATERIAL. ��� Burkina Faso ��� 1 subadult (SVL 23.8 mm); W National Park; Kabougou River (its protected part); 11.97060��N, 2.01627��E; 265 m a.s.l.; 4.II.2017; H. J. Ayoro leg.; hidden under a dead wood; hand capture ��� 4 subadults (SVL 18.0- 24.6 mm); Arly National Park, the edge of Bouliabou pond; 11.57892��N, 1.46397��E; 180 m a.s.l.; 2.III.2017; same collector as for preceding; in the cracks of dry mud; hand capture ��� 2 ♂ (SVL 26.7-29.6 mm); the edge of Bouliabou pond; 11.57892��N, 1.46397��E; 180 m a.s.l.; 11. VI.2018; same collector as for preceding; hand capture ��� 2 ♂ (SVL 26.4-27.4 mm); Arly National Park, the floodplain (500 m north of Bouliabou pond); 11.57852��N, 1.46004��E; 255 m a.s.l.; 16-17. VI.2018; same collector as for preceding; near water body; hand capture ��� 5 subadults (SVL 21.5-25.5 mm); Deux Bal��s National Park, the edge of ���Baignade des El��phants���; 11.71809��N, ��� 2.83793��W; 270 m a.s.l.; 8.XI.2017; same collector as for preceding; on dry grasses; hand capture ��� 5 individuals; Deux Bal��s National Park, the edge of Jonction pond; 11.56736��N, ��� 2.86041��W; 255 m a.s.l.; 9.XI.2017; same collector as for preceding; on the mud; visual report ��� 9♀ (SVL 28.5-30.9 mm), 3 subadults (SVL 25.0- 26.5 mm); the edge of Koh��na pond; 11.67662��N, ��� 2.83611��W; 270 m a.s.l.; 9.XI.2017; same collector as for preceding; under dead wood and on the mud; hand capture ��� 1♀ (SVL 29.0 mm); the edge of Tiona pond; 11.63241��N, ��� 2.85437��W; 265 m a.s.l.; 9.XI.2017; same collector as for preceding; under dry grasses; hand capture ��� 2 subadults (SVL 25.6 -27.0 mm); Deux Bal��s National Park, a flooded zone of Mouhoun River; 11.77879��N, ��� 2.91244��W; 245 m a.s.l.; 10.XI.2017; same collector as for preceding; on dry gasses; hand capture ��� 1♀ (SVL 37.1 mm); Baboro swampy valley; 9.92521��N, ��� 4.5739��W; 230 m a.s.l.; 4.VIII.2018; H. J. Ayoro & A. Ohler leg.; near small water body; hand capture ��� 1♀ (SVL 36.3 mm); flooded zone of the Como�� River; 9.93239��N, ��� 4.60655��W; 220 m a.s.l.; 5.VIII.2018; same collectors as for preceding; near water body; hand capture ��� 2♂; Kou Forest, in its savannah part; 11.18815��N, ��� 4.44397��W; 375 m a.s.l.; 9.VIII.2018; same collectors as for preceding; under flooded grasses on a trail; acoustic report ��� 3♂; Dori, Toroodi; 14.00554��N, ��� 0.02405��W; 280 m a.s.l.; 24.VIII.2018; H. J. Ayoro leg.; visual and acoustic reports ��� 1♀ (SVL 34.2 mm), 3 ♂ (SVL 28.2-30.2 mm); the edge of Dori pond, swampy valley (on the road out from Dori to Gorom-Gorom); 14.04736��N, ��� 0.05723��W; 270 m a.s.l.; 25.VIII.2018; same collector as for preceding; under small submerged grasses; hand capture and acoustic report ��� 1 ♂ (SVL 33.0 mm), 1 ♀ (SVL 32.7 mm); the edge of Pabr�� dam; 12.51597��N, ��� 1.59923��W; 290 m a.s.l.; 10-12. VII.2017; H. J. Ayoro & A. Ohler leg.; around small water body; hand capture ��� 1♀ (SVL 33.7 mm); the edge of ��� Ancien barrage���; 12.22843��N, ��� 1.34766��W; 280 m a.s.l.; 16.VII.2017; same collector as for preceding; in the paddy field; hand capture ��� 4 subadults (SVL 22.9-28.7 mm); the edge of Fafo dam; 11.33470��N, ��� 3.01947��W; 300 m a.s.l.; 10.XI.2018; H. J. Ayoro leg.; in a vegetable garden; hand capture ��� Djikologo, close to the Bontioli Reserve; 10.93686��N, ��� 3.13800��W; 260 m a.s.l.; 2011; J. Penner leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 76 specimens studied. Specimens studied 27.2-34.4 mm SVL in males and 29.0- 37.1 mm SVL in females; thus within the size range given by R��del (2000) and Lamotte (1967a). S pecies with a moderately pointed snout and a slender body. Limbs thin and long; webbing formula: I 1 - 2 II 1 - 2 III 1 �� - 2 IV 2 - 1 V. Supernumerary tubercles absent on toes. Three pairs of symmetrical and continuous ridges present on the back. A white, continuous lateral ridge extending from above the tympanum to the posterior part of the back. Dorsal colour pattern from brown, greenish to greyish with dark brown spots (Fig. 6B, C). Breeding males with greenish colour (R��del 2000). Presence of a wide white, green or red vertebral band in many specimens. HABITAT. ��� Ptychadena pumilio is ubiquitous in Guinean and Sudanese savannah and Sahelian habitats. It may even enter degraded rainforest (R��del 2000). We found it near water bodies among grasses and sometimes in the humid areas near water bodies in depressions, as well as in flooded zones in the rainy season. In Dori, we found calling males between flooded short grasses. In the dry season, P. pumilio was observed concealing under rocks, heaps of dry grass and cracks of dry mud around ponds and river banks, in Arly and W National Park from February to March 2017 and in Deux Bal��s National Park in November 2017., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 568-570, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","LAMOTTE M. 1967 a. - Le probleme des Ptychadena (Famille des Ranidae) du groupe mascareniensis dans l'Ouest Africain. Bulletin du Museum national d'Histoire naturelle, 2 eme serie 39: 647 - 656."]}

Ptychadena bibroni (Hallowell, 1845) COLLECTION MATERIAL. ��� Burkina Faso ��� 4 subadults (SVL 29.8- 33.9 mm a.s.l.); Houet province, Bobo-Dioulasso; 11.18801��N, ��� 4.29744��W; 280 m a.s.l.; 6.XI.1966; M. Lamotte leg.; MNHNRA-1996.8731, 1996.8732, 1996.8733, 1996.8734 ��� 1 ♀ (SVL 35.6 mm); Bobo-Dioulasso; 31.X.1970; M. Lamotte leg.; MNHN-RA-1996.607 ��� 2 subadults (SVL 22.3-24.4 mm); Nasso; 11.20954��N, ��� 4.43953��W; 340 m a.s.l.; 17.X.1968; M. Lamotte leg.; MNHN-RA-1996.638, 1996.643 ��� 7 ♀ (SVL 34.4-41.6 mm), 1 ♂ (SVL 28.7 mm); L��raba province, between Loumana and Sindou; 10.62052��N, ��� 5.29650��W; 330 m a.s.l.; 4.XI.1970; M. Lamotte leg.; MNHN-RA-1996.8741, 1996.8742, 1996.8743, 1996.8744, 1996.8745, 1996.8746, 1996.8747, 1996.8748 ��� 8 subadults (SVL 23.3-30.5 mm); L��raba province, north-east of Ferk�� (close to the Ivorian border); 10.XI.1970; M. Lamotte leg.; MNHNRA-1996.8749, 1996.8751, 1996.8738, 1996.8739, 1999.7315, 1999.7316, 1999.7318, 1999.7320 ��� 1 ♀ (SVL 47.3 mm); Como�� province, in Timp��rada River near Yend��r��; 10.20861��N, ��� 4.97900��W; 300 m a.s.l.; 28.X.1970; M. Lamotte leg.; MNHN-RA-1996.8737. FIELD MATERIAL. ��� Burkina Faso ��� 5 juveniles (SVL 16.1-19.8 mm); Koubri Nabazana dam; 12.20319��N, ��� 1.35105��W; 280 m a.s.l.; 17.VII.2017; H. J. Ayoro & A. Ohler leg.; under accumulated dry grasses; hand capture ��� 1 juvenile (SVL 24.3 mm); Deux Bal��s National Park, Ouahabou bouli; 11.60767��N, ��� 3.06035��W; 260 m a.s.l.; 8.XI.2017; H. J. Ayoro leg.; under dead wood near water body; hand capture ��� 1 ♀ (SVL 31.5 mm); downstream from Soumbou dam; 11.69872��N, ��� 2.79738��W; 260 m a.s.l.; 8.XI.2017; same collector as for preceding; under a stone near water body; hand capture ��� 3♀ (SVL 33.0- 35.5 mm), 10 subadults (SVL 28.2-30.5 mm); ��� Baignade des El��phants ���; 11.71816��N, ��� 2.83777��W; 260 m a.s.l.; 8.XI.2017; same collector as for preceding; on dry grasses near water body; hand capture ��� 1 juvenile (SVL 23.1 mm); Koh��na pond; 11.67601��N, ��� 2.83577��W; 265 m a.s.l.; 9.XI.2017; same collector as for preceding; around water body; hand capture ��� 3♀ (SVL 37.4-38.6 mm); Tiona pond; 11.63241��N, ��� 2.85437��W; 265 m a.s.l.; 9.XI.2017; same collector as for preceding; under dry grasses around water body; hand capture ��� 5 subadults (SVL 26.7-31.5 mm); the bank of Mouhoun River; 11.77890��N, ��� 2.91259��W; 245 m a.s.l.; 10.XI.2017; same collector as for preceding; on mud and under grasses; hand capture ��� 3♀ (SVL 33.1 -37.0 mm), 2 subadults (SVL 30.0- 30.5 mm); W National Park, the edge of Gnifagou pond; 12.02083��N, 2.20222��E; 280 m a.s.l.; 3.II.2017; same collector as for preceding; under the dead wood near water body; hand capture ��� 3♀ (SVL 31.7-38.8 mm), 1 subadult (SVL 29.1 mm); W National Park, the edge of Soanda pond; 12.02861��N, 2.25890��E; 260 m a.s.l.; 3.II.2017; same collector as for preceding; under stones near water body; hand capture ��� 1 ♀ (SVL 31.8 mm); M��krou River (close to the Beninese border); 11.89991��N, 2.40260��E; 240 m a.s.l.; 4.II.2017; same collector as for preceding; under rocks on the bank of the river; hand capture ��� 1 ♂ (SVL 38.7 mm); Kabougou River; 11.96860��N, 2.01259��E; 280 m a.s.l.; 18. VI.2018; same collector as for preceding; in paddy field; hand capture ��� 1 ♀; on the edge of PK-27 bouli; 11.88744��N, 2.11913��E; 265 m a.s.l.; 20.VI.2018; same collector as for preceding; around water body; hand capture ��� 1 juvenile (SVL 18.5 mm); W National Park, Nangouli swampy valley; 11.89793��N, 2.00091��E; 250 m a.s.l.; 21. VI.2018; same collector as for preceding; on a stone near water body; hand capture ��� 1 ♀ (SVL 40.3 mm), 2 subadults (SVL 26.1-29.3 mm); the bank of Arly River; 11.57048��N, 1.45218��E; 160 m a.s.l.; 3.III.2017; same collector as for preceding; in a crack of dry mud; hand capture ��� 1 ♂ (SVL 32.3 mm), 1 ♀ (SVL 32.0 mm); Bongola swampy valley; 11.53252��N, 1.48241��E; 170 m a.s.l.; 13. VI.2018; same collector as for preceding; near water body; hand capture ��� 3 ♂ (SVL 33.1 -36.0 mm); Arly National Park, on the floodplain; 11.57852��N, 1.46004��E; 255 m a.s.l; 16-17. VI.2018; same collector as for preceding; around water body; hand capture ��� 3 ♀ (SVL 44.8-49.5 mm); Como��-L��raba Forest, Baboro swampy valley; 9.92521��N, ��� 4.57391��W; 230 m a.s.l.; 4.VIII.2018; H. J. Ayoro & A. Ohler leg.; under a heap of dead branches and leaves in a fallow field; hand capture ��� 1 ♀ (SVL 32.5 mm); Tuy province, at the backwater of Fafo; 11.33470��N, ��� 3.01947��W; 300 m a.s.l.; 4.XI.2018; H. J. Ayoro leg.; in a vegetable garden; hand cpature ��� Ioba province, Dano; 11.16600��N, ��� 3.08688��W; 340 m a.s.l.; J. Penner leg. ��� Gbagba (close to Dano); 11.18133��N, ��� 3.05164��W; 300 m a.s.l.; 2011; J. Penner leg.; field observation. LITERATURE REPORTS. ��� Burkina Faso ��� Passor�� province, 80 km north-west Ouagadougou; 12.96744��N, ��� 2.26155��W; 330 m a.s.l.; 30.VII.1992 (B��hme et al. 1996) ��� Yatenga province, Roba near Zamioro; 13.66675��N, ��� 2.50001��W; 350 m a.s.l. (Lamotte & Ohler 1997) ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 76 specimens examined. Newly collected females 31.7-49.5 mm SVL, males 33.2-38.7 mm SVL. Smaller than Lamotte & Ohler (1997) vouchers: males 38-51 mm and females 40- 61 mm. Specimens with a pointed snout, distinct and long limbs. Webbing formula: I 1 - 2 II 1 - 2 �� III 1 - 3 IV 2 �� - 1 V. Presence of supernumerary tubercles on the toes III and IV. Three pairs of dorsal ridges, including a pair of sacral ridges, a pair of ridges visible on the head between the eyes and ending in the lower part of the back, a second pair of ridges from behind the eyes to the same level as the median ridges and a third pair of ridges visible approximately above the tympanum, near the eye to the level of the other two pairs. The second and third pairs of ridges discontinuous in some specimens. A lateral ridge usually absent or only present as a short row of discontinuous light warts. For further detailed descriptions see Lamotte & Ohler (1997) and R��del (2000). Dorsum a grey to pale brown with numerous tiny dark brown to black spots, restricted to the longitudinal ridges. Light reddish or yellow vertebral lines may be present or absent. HABITAT. ��� We encountered this species under grasses at the edges of ponds and on river banks. It was also found in paddy fields. In the dry season, we collected P. bibroni under rocks, deadwood and accumulated dry grass around ponds. The species was found in dry and wet savannah habitats, intermittent freshwater wetlands, rural gardens, heavily degraded forests (agricultural areas), channels and ditches (R��del 2000). It is a common species, widely distributed in West Africa (Lamotte & Ohler 1997; Nago et al. 2006; Segniagbeto et al. 2007; Mohneke et al. 2011; Onadeko 2016; S��nchez-Vialas et al. 2017)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 566-568, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","LAMOTTE M. & OHLER A. 1997. - Redecouverte de syntypes de Rana bibroni Hallowell, 1845, designation d'un lectotype et description d'une espece nouvelle de Ptychadena (Amphibia, Anura). Zoosystema 19: 531 - 543. http: // sciencepress. mnhn. fr / fr / periodiques / zoosystema / 19 / 4 / redecouverte-de-syntypes-de-ranabibroni-hallowell- 1845 - designation-d-un-lectotype-et-descriptiond-une-espece-nouvelle-de-ptychadena-amphibia-anura","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","WERNER F. 1908 (1907). - Ergebnisse der mit Subvention aus der Erbschaft Treitl unternommenen zoologischen Forschungsreise Dr. Franz Werner's nach dem agyptischen Sudan und Nord- Uganda. XII. Die Reptilien und Amphibien. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch- Naturwissenschaftliche Classe 116: 1823 - 1926.","BOULENGER 1882. - Catalogue of the Batrachia Salientia S. Ecaudata in the collection of the British Museum. Taylor & Francis, London, i-xvi + 1 - 503, pl. 1 - 30. https: // doi. org / 10.5962 / bhl. title. 8307","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90.","ONADEKO B. A. 2016. - Distribution, diversity and abundance of anuran species in three different vegetations habitats in southwestern Nigeria. Ethiopian Journal of environmental Studies and Management 9: 22 - 34. http: // dx. doi. org / 10.4314 / ejesm. v 9 i 1.3","SANCHEZ- VIALAS A., CALVO- REVUELTA M. & MARQUEZ R. 2017. - Ptychadena in Mauritania and the first record of Ptychadena schillukorum. Zookeys 673: 125 - 133. https: // doi. org / 10.3897 / zookeys. 673.10265"]}

Tomopterna milletihorsini (Angel, 1922) COLLECTION MATERIAL. ��� There are no voucher specimens available for this species from Burkina Faso although its presence was reported in the East and Central-East in this country in the literature. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke et al. 2010b; Mohneke et al. 2011). DESCRIPTION. ��� Species with a short head, a horizontal pupil, short and strong limbs. Inner metatarsal tubercles large and shovel-shaped, tarsal tubercles usually present. Frog with a feebly warty back skin and smooth ventral skin. Dorsal colour brown to beige or dark brown. Pale dorsolateral lines often present. A white supratympanic ridge. Numerous dark brown, greenish or reddish spots, often with black borders, distinct on head, back and flanks. REMARKS. ��� The genus Tomopterna Dum��ril & Bibron, 1841 still comprises various undescribed species and needs taxonomic revision (Padial & De La Riva 2004; Padial et al. 2013). There are 14 species known in this genus and these are also known to be highly cryptic and morphologically similar (Wasonga & Channing 2013; Wilson & Channing 2019). The commonly used name for West African populations was Tomopterna cryptotis (Boulenger, 1907) (Lamotte & Xavier 1981; R��del 2000; Padial & De La Riva 2004; Mohneke 2011; Padial et al. 2013; Sow et al. 2017). Ohler & Fr��tey (2008) investigated the status of Arthroleptis milletihorsini (Angel, 1922), the type of which was deposited in MNHN collection. According to their examination, the characters did not correspond to the morphological characters of the genus Arthroleptis Smith, 1849, but rather to the characters of the genus Tomopterna Dum��ril & Bibron, 1841. The holotype is a young specimen of 18 mm SVL. Ohler & Fr��tey (2008) allocated the nomen Arthroleptis milletihorsini to Tomopterna, as Tomopterna milletihorsini (Angel, 1922) with the type locality being Kati 12 km north to Bamako from Mali. We tentatively use this name for West African Tomopterna. HABITAT. ��� During our field sampling, the presence of Tomopterna milletihorsini has been reported from Dori and Yakouta zones where gardeners and fishermen reported a frog species which inflate their body as Pyxicephalus maltzanii and is living under the sand. In Koti women indicated its presence as common in their groundnut fields after the harvesting (mostly from November to December). Women reported that the species was found by young shepherds who search it in sand in dried backwaters or in holes of groundnut fields after harvesting. In West Africa, Tomopterna is found in semiarid to arid habitats on sandy soils in the Sahel zone (R��del 2000; Padial & De La Riva 2004; Mohneke 2011). The species has been also found in mountains zones from Mauritania (Padial et al. 2013; Sow et al. 2017)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 574, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","MOHNEKE M., ONADEKO A. B., HIRSCHFELD M. & RODEL M. - O. 2010 b. - Dried or fried: amphibians in local and regional food markets in West Africa. Traffic Bulletin 22: 117 - 128.","PADIAL J. M. & DE LA RIVA I. 2004. - Annotated checklist of the amphibians of Mauritania (West Africa). Revista Espanola de Herpetologia 18: 89 - 99.","PADIAL J. M., CROCHET P. A., GENIEZ P. & BRITO J. C. 2013. - Amphibian conservation in Mauritania. Basic and applied Herpetology 27: 11 - 22. http: // dx. doi. org / 10.11160 / bah. 13002","WASONGA V. D. & CHANNING A. 2013. - Identification of sand frogs (Anura: Pyxicephalidae: Tomopterna) from Kenya with the description of two new species. Zootaxa 3734: 221 - 240. http: // dx. doi. org / 10.11646 / zootaxa. 3734.2.7","WILSON L. & CHANNING A. 2019. - A new sand frog from Namaqualand, South Africa (Pyxicephalidae: Tomopterna). Zootaxa 4609: 225 - 246. http: // dx. doi. org / 10.11646 / zootaxa. 4609.2.2","LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","SOW A. S., GONCALVES D. V., SOUSA F. V., MARTINEZ- FREIRIA F., SANTAREM F., VELO- ANTON G., DIENG H., CAMPOS J. C., DIAGNE S. K., BORATYNSKI Z. & BRITO J. C. 2017. - Atlas of the distribution of amphibians and reptiles in the Diawling National Park, Mauritania. Basic and applied Herpetology 31: 101 - 116. http: // dx. doi. org / 10.11160 / bah. 87","OHLER A. & FRETEY T. 2008. - Statut du nom Arthroleptis milletihorsini, Angel, 1922 (Amphibia, Anura). Alytes 25: 173 - 175."]}

Sclerophrys xeros (Tandy, Tandy, Keith & Duff-MacKay, 1976) COLLECTION MATERIAL. ��� Burkina Faso ��� 2♀ (SVL 63.8-65.8 mm), 2 ♂ (SVL 59.3-66.9 mm); Passor�� province, Samba; 12.70141��N, ��� 2.40390��W; 300 m a.s.l.; 10.VII.1961; M. Lamotte leg.; MNHNRA-1979.643, 1979.645, 1979.644, 1979.646 ��� 2 ♀ (SVL 64.7- 79.6 mm), 3 ♂ (SVL 60.5-68.9 mm); Ouagadougou; 12.38011��N, ��� 1.51100��W; 300 m a.s.l.; V.1975; M. Lamotte leg.; MHNG 1537.1 - 5 ��� 3 ♂ (SVL 69.9-73.7 mm), 4 ♀ (SVL 78.1-81.1 mm), 1 subadult (SVL 38.5 mm); Bougouriba province, Di��bougou; 10.98411��N, ��� 3.25603��W; 300 m a.s.l.; V.1976; NA leg.; MHNG 1536.89 - 96. FIELD MATERIAL. ��� Burkina Faso ��� 3 ♀ (SVL 66.0- 74.1 mm); Deux Bal��s National Park, Tiona pond; 11.63245��N, ��� 2.85431��W; 270 m a.s.l.; 9.XI.2017; H. J. Ayoro leg.; edge of the pond; hand capture ��� 1 ♂ (SVL 77.4 mm); Saaba; 12.37210��N, ��� 1.40372��W; 300 m a.s.l.; 4.IX.2017; same collector as for preceding; in a small water body near a rubbish dump ��� 5 ♀ (SVL 58.1-68.3 mm); Djibasso; 13.12511��N, ��� 4.16030��W; 340 m a.s.l.; 30.IX.2017; same collector as for preceding; under a light hunting insects; hand capture ��� 2 ♀ (SVL 60.1-65.1 mm); same locality; 5.X.2017; same collector as for preceding; under a cattle hay; hand capture ��� 7 ♂ (SVL 71.8-85.3 mm), 1 ♀ (SVL 78.1 mm); W National Park, Kabougou village swampy valley (3 km north-west of the Kabougou forest post office); 11.96073��N, 1.99745��E; 280 m a.s.l.; 22. VI.2018; same collector as for preceding; in a millet field; hand capture ��� 10 ♀ (SVL 51.5-84.5 mm); N���Djomga village pond (on the way from Dori to Gorom-Gorom); 14.07222��N, ��� 0.04865��W; 270 m a.s.l.; 25.VIII.2018; same collector as for preceding; close to a millet field; hand capture ��� 2 ♀ (SVL 64.7-68.7 mm); Yakouta, downstream from the Yakouta dam; 14.07450��N, ��� 0.14291��W; 280 m a.s.l.; 26.VIII.2018; same collector as for preceding; under shrubs; hand capture ��� 1 ♂ (SVL 78.8 mm); Sebba; 13.43904��N, 0.52340��E; 250 m a.s.l.; 22.IX.2018; A. Antarou leg.; hand capture ��� 2 ♂; D��dougou, SONATUR city; 12.48972��N, 3.43611��W; 300 m a.s.l.; 23.VIII.2017; H. J. Ayoro leg.; in a drain; visual and acoustic report ��� 1 ♂; Arly National Park, Bongola swampy valley; 11.53252��N, 1.48245��E; 170 m a.s.l.; 13. VI.2018; same collector as for preceding; acoustic report ��� 1 ♂; Arly National Park, near Bouliabou pond; 11.57888��N, 1.46390��E; 170 m a.s.l.; 16. VI.2018; same collector as for preceding; acoustic report ��� 2 ♂; Folonzo village swampy valley; 9.95768��N, ��� 4.67454��W; 240 m a.s.l.; 6.VIII.2018; H. J. Ayoro & A. Ohler leg.; acoustic report ��� 1 ♂; Kou River savannah; 11.18814��N, ��� 4.44397��W; 375 m a.s.l.; 9.VIII.2018; same collectors as for preceding; acoustic report. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���Gourma; 12.06051��N, 0.34913��E; 300 m a.s.l.; IX.-XII.2004 (Brito et al. 2008) ��� Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 49 specimens examined. Toad, similar to S. maculata and S. regularis, but skin with very prominent warts bearing spinules, more pointed than those of the two other species. Red pattern on the rear parts of thighs (absent in three other Sclerophrys species) and smooth, kidney shaped parotid glands allowing easy identification in the field. For more information on S. xeros, see Tandy et al. (1976), Lamotte & Xavier (1981), Salvador (1996) and R��del (2000). HABITAT. ��� This species was generally recorded from disturbed areas, around water bodies and open wetter habitats. It was sometimes seen inside protected areas. The recent samples were generally from paddy fields and around village ponds. Lanza (1981) and Salvador (1996) reported S. xeros from dry savannahs and very arid zones, at the vicinity of dry rivers, oasis and cattle troughs., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 556, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["BRITO J. C., REBELO H., CROCHET P. A. & GENIEZ P. 2008. - Data on the distribution of amphibians and reptiles from North and West Africa, with emphasis on Acanthodactylus lizards and the Sahara Desert. Herpetological Bulletin 105: 19 - 27.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","TANDY M., TANDY J., KEITH R. & DUFF- MACKAY A. 1976. - New species of Bufo (Anura: Bufonidae) from Africa's dry savannahs. Pearce Sellards Series. Texas memorial Museum 4: 1 - 20.","LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","SALVADOR A. 1996. - Amphibians of northwest Africa. Smithsonian herpetological Information Service 109: 1 - 45. https: // doi. org / 10.5479 / si. 23317515.109.1","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","LANZA B. 1981. - A check-list of the Somali amphibians. Monitore zoologico Italiano, n. s., Supplemento 15: 151 - 186."]}

Ptychadena oxyrhynchus (Smith, 1849) COLLECTION MATERIAL. ��� Burkina Faso ��� 2 ♀ (SVL 52-57 mm); L��raba province, near Katiar�� (north-east Ferk��); 9.91200��N, ��� 4.94771��W; 250 m a.s.l.; 10.XI.1970; M. Lamotte leg.; MNHN-RA-1996.3873, 1996.3874 ��� 2♀ (SVL 48.5-64.2 mm), 1 ♂ (SVL 55.2 mm); Bobo-Dioulasso; 11.18801��N, ��� 4.29743��W; 280 m a.s.l.; 6.XI.1970; M. Lamotte leg.;MNHN-RA-1996.3869, 1996.3870, 1996.3871. FIELD MATERIAL. ��� Burkina Faso ��� 1 ♂ (SVL 47.8 mm), 1 ♀ (SVL 62.3 mm); Tuy province, Fafo dam; 11.33470��N, ��� 3.01947��W; 300 m a.s.l.; 8.XI.2018; H. J. Ayoro leg.; in a vegetable garden; hand capture. LITERATURE REPORTS. ��� Burkina Faso ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l.; (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Mogtedo; 12.31844��N, ��� 0.83601��W; 280 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 7 specimens studied. Five females 48.5-64.2 mm; SVL of two males 47.8-55.2 mm SVL, close to e.g. those given by Konan et al. (2016), male size as 46.0- 60.9 mm and females size as 58.0- 65.1 mm; and by R��del (2000), males 40-53 mm, females 51-64 mm. Snout very pointed and hind legs very long. Webbing formula: I 0 - �� II 0 - 1 III 0 - 1 IV 1 - 0 V. On the back presence of four pairs of longitudinal ridges; a pair of short ridges and white lateral ridges on the flanks. Dominant dorsal colour a greyish to brown. HABITAT. ��� During the dry season, the species can be found under stones on river banks, whereas they breed in savannah ponds during the rainy season (R��del 2000). This frog occurs in open landscapes, savannah-forest ecotone and the degraded part of rainforest (Hughes 1988; Konan et al. 2016; Onadeko 2016). Some authors also list more arid habitats (Amiet 1974). It is widely distributed in West Africa and in a large part of sub-Saharan Africa from Senegal to South Africa (R��del 2000; Channing 2001; Channing & Howell 2006)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 568, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","KONAN J. C. B. Y. N., KOUAME N. G., KOUAME A. M., ADEPO- GOURENE. A. B. & RODEL M. - O. 2016. - Feeding habits of two sympatric rocket frogs (genus Ptychadena) in a forest remnant of southern-central Ivory Coast, West Africa. Entomology, Ornithology and Herpetology 5: 176. https: // doi. org / 10.4172 / 2161 - 0983.1000176","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","HUGHES B. 1988. - Herpetology in Ghana (West Africa). British herpetological Society Bulletin 25: 29 - 38.","ONADEKO B. A. 2016. - Distribution, diversity and abundance of anuran species in three different vegetations habitats in southwestern Nigeria. Ethiopian Journal of environmental Studies and Management 9: 22 - 34. http: // dx. doi. org / 10.4314 / ejesm. v 9 i 1.3","AMIET J. L. 1974. - Voix d'amphibiens camerounais. IV. Raninae: genres Ptychadena, Hildebrandtia et Dicroglossus. Annales de la Faculte des Sciences du Cameroun 18: 109 - 128.","CHANNING A. 2001. - Amphibians of Central and Southern Africa. Cornell University Press, New York, USA, 470 p.","CHANNING A. & HOWELL K. M. 2006. - Amphibians of East Africa. Cornell University Press, New York, USA, 432 p."]}

Variations within Pedioplanis laticeps (Smith, 1845) A total of 36 individuals were examined (Table 2). Max. SVL: 67 mm (ZMB 23443, adult male, Warmbad, Namibia). Scalation: The 3 nasal scales on each side always bulging, the interior ones usually separated by the rostral and the frontonasal, only in 3 out of 36 specimens the nasals meet in a suture (which is shorter than in burchelli); 29 to 36 transverse ventral rows (median 32); 14 to 18 longitudinal ventral rows (median 16). The anterior 6 to 11 transverse rows (sometimes called pectoral scales) are arranged in a V-shape and scale numbers are substantially reduced around the armpits, after which they are aligned horizontally onto the flanks, often only gradually decreasing in size and merging into the dorsal scales. Midbody scales smooth, arranged in 46 to 64 rows (median 56) (48 to 62 in Branch 1998, Conradie et al. 2012). In nine individuals (25%) the prefrontals are in contact (mostly separated by 1 scale; 2 in one individual, or by a suture between the frontal and the frontonasal in one individual). Scales in the temporal region show a tendency to enlarge towards the ear opening (in case there is an enlarged scale at the anterodorsal edge of the ear opening, it is not curved and there is at least one additional row of scales separating it from the ear opening); tympanic shield and elongate scales (lobes) absent; upper labials 8 to 13 (usually 11), with 4 (in two individuals) to 7 (usually 5 or 6) in front of the subocular scale which borders the lip; usually 2 scales between the subocular and the freno-ocular; lower labials 5 (in one individual on one side only) to 9, usually 7 or 8; femoral pores 11 to 18 on each thigh; enlarged scales in collar 6 to 11; occipital scale present; interparietal usually longer than broad, in four individuals occipital and interparietal are separated by 1 additional scale or granule; supraoculars 2, bordered by small granules anteriorly, posteriorly and along the supraciliaries in all individuals. In 61% the anterior supraocular is also surrounded with granules along its inner border, separating it from the frontal, and in 28% both supraoculars have the inner border edged with granules. Usually 15 or more granules anterior to the supraoculars. Chin shields pairs usually 5 (exceptionally up to 7), in most cases the anterior 3 are entirely in contact along their interior edge (in three individuals only the anterior 2 chin shields). Freno-ocular 1, frenonasal 1, in one individual (the holotype of E. laticeps) they are fused on one side, in a second one there is one additional scale below the freno-ocular; supraciliaries 6 (in one individual) to 12, usually 9 to 10; lower eyelid covered with several small, semi-transparent scales. ……continued on the next page ……continued on the next page ……continued on the next page ……continued on the next page Fore limb scales mainly small, flat and smooth, anteriorly large in a longitudinal row extending to the toes; hind limb scales small and smooth posteriorly and externally, below larger and imbricate with a row of large shieldlike plates anteriorly; tibia scales small and not keeled as are the top of the feet; soles of the hind feet covered in granular scales, each with 3 spines (1 median large, 2 lateral small ones) directed towards the 5th toe. There are 28– 32 spiny lamellae (mainly bi- or tricarinate, sometimes additional diminutive carinae) beneath the 4th toe. Colouration is variable. The examined young are brownish-black above with up to 6–8 whitish stripes in the neck region. Shortly behind the occiput the median pair fuses, towards midbody the next pair fuses which extend to the tail base. The number of white streaks from the occiput to the tail base is hence 8, 7, 6, 5 (or 7, 6, 5 or 6, 5). Adults can display similar striping (with up to 9 stripes in the neck in two individuals). The median stripes can also be of a light creamish-brownish colour. Other individuals have indistinct light stripes. The back may be yellowish brown, with or without small dark spots or chevrons, sometimes with two dorsal bands of black spots. There can be white ocelli on the sides. Laterally, the brown spaces between the stripes can be speckled with white, yellow and orange. The limbs are dark brown with whitish, black-edged ocelli. Ventral surfaces of belly, throat and limbs are white but can turn orange at the tail and the thighs., Published as part of Sebastian Kirchhof, Johannes Penner, Mark-Oliver Rödel & Johannes Müller, 2017, Resolution of the types, diagnostic features, and distribution of two easily confused Sand Lizards, Pedioplanis laticeps (Smith, 1845) and P. burchelli (Duméril & Bibron, 1839) (Squamata: Lacertidae), pp. 82-109 in Zootaxa 4318 (1) on pages 90-97, DOI: 10.11646/zootaxa.4318.1.3, http://zenodo.org/record/886124, {"references":["Smith, A. (1845) Illustrations of the Zoology of South Africa. Reptilia. Part XXIII. Smith, Elder, and Co., London.","Branch, W. R. (1998) Field guide to the snakes and other reptiles of southern Africa. Third edition. Struik Publishers, Cape Town, 368 pp.","Conradie, W., Measey, J. G., Branch, W. R. & Tolley, K. A. (2012) Revised phylogeny of African sand lizards (Pedioplanis), with the description of two new species from south-western Angola. African Journal of Herpetology, 61, 91 - 112. https: // doi. org / 10.1080 / 21564574.2012.676079"]}

uploaded for EU BON by Plazi The EU BON project aims to Build the European Biodiversity Observation Network, and is the European contribution to the Group on Earth Observation Biodiversity Observation Network (GEO BON). This present deliverable, entitled “Strategies and business plan for regional and global biodiversity information infrastructures” (D7.4) fits under EU BON Work Package (WP) 7 “Implementation of GEO BON: strategies and solutions at European and global levels”, and provides an overview of the outputs resulting from two WP7 Tasks. The objectives of this deliverable were to develop (1) a strategy for a global GEO BON infrastructure with optimised functionality in terms of efficiency and operability (as part of Task 7.4), and (2) a business plan for sustaining the European Biodiversity Observation Network after the funded phase (as part of Task 7.5). Building on the outputs from other EU BON work packages and extensive consultation (including externally to the project consortium), these objectives have been met. The global and European-level map of the biodiversity informatics landscape (Task 7.4) has led to a better understanding of the landscape’s current structure and functioning. This will enable key players to establish or strengthen collaborations, avoid effort duplication, and facilitate access to the biodiversity data, information and knowledge required to support effective decision-making. The business plan for the European Biodiversity Observation Network (Task 7.5) summarises what the network has to offer, to identified end-users (from policy, science/research, and citizen-science), in terms of products, tools and services, including those that can potentially generate income. A possible organisational structure, potential sources of funding, along with entry points for the network in other projects and initiatives (established or upcoming), are also presented.

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

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

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